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Goeldiana Zoologia

Número 15: 2 de dezembro de 1992

A New Species of Capuchin Monkey, Germ^CeAus^fxIeben, 1777
(Cebidae: Primates) from Eastern Brazilian ^Aínazonia

Helder L. Queiroz 1

ABSTRACT — The Kaapor Capuchin Cebus kaapori sp.n. from the Municipality
of Carutapera, Maranhão, Brazil, is describcd. A comparison between this form
and other capuchin monkey species is presented. The limits of the geographical
distribution of the new species and its conservation status are discussed.

KEY WORDS — Cebus kaapori sp.n., Primates, Cebidae, Capuchin Monkey,
Brazilian Amazônia, Taxonomy and Systematics, Geographical Distribution,
Ka’apor Indians

RESUMO — O cairara caapor Cebus kaapori sp.n., do Município de Carutapera,
Estado do Maranhão, Brasil, é descrito. Uma comparação desta com as outras
espécies do género é apresentada e discutem-se ainda os limites da distribuição
geográfica e o status de conservação da nova espécie.

PALAVRAS-CHAVE — Cebus kaapori sp.n., Primatas, Cebidae, Cairara,
Amazônia Brasileira, Taxonomia e Sistemática, Distribuição Geográfica, índios
Ka’apor

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1 Museu Paraense Emílio Goeldi, Departamento de Zoologia. Caixa Postal 399, Belém. PA.
cA •IP. 66040 lira? "

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2

A ncw species of Capuchin Afonkey from easlem BraziUan Amazônia

TAXONOMY OF THE CAPUCHIN MONKEYS

The genus Cebus Erxleben, 1777, is generally seen as the taxonomically most
complex of thc New World primates (Lima 1945; Hill 1960; Mittermcier &
Coimbra-Filho 1981). During the pasl few decades the consensus of specialists
(Hershkovitz 1949, 1972; Eisenbcrg 1989; Emmons 1990) is that the genus
should be divided into four species. Cebus is usually separated into two groups
on the basis of the presence or absence of tufts of dark hair on the frontal
crown (top of head). The "untufted" species are C. albifl-ons, C. nigrivittatus and
C. capucinus, while the "tufted" group ineludes only C. apella (Mittermcier &
Coimbra-Filho 1981; Mittermcier, Rylands & Coimbra-Filho 1988). Some
authors question the species status of the untufted forms, especially given the
lack of sympatry between them, a characteristic usually explained by competitive
exclusion (see Torres de Assumpção 1983, for a more detailed discussion).

Whitc-throated capuchin monkeys, Cebus capucinus Linnaeus, 1758, have
black upper parts (except for the sides of ncck), shoulders and upper arms
sharply contrasting pale yellow to whitish vcntrally. The head is yellow with a
V-shaped black cap on the crown (Emmons 1990). C. capucinus ranges from
Belize to northern Colombia (Freese & Oppenheimer 1981). Brown capuchin
monkeys, Cebus apella Linnaeus 1758, while being the most morphologically
variable of the genus, is lhe only "tufted'’ species. With dark brown upper parts,
the shoulders of brown capuchin monkeys are paler than the back. The crown
is completely covered with a black or dark brown cap that extends down the
cheek, becoming a distinct dark bar in front of the car. This cap can also form
horns of hair above the ears in males. The hands, hindlimbs and feet are black
or dark brown, as is the tail, which is darker at the tip (Emmons 1990). C.
apella , the most widely distributed neotropical primatc species, ranges from
extreme northern South America lo northern Argentina (Brown & Colillas
1984). While-fronted capuchin monkeys, Cebus albifrons Humboldt 1812, have
pale smokey grey-brown upper parts, and yellowish brown or reddish brown
fore- and hindlimbs. The face is fringed with silvery white, with white hairs
circling the facial region. The crown has a Sharp, wedgc-shaped cap of dark
smokey brown, generally extending forwards as a thin stripe down thc ccnter of
the face. The tail is smokey silvery yellow, paler at the tip than at the base
(Emmons 1990). This species occurs from Southern Colombia and Venezuela
south through thc Amazon basin lo northern Bolivia (Eisenbcrg 1989). The
castern limit is formed by the Negro and Tapajós rivers (Emmons 1990). Wedge-
cappcd or wceping capuchin monkeys, Cebus nigrivittarus Wagner, 1848 (or
Cebus olivaceus Schomburgk, 1848), are a uniform lawny brown on thc upper
parts, sometimes being frosted with dirty yellow. The shoulders and upper arms
are silvery yellow and the back of thc head and ncck is reddish. The head is
brownish yellow with a sharp V-shaped dark brown cap, and a thin brown stripe

