Full text of "Studies in neotropical mammalogy : essays in honor of Philip Hershkovitz"

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Zoology

NEW SERIES, NO. 39

STUDIES IN NEOTROPICAL MAMMALOGY
Essays in Honor of Philip Hershkovitz

Edited by Bruce D. Patterson and Robert M. Timm

December 31, 1987
Publication 1382

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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STUDIES IN NEOTROPICAL MAMMALOGY
Essays in Honor of Philip Hershkovitz

Phiup Hershkovttz

FIELDIANA

Zoology

NEW SERIES, NO. 39

STUDIES IN NEOTROPICAL MAMMALOGY
Essays in Honor of Philip Hershkovitz

Edited by

Bruce D. Patterson

Division of Mammals

Field Museum of Natural History

Chicago, Illinois 60605-2496

Robert M. Timm

Division of Mammals

Field Museum of Natural History

Present address:

Museum of Natural History

Department of Systematics and Ecology

University of Kansas

Lawrence, Kansas 66045

Accepted for publication July 30, 1985
December 31, 1987
Publication 1382

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

HELDIANA: ZOOLOGY

New Series, No. 39

Studies in Neotropical Mammalogy:

Essays in Honor of Philip Hershkovitz

Bruce D. Patterson and Robert M. Timm, Editors

ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

Preface vii

A Biographical Sketch of Philip Hershkovitz, with a Complete Scientific Bibliography 1

Bruce D. Patterson

A History of the Recent Mammalogy of the Neotropical Region from 1492 to 1850 11

Philip Hershkovitz

A New Superfamily in the Extensive Radiation of South American Paleogene Marsupials 99

Rosendo Pascual and Alfredo A. Carlini

An Additional 14-Chromosome Karyotype and Sex-Chromosome Mosaicism in South American

Marsupials 1 1 1

Milton H. Gallardo and Bruce D. Patterson

Notes on the Black-Shouldered Opossum, Caluromysiops irrupta 117

Robert J. Izor and Ronald H. Pine

Feeding Habits of the Opossum {Didelphis marsupialis) in Northern Venezuela 125

Gerardo A. Cordero R. and Ruben A. Nicolas B.

Notes on Distribution of Some Bats from Southwestern Colombia 133

Michael S. Alberico

Distributional Records of Bats from the Caribbean Lowlands of Belize and Adjacent Guatemala

and Mexico 137

Timothy J. McCarthy

New Species of Mammals from Northern South America: Fruit-Eating Bats, Genus Artibeus

Leach 163

Charles O. Handley. Jr.

Seasonality of Reproduction in Peruvian Bats 173

Gary L. Graham

Tent Construction by Bats of the Genera Artibeus and Uroderma 187

Robert M. Timm

Comparative Ultrastructure and Evolutionary Patterns of Acinar Secretory Product of Parotid

Salivary Glands in Neotropical Bats 213

Carleton J. Phillips, Toshikazu Nagato, and Bernard Tandler

Distribution of the Species and Subspecies of Cebids in Venezuela 231

Roberta Bodini and Roger Perez- Hernandez

Host Associations and Coevolutionary Relationships of Astigmatid Mite Parasites of New World

Primates. I. Families Psoroptidae and Audycoptidae 245

Barry M. OConnor
Notes on Bolivian Mammals. 2. Taxonomy and Distribution of Rice Rats of the Subgenus Oli-

goryzomys 261

Nancy Olds and Sydney Anderson

New Records and Current Status of Euneomys (Cricetidae) in Southern South America 283

Jose L. Ydhez, Juan C Torres-Mura. Jaime R. Rau, and Luis C. Contreras

Morphological Variation, Karyology, and Systematic Relationships of Heteromys gaumeri (Ro-

dentia: Heteromyidae) 289

Mark D. Engstrom. Hugh H. Genoways, and Priscilla K. Tucker

Species Groups of Spiny Rats, Genus Proechimys (Rodentia: Echimyidae) 305

James L. Patton

An Assessment of the Systematics and Evolution of the Akodontini, with the Description of New

Fossil Species of Akodon (Cricetidae: Sigmodontinae) 347

Osvaldo A. Reig

Biogeography of Octodontid Rodents: An Eco-Evolutionary Hypothesis 40 1

Luis C. Contreras, Juan C. Torres-Mura, and Jose L. Ydnez

Population Dynamics and Ecology of Small Mammals in the Northern Chilean Semiarid

Region 413

Peter L. Meserve and Eric Le Boulenge

Demography and Reproduction of the Silky Desert Mouse (Eligmodontia) in Argentina 433

Oliver Pearson. Susana Martin, and Javier Bellati

Baculum of the Lesser Andean Coati, Nasuella olivacea (Gray), and of the Larger Grison, Galictis

vittata (Schreber) 447

Edgardo Mondolfi

Origin, Diversification, and Zoogeography of the South American Canidae 455

Annalisa Bert a

Comparative Cytogenetics of South American Deer 473

Angel E. Spot or no. Nadir Brum, and Mariela Di Tomaso

Faunal Representation in Museum Collections of Mammals: Osgood's Mammals of Chile 485

Bruce D. Patterson and Clare E. Feigl

Taxonomic Index 497

Subject Index 505

Preface

In the early 1 980s, we discussed the possibility
of a testimonial volume for Philip Hershkoviiz
with Larry Marshall, then with the Department of
Geology, Field Museum. As the senior mammal-
ogist at Field Museum and a student of South
American mammalogy for almost half a century,
Hershkovitz had generously provided invaluable
advice and assistance to each of us in the early
stages of our careers. We felt a Festschrift in his
honor might repay a portion of our debts to him
and, at the same time, serve as an independent,
lasting tribute to his life-work.

In the entire history of Field Museum, only
three testimonial volumes had been produced in
honor of museum scientists. Each recognized the
contributions of men who were both preeminent
scientists and museum administrators: Wilfred H.
Osgood, Chief Curator of Zoology, 1921-1941;
Karl P. Schmidt, Chief Curator of Zoology, 1941-
1956; and Rainer Zangerl, Chief Curator and
Chairman of Geology, 1962-1974. Although
Hershkovitz has never served in an upper-level
administrative capacity, his contributions to the
museum through distinguished and continuing re-
search clearly qualified him for this honor.

However, plans for a testimonial volume in
Fieldiana: Zoology did not materialize until No-
vember 1 983. By that time, Marshall had assumed
a new position at the University of Arizona and
Hershkovitz had just celebrated his 74th birthday.
Given realistic editing and publication schedules,
we were faced with the prospect of producing the
volume nearly midway between traditionally cel-
ebrated anniversary dates. Nevertheless, such tim-
ing is somehow fitting: Hershkovitz the man is
both extemporaneous and unconventional.

Another notable departure from the Festschrift
tradition is evident from the table of contents:
Hershkovitz himself is a contributor! On many
occasions Hershkovitz had lamented the lack of a
historical review of South American mammalogy.
During the present information explosion, scien-
tists are hard-pressed to keep up with current de-
velopments of direct relevance to their research;
much less are they afforded the occasion to amble
through historical records in Latin, German,
French, Spanish, and Portuguese, even though these
records are full of interesting and relevant infor-
mation. As a result of his 50 years in the discipline,
Hershkovitz may be unique in his broad knowl-
edge of both historical literature and current re-
search on Neotropical mammals. The editors

therefore prevailed upon him to write such a his-
torical survey to complement and enhance this
volume. We convinced him that, by assembling a
historical analysis of the subject, he would provide
a tremendous service to younger workers.

Other contributions to the volume came from
friends and colleagues of Hershkovitz. All share
an interest in the distribution, taxonomy, and nat-
ural history of Neotropical mammals, and each
one was inspired to honor Hershkovitz with their
contribution. Each of the contributions focuses on
those fields of Neotropical mammalogy to which
Hershkovitz has contributed most significantly.

We owe thanks to numerous persons connected
with this volume. First and foremost, Tanisse Be-
zin, Managing Editor of Field Museum Press, de-
serves recognition. Her keen eye for grammar and
style eliminated numerous editorial inconsisten-
cies forwarded by the volume editors. Graham
Harles, Field Museum Press copy editor, provided
skillful editing and proofreading. The Scientific
Editor for Fieldiana, Timothy Plowman, endured
countless interruptions during production of this
volume and served as corresponding editor for our
own papers. Translations of abstracts into Spanish
and Portuguese were kindly provided by Myriam
Ibarra (an Ecuadorean ichthyologist) and Debra
Moskovits (a Brazilian ecologist), who offered these
as their own tributes to Hershkovitz. Assistance
in assembling the indices was provided by Mary
Anne Rogers.

Finally, we are enormously indebted to a ded-
icated body of reviewers, who critically evaluated
papers in this volume. Their constructive advice
and recommendations made editorial tasks far
lighter. The editors gratefully thank: W. T. Atyeo,
P. V. August, K. Benirschke, W. A. Clemens, J.

A. Davis, W. B. Davis, M. R. Dawson, M. D.
Engstrom, J. Fooden, G. L. Forman, M. H. Ga-
llardo, A. L. Gardner, H. H. Genoways, W. E.
Glanz, M. S. Hafner, D. Hunsaker II, R. J. Izor,
J. A. W. Kirsch, K. F. Koopman, M. A. Mares,
R. E. Martin, T. J. McCarthy, G. G. Musser, P.
Myers, J. L. Patton, O. P. Pearson, R. H. Pine, W.

B. Quay, L. Radinsky, O. J. Reichman, D. S. Rog-
ers, R. W. Thorington, Jr., W. D. Tumbull, J. H.
Wahlert, S. D. Webb, C. Wemmer, J. O. Whitaker,
Jr., D. E. Wilson, R. G. Wolff, and A. E. Wood,
in addition to anonymous reviewers of our own
papers.

B. D. Patterson
R. M. TiMM
Chicago, Illinois

A Biographical Sketch of Philip Hershkovitz,
with a Complete Scientific Bibliography

Bruce D. Patterson

Philip Hershkovitz was bom October 12, 1909,
in Pittsburgh, Pennsylvania, to Aba Hershkovitz
and Bertha Halpem. He was the second of four
children and their only son. He attended primary
and secondary schools in Pittsburgh, graduating
from Schenley High School in February 1927. In
1929 he enrolled at the University of Pittsburgh
where he majored in zoology, serving as an Un-
dergraduate Assistant in that department during
1 930-1 931. Having exhausted Pittsburgh's course
offerings in zoology and seeking to pursue a career
in mammalogy, he was advised to transfer to
another school with an expanded curriculum (Har-
vard University, University of Michigan, or Uni-
versity of California, Berkeley). In his junior year
(1931), he transferred to the University of Mich-
igan at Ann Arbor because of its proximity to his
home. There he became an Undergraduate Assis-
tant in the Museum of Zoology, working under the
supervision of Professor and Curator Lee R. Dice
during 1931-1932. He supplemented the meager
earnings of this position with taxidermy jobs, which
supported him during the early years of the Great
Depression.

His first fieldwork was undertaken during the
summer of 1932. He went to the San Marcos re-
gion of Texas to collect blind cave salamanders
{Typhlomolge rathbuni) for Professor Uhlenhuth
of the University of Maryland Medical School.
Having a principal interest in mammals, he want-
ed to collect small mammals in areas surrounding
the caves, but Dice could spare no traps for him
and told him to purchase some in Texas.

While hitchhiking from Ann Arbor to Texas,
Hershkovitz stopped to visit friends in Chicago.
There, a chance visit to Field Museum of Nat-
ural History secured him the traps and supplies
he needed and seemingly set the course of his later

career. Colin Sanborn, then Curator of Mammals
during Wilfred Osgood's tenure as Chief Curator
of Zoology (1921-1941), befriended Hershkovitz
and loaned him the necessary supplies. As a con-
sequence, the mammals that Hershkovitz collect-
ed in Texas that first of many field seasons were
deposited in the Field Museum collections. He
now maintains that his chance visit to Field
Museum in 1932 indelibly fixed that institution
as the place at which to pursue his career goals.

Hershkovitz's formal education was delayed by
the worsening economic situation during 1 933. No
longer able to afford tuition, he sought advice on
subsistence during the Depression, and was told
that Ecuador and Paraguay were undoubtedly the
least expensive countries in this hemisphere in
which to live. Transportation costs decided the
issue, and in 1933 he set sail via the Grace Line
from New York to Guayaquil, Ecuador for the
whopping sum of $600, one-way.

He stayed in Ecuador until 1937. During this
period, he mastered Spanish and learned how to
live off the land in the Neotropics. His boots dis-
integrated after six months' time and thereafter he
went barefoot. He assembled a fine collection of
Ecuadorean mammals for the Museum of Zool-
ogy, University of Michigan, supporting his activ-
ities in part by selling horses bought on the Pe-
ruvian frontier.

He then returned to the University of Michigan
where he again enrolled as an undergraduate, grad-
uating in 1938 with a Bachelor of Science degree.
By this time. Dice had moved from the Museum
of Zoology to the Laboratory of Vertebrate Ge-
netics, and William H. Burt had assumed the cu-
ratorship in the Museum. Hershkovitz spent the
years 1938-1941 as a graduate student enrolled at
the University of Michigan, working on his Ecua-

PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ

dorean collection under Burt's direction. From
1939-1941, he was supported in this work by a
Graduate Assistantship. In 1940 he received his
Master of Science degree and immediately entered
the doctoral program.

Two years before the expected completion of
his doctoral program, the Curator of Reptiles and
Amphibians at the Museum of Zoology, Helen
Gage, told Hershkovitz about the Walter Rath-
bone Bacon Travelling Scholarship of the United
States National Museum. This program was cus-
tomarily reserved for postdoctoral support, but
Mrs. Gage strongly urged him to apply immedi-
ately. Thus encouraged, Hershkovitz submitted a
brief proposal for work in the Santa Marta region
of northern Colombia; his compliance with Mrs.
Gage's wishes in this matter was so perfunctory
that he failed to include a map of the proposed
itinerary. But Remington Kellogg at the National
Museum had long wished to obtain a Bacon Schol-
ar for the Mammal Division and asked Hersh-
kovitz to send the omitted material. Much to his
surprise, Hershkovitz was awarded the scholar-
ship and left Ann Arbor immediately for Wash-
ington. He spent two months there studying the
then very poor collection of Neotropical mam-
mals. Afterward he spent two years in Colombia
(1941-1943) collecting mammals, other verte-
brates, and ectoparasites. The resulting collection
was the National Museum's first large and repre-
sentative Neotropical mammal accession.

In 1943 Hershkovitz's work was interrupted by
World War II, and he returned to Ann Arbor to
enlist in the Armed Services. He was assigned to
the Office of Strategic Services (OSS) and served
from 1943-1946 in the European Theater. While
serving in France, he met Anne Marie Pierrette,
whom he married in 1946. The two returned to
the United States, where in 1946 and 1947 he
continued his Bacon Scholarship studies of Co-
lombian mammals in Washington. The first of
three children (Francine, Michael, and Mark) was
bom in 1947.

About this time, he was contacted regarding the
opening of a curatorial position at Field Museum
in Chicago, an opportunity he eagerly hailed for
several reasons: ( 1 ) The comprehensive collections
of Neotropical mammals at Field Museum would
be a tremendous resource for what he had already
decided would be his life's work; (2) he had the
highest regard for W. H. Osgood, who as a prin-
cipal authority on South American mammals
would be a great personal resource on which to
draw; (3) the press of family responsibibties made

continuation of his graduate studies untenable; and
(4) aspirations to a curatorial position had been
the raison d'etre of his graduate program; a cur-
atorial position made the graduate degree sujjer-
fluous. Thus he jumped at the offer of employment
at Field Museum, knowing full well that it marked
the end of his graduate program at Michigan. Like
many similar institutions, Michigan had a final
year-in-residence degree requirement. Unfortu-
nately, Osgood died in June of 1947, and what
might have been a remarkably productive ap-
prenticeship under Osgood never came to pass.

Upon his arrival at Field Museum, Hershkovitz
found an uncurated backlog of some four or five
years of accessions. Nevertheless, he wasted little
time in returning to the field, prompted in part by
postwar housing shortages in Chicago. (One can
almost hear him now, telling the Museum's Di-
rector Clifford Gregg that the nearest affordable
housing was in Bogota!) In 1 948 he and his family
moved to Colombia where he resumed his inven-
tory of the mammals of that country. He remained
in Colombia until the press of curatorial duties
and a gently delivered ultimatum from Sanborn
finally recalled him to Chicago in 1952.

The collections he made in Colombia, first for
the National Museum, then for Field Museum,
were to be the heart of all his subsequent research.
But unlike others studying the mammal faunas of
specific geographical regions, Hershkovitz found
it unsatisfying to assess the systematics of Colom-
bian mammals without following them across na-
tional boundaries. Studies of a species or species
group in Colombia led him to evaluate its context
within genera, families, and even orders; and the
remarkable diversity of Colombia's mammal fau-
na led him into most major groups and most Neo-
tropical subregions. In the course of his career, he
has published dozens of generic, tribal, and fa-
milial revisions, covering all 1 2 orders of Neo-
tropical mammals. Few spatial and temporal
boundaries have withstood the onslaught of his
studies of Neotropical mammals. As examples one
can point to the cosmopolitan Catalog of Living
Whales {\9()6)—2iiXtr all, most cetaceans do occur
in South American waters— and studies of Oli-
gocene and later fossils (1974, 1982).

One senses that the Department of Zoology dur-
ing Hershkovitz's early years was a stimulating,
harmonious one. Chief Curator Karl P. Schmidt
took an almost paternal interest in junior staff and
served as a confidant on the most personal of mat-
ters. In addition to Colin Sanborn, who was most
considerate of his junior curator's interests and

HELDIANA: ZOOLOGY

talents, Hershkovitz shared mammalogical prob-
lems and topics with Dwight Davis, Curator of
Anatomy, and Bryan Patterson, Curator of Ver-
tebrate Paleontology. During the early and mid-
1950s, Hershkovitz established a vigorous and
productive research program and participated in
all aspects of departmental affairs.

However, upon Schmidt's retirement in 1957,
Austin S. Rand became Chief Curator of Zoology,
and neither Rand nor Hershkovitz did much to
disguise their antipathy for one another. Over the
ensuing years, Hershkovitz increasingly detached
himself from museum operations, culminating with
Joseph Moore's appointment as Curator of Mam-
mals in 1961, and Hershkovitz's appointment that
year to Research Curator. No one before or since
has held this title at Field Museum. Hershkovitz
formally retired in 1971, although his work has
continued unabated as Curator Emeritus. During
his career, he assisted countless students in mam-
mal projects, but has served on only a single grad-
uate committee, that of Jack Fooden, now himself
a renowned biologist and primate specialist at Field
Museum.

Few scientists can claim the independence in
research that is indicated in Hershkovitz's bibli-
ography. Of his approximately 300 scientific, pop-
ular, and encyclopedia articles, only three repre-
sent collaborative efforts. The first, with William
P. Harris, an important benefactor of the Museum
of Zoology of the University of Michigan, was
suggested by Burt in recognition of Harris's inter-
ests in squirrels and in token repayment for his
patronage of the museum. The second, with Paul
Rode, came about one afternoon in the Museum
National d'Histoire Naturelle in Paris when
Hershkovitz offhandedly suggested that designat-
ing a lectotype might solve a nomenclatural prob-
lem that Rode had encountered in his research.
Rode insisted that Hershkovitz share authorship
on the resulting paper. Later, after further study
in the United States, Hershkovitz arrived at a con-
trary opinion and wrote a paper, with Rode as
coauthor, correcting their earlier one.

