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OCCASIONAL PAPERS ^ ^^q.

of the

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 145. PAGES 1-12 6 DECEMBER 1991

ANNOTATED CHECKLIST OF THE MAMMALS
OF CUZCO AMAZONICO, PERU

Neal Woodman", Robert M. Timm', Rosa Arana C.-, Victor
Pacheco-, Cheryl a. Schmidt\ Errol D. Hooper', and

Cecilia Pacheco A.-

Complete or nearly complete inventories of mammalian species are
available for few specific localities in the neotropics because of insufficient
sampling. Inferences about abundances of most of the species inhabiting an
area can be made for even fewer localities (e.g., Terborgh et al., 1984; Timm,
in press). This lack of information severely limits our ability to compare
mammalian faunas among sites or study broad patterns of mammalian
distributions in the neotropics. The situation is aggravated by the increasing
deterioration of natural habitats and the resulting extirpation of many species.

To increase our knowledge of neotropical rain forest mammals, the
mammal survey team of the neotropical Biological Diversity Program
(BIOTROP) developed and tested a formal protocol for surveying neotropical
mammals that allows direct comparisons to be made among sites. This study
was one part of the BIOTROP pilot project, a joint venture by The University
of Kansas Museum of Natural History (KU), Museo de Historia Natural,
Universidad Nacional Mayor de San Marcos (MUSM), Missouri Botanical
Garden, and Harvard University. The goal of this project was to develop and

'Museum of Natural History and Department of Systematics and Ecology, The
University of Kansas, Lawrence. Kansas 66045-2454, USA.

"Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado
14-0434. Lima-14, Peru.

^Department of Biology. University of South Dakota. Vemiillion, South Dakota
57069. USA.

© Museum of Natural History. The University of Kansas, Lawrence.

2 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 145

implement a strategy for rapidly inventorying neotropical sites to obtain data
on species richness and associations of species for selected groups of
organisms in relation to certain measurable environmental variables.

The study site. Reserva Cuzco Amazonico. is a national wildlife reserve
located along the north bank of the Rio Madre de Dios, 14 km east of Puerto
Maldonado. Provincia de Tambopata, Departamento de Madre de Dios.
southeastern Peru ( 1 2°33'S. 69°03'W). The Reserve is approximately 300 km
east-northeast of the city of Cuzco and about 50 km west of the Bolivian
border. The elevation of Cuzco Amazonico is approximately 200 m. Cuzco
Amazonico includes approximately 10,000 ha of relatively undisturbed
lowland rain forest under the management of Jose E. Koechlin. who operates
a tourist lodge ( Albergue Cuzco Amazonico. SA) located near the southwest
comer of the reserve. Two 500-m x 500-m study areas dedicated specifically
to BIOTROP field studies were established within the Reserve. One study
area. Zone 1, was in seasonally inundated swamp forest, whereas the second.
Zone 2. was located in lowland forest relatively free of standing water
throughout the year. A complete description of the study zones including
maps was presented in Duellman and Koechlin (1991). Amphibians and
reptiles from Cuzco Amazonico were described by Duellman and Salas
( 1 99 1 ); a list of the birds from the Reserve was provided by Davis et al. ( 1 99 1 ).

The list of mammals presented herein (Table 1 ) represents the preliminary
results of field work completed by the BIOTROP mammal survey team at
Reserva Cuzco Amazonico from 8 June-20 July 1989 and 19 January- 1
March 1990. More detailed accounts of the natural histories of the mammal
species we encountered are in preparation and will be presented in a forthcoming
paper. We also include records of specimens collected at this locality by J. E.
Cadle, J. L. Patton, E. D. Pierson, R. M. Warner, and E. Yensen at various
times between 1 979 and 1 984; these specimens are deposited in the collections
of the MUSM, the Museum of Vertebrate Zoology, University of California,
Berkeley, and the College of Idaho Museum of Natural History, Caldwell. All
species of marsupials, bats, and rodents that we captured are represented by
voucher specimens deposited at MUSM and KU.