Goeldiana Zoologia, mimem 15, 1992 3

clown ihc center of ihe facc as far as the nose. The hands, wrists and feet arc
dark brown. C. nigriviuatus is distributed in northern South America from
northern Venezuela across the Guyanas (Eisenberg 1989), with its western limit
at the Negro river and Southern limit at the Amazonas river.

A minor problem of nomenclalure exists in lhe case of the use of C. nigri-
vittatus and C. oUvaceus (Mittermeier & Coimbra-Filho 1981; Eisenberg 1989
and Emmons 1990). Frecse & Oppenheimer (1981) and Mittermeier &
Coimbra-Filho (1981) use nigrivit tatus, while Mittermeier, Rylandsand Coimbra-
Filho (1988) follow Robinson (1986) and Robinson & Janson (1987) in using
oUvaceus. Eisenberg (1989) and Emmons (1990) also prefer the latter, although
here 1 shall use nigriviuatus pending the review of the genus by Hershkovitz.

Cebas apella is sympatric over large arcas with C nigriviuatus and C. albifrons
(Hill 1960; Freese & Oppenheimer 1981). No other cases of sympatry between
two species of Ccbus have been reported.

In contrast with the consensus existing at lhe specific levei, there is little
agreement on the taxonomy of subspecific forms. Relatively small samples from
regions where populalions intergrade are responsible for much of the confusion
(Hershkovitz 1949; Torres de Assumpção 1983). Kinzey (1980) described five
subspccies of C. apella , while Hershkovitz (1949) recognized thirteen subspecics
of C. albifrons , five of C. nigriviuatus and five of C. capucinus. Hill (1960)
described a large number of subspccies for each one of the four species, but
further studies must be eonductcd to help resolve thesc questions (Mittermeier.
Rylands & Coimbra-Filho 1988). Studying the morphological characteristics of
more than 750 spccimens of C. apella, Torres de Assumpção (1983, 1986)
identified six geographica! arcas where the species showed some stability, and
suggested that the interpretation of her data in terms of lhe subspecies of C.
apella depends on the levei of interbreeding between thesc areas, which have
been geographically isolated in the pasl. Subspccies for the untufted group can
be considcred as provisional. In the case of C. nigriviuatus, for example, the lack
of any systematic intraspecific variation (as shown in Hershkovitz 1971) makes
taxonomy at the subspecific levei extremely questionable (John G. Robinson,
personal communication 1991).

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4

A ntv species of Capuchin Monkey frorn eastem Brazilian Amazônia

A NEW CAPUCHIN MONKEY SPECIES FROM BRAZILIAN AMAZÔNIA

An umlescribed form of untufted capuchin was rccently cncountcrcd during
field work among thc Ka’apor Indians of Gurupiuna Village, in the Alto Turiaçu
Indian Reservation of the Brazilian State of Maranhão. This site was previousiy
considered to be outside the geographical range of the untufted species group.
The first specimen observed had been hunted by the indians, who preserved its
skin and skull, which is described hcrc as the paratype. The ka’apor capuchin,
here dcsignated Cebus kaapori sp.n., was subsequcntly seen in thc wild in this
same reservation, in thc Caru Indian Reservation, also in Maranhão (wherc
these animais are raised as pets by lhe Guajá indians) and at othcr sites outside
these reservations in the northwest of Maranhão, wherc the holotype was
collected.

Cebus kaapori sp.n.

Holotype: Juvenilc female, skelcton and stuffcd skin preserved (Figure 1).
Collected at Quadram 7, 10 km Southwest of the Chega-Tudo Prospection
(02°30’S, 47°30’W) (Carutapera, MA), near the righl bank of the Gurupi river,
Maranhão, Brazil (Figure 2). Helder L. Queiroz coll. on 24' h of August 1991.
The holotype is deposited int the zoological collection of the Museu Paraense
Emílio Goeldi, Belém, Pará, Brazil, registered as specimen MPEG 22025.