Independent thought is also exemplified by the
sometimes heated debates in which Hershkovitz
has participated over the years. His published re-
views and the discussion sections of many of his
papers record his clearly enunciated views on such
topics as the role of penial morphology in rodent
taxonomy, the age and derivation of the South
American fauna, panbiogeography, evolution of
pelage coloration, and the systematic position of
certain species (e.g., Dolichocebus). While such

firmly held views brand him as something other
than conciliatory or diplomatic, they accurately
reflect his abiding passion and zest in science. Un-
fortunately, some acerbic exchanges had the effect
of stifling the scientific dialogue to which they were
offered (e.g., penial morphology).

Hershkovitz has focused his research on Neo-
tropical mammals, their origin, evolution, dis-
persal, classification, nomenclature, and system-
atics. Specialists in these fields are well aware of
his impact. However, he is perhaps most widely
known for his work on three general topics of
Neotropical mammalogy: faunal origins, meta-
chromism, and New World monkeys. It would be
folly to attempt to review all of his research, and
more definitive appraisals on selected topics can
be found scattered throughout this volume. How-
ever, some comments on these general issues seem
in order.

As late as his revision of phyllotine rodents
( 1 962), Hershkovitz adhered to traditional notions
of the derivation of certain South American taxa,
notably "cricetid" rodents, from North and Mid-
dle American stocks. This hypothesis of origins
has been advocated most articulately by George
G. Simpson, Bryan Patterson, and Rosendo Pas-
cual, and more recently by Larry G. Marshall and
S. David Webb. However, in the early 1960s,
Hershkovitz was approached by Rupert Wenzel,
Curator of Insects at Field Museum, who ques-
tioned him on the evidence for Plio-Pleistocene
origins of the Neotropical cricetids. Wenzel's stud-
ies of the ectoparasites of Panamanian mammals
suggested much earlier. South American origins.
His interest piqued, Hershkovitz reviewed avail-
able evidence, synthesizing continental drift (which
was then becoming established in geological cir-
cles) and neontological studies of mammals (es-
pecially those of Hooper and Musser, which
showed a relatively sharp dichotomy between sim-
ple and complex penis-types of cricetids). He con-
cluded that continental drift permitted a much
greater role for paleotropical stocks in South
American faunal origins than was allowed by the
Simpsonian school, which in turn pointed to a
much greater time period for independent evo-
lution. Interestingly, and perhaps even character-
istically, Hershkovitz concluded that South Amer-
ican rodents were not only not derived from North
American stocks, but instead gave rise to them.
These views were published in 1966, 1969, and
1972.

Hershkovitz's theory of metachromism, or de-
terministic evolution of pelage coloration through

PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ

the loss of one or the other or both classes of hair
pigments (eumelanins and phaeomelanins), was
first pubHshed in 1968. Since then he has used it
repeatedly in describing geographic variation in
platyrrhine monkeys (e.g., 1977). However, the
origins of this concept stem from his earlier work
on the Sciunds granatensis group in northern Co-
lombia where populations of squirrels thoroughly
isolated from one another show similar progres-
sions of pelage patterns. Few workers other than
Neotropical primatologists (and not all of these)
have accepted his interpretations, although the
theory is potentially applicable to a variety of oth-
er, mostly diurnal taxa showing pelage pattern
variations. While Timothy Lawlor detailed some
theoretical misgivings with the theory in a 1969
paper in Evolution (rebutted by Hershkovitz in
1970), to my knowledge it has not been substan-
tially refuted. The theory is eminently testable:
refutation would simply entail showing that pelage
pattern variation of taxa arranged by metachro-
mism is not congruent with well-established phy-
letic patterns.

Finally, some explanation seems warranted for
Hershkovitz's current devotion to primates. In-
deed, many recent workers unschooled in mam-
malian systematics think of him as a primatolo-
gist. Nothing could be further from the truth, as
he hastens to point out. He had published several
articles on primates in the course of working up
his Colombian collections, but gave these taxa no
special attention until the 1960s. Then govern-
ment funding for primate studies soared, largely
because of interest in biomedical applications, es-
pecially for the complex and taxonomically con-
fused Callitrichidae. For almost 20 years, Hersh-
kovitz has focused first on the Callitrichidae and
Callimiconidae, now on the Cebidae. His slower
progress through these groups is attributable to the
vast body of current knowledge about them; his
1977 and subsequent works serve as model
syntheses of skin and skull morphology with bio-
chemistry, karyology, ethology, serology, and ep-
idemiology. By his own estimation, monkeys do
not culminate his studies of Neotropical mam-
mals, but rather represent a large and complex
group to be covered in his attempt to treat all South
American mammals. After seven years of work on
Volume II of his primate monograph, he has near-
ly completed generic revisions of cebids lacking
prehensile tails and is beginning comparative stud-
ies of their organ systems. In 1 984 he submitted
another grant proposal for this work, totaling one-
half million dollars in direct costs. His is not a

modest work; it has been described by Pine ( 1 982;
Vol. 6, Spec. Publ. Ser., Pymatuning Lab. Ecol.)
as "the most heroically monumental revisionary
monograph ever devoted to a Neotropical group."
In 1984, Hershkovitz turned 75 years old. The
14 years he spent in the field in South America
have served him well, for he seems younger than
many men 15 years his junior. His tireless energy
is best indicated by his habitual use of stairs rather
than elevators (even his two divisional offices are
three floors apart), a continuing program of field-
work (most recently in Brazil during 1986 and
1987), and a museum workday that extends from
9 a.m. to 6 p.m., uninterrupted by coffee breaks
or even lunch. Visitors to his home, now within
walking distance of the Museum, know of his office
there which relieves the chronic insomnia of ad-
vancing years. He is an outstanding cook, a genial
host, a trusted and valued friend, and an awe-
somely productive scientist.

Publications of Philip Hershkovitz
1938

1. A new caecilian from Ecuador. Occasional
Papers, Museum of Zoology, University of
Michigan, 370:1-3.

2. Two new squirrels fi-om Ecuador. Occasion-
al Papers, Museum of Zoology, University
of Michigan, 391:1-6 (with W. P. Harris).

3. A review of the rabbits of the andinus group
and their distribution in Ecuador. Occasion-
al Papers, Museum of Zoology, University
of Michigan, 393:1-15.

1940

4. Four new oryzomyine rodents from Ecua-
dor. Journal of Mammalogy, 21:78-84.

5. Notes on the distribution of the akodont ro-
dent, Akodon mollis, in Ecuador with a de-
scription of a new race. Occasional Papers,
Museum of Zoology, University of Michi-
gan, 418:1-3.

6. A new spiny mouse of the genus Neacomys
from eastern Ecuador. Occasional Papers,
Museum of Zoology, University of Michi-
gan, 419:1-4.

1941

7. The South American harvest mice of the ge-
nus Reithrodontomys. Occasional Papers,
Museum of Zoology, University of Michi-
gan, 441:1-7.

FIELDIANA: ZOOLOGY

1944

8. A systematic review of the Neotropical water
rats of the genus Nectomys (Cricetinae). Mis-
cellaneous Publications, Museum of Zool-
ogy, University of Michigan, 58:1-88.

1945

9. Designation d'un lectotype de Callithrix
penicillatus (E. Geoffroy). Bulletin du Mu-
seum National d'Histoire Naturelle, Paris
17(3):22 1-222 (with P. Rode).

1947

10. A correction. Journal of Mammalogy, 28(1):
68 (with P. Rode).

1 1 . Mammals of northern Colombia. Prelimi-
nary report no. 1 : Squirrels (Sciuridae). Pro-
ceedings of the United States National Mu-
seum, 97:1-46.

1948

12. Mammals of northern Colombia. Prelimi-
nary report no. 2: Spiny rats (Echimyidae),
with supplemental notes on related forms.
Proceedings of the United States National
Museum, 97:125-140.

13. Mammals of northern Colombia. Prelimi-
nary report no. 3: Water rats (genus Necto-
mys), with supplemental notes on related
forms. Proceedings of the United States Na-
tional Museum, 98:49-56.

1 4. The technical name of the Virginia deer with
a list of the South American forms. Pro-
ceedings of the Biological Society of Wash-
ington, 61:41-48.

1 5. Names of mammals dated from Frisch, 1 775,
and Zimmermann, 1777. Journal of Mam-
malogy, 29(3):272-277.

1949

1 6. Technical names for the fallow deer and Vir-
ginia deer. Journal of Mammalogy, 30(1):
94.

1 7. Generic names of the four-eyed pouch opos-
sum and the woolly opossum (Didelphidae).
Proceedings of the Biological Society of
Washington, 62:11-12.

18. Technical names of the African muishond
(genus Zorilla) and the Colombian hog-nosed
skunk (genus Conepatus). Proceedings of the
Biological Society of Washington, 62: 13-16.

1 9. Mammals of northern Colombia. Prelimi-
nary report no. 4: Monkeys (Primates), with
taxonomic revisions of some forms. Pro-
ceedings of the United States National Mu-
seum, 98:323-427.

20. Mammals of northern Colombia. Prelimi-
nary report no. 5: Bats (Chiroptera). Pro-
ceedings of the United States National Mu-
seum, 99:429-454.

21. Status of names credited to Oken, 1816.
Journal of Mammalogy, 30(3):289-301.

22. Tapirs: Strange mammals native to Asia and
tropical America from Mexico south. Chi-
cago Natural History Museum Bulletin,
20(9):6-7.

1950

23. Mammals of northern Colombia. Prelimi-
nary report no. 6: Rabbits (Leporidae), with
notes on the classification and distribution
of the South American forms. Proceedings
of the United States National Museum, 100:
327-375.

1951

24. Mammals from British Honduras, Mexico,
Jamaica and Haiti. Fieldiana: Zoology,
31(47):547-569.

1953

25. Zorilla I. Geoffroy and Spilogale Gray, ge-
neric names for African and American pole-
cats, respectively. Journal of Mammalogy,
34(3):378-382.

26. Four years on a zoological expedition in Co-
lombia. Chicago Natural History Museum
Bulletin, 24(l):3-4.

27. The reindeer— Important to man in fact and
fancy. Chicago Natural History Museum
Bulletin, 24(12):3-4.

1954

28. Mammals of northern Colombia, Prelimi-
nary report no. 7: Tapirs (genus Tapirus),
with a systematic review of American species.
Proceedings of the United States National
Museum, 103:465-496.

29. What the groundhog undergoes to make a
"holiday." Chicago Natural History Mu-
seum Bulletin, 25(2):3-4.

30. Who's a cow? Chicago Natural History Mu-
seum Bulletin, 25(7):5.

PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ

3 1 . Some ecological aspects of natural versus ar-
tificial rehabilitation of a water basin area in
Bogota, Colombia. Boletin del Instituto de
U Salle, Bogota, 41(193/194):80-83.

1955

32. South American marsh rats genus Holochi-
lus with a summary of sigmodont rodents.
Fieldiana: Zoology, 37:639-673.

33. [Review] Mammals, a guide to familiar
American species. Chicago Natural History
Museum Bulletin, 26(7):7.

34. Notes on American monkeys of the genus
Cebus. Journal of Mammalogy, 36:449-452.

35. Status of the generic name Zorilla (Mam-
malia): Nomenclature by rule or by caprice.
Proceedings of the Biological Society of
Washington, 68:185-192.

36. On the cheek pouches of the tropical Amer-
ican paca. Agouti paca (Linnaeus, 1766).
Saiigetierkundliche Mitteilungen, 3(2):67-70.

37. Know your rabbits. Sports Afield, 134(6):
36-41,88.

1956

38. Comments on Galerella Gray, Herpestes II-
liger, Leucomitra Howell, Icticyon Lund,
Lutreola Wagner, Oryctogale Merriam,
Paracynictis Pocock. Opinion 384 Interna-
tional Commission of Zoological Nomen-
clature, 12(5):71-190(intext).

39. Critical remarks on the status of names in
Oken's "Lehrbuch." Opinion 417, Interna-
tional Commission on Zoological Nomen-
clature, 14(l):33-35.

1957

40. Comments on Canis dingo Meyer. Opinion
451, International Commission on Zoolog-
ical Nomenclature, 15(17):335-336.

41. Comments on the validation of Muntiacus
Rafinesque. Opinion 460, International
Commission on Zoological Nomenclature,
15(26):467-468.

42. Comments on the generic name Mormoops
Leach. Opinion 462, International Com-
mission on Zoological Nomenclature, 16(1):
8-9.

43. Comments on Sciurus gambianus. Opinion
464, International Commission on Zoolog-
ical Nomenclature, 16(3):36-39.

44. Comments on the validation of silvestris
Schreber, 1777 [Felis {catus) silvestris].

Opinion 465, International Commission on
Zoological Nomenclature, 16(4):49.

45. Comments on the validation of the name
Phacochoerus Cuvier. Opinion 466, Inter-
national Commission on Zoological No-
menclature, 16(5):67-68.

46. Comments on the validation of the name
Odobenus Brisson. Opinion 467, Interna-
tional Commission on Zoological Nomen-
clature, 16(6): 84-8 5.

47. The systematic position of the marmoset,
Simia leonina Humboldt (Primates). Pro-
ceedings of the Biological Society of Wash-
ington, 70: 1 7-20.

48. The type locality of Bison bison Linnaeus.
Proceedings of the Biological Society of
Washington, 70:31-32.

49. A synopsis of the wild dogs of Colombia.
Novedades Colombianas Museo de Historia
Naturale Universidad del Cauca (Popayan),
no. 3:157-161.

50. On the possible occurrence of the spectacled
bear, Tremarctos ornatus(F. Cuvier, 1825),
in Panama. Saugetierkundliche Mitteilun-
gen, 5(3): 122-1 23.

1958

5 1 . [Review] Biological investigations in the Sel-
va Lacandona, Chiapas, Mexico. Quarterly
Review of Biology, 33(1):67.

52. [Review] Mammals of the Anglo-Egyptian
Sudan, by Henry Setzer. Quarterly Review
of Biology, 33:82.

53. Technical names of the South American
marsh deer and pampas deer. Proceedings
of the Biological Society of Washington, 71:
13-16.

54. Type localities and nomenclature of some
American Primates, with remarks on sec-
ondary homonyms. Proceedings of the Bi-
ological Society of Washington, 71:53-56.

55. Stabilization of zoological nomenclature by
a "Law of prescription." Bulletin of Zoolog-
ical Nomenclature, 15B(20/21):630-632.

56. A critique of Professor Chester Bradley's
"Principle of conservation." Bulletin of Zoo-
logical Nomenclature, 15B(25/28):9 11-913.

57. The status of secondary homonyms and the
concept of permanent rejection. Bulletin of
Zoological Nomenclature, 15B(37/38):1242-
1243.

58. A geographic classification of Neotropical
mammals. Fieldiana: Zoology, 36(6):583-
620.

FIELDIANA: ZOOLOGY

59. The metatarsal glands in white-tailed deer
and related forms of the Neotropical region.
Mammalia, 22(4): 5 3 7-546.

1959

60. The scientific names of the species of ca-
puchin monkeys (Cebus Erxleben). Proceed-
ings of the Biological Society of Washington,
72:1-4.

6 1 . Two new genera of South American rodents
(Cricetinae). Proceedings of the Biological
Society of Washington, 72:5-10.

62. A new species of South American brocket,
genus Mazama (Cervidae). Proceedings of
the Biological Society of Washington, 72:
45-54.

63. A new race of red brocket deer {Mazama
americana) from Colombia. Proceedings of
the Biological Society of Washington, 72:
93-96.

64. The type locality of Felix concolor concolor
Linnaeus. Proceedings of the Biological So-
ciety of Washington, 72:97-100.

65. Nomenclature and taxonomy of the Neo-
tropical mammals described by Olfers, 1818.
Journal of Mammalogy, 40(3):337-353.

1960

66. Supposed ape-man or "missing link" of
South America. Chicago Natural History
Museum Bulletin, 31(4):6-7.

67. [Review] The Mammals of North America.
Chicago Natural History Museum Bulletin,
31(5):6-7.

68. Publication dates for names of the Anubis
baboon. Journal of Mammalogy, 41 (3):402-
403.

69. Mammals of northern Colombia. Prelimi-
nary report no. 8: Arboreal rice rats, a sys-
tematic revision of the subgenus Oecomys,
genus Oryzomys. Proceedings of the United
States National Museum, 1 10:513-568.

1961

70. On the South American small-eared zorro
Atelocynus microtis Sclater (Canidae). Field-
iana: Zoology, 39(44):505-523.

71. On the nomenclature of certain whales.
Fieldiana: Zoology, 39(49):547-565.

72. "This is a mammal." Chicago Natural His-
tory Museum Bulletin, 3 2(6): 3.

1962

73. Suriname zoological expedition. Chicago
Natural History Museum Bulletin, 33(4):3,
7-8.

74. Bats and their menus. Chicago Natural His-
tory Museum Bulletin, 33(8):2-3, 5-8.

75. Evolution of Neotropical cricetine rodents
(Muridae) with special reference to the phyl-
lotine group. Fieldiana: Zoology, 46:1-524.

1963

76. A systematic and zoogeographic account of
the monkeys of the genus Callicebus (Cebi-
dae) of the Amazonas and Orinoco River
basins. Mammalia, 27(l):3-79.

77. [Review] Primates. Comparative Anatomy
and Taxonomy. Vol. V, Cebidae, part B., A
Monograph; Edinburgh University Press.
American Journal of Physical Anthropolo-
gy, 21(l):92-98.

78. [Review] Primates. Comparative Anatomy
and Taxonomy. Vol. V, Cebidae, part B., A
Monograph; Edinburgh University Press.
American Journal of Physical Anthropolo-
gy, 2 1(3):39 1-398.

79. Notes on South American dolphins of the
genera Inia, Sotalia and Tursiops. Journal
of Mammalogy, 44(1):98-103.

80. The nomenclature of South American pec-
caries. Proceedings of the Biological Society
of Washington, 76:85-88.

81. The Recent mammals of South America.
Proceedings of the XVI International Con-
gress of Zoology, Washington, D.C., Aug.
20-27, 1963.

82. Comments on the proposed suppression of
Zorilla I. Geoffroy, 1826. Z.N.(S.) 758. Bul-
letin of Zoological Nomenclature, 20(4):242-
244.

1965

83. Primate research and systematics. Science,
147(3662):1 156-1 157.

84. The importance of taxonomy in primate re-
search and care. Illinois Society for Medical
Research— Chicago Branch— Animal Care
Panel Bulletin, 39:2 pp.

1966

85. Catalog of living whales. Bulletin of the
United States National Museum, 246: 1-259.

PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ

86. Taxonomic notes on tamarins, genus Sa-
guinus (Callithricidae, Primates), with de-
scriptions of four new forms. Folia Prima-
tologica, 4:381-395.

87. On the identification of some marmosets,
family Callithricidae (Primates). Mamma-
lia, 30(2):327-332.

88. What ever happened to hairy man? Science,
153:362.

89. Comments on the proposal for conservation
oi Pan Oken, 1816, and Panthera Oken,
1816. Bulletin of Zoological Nomenclature,
23(2/3):67-69.

90. [Review] Evolutionary and Genetic Biology
of Primates, vol. II; Academic Press. Amer-
ican Biology Teacher, 28(7):564-565.

91. Comments on the proposed suppression of
Meles montanus Richardson, 1829, and M.
jeffersonii Harlan, 1825. Z.N.(S.) 1639. Bul-
letin of Zoological Nomenclature, 22(5/6):
336-337.