METHODS

Small terrestrial and arboreal mammals were surveyed primarily by
trapping along four 500-m long trails that marked the boundaries of the two
BIOTROP study zones. Trap stations were located at approximately 20-m
intervals along each trail and consisted of two pairs of traps, each of which
included a Sherman live trap and a Victor rat trap. One pair of traps was placed
on the ground and the other located from 0.2-2 m above the ground on
branches, lianas, fallen trees, or in shrubs. Trapping was carried out for 12

MAMMALS OF CUZCO AMAZONICO 3

consecutive days on each trail. Traps were baited with peanut butter mixed
with rolled oats for 6 days and a suet-seed-fruit mixture for 6 days. Trap
stations along two trails (one trail per zone; a total of 200 traps) were
monitored simultaneously. Trapping was conducted in the zones once during
the dry season (June-July ) and once in the wet season (January-February), for
a total of 9.600 trapnights. This was augmented by additional trapping in areas
containing habitat types not represented in the two zones to sample other
species that might occur in the Reserve. Tomahawk live traps were used to
sample medium-sized mammals too large for Sherman or Victor traps. One
Tomahawk trap was placed every 1 00 m along traplines in the zones. Mist nets
were used to capture bats along forest trails, in clearings, and over streams.
We were unable to sample the forest canopy. Larger mammals were censused
by recording sightings of the animals or identifiable tracks. More detailed
accounts of our mammal sampling techniques and results will be published
at a later date.

We estimated abundances of small marsupials and rodents from the
absolute numbers of captures in the zones. Abundances of other mammals
were estimated from frequency of sightings and from previous knowledge
about their habits and habitat requirements (e.g.. nocturnal vs. diurnal; wide-
ranging vs. relatively sedentary). These estimates are influenced by the
likelihood of these animals to enter traps, their attraction to baits, and
possibly, seasonal and /or annual abundances in the habitats encompassed by
the study zones. Populations of a mammal species in the tropics can vary
greatly in numbers from year to year; however, our estimates provide a
baseline from which more intensive population studies can be undertaken.
Disturbance resulting from our activities in the study areas probably affected
the number of sightings of mid-sized and larger mammals. Because of this,
we undoubtedly have underestimated abundances of some species.

RESULTS AND DISCUSSION

Reserva Cuzco Amazonico supports a mammalian fauna of at least 100
species belonging to 9 orders and 28 families (Table 1). Included are 9
marsupials, 42 bats, 7 primates, 5 xenarthrans, 1 rabbit, 24 rodents, 9
carnivores, 1 tapir, and 2 artiodactyls. On the basis of our observations of
mammals in adjacent areas (e.g., Lago Sandoval), reports in the published
literature, and our knowledge of species' distributions and habitat requirements,
we estimate that an additional marsupial, 12 bats, 6 monkeys, 4 xenarthrans,
3 rodents, 6 carnivores, and 3 artiodactyls probably are occasional or scarce
members of the fauna. If a total of 135 mammal species is correct, our efforts
recorded 74% of the mammal fauna of Cuzco Amazonico.

Seven mammal species collected by us from Cuzco Amazonico represent

4 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 145

either extensions of known ranges or are among first records for Departamento
de Madre de Dios. Our specimens of Marmosops parvidens. Philander
andersoni. and Molossus coibensis are the first records of these species from
southeastern Peru, and the specimens of Neacomys tenuipes are the first
reported from Peru. Despite intensive effort, we were able to collect only a
single specimen of Moiiodelphis adusta. This is the first published record for
the species from Departamento de Madre de Dios. Two Peroptery.x leucoptera,
collected from a day roost in a hollow fomied by the struts of a large, fallen
tree, represent a considerable southern extension of the known range of this
bat; this is only the second record of this rare emballonurid from Peru. Our
record of Choloepus hoffmauui is the southernmost for the species. Recently,
Ascorra et al. (1991) reported the first specimens from Departamento de
Madre de Dios of Choeroniscus minor, Micronycteris minuta, Mimon
crenukitum, and Rhiuophylla piimilio from the Manu National Park area; we
captured individuals of these species at Cuzco Amazonico as well.