Paratype: Adult male, skull and unstuffed, preserved skin. Collected at
Gurupiuna village (02°40’S, 46°20’W) in the Alto Turiaçu Indian Reservation,
Maranhão, Brazil, on the right or eastern bank of the Gurupi river (see Figure
1). Helder L. Queiroz and Ferdinando C. Nascimento colls. on 5 ,h of March
1990. Deposited in the zoological collection of lhe Museu Paraense Emílio
Goeldi, Belém, Pará, Brazil, registered as specimen MPEG 21978.

Uiugnosis: The longest-bodied untufted species of capuchin monkey, with a
silvery agouti mantle and silvery grey shoulders and tip of tail. Arms and
hindlimbs agouti. Hands and fcet black and dark brown. Triangular black cap
at crown with a black stripe down to nose. Forehead and face silvery grey and
beige. Occurring south of Amazonas river, possibly restrictcd to thc area
between Gurupi and Pindaré rivers in Maranhão State, Brazil.

6

A new species of Capuchin Monkey frorn eastent Brazitian Amazônia

Figure 2. The proposed limits of lhe geographical distribution of Cebus kaaporí sp.n., lhe lype localily
and site where lhe animais were scen and observed. The protecled areas within thc geographical
distribution are also shown, as well as the highways, roads and railroad and lhe places where inter-
views were performed (numbered as follows):l-Gurupi; 2-Quatro Bocas; 3-Maracaçumé; 4-Garimpo
Limão; 5 -Encruzo; 6-Maranhãozinho; 7-Bela Visla; 8-Sanla Luzia do Paruá; 9-Nova Olinda;
10-z\raguanã; 1 1-Zé Doca; 12-Chapéu de Couro; 13-Bom Jardim; 14-Sanla Inôs; 15-Pindaré Mirim;
16-Sanla Luzia; 17-FJoresta; 18-Santo Onofre; 19-Buriticupu; 20-Fazcnda Varig; 21-Piquiá;
22-Açailãndia; 23-Imperatriz and 24-Gurupiuna Village.

Goeldiana Zoologia, número 15, IW2

7

Geographical Distribution: In order to determine the geographical distribution
of the new species, observations of animais in the wild and interviews with local
residents were carried out along the Pará-Maranhão highway (BR-316), the
Santa Inês-Imperatriz. highway (BR-222) and the Carajás railroad. The towns
visited and the sites at which Kaapor capuchins were observed are shown in
Figure 1. This may be one of the smallest geographical ranges of an Amazonian
ccbid primate, being restricted in the present day to the arca belween Gurupi
and Pindaré rivers, in Maranhão State. The northeastern limit of the new
species’s range is the border of the Amazonian lowland high forest within the
"Cocais" ecosystem of middle Maranhão, extending at some points to the right
or eastern bank of Pindaré river, in its middle reaches, west of Santa Luzia. The
southeastern limit is defined by the distribution of the Amazonian forest.
Although not all small fragments of disturbed forest have been surveyed, the
presence of the new species south or east of this ecosystem seems unlikely. The
northern limit of the geographical distribution lies south of Maracaçumé, in the
Maracaçumé river basin, and the Southern limit is the northernmost forests of
Buriticupu. The western limit of the distribution is still unknown, but it may be
the Gurupi river. Howevcr, some interviews indieate that the kaapor capuchin
exists 5-10 km west of this river. 1 believe that the geographical distribution of
kaapori extended as far as the right bank of Amazon river in the past (see
below), but is now mostly restricted to Maranhão State. Human colonization
from the west and northwest, and the transformation of habitat to "Cocais" (also
as a result of human activity) from the east has determined the present distribu-
tion of the species. C. kaapori sp.n. is sympatric with C. apella throughout its
present distribution, but always at lower population densities.

Habitat: C. kaapori sp.n. was only observed in undisturbed and slightly disturbed
dense lowland Amazonian high forest ("mata-alta-de-terra-firme") under 200 m
of altitude, in very low densities. The interviews indieate that lhe new species
can also be found in edge habitats between the Amazonian and the "Cocais"
forest, due to its ability to feed on palm nuts. The new species was seen in the
neighborhood of indian gardens, in old fallows ("capoeiras velhas") and pristine
high forest. Guajá indtans, who raise these animais as pets, reported that C.
kaapori sp.n. can feed on palm fruits and are thus able to live in "Cocais" forest.
Miners and hunters from northwestcrn Maranhão also reported that C. kaapori
forms mixed groups with C. apella , but no sueh groups were observed.