92. On the status of Procyon brachyurus Wieg-
mann and P. obscurus Wiegmann. Z.N.(S.)
1640. Bulletin of Zoological Nomenclature,
22(5/6):338.

93. South American swamp and fossorial rats of
the Scapteromyine group (Cricetinae, Mu-
ridae) with comments on the glans penis in
murid taxonomy. Zeitschrift fiir Saugetier-
kunde, 31(2):81-149.

94. Status of the black-footed ferret in Wyo-
ming. Journal of Mammalogy, 47(2):346-
347.

95. Comments on the proposal on Zorilla by Dr.
Van Gelder and the counter proposal by Dr.
China. Z.N.(S.) 758. Bulletin of Zoological
Nomenclature, 2 3(2/3): 74-7 5.

96. Museum taxonomy serves medical research.
Bulletin of the Field Museum of Natural
History, 37(9):4-7.

97. Mice, land bridges and Latin American fau-
nal interchange, pp. 725-751. In Wenzel, R.
L., and V. J. Tipton, eds.. Ectoparasites of
Panama. Field Museum of Natural History,
Chicago.

1967

98. (Review] Evolutionary and Genetic Biology
of Primates, vol. I; Academic Press. Amer-
ican Biology Teacher, Nov. 1967:665.

99. Reply to Mayr's comment on the proposed
preservation oi Pan from Oken, 1816.
Z.N.(S.) 482. Bulletin of Zoological Nomen-
clature 24(5): 1 p.

1 00. Dynamics of rodent molar evolution: A study
based on New World Cricetinae, family Mu-
ridae. Journal of Dental Research, Suppl. to
46(5):829-842.

1968

101. Metachromism or the principle of evolu-
tionary change in mammalian tegumentary
colors. Evolution, 22(3):556-575.

102. [Review] Dynamics of rodent molar evolu-
tion: A study based on New World Cricet-
inae, family Muridae. Oral Research Ab-
stracts, May 1968.

1969

103. Comments on Cynocephalus Boddaert ver-
sus Galeopithecus Pallas. Z.N.(S.) 1 792. Bul-
letin of Zoological Nomenclature, 25(6):202-
203.

1 04. The evolution of mammals on southern con-
tinents. VI. The Recent mammals of the
Neotropical Region: A zoogeographic and
ecological review. Quarterly Review of Bi-
ology, 44(1): 1-70.

1970

105. The decorative chin. Field Museum of Nat-
ural History Bulletin, 41(5):7-10.

106. Dental and periodontal diseases and abnor-
malities in wild-caught marmosets (Pri-
mates— Callithricidae). American Journal of
Physical Anthropology, 32(3):377-392.

107. [Review] Taxonomy and Evolution of the
Monkeys of Celebes (Primates: Cercopithe-
cidae); Bibliotheca Primatologica, No. 10;
Karger. Folia Primatologica, 13(l):75-76.

108. Metachromism like it is. Evolution, 24(3):
644-648.

1 09. Notes on Tertiary platyrrhine monkeys and
description of a new genus for the Late Mio-
cene of Colombia. Folia Primatologica, 12:
1-37.

110. Errata: Notes on Tertiary platyrrhine mon-
keys and description of a new genus for the
Late Miocene of Colombia. Foha Primato-
logica, 12:1-37(1970).

111. Cerebral fissural patterns in platyrrhine
monkeys. Folia Primatologica, 13:213-240.

112. [Review] The Squirrel Monkey; Academic
Press. Journal of Mammalogy, 51(4):836-
839.

113. Supplementary notes on Neotropical Ory-
zomys dimidiatus and Oryzomys hammondi
(Cricetinae). Journal of Mammalogy, 51(4):
789-794.

FIELDIANA: ZOOLOGY

1971

1977

1 14. Stapedial processes in tympanic cavities of
capuchin monkeys (Cebus). Journal of
Mammalogy, 52(3):607-609.

115. Basic crown patterns and cusp homologies
of mammalian teeth, pp. 95-150. In Dahl-
berg, A. A., ed., Dental Morphology and
Evolution. University of Chicago Press, Chi-
cago.

116. A new rice rat of the Oryzomys palustris
group (Cricetinae, Muridae) from north-
western Colombia, with remarks on distri-
bution. Journal of Mammalogy, 52(4):700-
709.

126. Comment: Pan and Panthera or Oken's
Lehrbuch? Z.N.(S.) 482. Bulletin of Zoolog-
ical Nomenclature, 33(3/4): 135-1 36.

127. [Review] Catalogue of Primates in the Brit-
ish Museum (Natural History). I. Families
Callitrichidae and Cebidae; British Museum
(Natural History). Folia Primatologica, 28:
315.

128. Living New World Monkeys (Platyrrhini).
With an Introduction to Primates. Volume
I. University of Chicago Press, Chicago,
xiv +1117 pp.

1972

1 1 7. Notes on New World monkeys. Internation-
al Zoo Yearbook, 12:3-12.

118. The Recent mammals of the Neotropical
Region: A zoogeographic and ecological re-
view, pp. 31 1-431. In Keast, A., F. C. Erk,
and B. Glass, eds.. Evolution, Mammals and
Southern Continents. State University of
New York Press, Albany.

1974

119. The ectotym panic bone and origin of higher
primates. Folia Primatologica, 22:237-242.

120. A new genus of Late Oligocene monkey
(Cebidae, Platyrrhini) with notes on post-
orbital closure and platyrrhine evolution.
Folia Primatologica, 21:1-35.

1975

121. [Review] Taxonomic Atlas of Living Pri-
mates; Academic Press. American Journal
of Physical Anthropology, 41:155-156.

122. The scientific name of the \tsstv Noctilio
(Chiroptera), with notes on the chauve-sou-
ris de la Vallee d'Ylo (Peru). Journal of
Mammalogy, 56(l):242-247.

123. Comments on the taxonomy of Brazilian
marmosets (Callithrix, Callitrichidae). Folia
Primatologica, 24:137-172.

1976

124. The taxonomic status of """Noctilio ruber
Rengger." Mammalia, 40(1): 164-166.

125. Comments on generic names of four-eyed
opossums (family Didelphidae). Proceed-
ings of the Biological Society of Washington,
89(23):295-304.

1979

1 29. Races of the emperor tamarin, Saguinus im-
perator Goeldi (Callitrichidae, Primates).
Primates, 20(2):277-287.

1 30. The species of sakis, genus Pithecia (Cebi-
dae, Primates), with notes on sexual dichro-
matism. Folia Primatologica, 31:1-22.

1981

131. Comparative anatomy of platyrrhine man-
dibular cheek teeth dpm4, pm4, ml with
particular reference to those oT Homunculus
(Cebidae), and comments on platyrrhine
origins. Folia Primatologica, 35:179-217.

132. Philander and four-eyed opossums once
again. Proceedings of the Biological Society
of Washington, 93(4):943-946.

1982

133. Supposed squirrel monkey affinities of the
late Oligocene Dolichocebus gaimanensis.
Nature, 298(5870):20 1-202.

134. Subspecies and geographic distribution of
black-mantle tamarins Saguinus nigricollis
Spix (Primates: Callitrichidae). Proceedings
of the Biological Society of Washington,
95(4):647-656.

135. Neotropical deer (Cervidae). Part I. Pudus,
genus Pudu Gray. Fieldiana: Zoology, n.s.,
11:1-86.

136. The staggered marsupial lower third incisor
(I3), pp. 191-200. In Buffetaut, E., P. Jan-
vier, J. C. Rage, and P. Tassy, eds., Phylo-
genie et Paleobiogeographie. Livre jubiliare
en I'honneur de Robert Hoffstetter. Geobios,
memoire special 6, Lyon.

PATTERSON: BIOGRAPHY OF PHILIP HERSHKOVITZ

1983

137. Two new species of night monkeys, genus
Aotus (Cebidae, Platyrrhini): A preliminary
report on Aott4s taxonomy. American Jour-
nal of Primatology, 4:209-243.

138. On the validity of the family-group name
Callitrichidae (Platyrrhini, Primates). Mam-
malia, 48:153.

1 39. Taxonomy of squirrel monkeys, genus Sai-
miri (Cebidae. Platyrrhini): A preliminary
report with description of a hitherto un-
named form. American Journal of Prima-
tology, 6:257-312.

140. [Review] Mammalian Biology in South
America. M. A. Mares and H. H. Genoways,
eds. Ecology, 65(6): 1944-1 945.

1985

141.

1986

142.

A preliminary taxonomic review of the South
American bearded saki monkeys, genus Chi-
roptes (Cebidae, Platyrrhini), with the de-
scription of a new subspecies. Fieldiana: Zo-
ology, n.s., 27:1-46.

[Review] Handbook of Squirrel Monkey Re-
search. L. A. Rosenblum and C. L. Coe, eds.
Quarterly Review of Biology, 61:286-287.

143. The piebald saki. Field Museum of Natural
History Bulletin, 57(2):coverplate + 24-25.

1987

144. Uacaries, New World monkeys of the genus
Cacajao (Cebidae, Platyrrhini): A prelimi-
nary taxonomic review with the description
of a new subspecies. American Journal of
Primatology, 12:1-53.

1 45. First South American record of Coue's marsh
rice rat, Oryzomys couesi. Journal of Mam-
malogy, 68(1): 152-1 54.

146. The titi. Field Museum of Natural History
Bulletin, 58(6): 11-15.

147. The taxonomy of South American sakis, ge-
nus Pithecia (Cebidae, Platyrrhini): A pre-
liminary report and critical review with the
description of a new species and a new sub-
species. American Journal of Primatology,
12:387-468.

In Press

148. More on the Homunculus Dpm4 and ml
and comparisons with Alouatta and Stirto-
nia (Primates, Platyrrhini, Cebidae). Amer-
ican Journal of Primatology.

HELDIANA: ZOOLOGY

A History of the Recent Mammalogy

of the Neotropical Region from 1492 to 1850

Philip Hershkovitz

ABSTRACTS

The history of Neotropical mammalogy began with the first voyage of Christopher Colum-
bus in 1492. The earliest notices were purely anecdotal, recorded by Spanish chroniclers from
the mouths of the sailors on their return from voyages of discovery during the 1 5th and 1 6th
centuries. Colonization of the Guianan and Brazilian coasts during the 1 7th century provided
opportunities for inventories and descriptions of the mammals by trained European naturalists
and physicians. The systematization and scientific naming of the known Brazilian species by
Carolus Linnaeus in 1758 were based primarily on the mammals described in the 17th century
monograph of Brazilian biota by Georg Marcgraf The actual collection and preservation of
mammals for study, however, began in the 18th century with the Brazilian-bom Alexandre
Rodrigues Ferreira. The 18th and first half of the 19th century was an explosive and romantic
period of independently or govemmentally sponsored scientific expeditions for field observa-
tions, collections, preservations, and taxonomic studies of the specimens shipped to European
museums and private collectors. Outstanding among the naturalists who made significant con-
tributions to mammalogy during this period are Alexander von Humboldt, Johann Baptist
Ritter von Spix (Brazil), Maximilian Prinz Wied-Neuwied (Brazil), Johann Natterer (Brazil),
Sir Robert Herman Schomburgk and Richard Schomburgk (Guyana), Claudio Gay (Chile),
Johann Jakob von Tschudi (Peru), Felix de Azara (Paraguay), Rudolph Rengger (Paraguay),
Alcide Charles Victor d'Orbigny (Argentina, Bolivia), and Charles Robert Darwin (Patagonia
and Galapagos). Their itineraries, collections of mammals, taxonomies, and some field notes are
included in the accounts of these and other noteworthy naturalists. By the middle of the 1 9th
century, the mammalian fauna of South America became the best known of any continent with
exception of the western European part of Eurasia. The problems of origins and distribution
of Neotropical mammals intrigued scholars from among the earliest chroniclers down to pre-
evolutionary Darwin. Their concepts on these subjects are briefly discussed.

La historia de mastozoolo^a neotropical empieza con el primer viaje transatlantico de Cris-
tobal Colon en 1492. Poco despues de desembarcarse de sus viajes de regreso los descrubridores
y conquistadores del Mundo Nuevo en los siglos quince y diez y seis contaron a los cronistas
espaiioles de las cosas curiosas que encontraron. Colonizacion de las costas guyanas y brasileras
durante el siglo diez y siete ofrecio oportunidades a los naturalistas y medicos europeos residentes
para le van tar inventarios de los mamiferos y anotar y hacer informes sobre sus observaciones.
La sistematizacion y el nombramiento cientifico de las especies brasileras conocidas por Carolus
Linnaeus en 1758 fueron basadas primariamente sobre los mamiferos descritos y figurados por
Jorge Marcgraf en su monografia del siglo diez y siete. La coleccion y preservacion efectiva de
mamiferos para el estudio empezo en comienzos del siglo diez y ocho con el "Viajem Filosofica"
de Alejandro Rodriguez Ferreira, brasilero de nacimiento.

From the Division of Mammals, Department of Zo-
ology, Field Museum of Natural History, Chicago, Illi-
nois 60605-2496.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 1 1

El siglo diez y ocho y las primeras decadas del siglo diez y nueve senalaron un periodo explosive
y romantico de expediciones cientificas fomentadas por gobiemos europeos, o por naturalistas
particulares con los objectos de hacer observaciones sobre la fauna, tomar notas de campo, y
recoger y preservar ejemplares para estudios taxonomicos en los museos extranjeros. Entre los
naturalistas europeos que hicieron contribuciones de consequencia a la mastozoologia neo-
tropical en este epoca, se cuentan Alejandro von Humboldt, Juan Baptista Ritter von Spix
(Brasil), Maximiliano Principe de Wied-Neuwied (Brasil), Juan Natterer (Brasil), Lord Roberto
Herman Schomburgk y Ricardo Schomburgk (Guyana), Claudio Gay (Chile), Juan Jacobo von
Tschudi (Peru), Felix de Azara (Paraguay), Rodolpho Rengger (Paraguay), Alcidio Carlos Victor
d'Orbigny (Argentina, Bolivia), y Carlos Roberto Darwin (Patagonia y Galapagos). Compren-
dido en este informe son los itinerarios, listas de mamiferos coleccionados y observados,
taxonomias, y algunas experiencias de campo de los naturalistas mentados, y de otros digno
de atencion.

A mediados del siglo diez y nueve, la fauna mamifera de Sud America llego a ser la mejor
conocida de todos los continentes del mundo menos Europa. Problemas de origen y reparticion
geografica de los mamiferos del Mundo Nuevo estimularon la imaginacion de sabios desde los
primeros cronistas del Descubrimiento hasta el joven Darwin pre-evolutionario. Los conceptos
sobre estos temas son brevemente discutidos.

A historia da mastozoologia neotropical, come90u com a primeira viagem transatlantica de
Cristovao Colombo, em 1492. Os primeiros relatorios, puramente anedotais, foram registrados
pelos cronistas espanhois, logo apos o regresso das viagens de descobrimento durante os seculos
XV e XVI. As colonizacoes da costa Guianense e Brasileira durante o seculo XVII, ofereceram
amplas oportunidades a naturalistas e medicos, de treinamento Europeu, para inventoriar e
descrever os mamiferos encontrados. A sistematica e a nomenclatura cientifica das especies
Brasileiras conhecidas por Carolus Linnaeus em 1758 basearam-se primariamente nos ma-
miferos descritos por Georg Marcgraf, em sua monografia do seculo XVII. No entanto, as
colecoes e preservacoes de mamiferos para estudos come^aram, efetivamente, no seculo XVIII,
com a "Viajem filosofica" do Brasileiro, Alexandre Rodrigues Ferreira.

Marcaram o seculo XVIII, e as primeiros decadas do seculo XIX, um periodo explosivo e
romantico nas expedicoes cientificas. Estas foram patrocinadas tanto por naturalistas indepen-
dentes, como por govemos Europeus, a fim de realizarem observacoes sobre a fauna e colecoes
para estudos taxonomicos nos museus Europeus. Entres os naturalistas Europeus que distin-
guiram-ce em suas contribuicoes aos estudos de mamiferos neotropicais durante esta epoca,
sobressaem: Alexandre von Humboldt, Juan Baptista Ritter von Spix (Brasil), Maximilian
Principe de Wied-Neuwied (Brasil), Johan Natterer (Brasil), Sir Robert Herman Schomburgk
e Richard Schomburgk (Guiana), Claudio Gay (Chile), Johan Jakob von Tschudi (Peru), Felix
de Azara (Paraguai), Rudolph Rengger (Paraguai), Alcides Charles Victor d'Orbigny (Argentina,
Bolivia) e Charles Robert Darwin (Patagonia e Galapagos). Os itinerarios, as listas de mamiferos
observados e colecionados, as taxonomias, e algumas notas de campo encontram-se incluidos
nos relatorios aqui apresentados sobre estes e outros naturalistas importantes.

Nas meadas do seculo XIX, a fauna mamifera sul-americana tomouse a melhor conhecida
de todos OS continentes, exceto a da Europa. Os problemas de origem e da distribuicao geografica
dos mamiferos neotropicais estimularam a imaginacao de varios estudiosos, desde os primeiros
cronistas ate o pre-evolucionario Darwin. Seus conceitos sobre estes temas sao brevemente
discutidos.

Organization II. Voyages of Discovery: 1 5th and

16th Centuries 14

I. Introduction 13 III. Spanish Chroniclers of New

The Neotropical Region Defined . . 14 World Discoveries 14

12 HELDIANA: ZOOLOGY

IV. First Mammals: Anecdotal Period

Island Mammals of the

Discoverers 16

Mainland Mammals of the

Discoverers 18

V. Brazil: Mammalogy Through

1 8th Century 21

Andre Thevet (1503-1592) 21

Georg Marcgraf (1610-1644) 21

Alexandre Rodrigues Ferreira

(1756-1815) 21

VI. Brazil: Mammalogy to Middle of

1 9th Century 27

Introduction 27

Johann Baptist Ritter von Spix
(1781-1826) and Carl Friedrich

von Martius (1794-1866) 27

Maximilian Prinz von Wied-Neu-

wied (1782-1867) 31

Johann Natterer (1787-1843) .... 34
VII. GuiANAs: Mammalogy to End of

1 8th Century 38

Pierre Barrere (1690-1755) 38

Jose Gumilla (d. 1750) 38

Jacques Nicolas Bellin (1703-1772)

Edward Bancroft (1744-1821) .... 38
Philippe Fermin (1720-1790) .... 39

Monsieur Bajon (1763?) 40

John Gabriel Stedman (1744-1797)

VIII. GuiANAs: Mammalogy of First

Half of 19th Century 43

Sir Robert Herman Schomburgk
(1804-1865) and Richard

Schomburgk (181 1-1891) 43

IX. Alexander von Humboldt ( 1769-

1859) AND AlME BONPLAND

(1773-1858) 51

X. Paraguay 57

Felix de Azara (1746-181 1) 59

Johann Rudolph Rengger

(1795-1832) 64

XI. Chile 64

Giovanni Ignazio Molina

(1737-1829) 64

Eduard Friedrich Poeppig

(1798-1868) 65

Claudio Gay (1800-1873) 65

XII. Peru 65

Johann Jacob von Tschudi

(1818-1889) 65

XIII. Patagonia 71

Alcide Charles Victor d'Orbigny

(1802-1857) 71

Charles Robert Darwin (1809-1882)
77

XIV. Georges Louis Leclerc de Buffon

(1707-1788) 87

XV. Faunal Origins and Distribution

Jose de Acosta (1539-1600) 87

Antonio Vazquez de Espinosa

(1560/1575-1630) 90

Carolus Linnaeus (1707-1778) ... 90
Georges Louis Leclerc de Buffon

(1707-1788) 90

Johann Andreas Wagner

(1797-1861) 91

Maximilian Prinz von Wied-

Neuwied (1782-1867) 91

Johann Jacob von Tschudi

(1818-1889) 91

Charles Robert Darwin (1809-1882)

XVI. Inventories to Middle of 1 9th

Century 91

System Naturce of Linnaeus,

1758, 1766 91

Histoire Naturelle of Buffon,

1750-1789 92

Synopsis Mammalium of Schinz,

1844 92

XVII. Summary 92

XVIII. Acknowledgments 94

XIX. Literature Cited 94

I. Introduction

The gradual accumulation of knowledge of
Neotropical mammals is recorded here from the
time of the first voyage of discovery by Christo-
pher Columbus in 1492 to the middle of the 19th
century, or just before the Darwinian revolution
in biological thought. The knowledge was mainly
of species or kinds, the numbers of kinds, their
behavior, habitat, geographic distribution, and re-
lationship to man. Early voyagers to the New World
followed by naturalist-travelers gathered the data
used later by philosophers and scientists for the
development of biological principles. Only the
most important and better-known contributors are
discussed here. At least as many more personages
could be included in a more extended account.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

The Neotropical Region Defined

The Neotropical Region, as defined by its mam-
malian fauna, includes all South America, Middle
America except the dry and temperate zones of
Mexico, continental islands of coastal Middle and
South America, and the oceanic Bahamas, West
Indies, Galapagos, and Falklands (Hershkovitz,
1972, p. 326).