Small terrestrial rodents and marsupials at Cuzco Amazonico are extremely
abundant and diverse, in contrast to other lowland tropical sites. Twenty-three
rodent species (exclusive of introduced murids) are confirmed from Cuzco
Amazonico, in contrast to 1 5 species known from moist, lowland forest at La
Selva, Costa Rica (Timm et al., 1989; Wilson, 1990), 15 species from Barro
Colorado Island, Panama (Glanz, 1990), and 16 species from forest north of
Manaus, Brazil ( Malcolm. 1 990). The rodent fauna is most comparable to that
at Cocha Cashu. Peru, from which 24 species are known (Janson and
Emmons, 1990). Bats, which are generally abundant at other lowland tropical
sites where we have worked (see also Janzen and Wilson, 1983), were much
less common at Cuzco Amazonico. For example, in the Caribbean lowlands
of Costa Rica, we typically could capture 10-15 individuals or more in a 3-hr
period. Atypical catch at Cuzco Amazonico during the same time period was
four to six individuals. Bat diversity also was somewhat low (42 confirmed
species) in contrast to La Selva (63 species; Wilson, 1990), but in part, this
may reflect our inability to sample in the canopy. Additional field work
concentrating on bats undoubtedly will increase the number of species known
from Cuzco Amazonico. We found that white-tailed deer, peccaries, tapirs,
larger monkeys, and other large "game" animals were uncommon to rare.
Although wildlife at Reserva Cuzco Amazonico has protected status, there are
no full-time game wardens, and poaching continues to occur.

Populations of some mammals at Cuzco Amazonico have undergone
changes just within the last few years. When Eric Yensen and Richard Warner
visited the Reserve in 1 984. Mctachirits nudicaiidatiis and Philander andersoni
were quite common and Didelphis niarsupialis were so rare that they saw only
a single individual (E. Yensen. pers. comm.). During our survey of the fauna
5 years later, Metachirus and Philander were uncommon, whereas Didelphis
were very abundant.

MAMMALS OF CUZCO AMAZONICO 5

Until quite recently, most of the smaller, long-tailed mouse-opossums of
the family Didelphidae from Central and South America were included
together under the single generic name Mannosa. We concur with Gardner
and Creighton (1989) that this diversity is better represented taxonomically
by the recognition of several genera. Hence, herein we use Mannosa for the
species M. mitrina. Mavmosaps for M. noctira^us and M. parvidens. and
Micoureiis forM. vegina. We follow Hershkovitz (pers. comm.) in recognizing
Philander andersoni as specifically distinct from P. opossum.

Taxonomy of the fruit-eating bats of the gQnw:^ Artiheus is currently under
debate and much of it is in need of revision. For the purposes of this paper, we
follow Koopman (1978) in considering the large, widely distributed form in
the Amazon Basin, A. planirostris. as a distinct species; it sometimes is
considered a subspecies of A. jamaicensis. We follow Handley (1989) in
recognizing the large, blackish /\/Y/7k^z/5 of the Amazon Basin as A. obscurus;
this species previously has been reported under the names A. "'daxisir A.
fuliginosus, andAJamaicensisfullginosus. Koopman (pers. comm.) considers
A. obscurus to be a subspecies of A. jamaicensis. We follow Owen (1987) in
recognizing the smaller members of this group as a separate genus, Dermanura.
Taxonomy of the small, fruit-eating bat, Dermanura cinerea, is problematic.
Handley ( 1 989) restricted this species to the Atlantic and Caribbean coastline
of South America, from Brazil to Venezuela. Our specimens clearly represent
a single species, and they are distinct from D. anderseni, D. glauca. and D.
glioma. The only appropriate name available for the specimens from Cuzco
Amazonico isD. cinerea, 2ind they match Handley 's description and measure-
ments fairly well.

In the checklist we follow Hall ( 1981 ) and Gardner and Ferrell (1990) in
recognizing Platyrrhinus as the senior synonym of Vampyrops.