8

A ncw species of Capuchin Monkey from eastem Brazilian Amazônia

Description: General aspect grayish agouti-brown, with the brown stripe on the
dorsal mid-line less marked than in other speeies of the genus. Grayish light
brown at sides. Shoulders silvery grey and beige extending to the proximal part
of the arm. The distai part of the forelimb is agouti turning dark brown and
black on wrists. Hands dark brownish black. Legs are equally agouti and beige
proximally and brownish black distally. Feel are blackish, like the hands. Tail is
grayish agouti at base, turning silvery grey and beige at tip. Excepl for dorsal
mid-line hairs, therc are agouti bands at tip and silvery bands at base of mantle
hairs. Hairs from the dorsal mid-line are entircly dark brown agouti. Close to
hands and fcet, hairs are agouti at tip and black at base, but on the arm, hairs
are banded with black, brown and silvery grey and beige; bands browner close
to wrists and more beige near shoulders. Hairs from the shoulders are brown at
base and silvery grey or beige at tip. Tail hairs are black or dark brown at base
and brown agouti and beige at tip, turning to silvery grey and beige on hair tip
distally. Hairs completely silvery grey and completely beige are found ventrally
along lhe tail and at its tip on both surfaces. Similar hairs are also found on the
neck (ventrally), and mixed with entircly beige hairs in the lhoracic and abdomi-
nal regions. Inner parts of legs and arms have cntirely light brown agouti or
black hairs. The brown mid-line hairs turn dark brownish and brownish black
to black, forming longcr, thinner fur on the cap, continuing onto the crown and
then shorter when reaching the face. The face is pink or flesh-like in nature,
with small white and silvery grey and beige hairs on the chccks, moustache and
chin. Forehead and regions between chccks and ears are silvery grey and beige,
and the hairs have small black bands. Ears are almost naked and pink in nature.
Lips and nosc are also naked, but almost black in nature with small black hairs.
Eyebrows are silvery grey and beige, but some black hairs are found. The cap
starts back of the head, but is only prominent on the crown. Composed of long
black hairs, the cap is triangular in shape, with a thin black stripe of very small
hairs strctching down to the nose, Crossing forehead between the eyes.

Discussion: In general appearance, C. kaapori sp.n. is most similar to C.
nigrivittatus. but the kaapor capuchin’s geographical range is separated from that
of the wecping capuchin by more than 4(X) km. Cebus kaapori sp.n. has a silvery
agouti mantle, and its tail is brown turning to silvery grey at tip, while the forms
of nigrivittatus from castcrn Venezuela and Guyanas have a brown rcddish or
coppcr mantle, and those from the region of the Branco river of Amazonas State
have grayish or dark grayish mantles. Tails are always darker in these forms. The
nape is blackish dark brown, contrasting with the copper nape of C. nigrivittatus.
The shoulders of kaapori are silvery grey and the arms and hindlimbs are agouti.

Gocldiana Zoologia, número 15, 1992

9

MEASUREMENTS AND COMPARISONS WITH OTHER CAPUCHIN
SPECIES

With only a single adult specimen available (the paratype), it is impossible
to analyze intraspecific variation in Cebus kaapori sp.n., and difficult to draw
detailed conclusions on interspecific variation within the genus. The ncw form
appears to be longer-bodied and less robust than other untufted spccies, but
there is little to separate the species on the basis of most craniomelric and
morphological characteristics. Externai and craniomelric measurements from the
paratype of Cebus kaapori sp.n. are presented in Tables 1 and 2, respectively,
and compared with valucs taken from lhe literature for the remaining four
species of the genus. No cranial measurements were available for C. nigrivil tatus
in the literature, and five nigrivittatus crania from the Gocldi Museunt collection
(MPEG 993, MPEG 994, MPEG 1235, MPEG 1284 and MPEG 1633) are given
in Table 2. Because of the marked sexual dimorphism of the genus (Torres de
Assumpção 1986), only males specimens are used here. Table 1 shows that C.
kaapori is longer-bodied than all other untufted species, overlapping with only
C. apella.

The new species is undoubledly similar to C. nigrivittaius, but the distance
between the geographical ranges of C. nigrivittatus and C. kaapori, and the
resulting absence of inlerbreeding between these populations suggest that other
speciation processes have occurrcd, and that C. kaapori is a true species. In
addition, Table 2 suggests that C. nigrivittatus has a longer maximum cranial
length than all other species of the genus, including C. kaapori, suggesting a
degree of divergence of these two forms.