With few exceptions, modem names for Neo-
tropical countries and geographic features are used
throughout the text. The map (fig. 1) shows the
South America of the colonial period with colonial
or precolonial names for political subdivisions and
geographic features.

II. Voyages of Discovery:
15th and 16th Centuries

The inhabited islands found by Columbus on
his first voyage across the Atlantic Ocean in 1492
were thought to be near the mainland of China or
India. The islanders welcomed the ships' crews
with food and drink, but the great stores of pre-
cious metals, stones, and artifacts the travelers ex-
pected to find were not seen. Nevertheless, the
voyagers claimed the islands for the Spanish crown
and returned with accounts told to awaiting re-
porters of their discoveries, including their de-
scriptions of plants and animals of economic value
or imputed medicinal virtues.

Zoological results of the four transatlantic voy-
ages commanded by Christopher Columbus— the
first (1492-1493) and second (1493-1496) to the
Antilles, the third (1498) to the Antilles and Ven-
ezuela, and the last (1 502-1 504) to Middle Amer-
ica—included reports of a variety of mammals.
The kinds seen were identified with such familiar
Old World forms as lion, tiger, bear, fox, dog,
ferret, rabbit, deer, boar, goat, sheep, rodent, mon-
key, and ape. Characterizations given were less
descriptions of external morphology than of gen-
eral mien, gross habitat, behavior in response to
human confrontations or predation on human
property, gastronomic qualities, and use, if any,
in medical treatment, ceremonial rites or magic,
or as household pets.

Those who followed Columbus in the discovery
and exploration of the mainland returned with
additional bits of information on mammals noted
by the attendant Spanish chroniclers. Among the
more important of these voyagers of discovery

were Pinzon, who followed Columbus to the Ven-
ezuelan coast in 1 500, and Amerigo Vespucci, who
sailed first with Ojeda to Brazil in 1499 and in-
dependently again in 1 502 and 1 503 in the service
of Portugal. Pedro Cabral, however, had already
claimed Brazil for Portugal in 1 500 on his way to
India. In 1516 Juan Diaz de Solis discovered the
estuary of the Rio de la Plata, and Sebastian Cabot,
in the service of Spain, sailed in 1526 up the Rio
Parana. Vasco Nunez de Balboa accompanied En-
ciso to Panama in 1510, and in 1513, with Fran-
cisco Pizarro, crossed the isthmus to behold the
vast Pacific Ocean. Pizarro visited Panama again
in 1531, recrossed the isthmus, and sailed south
along the west coast of South America to the dis-
covery and conquest of Peru. Cabeza Alvarez Nu-
nez de Vaca arrived in Buenos Aires in 1541 and
continued overland into Paraguay. Pedro de Val-
divia visited Venezuela in 1 530, Peru in 1 532, and
Chile in 1540, 1541, and 1552. The explorations
of Colombia by Gonzalez Jimenez de Quesada
from 1536 to 1539 and again in 1569 to 1571
signaled the end of the period of discovery and
conquest.

III. Spanish Chroniclers of New World
Discoveries

The recorders or chroniclers of New World dis-
coveries, conquests, happenings, and natural phe-
nomena were the clerics and scribes who accom-
panied the explorers or awaited their return to
Spain for recording the news. Most of the accounts
or records remained unpublished, but some of the
manuscripts are reportedly preserved in the ar-
chives of Spain or the Vatican. The chroniclers
whose published narratives contain interesting in-
formation on mammals include the following.

Peter Martyr of Anghiera (1455-1526), Italian
by birth, and the first and most prestigious chron-
icler of the Discovery, was a member of the Royal
Spanish Council of the Indies, Prothonotary of the
Catholic Church, correspondent of Popes, confi-
dant of Christopher Columbus, and friend of sea
captains, clergymen, and other contemporary voy-
agers to the New World. News he received from
his informants constitutes the first records of New
World discoveries. His chronicles, known as the
Decades and addressed to the Pope, began to ar-
rive at the Vatican in 1 494. The first Decade de
Orbe Novo, with first notices of American mam-
mals, was published in 1516, but pirated Italian

HELDIANA: ZOOLOGY

Fig. 1. Map, South America of the Colonial period from the Stevens (1726) translation of Herrera y Tordesillas.

editions appeared in 1504 and 1507. The fourth
Decade was published in 1521, and the complete
set of eight of the projected 10 appeared posthu-
mously in 1587.

Gonzalo Fernandez de Oviedo y Valdes (1478-
1557) was appointed royal chronicler of news sent

directly to him by provincial governors and other
New World officials. Included were Oviedo's own
observations and results of investigations during
his residence as representative of the Spanish
Crown in the Provinces of Darien, Panama, Gua-
temala, Cuba, and Santo Domingo. He published

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

the first part of his Historia de las Indias in 1 526,
other parts in 1535 and 1 547. The entire work was
printed between 1851 and 1855 in Madrid.

The Spanish Jesuit Jose de Acosta ( 1 539-1 600)
wrote his Historia Natural y Moral de las Indias
during a residence in Peru from 1571 to 1 587 and
saw it published in 1 590. Acosta's philosophical
inquiries extended to all asjjects of nature in the
New World and greatly influenced the thinking of
his contemporaries.

Antonio Vazquez de Espinosa (b. between 1560
and 1575, d. 1630), a Carmelite missionary, lived
many years in Spanish America, most of them in
Peru and Mexico. His natural history notes are
compiled from many sources, including his per-
sonal observations and testimony of people he met
in travels connected with his clerical duties. The
forgotten manuscript of his Compendium was dis-
covered in the Vatican library by Charles Upson
Clark in the early part of the 20th century. Clark's
English translation of the work was published in
1 942 by the Smithsonian Institution, and his tran-
scription of the original Spanish in 1 948 by the
same institution.

Antonio de Herrera y Tordesillas (1559-1625),
historiographer to the King of Spain, compiled
the General History of the Vast Continent and Is-
lands of America from archived reports by the
New World discoverers and conquistadores, gov-
ernors, clergy, colonists, and travelers. He also
borrowed heavily from published accounts, in-
cluding those of other chroniclers. There is no in-
dication that his notices on mammals were based
on personal observations. The first edition
of Herrera's History was published in 1601,
another in 1 60 1-1 6 1 5. These and a 1 728 Spanish
edition in the Library of Congress are cited in the
bibliography. I have not seen these works. The
Stevens translation, published 1725-1726, was
used here. Whatever the quality of the translation,
I find no fault with the descriptions of mammals,
and the stories about them are in line with similar
accounts in other sources.

IV. First Mammals: Anecdotal Period

Island Mammals of the Discoverers

The first Columbian voyage, in 1492, resulted
in the discovery of the Antillean islands of Cuba,
Hispaniola, and part of the Bahaman Archipelago.

According to Peter Martyr, who reported results
of the voyage in his first Decade (1504, 1516),
quadrupeds were not seen, but three kinds of
"rabbits" were said to occur in Hispaniola (Haiti
and Dominican Republic). The same animals, ac-
tually caviomorph rodents, were described later
by Oviedo during his residence in Santo Domingo.
The following accounts are freely translated or
paraphrased from the Spanish of the Paraguayan
(1944-1945) edition of Oviedo's work.

Hutia, the first "rabbit" (1944, libro XII, cap.
I), is smaller than the ordinary rabbit, its ears
smaller and tail ratlike. The hutia is said to be
dark grayish in color and very good eating. It was
hunted and killed by the barkless dogs of the na-
tives, but is no longer found, except rarely.

Gerrit S. Miller (1929, p. 12), studied the re-
mains of mammals in kitchen middens of the Sa-
mana Bay region, Dominican Republic, and con-
cluded that the original description of the hutia
"would apply as well to the species of Plagiodon-
tia, and presumably also to the Isolobodons [sic]
that there seems to be no reason to doubt that
these were the animals Oviedo had in mind."

The quemi, second of the "rabbits" (1944, libro
XII, cap. II), is said to be blackish like the hutia
and similar in form, but larger like an ordinary
hound. Natives of the island who saw and ate the
animal found it savory. Oviedo believed them ex-
tinct.

All attributions to the quemi, according to Mil-
ler ( 1 929, p. 13), agree with those of a "large rodent
whose remains I found in the caves near St. Mi-
chel, Haiti, in 1925. Consequently, I proposed for
it the generic name Quemisia. The presence of the
same creature in the Boca del Infiemo kitchen
midden appears to confirm my guess."

The mohuy "rabbit" (1945, libro XII, cap. Ill),
is smaller than the hutia, a paler brown or grayish
in color, its flesh highly esteemed by the island's
caciques and noblemen. The pelage, unlike that of
the hutia, is stiff", sharply pointed, and erect. Ovie-
do saw no mohuy, but knew persons on the island
who did and reportedly regarded its flesh as better
than that of the other "rabbits."

"There be little if any doubt," says Miller ( 1 929,
p. 13), "that the animal Oviedo thus described
was Brotomys voratus ... its remains have been
found in every kitchen midden that has been ex-
amined in the Dominican Republic. . . . The ac-
count of stiff", pointed, erect-standing hairs of the
back seems especially applicable to a relative of
the South American spiny-rats."

The cori, a fourth "rabbit," described by Oviedo

HELDIANA: ZOOLOGY

(1945, libro XII, cap. IV), is almost certainly the
domestic guinea pig. Miller (1929, p. 14) ques-
tioned whether the guinea pig was pre-Columbian
or a Spanish introduction. He inclined to the sec-
ond alternative "chiefly because remains of the
animal have been found in only one midden." It
appears, however, that one Simone Verde, who
accompanied Columbus on his first voyage, men-
tioned in a letter dated 20 March 1494 (cf. Martyr
in Gaffarel, 1907 trans, p. 12, footnote 2; p. 14,
footnotes 1 , 2) the existence on the island of a
black and white dormouse-like animal without tail.
The guinea pig or cui, domesticated in Peru, was
carried by pre-Columbian Indians for food and
barter and introduced into islands and many parts
of mainland South America where cavies do not
naturally occur. Many of them, such as completely
isolated colonies I saw in Colombia near Bogota,
had become feral, their coloration having reverted
to the wild or agouti pattern.

Other Hispaniolan mammals mentioned by
Oviedo are the barkless domestic dogs and house
rats, the latter certainly brought by the Spaniards.
Apart from the extinct insectivore Nesophontes,
Miller found no remains of mammals the size of
mice or rats in kitchen middens or owl pellets.

Two additional native West Indian mammals
observed by Oviedo in 1 523 or 1 524 in Cuba differ
from those of Hispaniola. My paraphrased trans-
lation of Oviedo's Spanish descriptions follows.

The guabiniquinax is somewhat larger than a
rabbit, its feet similar, the tail long and ratlike, the
pelage smoother than that of a badger, the skin
white, the flesh savory. It lives and breeds in the
mangroves along the coast. To capture it, the In-
dians position their canoes beneath the mangroves
where it nests, then shake the tree to cause the
animal to fall into the water where it is seized.

The animal as described above is certainly a
form of Capromys, but Oviedo continues as fol-
lows: "The animal is the size of a hare, looks like
a fox, its color is dark brown mixed with reddish,
the tail hairy and the head shaped like that of a
ferret. It abounds along the Cuban coast." The
characterization and habitat are obviously out of
place and probably were meant to be included with
the description of the ayre, the second of the Cu-
ban mammals reported by Oviedo. Herewith my
paraphrased translation of his description of that
animal.

The ayre is reddish brown, the size of a rabbit
with pointed muzzle, its flesh exceedingly tough.
Notwithstanding, the natives cook or roast as many
of the animals as they can capture, for they are

abundant. But no matter how long the meat may
be cooked or roasted, it is no less tough to chew.

This characterization seems to fit the insectivore
Solenodon. On the other hand, the flesh of Cap-
romys, as of most if not all caviomorphs, is tender
and, as a rule, delectable.

From his correspondents Oviedo received no-
tice of still another mammal, the guacabitinax, an
inhabitant of the islands near those of Las Perlas
in the Golfo de San Miguel and the Isla de las
Culebras or Gorgona, off" the southwest coast of
Colombia. The name, not to be confused with the
preceding, and the description and details of the
animal's habits, are unmistakably those of the paca
{Agouti paca Linnaeus).

Manatees sighted at sea at various times by Co-
lumbus and his men were believed to be mer-
maids, albeit ugly ones. Martyr's narrative of a
captive manatee as given in the available French
translation of his third Decade (Gaffarel, 1907) is
composite. The account by Herrera of the same
manatee (in Stevens's translation, 1725, vol. 1, p.
278) appears to hew closer to the original source
of information:

The Spaniards at this Time found a new
Sort of Fish, which was a considerable ad-
vantage to them; tho' in those Parts there is
much Variety. It is call'd Manati, in shape
like a Skin they use to carry Wine in, having
only two Feet at the Shoulders, with which
it swims, and it is found both in the Sea and
in Rivers. From the Middle it sharpens off"
to the Tail, the Head of it is like that of an
Ox, but shorter, and more fleshy at the Snout;
the Eyes small, the Colour of it grey, the Skin
very hard, and some scattering Hairs on it.
Some of them are twenty Foot long, and ten
in Thickness. The Feet are round, and have
four Claws on each of them. The Females
bring forth like the Cows, and have two Dugs
to give suck. The Taste of it is beyond Fish;
when fresh it is like Veal, and salted like
Tunny-Fish, but better, and will keep longer;
the Fat of it is sweet, and does not grow
rusty. Leather for Shoes is dress'd with it.
The Stones it has in the Head are good against
the Pleurisy and the Stone. Sometimes they
are taken ashore, grazing near the Sea, or
Rivers, and when young they are taken with
Nets. Thus the Cazique Caramestex took
one, and fed it twenty-six Years in a Pond,
and it grew sensible and tame, and would
come when call'd by the Name of Mato,

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

which signifies Noble. It would eat what-
soever was given it by Hand, and went out
of the Water to feed in the House, would
play with the Boys, let them get upon him,
was pleas'd with Musick, carry'd Men over
the Pool, and took up ten at a Time, without
any Difficulty.

Martyr's third Decade mentions many "rab-
bits" and deer encountered in 1 5 1 6 by Andre Mo-
rales on the forested Isla Rica (now San Jose) of
the Archipielago de las Perlas in the Golfo de Pa-
nama. The deer, most likely Mazama gouazoubira
permira Kellogg, 1 946, and rabbits (Sylvilagus sp.)
were said to be so abundant that Spaniards could
shoot them with arrows from horseback. The re-
tiring tapeti, Sylvilagus brasiliensis, the only rabbit
known from mainland Panama and the Pearl Is-
lands, would have been an unlikely target for the
equestrian Spaniards.

Mainland Mammals of the Discoverers

First knowledge of mainland American mam-
mals was contained in reports of the Paria Pen-
insula, Venezuela, by Columbus on his third trans-
atlantic voyage in 1498 and Vicente Yaiiez Pinzon,
who followed in the tracks of Columbus. Martyr's
first Decade carried the news of their encounters
with the common opossum (Didelphis marsupi-
alis), sloths, armadillos, anteaters, deer (Odocoi-
leus, Mazama), peccaries, tapir, kinkajou (Potos
flavus), barkless dog {Canis familiaris), jaguar,
puma and their color varieties, vampire(?) bats,
and red howler monkey (Alouatta seniculus).

On his fourth and last voyage (1 502-1 504), Co-
lumbus explored the Atlantic coast of Middle
America from the Golfo de Uruba to Guatemala.
Spanish emissaries charged with establishing set-
tlements followed quickly. Mammals reported by
them and noted in Martyr's second and third
Decades, and by Oviedo, include the common
opossum, bats, monkeys, three-toed sloths, ant-
eaters, armadillos {Dasypus novemcinctus), white-
tail deer, red brocket, collared and white-lipped
peccaries, squirrels, a composite of carnivorous
species identified as raposas (including Didel-
phis?), zorros (foxlike Camivora), lobos (Dusicyon
or Lutra), rabbits (Sylvilagus brasiliensis), "hares"
(Dasyprocta sp.. Agouti paca, and perhaps the newly
introduced European hare or rabbit). The domes-

tic dog, like that first seen in the Antilles, was
barkless.

An encounter with vampire bats by Pinzon's
men is reported by Martyr in the first Decade.
Vampires are also mentioned in the second De-
cade in connection with Enciso's disastrous ex-
periences in the Darien and in the third Decade
in accounts of the animals of the Golfo de Uruba.
The following characterization of a vampire bat
by Herrera (in Stevens's translation, 1726, vol. 2,
p. 7 1 ) is a translation from the original sources in
Spanish.

"This venomous Creature has one quality that
tho' it bites one man among an hundred one Night,
the next Time it only bites in the very same Place,
tho' the Person bit be among two hundred; which
it does either on the Toes, the Fingers, or the Head,
and much Blood runs from it."

That the same vampire bat should visit the same
person sleeping in the same place on successive
nights may not be unusual. An experience of mine
in 1935 on the Rio Napo in Ecuador is of interest
in this regard. Two Indian families and I, alto-
gether 10 persons including a five-year-old girl,
traveled three days upstream in a large dugout
canoe. The river was low and we could bivouac
on sandbars at the end of each day's travel. On
each of the three nights, a vampire bat visited the
little girl, scraped the skin of her nose, and fed on
the trickling blood. No other member of the party
was attacked. It seems improbable that the same
bat should have found the same victim at each of
the three different bivouacs. Perhaps the child slept
more soundly than the others of the party, or her
blood was more attractive to the vampires which
abounded in the region.