Spiny mice of the genus Neacomys are poorly known, and the taxonomy
of these diminutive, spiny cricetids is greatly in need of revision. We refer
larger members of the genus (total length of skull [GLS] > 21.5; maxillary
toothrow [MTR] > 2.7 ) from Cuzco Amazonico to the widespread species. A'.
spinosus. The smaller Neacomys (GLS < 21.5; MTR < 2.7) are refeired
tentatively to N. tenuipes. although their average dimensions are somewhat
smaller than those previously reported for this species (Lawrence, 1 94 1 ). This
is the first time that two species of these spiny mice have been reported from
the same site in Peru.

Specimens of Rhipidomys from Cuzco Amazonico clearly represent two
species, the larger of which (GLS > 36.0; head and body length [HB] > 140)
we refer to R. couesi. Although the genus is being revised (C. Tribe, pers.
comm.). species of Rhipidomys currently are poorly defined and there is a
confusing array of available names. Because of this, we refer to the smaller
species from Cuzco Amazonico as Rhipidomys sp. 2. Inspection by one of us
(Timm) of specimens of Rhipidomys from Manu National Park, deposited at

6 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 143

the U.S. National Museum, indicates that these are conspecific with our R.
couesi from Cuzco Amazonico. Previously, these specimens were reported as
R. leucodactylus by Terborgh et al. (1984).

In Table 1 mammalian orders are aiTanged following Hall ( 198 1 ). Families
within orders, genera within families, and species within genera are listed
alphabetically.

Table 1 . Mammals of Cuzco Amazonico, Peru. Abbreviations. — Abundance: A
= abundant, often observed and /or captured in appropriate habitats; small teirestrial
/arboreal mammals represented by more than 15 captures in zones. C = common,
frequently observed and /or captured in appropriate habitats; 10-15 captures in zones.
U = uncommon, occasionally observed and /or captured in appropriate habitats; 5-10
captures in zones. R = rare, few sightings /captures in appropriate habitats; <5 captures
in zones. P = probable, likely to occur in this area, but not observed or captured; in
some cases, this may be the result of seasonal movements, rarity, or difficulty in
discovery owing to habits and /or size. L = found at nearby Lago Sandoval, but not
known at present from Cuzco Amazonico. An asterisk (* ) signifies that the species was
observed, but is not represented by voucher specimens. Macrohabitats: A = arboreal;
C = commensal with human habitations; T = terrestrial; V = volant; W = aquatic or
aquatic margin.

Taxon

Abundance

Macrohabitat

Marsupicarnivora

Didelphidae

Caluromys lauatus
Chironectes minimus
Didelphis marsiipialis
Marmosa mwina
Marmosops noctivagiis
M. parviclens
Metachirus nmlicaiidatus
Micoweus regina
Mouodelphis adusta
P Inlander andcrsoni
Chiroptera