10

A new species of Capuchin Monkey from eastem tíraziban Amazônia

Table 1. Externai measurements of the paratype of Ccbus kaapori sp.n. and the range of measure-
ments for other species of the genus (from Emmons 1990).

Species

I lead/Hody
(mm)

Tail

(mm)

Foot*

(mm)

Ear

(mm)

Weight

(kg)

Ccbus capucinus

335-453

350-551

110-150

21-42

1. 4-3.9

Cebus apcllu

350-488

375-488

107-132

28-43

1. 7-4.5

Ccbus albifrons

358-460

401-475

112-136

32-45

1. 2-3.6

Cebus olivaccus +

374-460

400-554

120-143

35-51

2.3-42

Cebus kaapori

465

510

122/123

33

3.05

+ Sensu Robinson (19K6), or C. nigrivittatus.
* Foot without and with nail, respectively

Table 2. Craniometric measurements of the paratype of Cebus kaapori sp.n. and of other species of
the genus.

Measurements

C. capuc

(1 Cf)

C. albifr *
(18 efef)

C. nigriv
(5 tf cf)

C. kaapori
(1 tf)

C. apcUa'

(1 tf)

Maximum Cranial Fength

95.3

92.8

100.6

94.5

94.5

Maximal Cranial Breadth

56.0

50.8

56.2

52.7

53.1

Bizygomatic

61.0

61.8

67.1

61.5

60.5

Biorbital

52.4

51.8

48.8

45.5

44.8

Uppcr check-tceth

21.7“

26.3

29.8

21.0

27.8

I.ower cheek-teeth

25.4”

29.0

33.9

293

31.5

From Hill 1960.
Without canines.

Gocldiana Zoologia, número 15, 1992 1 1

DISTRIBUTION AND CONSERVATION STATUS OF THE NEW SPECIES

Six untufted capuchins were collected in 1897 on the Acará and Capim rivers
(south of Belém, Pará) and deposited in the zoological collection of the Goeldi
Museum by Goeldi & Hagmann (1906). While the specimens were denominated
Cebus capucinus, their description was highly similar to that of Cebus kaapori
sp.n. reported above. At the time, the name capucinus was applied also to nigri-
vittatus. Unfortunately, these specimens were not found in the Goeldi Museum,
and have probably becn lost, but their description suggests that the western limit
of the distribution of Cebus kaapori was much closer to Belém in the past.
Surprisingly, this report has bcen overlooked in all subsequent revisions of the
gcnus, and even in studies of C. nigrivittatus. lf the present geographical distribu-
tion of C. kaapori sp.n. is confirmed as describcd here, this new species would
range over a total area of approximately 15, 000 knr, one of the smallcst ranges
of an Amazonian cebid.

As shown in Figure 2, this area encompasscs the Gurupi Biological Reserve,
and lhe Caru and Alto Turiaçu Indian Rescrvations, where Cebus kaapori sp.n.
is a game species for local indians. In addition, the Gurupi reserve is subject to
conlinuous invasion by loggers. All three protected areas, representing almost
1,000,000 ha, are threatcned not only by loggers, but also by gold miners,
squatters and ranchers.

There are also highways and minor roads in the area (Figure 2), and the
Carajás railroad has had a significam influence on colonization in the region.
Innumerable small towns have thus been established in the vicinity of these
protected areas and their inhabitants usually invade these areas to hunt, fish or
collect forest resources.

Considering all these problems, Cebus kaapori sp. n. may be seen as a
threatcned species. The contiguous protected areas seen in Figure 2 may
nevertheless represem a viable alternative for the protcction of the new species
unlil a systematic conscrvation program can be formulated.