The last of Martyr's eight Decades includes de-
scriptions of Spanish settlements in the Golfo de
Paria, Venezuela. In addition to those mammals
previously mentioned by Martyr (above) are the
lesser anleater (Tamandua tetradactyla), capuchin
monkey (Cebus apella or C. nigrivittatus), peccary,
deer (Odocoileus virginianus), jaguar {Felis onca),
spotted cats {Felis pardalis and/or F. wiedii), wea-
sels (Mustela frenata), skunk (Conepatus chinga),
porcupine (Coendou prehensilis), and manatee
{Trichechus manatus). Oviedo described the same
animals of the region in greater detail, but with no
additions of sp>ecies. Vazquez de Espinosa, who in
1628 presumably visited the northern Venezuelan
coast and the town of Santo Tomas above the
mouth of the Rio Orinoco, reported the same
mammals as well as squirrels (Sciurus aestuans)

FIELDIANA: ZOOLOGY

and many kinds of monkeys. He claimed that Isla
Margarita, off the Venezuelan coast, was overrun
with rabbits {Sylvilagus floridanus).

Many of the larger mammals of Colombia in
the territories of the Muso and Colima Indians
north of Bogota were already known by 1544.
With bats and other small mammals omitted, more
kinds were reported by Herrera than could be re-
corded today from the same region on the basis
of extant specimens preserved in museums. Her-
rera, in the English version by Stevens (1726, vol.
6, p. 191), states:

There are a great number of grey Swine
[Tayassu pecan] that have the Navel on the
Back, and a smaller sort of several Colours
[Tayassu tajacu] much like wild boars. Ti-
gers (Felis onca) not numerous but very
fierce; Lions (Felis concolor) that do no harm,
except only among the Cattle and two other
sorts of Tigers that were inoffensive besides
another sort that are always in the water,
like Greyhounds, and all their four feet are
like those of a Goose [Lutra annectens]. The
black wild cats [Felis yagouaroundi] seize
the Hens, carry them away under one of
their front legs and run away on the other
three. The black Bears [Tremarctos ornatus]
like those in Spain, do no hurt but only to
the small Cattle. The Ant-Bears [Myrme-
cophaga tridactyla] when they go, lay their
Tail, which is long, on their Heads, winding
them about their Necks, and so walk from
Ant-hill to Ant-hill, stretch out their Tongues
near half a Yard which are soon cover'd with
Pismires, then they draw them back and eat
them. There are Dantas [Tapirus pinchaque
or T. terrestris]. Deer [Odocoileus virgini-
anus] like ours in Europe, and others red
like wild Goats [Mazama rufina or Mazama
americana], and the Bezoar stones found in
them are best. The Guadatinajas [Agouti
paca] are like Hares; and the Zorillas [Di-
delphis marsupialis] or little foxes, that have
a purse under their Belly, in which they carry
their Cubs, the ever so many, are very mis-
chievious to the Henroosts. The little Crea-
tures call'd Umazia [Marmosa] have a dug
growing out for every one of their young,
and they stick to it till bred up. The Ar-
madillo [Dasypus novemcinctus] which has
been spoken of having five claws in each
Forefoot, with which it throws up the Earth,

is tame and eaten. The Perico Ligero [Bra-
dypus variegatus] is three hours climbing a
Tree, goes about in the Night, gives a cry
every time it lifts a Foot, and is half an Hour,
between every Step, is as big as a Barbary
monkey, and fierce, yet does no harm. There
are cats (?) that sleep all the Day, and all the
Night catch Birds and Mice. The Pizma [Na-
sua nasua] about as big as a large Lap Dog,
has a bad countenance, a long Snout, its voice
like a Bird, defends itself against Dogs, and
the Spaniards call them Badgers. The
Hedgehogs [spiny echimyids] are like those
in Spain, the largest like Porcupines [Coen-
dou sp.] darting out their Prickles. There are
many sort of Apes, squirrels.

Elsewhere, in the Province of Bogota, Herrera
(in Stevens's translation, 1726, vol. 6, p. 77) notes
"innumerable apes, monkeys, ferrets [marsupi-
als?], squirrels, weasels [Mustela frenata], deer
[Odocoileus virginianus], roebuck [Mazama rufi-
na] and Rabbits [Sylvilagus brasiliensis] . . . but
not Hares." Manatees were reported from the Rio
Magdalena.

From coastal Colombia, at Zaragoza, 30 leagues
from Caceres in the lower Rio Cauca Valley, Vaz-
quez de Espinosa records jaguar, puma, danta
(Tapirus terrestris), oso (Myrmecophaga or Ta-
mandua), cuchumbi (Nasua), armadillo (Dasy-
pus), raposa (Dusicyon thous), chucha (Didelphis
marsupialis), "three" species of sahinos or pec-
cary, perico ligero (Bradypus variegatus), nutria
(Lutra or Chironectes), and guadatinaja (Agouti
paca).

Acosta's long residence in Peru made him fa-
miliar with some of the mammals in the vicinities
of Cuzco and Lima and others about which he
may have learned from travelers or records. He
described sahinos (peccaries), dantas (tapir), ar-
madillos, perico ligero (three-toed sloth), osos
(anteaters), otoronco (bear), chinchilla, vizcacha,
cui (guinea pig). The "liebres verdaderas" or true
hares are certainly the introduced European hare.
He affirmed that conejos or rabbits (Sylvilagus
brasiliensis) occur in the Reino de Quito (Ecua-
dor).

Acosta declared there were monkeys of all kinds
throughout America, but those he described were
Middle American. At Capira near Nombre de Dios,
Panama, he saw monkeys (presumably spider
monkeys) swing by their tails from a tree on one
side of a stream to another tree on the opposite

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

side. Where the river was too wide for this ma-
neuver, the monkeys of the troop, he related,
crossed by forming a hanging chain holding on to
each other's tail, then swinging so that the endmost
could grab the branch of a tree on the other side
of the river and let all the others clamber up.

The anecdote is less a fabrication than an ex-
aggeration. Individual howlers and spider mon-
keys, usually the alpha male or an old suckling
female, may bridge a narrow gap in the canopy
pathway by holding with its prehensile tail the
branch on one side of the gap and with it swinging
the body to catch, with outstretched arms, the
nearest branch of the far side. Monkeys too small
or weak to hurdle the gap run or scramble over
the bridging back of their elder. I have seen strong
young adults take advantage of the same conve-
nience.

Acosta also narrated the tale of a monkey that
resided in the palace of a provincial governor. As
related to him, the simian was trained to fetch
wine from the town tavern. The animal would set
off with the empty wine pitcher in one hand, the
wine money gripped in the other. Not before the
pitcher was filled to the brim did the sage monkey
release his coins to the tavern keeper. There were
times on these errands when taunting street ur-
chins chased and hurled stones at the monkey.
Annoyed by this sport, the simian halted, set down
the pitcher, and returned the stones with sufficient
force and accuracy to rout his tormentors. Retriev-
ing his pitcher, he moved on serenely to deliver
the wine at the palace.

Peruvian "sheep" or camelids were greatly ad-
mired by the Spaniards when first seen. Acosta's
interesting account of them was suitably appre-
ciated by Herrera, and the English translation by
Stevens (1726, vol. 4, p. 36) is quoted herewith.

There are no such Vicunas and Sheep in
New Spain [Mexico] As those of Peru, and
those Sheep are Tame, and very serviceable;
but the Vicunas are wild, and have no Horns,
the like of them not to be seen in the whole
World, but only in Peru and Chile, bigger
than Goats, but smaller than Calves, their
Colour almost Murrey, breeding on the
highest Mountains, in cold and desert Places,
which they call Punas. They go in flocks,
run swiftly, and when they see any Men, fly
and drive their Young before them. Of their
Wooll are made very valuable Mantles,
which never lose their Colour, because it is
natural; they are said to be good for Inflam-

mations in the Kidneys, as are Quilts made
of the Wooll, because they moderate the
Heat, and the same in the Gout; and in them
the Bezoar Stones are found.

The abundance and ubiquity of llamas may have
inspired some Spaniards to attempt to raise Old
World camels in Peru. According to Acosta, some
brought from the Canary Islands were bred for a
while.

Sebastian Cabot's journal of conquest and ex-
ploration of the Province of Rio de La Plata, then
consisting of modem northern Argentina, cisan-
dean Bolivia, and southeastern Brazil, included
data on natural history. As recorded by Herrera,
the mammals seen were the hairy armadillo
{Chaetophractus sp.) and several other kinds, ca-
vies {Cavia), swamp deer (Blastocerus dichotomus),
pampa deer {Blastoceros bezoarticus), brockets
(Mazama sp.), tapirs {Tapirus terrestris), peccaries
(any or all of the known species), howler monkeys
(Alouatta), canids (Dusicyon), lesser anteater (7a-
mandua tetradactyla), jaguar (Felis onca), and
puma (Felis concolor). Southern Brazilian mam-
mals in particular included deer, peccary, tapir,
"rabbits" with small, round ears {Dolichotisl), paca
{Agouti paca), armadillo, sloth (Bradypus torqua-
tus), opossum {Didelphis albiventris), monkeys, and
coastal seals, most likely Arctocephalus australis.

Vazquez de Espinosa adds capybaras, armadil-
los (tatu and quirquincho specified), and guanacos.
In the vicinity of Chuquisaca (La Plata), Bolivia,
the missionary notes brockets {Mazama), vicufia,
guanaco, dark gray wildcats known as oscollos,
jaguar called "otorongo," puma locally called
poma, a large beast called lilisto with a horselike
head that lures cattle and humans, a ferret called
siqui {Mustela frenatal), skunks or anatiria {Co-
nepatus), bear {Tremarctos ornatus), antbears
(probably Tamandua), vizcacha {Lagidium), and
cuis {Cavia porcellus).

The occurrence of sea lions {Otaria flavescens)
and fur seals {Arctocephalus) on both southern
continental coasts was mentioned by Vazquez de
Espinosa. The sea lions along the coast of Are-
quipa, Peru, he reported come out of the water
onto the rocks and make low sounds at night. The
animals were hunted by the Indians for their hides.
In northern Chile, the natives of Arua and Ata-
cama converted the hides into balloon-like floats
for support of their seagoing fishing rafts.

The conquest of Chile by Pedro de Valdi via in
1 54 1 provided the chroniclers with additional in-
formation on mammals. Vazquez de Espinosa re-

HELDIANA: ZOOLOGY

ported huemul {Hippocamelus bisulciis), "fallow
deer" (spotted fawns of huemul), guanaco, and
vicuna in the vicinity of Osomo. According to the
same authority, the Rio Guasco valley (29°S) har-
bored "squirrels" (chinchillas) with very fine fur.

V. Brazil: Mammalogy Through
18th Century

Andre Thevet (1503-1592)

The French missionary Andre Thevet arrived
in 1555 in Rio de Janeiro, the principal port of a
French colony in the ephemeral France Antarc-
tique. Thevet returned to France via the Antilles
a year later, and the accounts of his travels were
published in 1557 or 1558. Father Thevet's cu-
riosity about all he saw in the New World knew
no bounds, and he became an avid collector of
Indian artifacts, local birds, and insects. Not all
objects and events described in his book con-
formed to popular European prejudices or gen-
erally accepted misconceptions. The work stirred
up considerable debate and was rejected by many
not prepared to accept the realities that opossums
had pouches or that the bodies of American In-
dians were not densely furred.

The Brazilian mammals described or men-
tioned by Thevet include the locally common
opossum (Didelphis albiventris), tapeti (Sylvilagus
brasiliensis), agouti {Dasyprocta leporina, declared
good eating), peccaries, deer (probably Mazama),
coati {Nasua nasua), tapir (Tapirus terrestris), ca-
puchin monkey {Cebus apella), golden tamarin
{Leontopithecus rosalia), armadillos, jaguar (Felis
onca), and deer-hunting canids (Speothos"?), but
no lions or wolves. The three-toed sloth was abun-
dant, but never observed eating or drinking. The-
vet adds, however, that there are those who believe
the beast sustains itself solely by the small, slender
leaves of a very high tree called amahut.

Georg Marcgraf (or Marggrav or Marggraf]
(1610-1644)

Most illustrious of the pre-Linnaean naturalist-
explorers of Brazil was Georg Marcgraf Bom in
Liebstad, Saxony, educated in Holland with em-
phasis on astronomy and botany, he sailed for
Brazil in 1638 on a scientific expedition led by
Johann Moritz, Count of Nassau-Siezen. The par-

ty, which included the young physician Piso (1611-
1678), landed in Pemambuco. Explorations were
restricted to northeastern Brazil in the present states
of Pemambuco, Paraiba, and Rio Grande do
Norte. Among MarcgraPs accomplishments were
the construction of an astronomical observatory,
the first of its class in the New World, and a mono-
graphic study of the plants and animals of the
region. After turning over his notes and illustra-
tions to Moritz, for preparation and publication,
the naturalist sailed for Africa, where he died
shortly after arrival. MarcgraPs monumental His-
toriae Rerum Naturalia Brasiliae, a part of Willen
Piso's Historia Naturalis Brasilia, was published
in 1648 in Amsterdam.

Of the mammals of the northeastern region of
Brazil described by Marcgraf, 32 were native
species, the others introduced. Their detailed de-
scriptions and life history notes, together with crude
but useful woodcuts (fig. 2), were among the pri-
mary references on which Linnaeus based bino-
mials in the 10th (1758) and 12th (1766) editions
of his Systema Naturce.

The mammals are listed in Table 1 by the in-
digenous names used by Marcgraf and their cur-
rent scientific names. Provenance of the forms
which served as types for binomialists, mainly
Linnaeus, was restricted for taxonomic purposes
to Pemambuco by Thomas (1911).

Alexandre Rodrigues Ferreira (1756-1815)

The first Brazilian naturalist of European ex-
traction, Alexandre Rodrigues Ferreira, was bom
in Salvador, Bahia. He pursued higher studies in
Portugal, received his doctorate in 1779 from the
University of Coimbra, and was then appointed
Naturalist of the Museu Real d'Ajuda in Lisbon.
He retumed to Brazil in 1783 commissioned by
the museum to collect samples of plants, animals,
and minerals and to record all matters of scientific
and political interest within his scope. The expe-
dition, or "Viagem Filosofica," explored the prov-
inces of Grao Para, Rio Negro, Mato Grosso, and
Cuiaba from 1 783 to 1 792 (fig. 3). Rodrigues Fer-
reira retumed to Lisbon the following year.

The scientific materials collected in Brazil, with
notes and illustrations, were deposited in the Mu-
seu d'Ajuda. Included were 4 1 7 species of animals
represented by 592 specimens. Of these, 76 spec-
imens represented 65 species of mammals. The
whole collection was confiscated by the invading
armies of Napoleon and taken to Paris for study

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

•c

2-*

03 so

<N ea

O u
Cl, a

HELDIANA: ZOOLCXJY

Table 1. The Brazilian mammals of Marcgraf (1648) and their current Linnaean names.

Page

Vernacular name

Linnaean name

Figure

221
222
223
223
223
224
224
224
225
225
226

226

227
227
228
228
229
229
229
230

230
231
231
232
233
233

234
235
235
235
235
235

Ai sive Ignavus

Carigueya, female

Tai-ibi (male)

Aperea, type of the species

Tapeti, type of the species

Cavia Cobaya

Paca, type of the sjsecies

Aguti vel Acuti

Tamandua guacu, type of the species

Tamandua-i, type of the species

Guariba (fig. misplaced on p. 228), type of the

species
Lower figure only of caitaia misplaced with

text of the guariba
Cagui minor, type of the species
Caitaia

Coati, type of the species
Coatimondi

Tapiierete, type of the species
Mus araneus, type of the species
Tajacu Caaigoara
Capybara, Rio Sao Francisco, type of the

species
Scyurus
Tatu
Tatu-ete
Tatu Apara

Maraguo sive Maracaia
Cuandu, type of the species

Ibiya, type of the species

Cuguacu-ete (female), type of the species

Cuguacu-apara (male)

Jaguara, type of the species

Jaguarete

Cuguacuarana

Bradypus variegaltds Schinz, IS25 2

Didelphis albiventris Lund, 1 840 2
Didelphis albiventris Lund, 1 840

Cavia aperea Erxleben, 1777 2

Sylvilagus brasiliensis Linnaeus, 1758* 2

Cavia porcellus Linnaeus, nSS 2

Agouti paca Linnaeus, 1 766 2

Dasyproct a leporina Linnaeus, 175% 2

Myrmecophaga tridactyla Linnaeus, 1758 2

Tamandua tetradactyla Linnaeus, 1758 2

Alouatta belzebul Linnaeus, 1 766 2

Cebus apella libidinosus Spix, 1 823 (fig. only) 2

Callithrix jacchus Linnaeus, 1758 2

Cebus apella libidinosus Spix, 1823 2

Nasua nasua Linnaeus, 1766 2

Nasua nasua Linnaeus, 1 766

7a/7/ru5 r^rrw/m Linnaeus, 1758 2

Monodelphis americana Miiller, 1776

ra>'a$5M /a/acM Linnaeus, 1758 2

Hydrochaeris hydrochaeris Linnaeus, 1 766 2

Sciurus aestuans Linnaeus, 1 766

Dasypus septemcinctus Linnaeus, 1758

Dasypus novemcinctus Linnaeus, \1 5% 2

Tolypeutes trici net us Linnaeus, 1758 2

Felis tigrina Schreber, 1775

Coendou prehensilis prehensilis. Linnaeus,

1758 2

Pteronura brasiliensis Gmelin, 1 788 2
Blastoceros bezoarticus Linnaeus, 1 758
Blastoceros bezoarticus Linnaeus, 1758

F(e//5 onca Linnaeus, 1758 2

Felis onca (melanistic) 2
Felis concolor Linnaeus, 1771

* Editors' Note: Here and elsewhere in this paper. Article 51(c) of the International Code of 2kx)logical Nomen-
clature, governing the use of parentheses in scientific names, is not followed.

by Etienne Geoffroy St.-Hilaire of the Museum
National de Histoire Naturelle in Paris.

Monkeys constituted a sizeable part of the loot,
and the following were described as new by Etienne
Geoffroy St.-Hilaire in 1812 and by others as not-
ed in brackets; the current form of each name is
used: Callithrix jacchus penicillatus, Callithrix
jacchus geoffroyi [Humboldt], Callithrix jacchus
aurita, Callithrix humeralifer, Callithrix argentata
melanura, Saguinus labiatus, Saimiri ustus [I.
Geoffroy], Callicebus amictus, Callicebus person-
al us, Pithecia monachus, Alouatta fusca, Cebus
apella cirrifer. Cebus flavus, and Lagothrix la-
gothricha canus. Mounted specimens of previ-
ously named forms also brought to Paris from the
Lisbon museum included Callithrix jacchus Lin-
naeus, Leontopithecus rosalia Linnaeus, Chiro-
potes satanas Hoffmannsegg, Brachyteles arach-
noides E. Geoffroy, Inia geoffrensis Blainville, and
probably others lost or discarded.

Except for the descriptions by the French zo-
ologist, the specimens and manuscripts of Rod-
rigues Ferreira were largely neglected during the
naturalist's lifetime. The several portions of the
memoirs published posthumously were heavily
edited. In 1972, however, the entire Viagem Fi-
losofica, in two text volumes and two of colored
plates, was published by the Conselho Federal de
Cultura of the Brazilian Ministry of Education and
Culture.

Treatment of mammals in the zoological mem-
oir was a model of its kind for the times. Each
species was described, with bibliographic refer-
ences for the ones better known, external char-
acters and what was learned of habitat, habits,
reproduction, utilization by man, and gastronomic
rating. With respect to the last, Rodriguez Ferreira
grouped the Brazilian mammals according to those
used most widely for food (peccary, deer, tapir,
paca, agouti), those eaten only by Indians and some

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

' W'

KOIFIRO |)A AIAC.KM l'll<IS<^nc:A RF.AI.I/AI>A POR AlfXANDRK RODRU.I Is
l-F.RRHRA. \l MA hISI ANCIA APROXIMADA l>E Vf J7'.' kM (17M \-<K>

Fio. 3. Map of Brazil showing routes (bold lines) of Alexandre Rodrigues Ferreira, during the " Viagem Filosofica,
1783-1792; from Rodrigues Ferreira (1972).