Emballonuridae

Peroptevyx leucoptera
Rhynchonyctcris naso *
Saccopteryx hdineata
S. leptura

Molossidae

Molossus coihensis

Noctilionidae

Noctilio alhiventris
N. leporinus

R

A,T, W

P

W

A

A.T,W

R

A,T

A

A,T

U

A,T

u

T

A

A,T

R

T

U

W,T

U

V

L

V

U

V

U

V

R

C
P

V

V
V

MAMMALS OF CUZCO AMAZONICO

Table 1 . Continued

Taxon

Abundance

Macrohabitat

Phyllostomidae

Anoiira caudifer

P

V

Artiheus lituratus

A

V

A. ohscuvus

A

V

A. planiwstris

A

V

Carol I ia hrevicaucla

A

V

C. castanea

A

V

C. perspiciUata

C

V

Chirodenna salviiii

R

V

C. villositni

R

V

Choeroniscus minor

R

V

Chrotopterus awitiis

R

V

Dermauura cinerea

U

V

D. hart a

P

V

Desmodus rotundus

R

V

Glossophaga soricina

C

V

Lionycteris spurrelli

P

V

Lonchophylla thomasi

R

V

MacrophyUiim macrophyllum

P

V

Micronycteris inegalotis

R

V

M. niiniita

R

V

Mimon crenulatum

R

V

Phylloderma stenops

P

V

Phyllostomus elongatiis

U

V

P. hastatus

c

V

Platyrrhimis braclncephalus

u

V

P. dorsalis

p

V

P. infiisciis

R

V

Rhiiiophylla puniilio

C

V

Sphaeronycteris toxophylluni

P

V

Sturuira liliiuu

C

V

S. magna

P

V

S. tildae

R

V

Tonafia hidens

P

V

T. silvicola

R

V

Trachops cirrhosus

R

V

Uroderma bilohatum

C

V

U. magnirostritm

R

V

Vampyressa macconnelli

C

V

V. piisilla

C

V

Vampyrodes caraccioli

P

V

\ dmpyruni spectrum

R

V

Thyropteridae

Tliyroptera tricolor

R

V

8 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 145

Table 1 . Continued

Tax on

Abundance

Macrohabitat

Vespertilionidae

Eptesicus brasiliensis

R

V

Lasiiinis ega

R

V

Myotis albescens

R

V

M. nigricans

U

V

M. riparius

R

V

Primates

Callithrichidae

Callimico goeldii

P

A.T

Cebuella pygmaea

P

A

Saguinus fuscicoUis *

C

A

S. imperator

P

A

Cebidae

Alouatta seniculus *

R

A

Aotus nigriceps *

C

A

Ateles paniscus

P

A

Callicebus moloch

P

A

Cebus albifrons *

R

A

C. apella *

R

A

Lagothrix lagothhcha *

R

A

Pithecia monaclms

P

A

Saimih boliviensis *

C

A

Xenarthra

Bradypodidae

Bvadypus variegatus

U

A

Dasypodidae

Cabassous unicinctus

P

T

Dasypus kappleri

P

T

D. novemciuctus

u

T

Priodontes maximus

p

T

Megalonychidae

Choloepiis hojfmanni

u

A

Mymiecophagidae

Cyclopes didactyliis

p

A

Myrmecophaga tridactyla *

R

T

Tamandua tetradactyla *

u

A,T

Lagomorpha

Leporidae

Sylvilagus brasiliensis *

u

T

Rodentia

Agoutidae

Agouti paca *

u

T

MAMMALS OF CUZCO AMAZONICO

Table 1 . Continued

Taxon

Abundance

Macrohabitat

R

T

R

T

R

W

A

A

U

A

R

A

U

C

A

T

A

T

U

T

u

A

u

A

A

T

R

T

Cricetidae

Neacomys spiuosus

N. temiipes

Nectomys squamipes

Oecomys hicolov

O. tapajinus

O. super a ns

Oligoryzomys microtis

Oiyzomys capita

O. niditus

O. yunganus

RJiipidomys couesi

Rlupidomys sp. 2
Dasyproctidae

Dasyprocta variegata

Myoprocta pratti
Dinomyidae

Dinoniys hnuuckii
Echimyidae

Dactylomys dactylinus

Isothrix bistriata

Mesomys hispidus

Proechimys brevicauda

P. simonsi

P. steerei
Erethizontidae

Coendoii bicolor
Hydrochaeridae

Hydrochaeris hydrochaeris
Muridae

Rattus rattus
Sciuridae

Sciurus ignitus

S. spadiceiis
Carnivora
Canidae

Atelocynus microtis *

Speothos venaticus
Felidae

Felis concolor *

F. pardalis

F. wiedii

F. yagoiiaroundi *

P
R
A
C
U
C

R

P

U

C

A

R
P

R
R
P
R

A,T

A
A
A
T
T
T

A,T

A

C

A
A

T

T

T

T

T,A

T

10 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 145

Table 1 . Continued

Taxon

Abundance

Macrohabitat

Panthera oiica *

R

T

Mustelidae

Galictis virtata *

R

T.W

Eira harhara *

R

T,A

Luira hngicaudis *

L

W

Miistela sp.*

R

T

Pteronura hrasiliensis *

L

W

Procyonidae

Bassaricyon gahhii

P

A

Nasua nasiia

P

T,A

Potosflavus *

R

A

Perissodactyla