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A new specics of Capuchin Monkcy from eastern Brazilian Amazônia

ACKN OWLEDGEMENTS

I woulcl likc to thank lhe Noyes Foundation, which supported the initial field
and laboratory work. The collection of the holotype was made possible by
support from the John D. and Catherine T. MacArthur Foundation. Wildlifc
Conservation International-Brazil (WCI-Br) also supported my field work.
Ferdinando Nascimento was my field assistam at the time of the collection of
the paratype. The community of Chega-Tudo helped in collecting the holotype.
I am very indebted to José S. Silva Jr., who first called my attention for the new
form. I also thank Dr. J.M. Ayres, Dr. S.F. Ferrari, Dr. J.G. Robinson, Dr. I.
Gorayeb, Dr. D. Oren, Dr. A.B. Rylands and many others for giving me
extrcmely helpful additional information and suggestions, even when we did not
share the same taxonomic opinions. The collection of specimcns was authorized
byJBAMA Liccnce number 031/91 -DEVIS. The owncrs of captive C. kaapori
at'chega-Tudo kindly gave me pcrmission to collecl blood from their animais.
Drs. M. Lima, M.I. Sampaio, L. Harada and M.A. Medeiros, from the Depart-
ment of Genetics of the Federal University of Pará (UFPA) helped me collect
blood and liver samples for further cytogenctic, imunogenetie and DNA studies.
I thank photographer Amilton Trindade for photographs of the holotype and Sr*
Ana Maria Oliveira for revision of the bibliographic references.

REFERENCES

BROWN, A.D. & COLILLAS, O.J. 1984. Ecologia dc Cebus apella. In:
MELLO, M.T. (cú.) A Primatologia no Brasil. Brasília, Sociedade Brasileira
de Primatologia, v.l.

E1SENBERG, J.F. 1989. Mammals of the Neotropics. Chicago, The University
of Chicago Press, v.l.

EMMONS, L.H. 1990. Neotropical Rainforest Mammals - A Field Guide. Chica-
go, The University of Chicago Press.

FREESE, C. & OPPENHEIMER, J.R. 1981. The Capuchin Monkeys, Genus
Cebus. In: MITTERMEIER, R.A. & COIMBRA-FILHO (eds). Ecology and
Behavior of Neotropical Primates. Rio dc Janeiro, Academia Brasileira de
Ciências, v. 1 .

GÒELDI, E.A. & HAGMANN, G. 1906. Prodromo dc um catálogo crítico,
commentado, da colecção dc mammífcros no Museu do Pará (1894-1903).
Boi Mus. Goeldi (Mus. Para.) Hist. Nat. Ethnogr. 4: 38-122.

HERSHKOVITZ, P. 1949. Mammals of northern Colombia. Prcliminary report
no. 4: Monkeys (Primates), with taxonomic revisions of some forms. Proc.
U.S. Natn. Mus., Washington. 3232: 52-59.

HERSHKOVITZ, P. 1971. Slapcdial processes in tympanic cavities of capuchin
monkeys (Cebus ). ./. Mammalogy 52(3): 607-609.

Goeldiana Zoologia, número 15, 1992

13

HERSHKOVITZ, P. 1972. The Recent Mammals of Neotropical Region: A
Zoogeographic and Ecological Review. In: KEAST, A.; ERK, F.C. &
GLASS, B. (eds). Evolution, Mammals, and Southern Continents. New York,
State University of New York.

HILL, W.C.O. 1960. Primates. Comparative Anatomy and Taxonomy. Cehidae.
Edinburgh, Edinburgh University Press, v.4. part A.

KINZEY, W. 1980. Distribution ofsome Neotropical primates and the model
of Pleistocene forest refugia. INTERNATIONAL SYMPOSIUM, ASSOCIA-
TION OF TROPICAL BIOLOGY, 5, PROCEEDINGS, Caracas. New York,
Columbia University Press.

LIMA, E.C. 1945. Mammals. of Amazônia. Belém, Museu Paraense Emílio
Goeldi de Historia Natural e Etnografia, v.l.

MITTERMEIER, R.A. & COIMBRA-FILHO, A.F. 1981. Systematics: Species
and Subspecies. In: MITTERMEIER, R.A & COIMBRA-FILHO, A.F. (eds)
Ecology and Behavior of Neotropical Primates. Rio de Janeiro, Academia
Brasileira de Ciências, v.l.

MITTERMEIER, R.A.; RYLANDS, A.B. & COIMBRA-FÍLHO, A.F. 1988.
Systematics: Species and Subspecies - An Update. In: MITTERMEIER, R.A.;
RYLANDS, A.B.; COIMBRA-FILHO, A.F. & FONSECA, G.A.B. (eds)
Ecology and Behavior of Neotropical Primates. Washington, WWF Press, v.2.