FIELDIANA: ZOOLOGY

Table 2. Mammals illustrated in the Viagem Filosoftca by Rodrigues Ferreira (1971),

Plate no.

Brazilian name

Current scientific name F

gure

118

Gamba

Didelphis marsupialis Linnaeus

119

Macaco-da noite

Aotus sp.

120

Zogue-zogue; uapuca

Callicebus moloch Hoffmannsegg

121

Parauacu

Pithecia monachus E. Geoffroy

122

Cuxiu

Chiropotes satanas chiropotes Humboldt

123

Cuxiu-preto

Chiropotes satanas satanas Hoffmannsegg

124

Guariba-vermelho

Ahuatta seniculns Linnaeus

125

Guariba-da-mao-ruiva

Alouatta belzebul Linnaeus

126

Mico-de-cheiro

Saimiri ustus \. Geoffroy

127

Quata-de-cara-vermelha

Ateles paniscus Linnaeus

128

Barrigudo-cinzento

Lagothrix lagothricha Humboldt

129

Sauitinga

Callithrix argentata argentata Linnaeus

130

Saui dourado

Callithrix humeralifer chrysoleuca Wagner

131

Saui

Callithrix jacchus penicillata E. Geoffroy

132

Saui-de-mao-ruiva

Saguinus midas midas Linnaeus

133

Tamarin

Saguinus midas tamarin Link

134

Saui-de-bigode-branco

Saguinus labiatus labiatus E. Geoffroy

135

Tamandua-mirim

Tamandua tetradactyla Linnaeus

136

Tamanduai

Cyclopes didactylus Linnaeus

137

Tamanduai

Cyclopes didactylus Linnaeus

138

Preguifa-de-tres-dedos

Bradypus variegatus Schinz

139

Tatu-galinha

Dasypus novemcinctus Linnaeus

140

Tatu peba

Euphractus sexcinctus Linnaeus

141

Guaraxaim

Procyon cancrivorus F. Cuvier

142

Janauira

Speothos venaticus Lund

143

Guara

Chrysocyon brachyurus Illiger

144

Quati

Nasua nasua Linnaeus

145

Jupara

Potosflavus Schreber

146

Furao

Galictis vittata Schreber

147

Irara

Eira barbara Linnaeus

148

Ariranha

Pteronura brasiliensis Gmelin

149

Maracaja

Felis geoffroyi d'Orbigny and Gervais

150

Jaguartirica

Felis pardalis Linnaeus

151

Su9uarana

Felis concolor Linnaeus

152

Jaguar

Felis onca Linnaeus

153

On9a preta

Felis onca Linnaeus

154

Peixe-boi, male & female

Trichechus inunguis Natterer

155

Caitetu

Tayassu tajacu Linnaeus

156

Veado vermelho

Mazama americana Erxleben

157

Cariacu

Odocoileus virginianus cariacou Boddaert

158

Quatipuru- vermelho

Sciurus igniventris Wagner

159

Quatipuru-preto

Sciurus spadiceus Olfers

160

Quatipuru-louro

Sciurus igniventris Wagner

161

Rato-d'agua

Nectomys squamipes Brants

162

Prea

Cavia aperea Erxleben

163

Cutia-vermelha

Dasyprocta leporina Linnaeus

164

Cutia-preta

Dasyprocta fuliginosa Wagler

165

Acutiuaia

Myoprocta exilis Wagler

166

Paca

Agouti paca Linnaeus

167

Cuandu

Coendou prehensilis Linnaeus

168

Uiara

Inia geoffrensis Blainville

169

Tucuxi

Sotalia fluviatilis Gervais and Deville

white residents (anteaters, armadillos, sloths, por-
cupines, monkeys, jaguar), and animals not eaten
by humans (marsupials, melanistic felids, squir-
rels, capybara). Bezoar stones and certain parts of
the animal, usually tegumentary, were also cited
for their medicinal merits, particularly as anti-

venins for headaches and female sterility, or as
aphrodisiacs.

A memoir on the peixe boi or river manatee
(Tricheciis inunguis Natterer) provides detailed in-
formation on such topics as hunting, harpooning,
reproduction, size, weight, blubber, butchery,

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Fio. 4. Four monkeys of the "Viagem Filosofica" collections: upper left, parauaco (Pithecia monachus E. Geoffroy),
possibly the holotype; upper right, saui-de-bigode-branco {Saguinus labiatus labiatus E. GeofTroy), possibly the
holotype; lower left, mico-de-cheiro (Saimih ustus I. Geoffroy), possibly the holotype; lower right, saui (Callithrix
jacchus penicillata E. Geoffroy), possibly the holotype; from Rodrigues Ferreira (1972).

HELDIANA: ZOOLOGY

preservation, and market value of the flesh. The
author decried the slaughter of the young and not-
ed the disappearance of manatees in certain lakes.
Of all Brazilian mammals described or merely
listed in the Viagem Filosofica, those depicted in
color in the 50 plates (each 1 9 x 29 cm) are rep-
resentative. They are listed in Table 2 by plate
number with their Brazilian and current scientific
names. The animals were postured as prepared by
taxidermists (fig. 4). Many of the monkeys are
those later described by E. Geoffroy.

VI. Brazil: Mammalogy to Middle of
19th Century

Introduction

Growth of science in South America during the
first third of the 19th century shifted from the
Spanish colonies, with their wars for independence
and internal political turmoil, to the relatively sta-
ble Portuguese colony of Brazil. Following the in-
vasion of Portugal by the Napoleonic armies, the
royal family fled to Brazil and made Rio de Janeiro
its capital and center of cultural activities. During
previous years Brazil had been closed to foreigners
to prevent the mines of precious metals and min-
erals from passing out of control of the ruling Por-
tuguese. Dom Joao VI, however, opened the ports
and changed the environment to one befitting an
enlightened monarch in residence. Cultural insti-
tutions, including museums, libraries, and uni-
versities, were built, and scientific investigations
were promoted. Betrothal of the Archduchess Leo-
poldina, daughter of the Emperor of Austria, with
Dom Pedro, Crown Prince of Portugal and Brazil,
became the most important single factor in the
advancement of science in the New World during
the first half of the 1 9th century. The entourage
of the bride on her voyage to Brazil included some
of the best and most adventurous of the younger
scientists of Austria and Bavaria.

The Viennese naturalists of the party included
the field collector Johann Natterer, and from the
court of Munich, the zoologist Spix and the bot-
anist Martins. Two years earlier, in 1815, the most
accomplished of the naturalist-travelers, Maxi-
milian Prinz Wied zu Neuwied of Prussia, arrived
on the scene.

Modem Brazilian mammalogy begins with the
scientific accounts of the collections and travels of
these naturalists.

Johann Baptist Ritter von Spix (1781-1826)
and Carl Friedrich von Martins (1794-1866)

The German naturalist Johann Baptist Ritter
von Spix first studied for the priesthood, but after
two years his attention turned to medicine and
natural history. His doctorate was earned in 1 806.
That same year he was appointed assistant in the
Museum of the Munich Academy of Science, with
responsibility for the organization of the zoolog-
ical collections. In 1816 he was ordered by the
King of Bavaria to undertake a two-year scientific
expedition to Brazil, together with the museum's
assistant in botany, Carl Friedrich von Martins.
The two departed on 10 April 1817 through the
port of Trieste, and after considerable delay, they
arrived in Rio de Janeiro on 15 July 1817.

The exuberance and variety of the native plant
life in eastern Brazil at first awed and bewildered
the two young naturalists. Everything they saw was
new to them, and all they could possibly collect
and preserve was easily reached along the trails
they traveled from Rio de Janeiro to Minas Gerais
and beyond. Real or fantasized dangers lurking in
what they imagined as dark, brooding, impene-
trable forests restrained their urges for stepping ofl"
the beaten path. The strange and wonderful wild-
life encountered on the roads was enough to gratify
their utmost expectations and inspired them to
record their impressions in ecstatic prose. On the
trip from Ipanema, Sao Paulo, to Vila Rica, Minas
Gerais, they described, as translated into English
by Lloyd in equally romanticized and tortured
prose, the

numerous flocks of little monkeys [that] run
whistling and hissing to the recesses of the
forest; the cavies, running about on the tops
of the mountains, hastily secrete themselves
under loose stones; the American ostriches
(Emas), which herd in families, gallop at the
slightest noise, like horses through the bush-
es, and over hills and valleys, accompanied
by their young; the dicholopus {Seriemas),
which pursues serpents, flies, sometimes
sinking into the grass, sometimes rising into
the trees, or rapidly climbing the summits
of the hills, where it sends forth its loud
deceitful cry, resembling that of the bustard;
the terrified armadillo {Tatu Canastra, Peba,
Bola) runs fearfully about to look for a hid-
ing place, or, when the danger presses, sinks
into its armour; the ant-eater {Tamandud,
Bandeira mirim) runs heavily through the

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

^^ ^'Salvador
(Bahia)
Januaria
liPbrto de Salgado)
aOiamantina

\>^

^-■^buro Preto
(Villa ricQ)

Rio de Janeiro
Sao Poulo

Kane von Brasilien mit dem eingezeichneten Reiseweg von Johann Baptist von Spix und
Carl Friedrich Philipp von Martius anlaBlich ihrer Expedition in den Jahren 1817-1820.

Fig. 5. Map of Brazil showing routes of the Spix and Martius expedition (1817-1820); only principal stations
plotted; from Tiefenbacher(1983).

plain, and, in case of need, lying on its back,
threatens its pursuers with its sharp claws.
Far from all noise, the slender deer, the black
tapir or a pecari, feed on the skirts of the
forest. Elevated above all this, the red-head-
ed vulture (urubii) soars in the higher re-
gions; the dangerous rattle-snake {Casca-
vel), hidden in the grasses, excites terror by
its rattle; the gigantic snake sports suspended
from the tree with its head upon the ground;
and the crocodile resembling the trunk of a
tree, basks in the sun on the banks of the
pools. After all this has passed during the
day before the eyes of the traveler, the ap-
proach of night, with the chirping of the
grasshoppers, the monotonous cry of the
goat-sucker {Jodo corta pdo), the barking of

the prowling wolf, and of the shy fox, or the
roaring of the ounces, complete the singular
picture of the animal kingdom in these
peaceful plains.

For the next three years, the zoologist and bot-
anist explored the eastern states of Brazil from Sao
Paulo and Minas Gerais north to Para. Most of
July and August of 1 8 1 9 was spent in Belem (Para).
On 2 1 August they shipped up the Rio Amazonas,
making stopovers at the mouth of the Rio Tocan-
tins, the Rio Xingu ( 1 September), Santarem on
the Rio Tapajos ( 1 8 September), Obidos (23 Sep-
tember), Parintins, and Vila Nova da Rainha (1
October). The mouth of the Rio Madeira was
passed 1 5 October, and on 22 October they landed
at Barra do Rio Negro (Manaus). Travel upstream

HELDIANA: ZOOLOGY

continued in November with a stop at Tefe (for-
merly Ega) on 26 November. Spix then traveled
alone up the Solimoes to Tabatinga at the Peru-
vian border, arriving 9 January 1820. Martius, for
his part, ascended the Rio Japura to Araracuara
in eastern Colombia.

Spix returned to Manaus on 3 February 1820.
On 1 1 February he ascended the Rio Negro to
Barcelos and was back again in Manaus 28 Feb-
ruary to continue his travels downstream to Be-
lem, where he arrived on 16 April. He embarked
on 14 June 1820 for Europe from Rio de Janeiro
(fig. 5).

In the Reise. Spix and Martius (1828, p. 541)
made up an impressive list of the mammals of the
sertao (scrub country) of Campos Gerais de Sao
Felipe in the angle between the Rio Sao Francisco
and its eastern tributary, the Rio Verde Grande,
northern Minas Gerais. The data were evidently
compiled uncritically from a number of sources,
including local informers, personal observations,
and publications based on the Wied-Neuwied
(1826) collections. Their use and misuse of names
are too involved to unravel here. Except for the
missing bats (given elsewhere by Spix, 1823) and
some small rodents, it is unlikely that a similar or
larger number of mammalian species of the area,
based on actual specimens, could be made today.
The sertao mammals of the Spix and Martius ex-
pedition are listed in Table 3 by current scientific
names of the species only, with the Spix and Mar-
tius equivalents omitted.

In his journey up the Amazon, Spix noted habits
of the inia {Inia geoffrensis) (Spix &, Martius, 1831,
p. 1 1 1 9) and of the manatees (Trichechus inunguis)
(Spix & Martius, 1831, p. 1122).

The results of the expedition are recorded in
several publications, including the Simiarum et
Vespertilionum Brasiliensium by Spix (1823). The
account of the nearly three-year journey or Reise
in Brazil by Spix and Martius (1823-1831) is re-
plete with observations on the biology, geography,
geology, paleontology, mineralogy, meteorology,
and the various human cultures and industries of
the parts of the country they traveled. Many kinds
of mammals are mentioned, but except for bats
and monkeys, few of them were collected.

The zoological material actually collected con-
sisted of thousands of invertebrates and 498 species
of vertebrates, of which 34 were monkeys and 15
bats. Altogether, according to Avila Pires (1974,
p. 139), 85 species of mammals were collected.
Spix (1823) reported only on the monkeys and bats
and illustrated in color the types of all species.

Table 3. Mammals of the sertao of Campos Gerais
de Sao Felipe, Minas Gerais, recorded by Spix and Mar-
tius (1828, p. 541, footnote 3). Current scientific names
to species only are used. The Spix and Martius usage of
local, German, and scientific names is too confused for
tabulation. The arrangement is phylogenetic.

Marsupialia
Caluromys philander Linnaeus
Didelphis marsupialis Linnaeus

Primates
Callithrix jacchus Linnaeus
Cebus apella Linnaeus
Alouatta fusca E. GeofTroy
Alouatta caraya Humboldt

Edentata
Tamandua tetradactyla Linnaeus
Myrmecophaga tridactyla Linnaeus
Bradypus torquatus Desmarest
Bradypus variegatus Schinz
Dasypus novemcinctus Linnaeus
Tolypeutes tricinctus Linnaeus
Priodontes maximus Kerr
Euphractus sexcinctus Linnaeus

Carnivora
Dusicyon thous Linnaeus
Chrysocyon brachyurus Illiger
Nasua nasua Linnaeus
Procyon cancrivorus G. Cuvier
Conepatus chinga Molina
Eira barbara Linnaeus
Pteronura brasiliensis Gmelin
Felis wiedii Schinz
Felis tigrina Schreber
Felis pardalis Linnaeus
Felis concolor Linnaeus
Felis onca Linnaeus
Felis yagouaroundi E. Geoffroy

Perissodactvla

Tapirus terrestris Linnaeus
Artiodactyla

Mazama gouazoubira Fischer

Mazama americana Erxleben

Blastoceros bezoarticus Linnaeus

Lagomorpha
Sylvilagus brasiliensis Linnaeus

RODENTIA

Sciurus aestuans Linnaeus
Wiedomys pyrrhorhinos Wied-Neuwied
Echimys and/or Proechimys species?
Myocastor coypus Molina
Kerodon rupestris Wied-Neuwied
Cavia aperea Linnaeus
Dasyprocta leporina Linnaeus
Agouti paca Linnaeus
Coendou insidiosus Kuhl
Chaetomys subspinosus Olfers

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

c 5?
11

c 5

2 i"
•S

HELDIANA: ZOOLOGY

Table 4. Monkeys (Primates) of the Spix and Martius Expedition described by Spix (1823); the arrangement is
phyiogenetic.

Current name

Spix and Martius synonym

Figure

Cebuella pygmaea Spix, 1 823

Callithrix jacchus jacchus Linnaeus, 1758

Callithrix jacchus penicillatus E. Geoffroy, 1812

Saguinus bicolor bicolor Spix, 1823

Saguinus fuscicollis fuscicollis Spix, 1823

Saguinus mystax mystax Spix, 1823

Saguinus nigricollis nigricollis Spix, 1 823

Saguinus oedipus geoffroyi Pucheran, 1845

Callicebus cupreus Spix, 1823

Callicebus personatus personatus E. Geoffroy, 1812

Callicebus personatus nigrifrons Spix, 1 823

Callicebus personatus melanochir Kuhl, 1820

Callicebus torquatus torquatus Hoffmannsegg, 1 807

Callicebus cinerascens Spix, 1823

Aotus vociferans Spix, 1 823

Actus azarae infulatus Kuhl, 1820

Pithecia monachus monachus E. Geoffroy, 1812

Pithecia pithecia pithecia Linnaeus, 1 766

Chiropotes satanas chiropotes Humboldt, 1812

Cacajao melanocephalus ouakary Spix, 1823

Alouatta caraya Humboldt, 1812

Alouatta belzebul discolor Spix, 1823

Alouatta fusca Spix, 1823

Alouatta seniculus stramineus Humboldt, 1812

Cebus albifrons unicolor Spix, 1 823

Cebus apella libidinosus Spix, 1823

Cebus apella macrocephalus Spix, 1823

Cebus apella xanthosternos Wied-Neuwied, 1 820

Lagothrix lagothricha lagothricha Humboldt, 1812
Lagothrix lagothricha carta E. Geoffroy, 1812
Brachyteles arachnoides E. Geoffroy, 1 806

Jacchus albicollis Spix, 1 823

Midas oedipus (varietas), Spix, 1823

Callithrix gigot Spix, 1823
Callithrix amicta E. Geoffroy, 1812

Nyctipithecus felinus Spix, 1823
Pithecia hirsuta Spix, 1823;
Pithecia inusta Spix, 1823
Pithecia capillamentosa Spix, 1823
Brachyurus israelita Spix, 1 823

Mycetes barbatus Spix, 1823

Cebus gracilis Spix, 1823

Cebus cucullatus Spix, 1823;

Cebus xanthocephalus Spix, 1823
Gastrimargus infumatus Spix, 1823
Gastrimargus olivaceus Spix, 1 823
Brachyteles macrotarsus Spix, 1 823

most life-size. Separate reports on all groups of
animals collected by Spix have been brought to-
gether in a Festschrift in his honor edited by Tie-
fenbacher (1983). The mammals are treated by
Kraft (1983).

The 31 presently recognized species and sub-
species of monkeys ( 1 5 new) and the 1 4 recognized
species of bats (six new) are listed in Tables 4 and
5 by current names with synonyms in parentheses.

Maximilian Prinz von Wied-Neuwied (1782-1867)

Maximilian Prinz von Wied-Neuwied was bom
in Prussia and studied biological sciences at the
University of Gottingen under the famous natu-
ralist-anthropologist Blumenbach. His ambition
to travel and study nature in South America was
realized when he sailed for Rio de Janeiro from
England the first week of May 1815, and arrived
on 17 July.