Tapiridae

Tapinis terrestris *

R

W

Artiodactyla

Cervidae

Mazama americaua

P

T

M. gouazouhira

P

T

Odocoileiis virginianiis *

R

T

Tayassuidae

Tayassu pecari *

P

T

T. tajacn

U

T

ACKNOWLEDGMENTS

Field work was supported by National Geographic Society grant 40 1 6-89.
Funds for travel by Peruvian personnel were granted by the Consejo Nacional
de Ciencia y Tecnologia (CONCYTEC) to the Asociacion de Ecologia y
Conservacion (ECCO). We are grateful to William E. Duellman. Director of
BIOTROP, for his skillful coordination of held work at Cuzco Amazonico.
Philip S. Humphrey, Director of the Museum of Natural History. The
University of Kansas, provided considerable moral and financial support.
Gerardo Lamas M.. Director of the Museo de Historia Natural. Universidad
Nacional Mayor de San Marcos, arranged for interinstitutional collaboration.
B.Anthony Luscombe. Vice President of ECCO, was instrumental in providing
outstanding logistical support. We are particularly indebted to Jose E.
Koechlin, owner of the Cuzco Amazonico Lodge, whose good will and
interest provided impetus, as well as excellent facilities, for our work. Blgo.
Jose Purisaca P. of the Direccion General Forestal y de Fauna. Ministerio de
Agricultura. Lima, issued permits. James L. Patton. Barbara R. Stein, and Eric
Yensen graciously allowed us access to specimens from Cuzco Amazonico

MAMMALS OF CUZCOAMAZONICO 11

under their care and freely shared information with us. Michael D. Carleton,
Alfred L. Gardner, and Bruce D. Patterson pennitted us to examine South
American comparative material in collections under their stewardship. Michael
Carleton. Guy G. Musser, Karl F. Koopman, and Ronald H. Pine aided us with
the respective taxonomies of Oecomys; Oryzoinys: bats of the genera Artlheus,
Dcnnaniira, and Myotis: and mouse opossums. We thank our North American
and Peruvian colleagues who worked with us at Cuzco Amazonico and whose
contribution of specimens and /or observations of mammals aided our work
there. Daixel R. Frost assisted with specimen preparation in the field: Bryant
W. Buchanan and Erik R. Wild contributed significantly to our knowledge of
the mammals of Cuzco Amazonico, and John J. Weins provided masterful
assistance with armadillos. Thor Holmes skillfully prepared the skeletal
material. Cesar Ascorra. William Duellman, Philip Humphrey. Philip Myers,
James Patton. and Linda Trueb critiqued earlier versions of this manuscript,
providing comments that improved it substantially.

RESUMEN

El trabajo de campo realizado por el equipo mastozoologico del Programa
de Biodiversidad Biologica Neotropical (BIOTROP) en la Reserva Cuzco
Amazonico, Departamento de Madre de Dios, Peru, dio por resultado un
inventario de las especies de mamiferos que habitan en esta localidad. La
fauna de mamiferos de Cuzco Amazonico consta de un minimo de 100
especies pertenecientes a 9 ordenes. Esta fauna incluye 9 marsupiales, 42
quiropteros, 7 primates, 5 xenarthros, I lagomorfo, 24 roedores, 9 camivoros,
1 perisodactilo, y 2 artiodactilos. Siete especies representan nuevos registros
que extienden los ranges de distribucion conocidos. Los pequeiios roedores
y marsupiales ten-estres de Cuzco Amazonico son mas abundantes y diversos
en comparacion a los de otras localidades de la selva baja neotropical. Los
murcielagos, normalmente muy abundantes en estos liltimos sitios, son
menos comunes en Cuzco Amazonico.

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Davis, T. J., C. Fox. L. Salinas, G. Ballon, and C. Arana 1 99 1 . Annotated checklist

of the birds of Cuzco Amazonico. Peru. Occas. Pap. Mus. Nat. Hist., Univ.

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12 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 145

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