ROBINSON, J.G. 1986. Seasonal variation in use of time and space by the
wedge-capped capuchin monkey, Cebus olivaceus: implications for foraging
theory. Smithson. Contri. Zool. 431: 1-160.

ROBINSON, J.G. & JASON, C.H. 1987. Capuchins, Squirrel Monkeys, and
Atelines: Socioecological Convergence with Old World Primates. In: SMUTS,
B.B.; CHENEY, D.L.; SEYFARTH, R.M.; WRANGHAM, R.W. &
STRUHSAKER, T.T. (eds). Primate Societies. Chicago, University of Chicago
Press.

TORRES DE ASSUMPÇÃO, C. 1983. An Ecological Study of the Primates of
Southeastem Brazil, With a Reappraisal of Cebus apella Races. Edinburgh,
University of Edinburgh. Ph.D. Thesis.

TORRES DE ASSUMPÇÃO, C. 1986. Resultados Preliminares da Reavaliação
das Raças do Macaco-Prego Cebus apella (Primates, Cebidae). In: MELLO,
M.T. (ed) A Primatologia no Brasil. Brasília, Sociedade Brasileira de Primato-
logia. v.2.

ypt»

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SCT / CNPq

MUSEU PARAENSE EMÍLIO GOELDI

Campus de Pesquisa — Av. Perime trai. Guamá
Caixa Postal 399. Telex: (091) 1419. Telefones: Parque (091) 224-9233
Campus: (091) 228-2341 e 228-2161
66.040 Belém, Pará, Brasil

GOELDIANA ZOOLOGIA é uma publicação do Departamento de Zoologia do Museu Paraense
Emílio Gocldi — CNPq.

N° 1. A reevaluation of Serpophaga araguayae Snetlilage, 1928 (Aves: Tyrannidae).

José Maria Cardoso da Silva

N° 2. Notes on Cyranotermes Araújo, wilh Description of a New Species (Isoptera,
Termitidae, Nasutitermitinae). Reginaldo Constantino
N° 3. Anhangatennes macarthuri, a New Genus and Species of Soil-feeding Nasute Termite
from Amapá, Brazil (Isoptera, Termitidae, Nasutitermitinae). Reginaldo Constantino
N° 4. New and reconfirmed bird records from the State of Maranhão, Brazil. David C. Oren
N° 5. Resultados de uma excursão ornitológica à ilha de Maracá, Roraima, Brasil. José Maria
Cardoso da Silva & David C. Oren

N° 6. Priority Areas for New Avian Collections in Brazilian Amazônia. David C. Oren &
Haroldo Guerreiro de Albuquerque

N° 7. Notes on Neocapritennes Holmgren, witli description of two new species from the
Amazon Basin (Isoptera, Termitidae, Termilinae). Reginaldo Constantino
N° 8. Ercymatermes roamdiceps, new genus and species of termite from the Amazon Basin
(Isoptera, Termitidae, Nasutiterminae). Reginaldo Constantino
N° 9. Aves do Estado do Maranhão, Brasil. David C. Oren

N° 10. Resumo da Classificação e Bibliografia dos Mutilídeos da América do Sul (Hymenop-
tera: Mutillidae). William Leslie Overal

N° 11. New Data on the Dislribution of Primales in the Region of the Confluence of the

Jiparaná and Madeira Rivers in Amazonas and Rondônia, Brazil. Stephen F. Ferrari &
Maria Aparecida Lopes

N° 12. A New Species of Marmoset, Genus Callithnx Erxleben, 1777 (Cailitrichidae, Primates),
from Western Brazilian Amazônia. Stephen F. Ferrari & Maria A Lopes
N° 13. Notes on the Neotropical genus Acanthocera Macquart (Diptera: Tabanidae) with
description of four new species. Augusto L. Henriques & José A. Rafael
N° 14. A New Species of Marmoset, Genus Calliihrix Erxleben, 1777 (Cailitrichidae, Primates)
from the Rio Maués Region, State of Amazonas, Central Brazilian Amazônia. Russcll
A. Mittermeier, Marco Schwarz & José Márcio Ayres
N° 15. A New Species of Capuchin Monkey, Genus Ccbus Erxleben, 1777 (Cebidae: Primates)
from Eastern Brazilian Amazônia. Helder L. Queiroz

Este número foi publicado com o apoio de:

The John D. and Catherine T. MacArthur Foundation

&

World Wildlife Fund — US

WWF

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