After a few excursions in the surroundings of

Rio de Janeiro, Wied-Neuwied left for Cabo Frio
on 15 August 1815, stopping at many fazendas
and villages along the way. He left Cabo Frio on
8 September for Sao Salvador dos Campos dos
Goitacazes (now simply Campos) on the Rio Pa-
raiba, and arrived on 25 September. After more
excursions and more collections in the state of Rio
de Janeiro, he crossed the Rio Itabapoana on 26
November into the state of Espirito Santo. A con-
siderable amount of time was devoted there to
explorations of the Rio Doce region. February 1816
saw Wied-Neuwied in Bahia, where he occupied
himself until May 1817. The coastal town of Bel-
monte, where he arrived in August 1816, was the
base for explorations of Botocudo Indian territory.
In December 1816 Wied-Neuwied established II-
heus as center for travel westward to Sao Pedro
de Alcantara, now Itabuna, and the border of Mi-
nas Gerais. On 1 May Wied-Neuwied embarked
at Salvador for Lisbon, then transshipped to Ger-
many through an English port.
Wied-Neuwied's itinerary is difficult to track be-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 5. Bats (Chiroptera) of the Spix and Martius Expedition described by Spix (1823); the arrangement is
phylogenetic.

Current name

Spix and Martins synonym

Rhynchonycteris naso Wied-Neuwied, 1820

Noctilio albiventris albiventris Desmarest, 1818

Noctilio leporinus leporinus Linnaeus, 1 758

Tonatia bidens Spix, 1823

Trachops cirrhosus Spix. 1823

Glossophaga sohcina Pallas, 1 766

Carollia perspiciUata Linnaeus, 1758

Artibeus planirostris Spix, 1 823

DiphyUa Spix, 1823

Diphylla ecaudata Spix, 1 823

Thyroptera Spix. 1 823

Thyroptera tricolor Spix, 1823

Eptesicm brasiliensis Desmarest, 1823

Promops nasutus Spix

Molossus ater E. Geoffroy, 1 805

Proboscidea rivalis Spix, 1 823;

Proboscidea saxatilis Spix, 1 823
Noctilio albiventer Spix, 1823
Noctilio rufiis Spix, 1823

Glossophaga amplexicaudata Spix, 1823
Vampynts soricinus Spix, 1823

Molossus fumarius Spix, 1823
Molossus ursinids Spix, 1823

cause of his many roundabout journeys and short
excursions with too few dates for fixing comings
and goings. To add to the difficulty, the names of
many localities he visited no longer exist or were
never plotted on any official map; a few names
have changed. Bokermann's (1957) gazetteer of
nearly all localities of the Reise, with page refer-
ences to their mention in Wied-Neuwied's works,
is indispensable for study of the naturalist's op-
erations in Brazil.

Wied-Neuwied was interested in all aspects of
nature, but the fauna and Indians engaged most
of his attention. His species accounts are models
of precision, his descriptions detailed, and com-
parisons where needed are made with published
descriptions by Humboldt, Azara, Buffon, and
others. The bibliographic references to the species
are complete. Observations of habitats and repro-
duction are carefully recorded, and geographic
range is usually given with circumspection. Wied-
Neuwied's account of Geoffi"oy's tufted-ear mar-
moset (his Hapale leucocephalus) is an example
(my translation):

I found it in the state of Espirito Santo. I
am unable to determine if it extends north
of the Rio Doce or beyond as I could not
hunt often in the dark forests of this river
because of the Botocudo Indians. I can
therefore state that the habitat of this species
lies between 20° and 21" south latitude. The
animal is common in the forests of the Rio
Espirito Santo, especially in the outlying bush
and the mangue bush {Conocarpus and Av-
icennis) bordering the river, as well as in the

low palm {Allagoptera pumila and others)-
covered sandy coastal districts not far from
the mouth of the Espirito Santo. . . .

The following excerpt of Wied-Neuwied's
(1826, p. 161) observations on the golden lion
tamarin (Leontopithecus rosalia rosalia Linnaeus)
brings together his observations on distribution,
habits, habitat, food, and reproduction:

The sahuim vermelho is nowhere abundant;
we saw only single individuals or family
groups, particularly in the Serra da Inua, the
forests of Sao Joao, and in the hilly forest
surrounding Ponta Negra and Gurupina. The
animal lives just as well on bushy sandy
plains as in the high mountain forests. It
feeds on fruits and insects and hides from
strangers by disappearing into the leafy tree-
tops. One or two young are produced at a
birth. The female carries the offspring on her
back or at her breasts [when suckling] until
they are strong enough to follow her on their
own. . . . Any excitement causes them to erect
the long hair surrounding their faces. In gen-
eral, however, their habits are similar to those
of other sahuis.

Wied-Neuwied also accurately delimited the
distribution of the subspecies Leontopithecus ro-
salia chrysomelas and added information on hab-
its and reproduction. Wied-Neuwied notes (1826,
p. 1 59) that "sahuis bom in Europe are carried by
the father but I have never seen this here."

Although generally careful in interpreting his

HELDIANA: ZOOLOGY

". ^....

Fig. 7. Some animals of the Wied-Neuwied Brazilian expedition: upper left, Hapale chrysomelas Wied-Neuwied
(= Leontopithecus rosalia chrysomelas), possibly the holotype; upper right, Mus pyrrhorhinos Wied-Neuwied (=
fViedomys pyrrhorhinos), possibly the holotype; lower left, Desmodus rufus Wied-Neuwied (= Desmodus rotundus
E. Geoffroy); lower right, Felis macroura Wied-Neuwied (= Felis wiedii Schinz), possibly the holotype; from Wied-
Neuwied (1822-1831).

data, Wied-Neuwied could arrive at unwarranted
conclusions. Among the bats collected, the leaf-
nosed Phyllostomus hastatus was largest and for
this reason was regarded as a blood-sucking vam-
pire, although Wied-Neuwied found only insects
and no blood in the stomach of this or any other
bat he had examined. After confessing he had nev-
er seen a bat feed on blood, he correctly blamed
the large bats seen fluttering around the pack mules
at night for causing them to appear next morning
covered with blood. Convinced in his judgment,
he described the wartlike excrescences around the

mouth of innocent phyllostomine bats as adap-
tations for blood-sucking. Ironically, Wied-Neu-
wied (1824, 1826) later described and figured the
external and dental characters of a bat he named
Desmodus rufus, unaware it was a real blood-suck-
ing vampire. Wied-Neuwied noted, however, that
he had no opportunity to observe the live animal,
because it had been captured and prepared as a
specimen by assistants during his absence. The
food and habits of this bat, he believed, were no
different from those of other bats.
The mammals of Wied-Neuwied's Brazilian ex-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

pedition are described or recorded in several pub-
lications. Those under Wied-Neuwied's own name
are found in Isis ( 1 820, 1821), the Reise nach Bra-
silien in two volumes ( 1 820, 1821), the Abbildun-
gen zur Naturgeschichte Brasi liens ( 1 822-1 83 1 , see
fig. 7 for some samples), and the four-volume Bei-
trdge zur Naturgeschichte von Brasilien. The first
volume of the last title is on reptiles, the second
on mammals ( 1 826), the third and fourth on birds.

Some diagnoses and binomials that Wied-Neu-
wied proposed for new forms received duly ac-
knowledged advance publication by Kuhl (1820)
and Schinz (1821). Authorship of such newly
named forms continues to be attributed to Wied-
Neuwied, according to Articles 1 1 and 50 of the
International Code of Zoological Nomenclature.
In the few cases where Kuhl or Schinz proposed
names other than those used by Wied-Neuwied,
authorship is determined by priority.

The actual number of mammals collected by
Wied-Neuwied is unknown. According to him, they
represented 82 species, but the number recognized
today as valid is 7 1 . The specimens were preserved
in his private museum, but duplicates were dis-
tributed to the natural history museums of Berlin,
Frankfurt, Leiden, and Paris. After Wied-Neu-
wied's death, the remainder of the collection was
sold, and the American Museum of Natural His-
tory in New York acquired a part in 1869. Avila
Pires ( 1 965, p. 3) affirms that fewer than 600 spec-
imens of the original collection are registered in
the catalogue of mammals of the New York in-
stitution. Of these, only 38 skins and 16 skulls are
of South American origin. Included are holotypes
(or syntypes) of Didelphis aurita Wied-Neuwied,
Didelphis cinerea Temminck, Molossus plecotus
Wied-Neuwied, Phyllostoma brevicaudum Wied-
Neuwied, Vespertilio leucogaster Wied-Neuwied,
Vespertilio naso Wied-Neuwied, Hypudeus dasy-
trichos Wied-Neuwied, and Mus pyrrhorhinos
Wied-Neuwied.

Table 6 lists all mammalian species recorded by
Wied-Neuwied. Current names are used; syn-
onyms used by Wied-Neuwied are included.

Johann Natterer (1787-1843)

Johann Natterer, bom near Vienna, was well
schooled in the sciences, especially biology, and
in modem languages and illustration. Natterer's
father, the imperial falconer and collector of birds
and insects, taught him to hunt and preserve an-

imals as museum specimens. In 1 8 1 6 he was em-
ployed as assistant in the Imperial Natural History
Museum of Vienna and in 1817 was appointed
member of an expedition to investigate the Bra-
zilian biota. He arrived in Rio de Janeiro on 5
November accompanied by Mikan and Pohl, both
naturalists, and Schott, a botanist. Within a year
Mikan, Sochor, a hunter, and two artists who were
to accompany Natterer, retumed to Europe. Pohl
and Schott retumed in 1821.

Natterer was primarily a bird collector, but his
interest in collecting extended to mammals, other
vertebrates, insects, and parasitic helminths. He
traveled light and, as a rule, worked alone or with
few native helpers (Ihering, 1 902). He collected in
most of the eastem coastal states and in Mato
Grosso and the Amazonian region between the
Rios Tapajos and Madeira and in the Rio Negro
basin north of the Rio Amazonas (fig. 8). His main
base for the first five years was Ipanema, Sao Pau-
lo. His itinerary— with goings and comings, side
trips, short stopovers in some sites, long delays in
others— was arranged chronologically by Pelzeln
(1871,1883) into "Reisen" (or journeys), with dates
given for all points visited, and is summarized
below. Only general areas or terminal points and
inclusive dates are given.

Johann Natterer's Brazilian Reisen, 1817-1835.
I. Rio de Janeiro, 5 November 1817 to 1 No-
vember 1818.
II. Eastem Sao Paulo, 2 November 1818 to
March 1820.

III. Southern Sao Paulo to boundary between
Rio Grande do Sul and Rio de Janeiro, July
1820 to 1 February 1821.

IV. Rio de Janeiro, Sao Paulo, 1 February to
September 1822.

V. Northern Sao Paulo, Goias, eastem Mato
Grosso, Minas Gerais, October 1822 to 31
December 1824.
VI. Mato Grosso, January 1825 to July 1829.
VII. Mato Grosso, Rio Madeira, and upper trib-
utaries to Borba in Amazonas (Capitania
Rio Negro), 15 July 1829 to June 1830.
VIII. Borba to Rio Negro, Rio Casiquiare, Ven-
ezuelan border, retum to Barcelos and Bor-
ba, June 1830 to 31 August 1830.
IX. Rio Negro from Barcelos to Rio Branco, 5
September 1831 to 2 July 1832; Barra do
Rio Negro, 29 August 1832 to 7 July 1834;
Rio Tapajos, August 1834.
X. Para, Maranhao, Rio Grande, Paraiba, Per-

FIELDIANA: ZOOLOGY

■ ■ ■ /yj/__/"/»«-»r ornt .\tm^ni6er A*!// <»«.»<'

' 6fx Ar^rintr fS'2/
/«y.//r/«- /«;'/< 17'f/l'ri.

./mm /^ifA/jt . 1ntft,*l /xV

1/:^/- /S'y? .-r -IZ

Fig. 8. Map of Brazil showing routes of Johann Natterer (bold line); from [brother of Johann] Natterer (1833,
Oken's Isis, heft VI, pi. 14).

nambuco, Bahia, Rio de Janeiro, September Natterer's enormous collections were sent to the

1834 to September 1835 (no mammal col- Vienna museum and, except for the birds and

lections). mammals, were never fully reported. His friend

Sailed for Europe 15 September 1835. Andreas Wagner (1797-1861) described most of

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 6. Brazilian mammals recorded by Wied-Neuwied (1826) with some figured in the Abbildungen (1822-
1831); the arrangement is phylogenetic.

Current name

Wied-Nenwied synonym

Figure

Marsupiaua

Afarmosa murina Linnaeus, 17S8

Marmosa cinerea Temminck, 1 824

Philander opossum frenata Olfers, 1818

Didelphis marsupialis aurita Wied-Neuwied, 1826

Chiroptera

Rhynchonyaeris naso Wied-Neuwied. 1820 (Reise)

Centronyaeris maximiliani Fischer, 1 829

Peropteryx macrotis Wagner. 1843

Diclidurus albus Wied-Neuwied, 1819

Noctilio leporinus Linnaeus, 1 758

Xfacrophyllum macrophyllum Wied-Neuwied, in

Schinz, 1821
Phyllostomus hastatus Pallas, 1867
PhvUostomus obscunis Wied-Neuwied, in Schinz,

i821
Glossophaga soricina Pallas, 1 766
Anoura caudifera E. Geoffrey, 1818
CaroUia brevicauda Wied-Neuwied, 1821

Carollia perspicillata Linnaeus, 1 758

Artibeus liturcaus 0\fcT%, 1818

Desmodus rotundus E. Geoffiroy, 1810

Myotis albescens E. Geoflfroy, 1806

Myotis nigricans Wied-Neuwied, in Schinz, 1821
Eumops perotis Wied-Neuwied, in Schinz, 1 82 1

Primates

Callithrix jacchus penicillatus E. Geoffix)y, 1812

Callithrix jacchus geoffroyi Humboldt, 1812

Leontopithecus rosalia chrysomelas Kuhl, 1 820
Leontopithecus rosalia rosalia Linnaeus, 1758
Callicebm personatus persoruUus E. GeoflSroy, 1812
Callicebus personatus melanochir Wied-Neuwied,

1820 (Reise)
Alouatta caraya Humboldt, 1812
Alouatta fusca E. Geoffroy, 1812

Cebus apella nigritus Goldfiiss, 1 809

Cebus apella robustus Kuhl, 1 820

Cebus apella xanthostemos Wied-Neuwied, 1820

(Reise)
Brachyteles arachnoides E. Geoflfroy, 1806

Edentata

Tamarulua tetradactyla Linnaeus, 1758

Myrmecophaga tridactyla Linnaeus, 1758

Edentata

Bradypus torquatus Desmarest, 1816

Didelphys myosuros Temminck. 1 825
Didelphis marsupialis. Wied-Neuwied, 1826, not
Linnaeus

Vespertilio caJcaratus Wied-Neuwied, in Schinz,

1821, not Rafinesque, 1818
Vespertilio caninus Wied-Neuwied, in Schinz,

1821, not Blumenbach, 1797
Diclidurus freyreissii Wied-Neuwied

1822, Abbild.

Noctilio dorsatus Desmarest, 1818; Noctilio
unicolor Desmarest, 1818

Artibeus planirostris Spix, 1823

Glossophaga amplexicaudata E. Geoflftoy, 1818

Phyllostoma bernicaudum {sic) Wied-Neuwied,

in Schinz, 1821
Phyllostoma brachyotos (sic) Wied-Neuwied,

in Schinz, 1821
Phyllostoma superciliatum Wied-Neuwied,

in Schinz, 1821
Rhinolophus ecaudatus Wied-Neuwied, in Schinz,

1821; ZJesmorfus rw^ Wied-Neuwied, 1824
Vespertilio leucogaster Wied-Neuwied, in Schinz,

1821

Hapale penicillatus kuhlii Wied-Neuwied, 1826

p. 142)*
Hapale leucocephalus Kuhlii (sic), Wied-Neuwied,

1826t

Mycetes niger Kuhl, 1820

Mycetes ursinus Humboldt, 1812, not Humboldt,

1805
Cebus cirrifer E. Geoffroy, 1812, not Cebus fatuel-

lus Linnaeus

? Cebus flavus E. Geoffroy, 1812
Ateles hypothanthus Kuhl, 1820

Myrmecophaga jubata Linnaeus, 1766

Bradypus tridactylus Wied-Neuwied, 1826, not
Linnaeus, 1758

HELDIANA: ZOOLOGY

Table 6. Continued.

Current name

Wied-Neuwied synonym

Figure

Cabassous unicinctus Linnaeus, 1758
Euphractus sexcinctus Linnaeus, 1758

Dasypus novemcinctus Linnaeus, 1 758
Priodontes maximus Kerr, 1 792

Carnivora

Dusicyon thous brasiliensis Wied-Neuwied, in

Schinz, 1821
Chrysocyon brachyurus Illiger, 1815
Nasua nasua solitaria Wied-Neuwied, in Schinz,

1821
Procyon cancrivorus G. Cuvier, 1 798
Potosflavus nocturnus Wied-Neuwied, 1826
Eira barbara Linnaeus, 1758
Pteronura brasiliensis Gmelin, 1 788
Felis wiedii Schinz, 1821
Felis pardalis mitis F. Cuvier, 1820
Felis yagouaroundi eyra Fischer, 1814
Felis concolor Linnaeus, 1 77 1
Felis onca Linnaeus, 1758

SiRENIA

Trichechus manatus lAnnditns, 1758

Perissodactyla

Tapirus terrestris lArmaitxis, 1758

Artiodactyla

Tayassu tajacu Linnaeus, 1758

Tayassu pecari Link, 1795

Mazama gouazoubira Fischer, 1814

Mazama americana Er\\ehcn, Mil

Blastoceros bezoarticus Linnaeus, 1758

Blastocerus dichotomus Illiger, 1815

Lagomorpha

Sylvilagus brasiliensis Linnaeus, 1758

RODENTIA

Sciurus aestuans Linnaeus, 1 766

Wiedomys pyrrhorhinos Wied-Neuwied, 1821

(Reise)
Oxymycterus rufus dasytrichos Wied-Neuwied,

in Schinz, 1821
Proechimys myosuros Lichtenstein, 1818
Cavia aperea Erxleben, 1 777
Kerodon rupestris Wied-Neuwied, 1820 (Isis)
Hydrochaeris hydrochaeris Linnaeus, 1 766
Dasyprocta leporina aguti Linnaeus, 1 766
Agouti paca Linnaeus, 1 766
Coendou insidiosus Olfers, 1818
Chaetomys subspinosus Olfers, 1818

Dasypus setosus Wied-Neuwied, 1 826; Dasypus

gilvipes Illiger, 1815
Dasypus longicaudus V^icd-Neuwied, 1826
Dasypus gigas Cuvier, 1822

Canis azarae Wied-Neuwied, 1 823

Canis campestris Wied-Neuwied, 1826
A^asMfl 50c/a//5 Wied-Neuwied, 1826

Mustela gulina Wied-Ncuwied, 1821

Felix macroura Wied-Neuwied, 1 823
Felis pardalis. Wied-Neuwied, 1826
Felis yaguarundi, Wied-Neuwied, 1 826

Felis brasiliensis Wied-Neuwied, 1 82 1

Manatus americanus Link, 1795

Tapirus americanus Gmelin, 1788

Dicotyles torquatus Cuvier, 1817

Cervus simplicicornis Illiger, 1815

Cervus rufus Cuvier, 1817

Cervus campestris 'Wied-Nexxwied, 1826,

not Cuvier, 1817
Cervus paludosus Desmarest, 1 822

Hypudeus dasytrichos Wied-Neuwied, 1826

* The name is a correctly formed trinomial but this form was not in use at the time, and Wied-Neuwied used no
trinomials elsewhere in his publications on Brazilian mammals.

t The name appears to be a trinomial although the patronymic, properly in the genitive, is not italicized. Most
likely Wied-Neuwied meant to cite Kuhl for this and the preceding taxon as authority for his use of the names in
question. It was common practice at the time to cite the author who replaced an earlier generic name with a different
one.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

the new mammalian species in a series of reports
published in the Archivfur Naturgeschichte ( 1 842,
1843), in the Abhandlungen der Akademie .Vfiin-
chen ( 1 847-1 849), and his supplementary volumes
of Schreber's Sdugethiere (1840-1855). Finally,
Pelzeln (1883) brought together most, if not all,
available taxonomic, descriptive, and geographic
data in a single report. Natterer intended to work
up the entire collection himself, but died within a
few years of his return to Europe. His journal, with
notes on habits, reproduction, and anatomy of the
Brazilian animals collected, was lost.

Natterer collected 78 1 specimens of mammals,
representing more than half (58%) of the currently
known Brazilian genera and nearly as many (44%)
of the species (table 3). Most poorly represented
are bats, mice, and mouse opossums. Had Natterer
been equipped with suitable traps and trammel
nets known at the time but not used in fieldwork,
he might have collected nearly all the mammalian
genera and species now known to occur in Brazil.
Still, his collection represented more species and
included t\pes of more new species than had been
collected in Brazil by anyone else in the century,
or p)ossibIy at any time.

The numbers of genera and species of mammals
collected by Natterer, as identified by Pelzeln
(1883), are listed in Table 7. The totals are com-
pared with the numbers currently recognized, some
genera having been increased and some species
eliminated by synonymy. The revised numbers of
genera and species are shown, in turn, as percent-
ages of the estimated total numbers of currently
known Brazilian genera and species of mammals.

VII. Guianas: Mammalogy to End of
18th Century

Pierre Barrere (1690-1755)

The physician, botanist, and correspondent of
the French Royal Academy of Sciences, Pierre
Barrere, resided three years (1752-1755) in Cay-
enne, with instructions to prepare a detailed report
on the natural history of French Guiana. The
work he finally published in 1 74 1 , however, is no
more than an abbreviated glossary- of Guianan
minerals, plants. moUusks, fishes, reptiles, birds,
and mammals. The list of mammals was uncrit-
ically compiled from Marcgraf and others. Species
previously recorded by early chroniclers from the
lower Rio Orinoco region which occur throughout

the Guianas but were not mentioned by Barrere
are the golden handed tamarin {Saguinus midas),
red brocket (Mazama americana), red acuchi (A/y-
oprocta exilis), tayra (Eira barbard), white-lipped
peccary (Tayassu pecari), and silky anteater (Cy-
clopes didactylus).

Jose Gumilla (d. 1750)

A natural history and geography of the Rio Ori-
noco region in Spanish, published by Father Jose
Gumilla, provides interesting, but largely erratic,
descriptions of the countryside and human inhab-
itants, but nothing of interest regarding native
mammals. Gumilla's explorations of the interior
led him to deny the reported existence of a con-
nection between waters of the Orinoco and Negro
rivers.

Jacques Nicolas Bellin (1703-1772)

The description of the Guianan possessions of
France, Spain, Holland, and Portugal, from the
Orinoco River to the Amazonas River, by Jacques
Nicolas Bellin, published in 1763, contains infor-
mation on natural history, but adds nothing note-
worthy to the then-known mammalian fauna.

Edward Bancroft (1744-1821)

The English physician Edward Bancroft lived
three years in Dutch Guiana, now Suriname, prac-
ticing medicine and gathering notes for his Essay
on the Natural History of Guiana. The work, pub-
lished in 1 769, deals broadly with plants and an-
imals, but the author's knowledge of mammals
was mostly limited to hearsay, although he also
made some observations on animals brought to
him by natives or seen in captivity or during short
walks into the countryside. Persistent reports of
the existence of apes or ape-men in South America
were recounted by Bancroft (p. 1 30) in these terms:

The Orang-Outang of Guiana is much larger
than either the African or the Oriental, if the
accounts of the natives may be relied on; for
I do not find that any of them have been
seen by the White inhabitants of this coast,
who never penetrate far into the woods.
These animals, in all the different languages
of the Natives, are called by names signi-

HELDIANA: ZOOLOGY

Table 7. Numbers of mammalian genera and species
collected by Johann Natterer in Brazil, 1817-1835, based
on Pelzeln (1883), and compared with currently known
totals.

Table 7. Continued.

Total

Taxon

Number
reported

by
Pelzeln
(1883)

Current
equiva-
lent
number

cur-
rently
known

for
Brazil

(esti-
mated,
1984)

Percent-
age of
current
total col-
lected by
Natterer

Marsupialia

Genera
Species

2
18

6
15

8
30

75%
50%

Chiroptera

Genera
Species

10
48

28
40

60
125

47%
32%

Primates

Genera
Species

16
50

87%
56%

Edeimtata

Genera
Species

10
16

10
12

12
15

83%
75%

Carnivora

Genera
Species

11
17

10
14

71%
56%

PiNNIPEDIA

Genera
Species

2
2

0%
0%

SlRENlA*

Genera
Species

1
1

1
2

100%
50%

Perissodactyla

Genera
Species

1
1

100%
100%

Artiodactyla

Genera
Species

4
6

80%

86%

Lagomorpha

Genera
Species

1
1

100%
100%

RODENTIA

Sciuropmorpha

Genera 1
Species 5

1
3

3
6

33%
50%

Myomorpha (Murinae
Genera 3
Species 1 7

excluded)

20
45

25%
24%

Caviomorpha

Genera
Species

23
47

65%
47%

Cetacea*

Genera
Species

2
2

100%
100%

Taxon

Total

cur-

rently

known

Percent-

Number

for

age of

reported

Current

Brazil

current

equiva-

(esti-

total col-

Pelzeln

lent

mated,

lected by

(1883)

number

1984)

Natterer

Totals

Genera

170

58%

Species

202

156

358

44%

* Fresh water only.

fying a Wild Man. They are represented by
the Indians as being near five feel in height,
maintaining an erect position, and having a
human form, thinly covered with short black
hair; but I suspect that their height has been
augmented by the fears of the Indians, who
greatly dread them, and instantly flee as soon
as one is discovered, so that none of them
have ever been taken alive, much less at-
tempts made for taming them. The Indians
relate many fabulous stories of these ani-
mals; and, like the inhabitants oi Africa and
the East, assert, that they will attack the
males, and ravish the females of the human
species.

Philippe Fermin (1720-1790)

Philippe Fermin, the author of an account pub-
lished in 1769 of the history, geography, and nat-
ural objects of colonial Suriname, was one of those
European men who "never penetrate far into the
woods." Indeed, Fermin believed that all Euro-
[)eans and Creoles were physically incapable of
coping with the difficulties of surveying the natural
fauna of the countryside, let alone the wilderness,
or resisting the diseases generated by the "foul"
air of forests and swamps. Notwithstanding this,
Fermin compiled a fair list of the mammals. The
didelphids included Didelphis marsupialis, Phi-
lander opossum, and Marmosa spp. All three kinds
of anteaters and the two- and three-toed sloths are
mentioned. The two native squirrels, Sciurus aes-
tuans and Sciurillus pusillus, are distinguished.
Other rodents are the capybara, paca, a porcupine,
cavy, spiny rats or echimyids (most likely of the
genera Proechimys and Echimys), and a water rat

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

(probably Nectomys squamipes). The carnivores
include tayra, otter, jaguar, puma, margay, ocelot,
and two kinds of bush dogs (Dusicyon thous and
possibly Speothos venaticus). Monkeys are Sa-
guinus midas, Saimiri sciureus, Pithecia pithecia,
Chiropotes sat anas, Cebus apella, Alouatta seni-
culus, Ateles paniscus. and variants of some of
them regarded as distinct species. African simians
introduced with the slave trade and mentioned by
Marcgraf are included. The ungulates are tapir,
brocket {Mazama americana), and the collared
and white-lipped peccaries.

Regarding white-lipped peccaries, Fermin af-
firms they form herds of as many as 300 individ-
uals. Hunters, he states, tremble when they hear
the sound of their clicking tusks. When attacked,
only two avenues of escape are open: The first is
a tree, if it can be climbed; the second and surest
is standing ground and urinating, the odor of the
urine, he affirms, being a powerful peccary repel-
lant.

Monsieur Bajon (1763?)

The French physician, surgeon, and anatomist
Bajon, with 1 2 years' residence in French Guiana,
investigated climate, agriculture, natural history,
and human diseases. The knowledge he gained was
acquired firsthand, much of it new or supplemen-
tary to what was already contained in the ency-
clopedic volumes on natural history by Buffon and
Daubenton.

In the second of his two-volume work, Bajon
(1778, p. 178) declared that, contrary to popular
belief, the jaguar feared man and did not attack
without provocation. His accounts of habits and
detailed descriptions of intestinal morphology and
female genitalia of peccaries supplement Dauben-
ton's (in Buffon) gross anatomy of a male collared
peccary. Bajon clarified the differences between
the agouti (Dasyprocta leporina) and acouchi {My-
oprocta exilis). He described the male agouti penis,
with its peculiar complement of spines, erectile
spears, and sharp blades. Descriptions with life
history notes are given for the chien sauvage {Du-
sicyon thous), eira {Eira barbara), and chien cra-
bier (Procyon cancrivorus). Marsupials fascinated
him, particularly the role of the pouch in females
of the pean {Didelphis marsupialis), quatre-ouel
(Philander opossum), and also the pouchless rat
de bois {Marmosa sp.). The commonly held belief
that each didelphid young is bom and develops at
the end of a teat was rejected by Bajon, but despite

numerous observations and dissections, he failed
to solve the mystery of marsupial birth.

Bajon's monographic account of the tapir {Tap-
irus terrestris) includes detailed, but not always
accurate, descriptions of anatomy, reproduction,
development, behavior, food, vocalization, hunt,
and human utilization.

John Gabriel Stedman (1744-1797)

A soldier of the Scots Brigade of the Nether-
lands, John Gabriel Stedman arrived in Suriname
in 1773 to help subdue the uprising of the African
slaves. Most of the fighting was already over when
he landed, so Stedman devoted much of his time
to recording his observations of life in the country
and wilderness. His Narrative, published in 1 796
in two volumes, contains much on the natural
history of Suriname, with illustrations by his own
hand (fig. 9). The mammals, some only listed, oth-
ers described, often with anecdotes, are the fol-
lowing. Stedman used local names, current bino-
mials are in parentheses.

Volume I, p. 14. Narwhal (Monodon monoceros).
Sighted from shipboard at Devil's Island off
Cayenne. ". . . appeared but six or eight feet
in length, and its horn about four. . . . The
narwhal ... is more frequently found in cold
than warm climates. The female is said to be
unprovided with that protuberance so re-
markable in the male. It appears that some
authors have confounded this animal with the
sword-fish, to which however it does not prove
to have the very smallest resemblance."
The locality record for the circumpolar narwhal
is unexpected, and no doubt erroneous.
Nevertheless, Stedman's description is accu-
rate albeit the dimensions given seem small.
At the same time, Stedman provided a de-
tailed description and good figure of a sword-
fish or sawfish to prove it was not a sawfish
he saw!

Volume I, p. 153, pi. 16. Sicapo (Bradypus tri-
dactylus).

Volume I, p. 153, pi. 16. Dago luyaree (Choloepus
didactylus).

Volume I, p. 153. Ourang-outang.
"I have never seen, nor heard described, while
I was in this country. . . ."

Volume I, p. 166, pi. 18. Micoo or mecoo {Cebus
apella) (fig. 9).

HELDIANA: ZOOLOGY

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HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Volume I, p. 166, pi. 18. Kessee-keesee or kishee
kishee (Saimiri sciureus) (fig. 9).

Volume I, p. 167. Monkee-monkee {Saguinus mi-
das).
"One morning I saw from my barge a monkey
of this kind come down to the water's edge,
rinsing his mouth, and appearing to clean his
teeth with one of his fingers."

Volume I, p. 168. Tavous (Lutra enudris).

Volume I, p. 221. Sea-cow (Trichechus manatus).
About three in the morning while asleep in a
boat, Stedman and a companion were sud-
denly thrown from their bunks.
"By the account of the negroes [a manati had]
slept under the boat, which, by the creature's
awakening, had been lifted up and thrown
upon one side, and again replaced when the
manati made its escape from underneath. I
did not so much as see the creature, nor in-
deed hardly had the negro, owing to the dark-
ness of the night."

Volume I, p. 222, pi. 24. Capasce (Dasypus no-
vemcinctus); largest armadillo (Priodontes
maximus).

Volume I, p. 223, pi. 24. Adjora (Coendou pre-
hensilis).

Volume I, p. 224. Hedge-hog (spiny rat, family
Echimyidae).

Volume I, p. 308, pi. 33. Bajew {Odocoileus vir-
ginianus, adult male and spotted fawn); boo-
see-cabritta (Negro), wirrebocerra (Indian)
(Mazama americana, large spotted female,
smaller spotted fawn).

Volume I, p. 347. Coney coney (Negro), puccarara
{Dasyprocta leporina).

Volume I, p. 355, pi. 37. Pingo (Tayassu pecari).
"They live in herds of sometimes above three
hundred and run always in a line, the one
closely following the other; when the foremost
or leader is shot, the line is instantly broken,
and the whole herd is in confusion, for which
reason the Indians take care (if possible) to
knock their captain on the head before the
rest; after this the others even often stand still,
stupidly looking at one another, and allowing
themselves to be killed one by one, of which
I have been a witness. They do not attack the
human species, or make any resistance at all,
like the European wild-boar, when wounded,
as has been by some authors erroneously as-
serted."

Volume I, p. 355. Peccary (Tayassu tajacu).

Volume I, p. 356. Cras pingo {Sus scrofa).

Volume II, p. 10, pi. 42. Quata or Quato (Ateles
paniscus).
"Their throwing short sticks and excrements
seems to be no more than a mimicking of the
human actions without any purpose, as they
neither have strength to throw far, nor dex-
terity to hit their objects, and if they befoul
them it is by accident only. But what appears
to be peculiarly remarkable is, that when one
is hurt by a musket or arrow, the poor animal
instantly claps its hand on the wound, looks
at the blood, and with the most piteous lam-
entations ascends to the very top of the tree,
in which he is assisted by his companions;
where, hanging by the tail, he continues to
bewail his fate, till by the loss of blood he
grows totally faint, and drops down dead at
the feet of his adversaries.
"It is not so extraordinary that one of this species,
when wounded, should be assisted by one of
his companions in climbing; but that they
should have so much knowledge in botany,
as to procure vulnerary herbs, and chew and
apply them to the wound, is what I cannot
credit, though it is so confidently asserted by
a late traveller; and as to the assistance they
give in passing a river, by holding each others
tails, and swinging till the lowermost is thrown
up to the branch of a high tree ... I must take
the liberty to doubt this fact. . . .

Volume II, p. 12. Wanacoe (Pithecia pithecia,
male).
"This monkey is the only one of the species
[monkeys] that is not sociable, being constant-
ly found alone, and so despicable is this sol-
itary animal, that he is continually beaten and
robbed of his food by all the others, from
whom he is too lazy to escape, though too
cowardly to fight."

Volume II, p. 12. Saccawinkee {Callithrix jacchus
jacchus).
[Common marmosets were brought from Brazil
for the pet market. They are not native to the
Guianan region.]

Volume II, pp. 16-17. Brown squirrel (Sciurus
aestuans); white squirrel (Sciurus aestuans,
albinotic); flying squirrel (probably mistaken
impression of a leaping pygmy squirrel, Sci-
urillus pusillus).

Volume II, p. 40, pi. 46. Taibo, woodrat (My-
oprocta exilis).
The description is better than the figure which
suggests a doglike marsupial.

HELDIANA: ZCX)LOGY

Volume II, p. 41, pi. 46. Crabbo-dago {Galictis
vittata).

Volume II, p. 49, pi. 48. Tyger (Felis onca).
"It has even happened that Ihe jaguar has car-
ried off young negro women at work in the
field, and too frequently their children."

Volume II, p. 50. Red tyger (Felis concolor).

Volume II, p. 50, pi. 48. Tyger-cat (Felis pardal is).

Volume II, p. 5 1 . Jaguaretta (melanistic Felis onca).
"I have never seen one."

Volume II, p. 135. Cabiai (Hydrochaeris hydro-
chaeris).

Volume II, p. 142, pi. 57. Vampire or spectre
(Vampyrum spectrum).
Figured are a flying bat and a side view of a
truncated head that had been preserved in
spirits. Stedman, while asleep, had been bitten
on his toe by a true vampire bat, likely Des-
modus rotundus. He had not seen his attacker,
but like others believed most bats were vam-
pires, particularly the larger species, most cer-
tainly the largest, Vampyrum spectrum.

Volume II, p. 144, pi. 47. Murine or mouse opos-
sum (Philander opossum).

Volume II, p. 152, pi. 58. Paca (Agouti paca).
"Nothing can be better eating than the Paca or
spotted Cavey."

Volume II, p. 153, pi. 58. Agouti pacarara, Indian
coney (Dasyprocta leporina).

Volume II, p. 153. Indian rat-coney (Myoprocta
exilis).
"This I never saw, unless it is the same animal
. . . that I have described under the name of
bush-rat."

Volume II, p. 175, pi. 59. Sea-cow or manatee
(Trichechus manatus) (fig. 9).

Volume II, p. 1 76, pi. 59. Tapir (Tapirus terrestris)
(fig. 9).

Volume II, p. 176. Mermaid.
"Major Abercromby . . . declared that a mer-
maid was lately seen in the River Surinam.
Lord Monboddo also positively affirms the
existence of sea-women and sea-men, while
he asserts that they were seen so late as 1 720.
But, however respectable his lordship's judge-
ment and authority may be on other subjects,
I can no more agree with him, as to men and
women, having fins and scales, than to their
having tails."

Volume II, p. 235. Howling baboon (Alouatta se-
niculus).

Volume II, pp. 325-326. Awaree (Didelphis mar-
supialis).

Volume II, p. 327, pi. 74. Quacy-quacy (Nasua

nasua).
Volume II, p. 328, pi. 74. Great ant-eater (Myr-

mecophaga tridactyla).
Volume II, p. 329. Tamandua (Tamandua tetra-

dactyla).
Volume II, p. 329. Fourmillier (Cyclopes didac-

tylus).

VIII. Guianas: Mammalogy of First
Half of 19th Century

Sir Robert Herman Schomburgk (1804-1865)
and Richard Schomburgk (1811-1891)

Robert Herman Schomburgk was bom in Frei-
burg, Germany, son of a Protestant minister. In
1829 he went to the United States and in 1830 to
Anegada of the Virgin Islands. His survey of the
island, submitted to the Royal Geographical So-
ciety of London, won him the command of an
exploring expedition to British Guiana (Guyana)
in 1835.

Robert Schomburgk's accounts of his travels in
the colony and bordering parts of Brazil and Ven-
ezuela during 1835-1839 were published by the
Royal Geographical Society in its Journal for vol-
umes 6 (1836), 7 (1837), and 10 (1840). The re-
ports were translated into German by O. A.
Schomburgk and published in 1841 as a single
volume. This, in turn, was translated back into
English by Roth (1931). A brief description of the
colony by Robert Schomburgk was published in
1840. Some notes on natural history by Schom-
burgk were included in his reports to the Geo-
graphical Society; others appeared in several num-
bers o^ the Annals of Natural History (London) for
1840.

Upon the successful conclusion of his explora-
tions in 1839 and return to England, Robert
Schomburgk was commissioned in 1840 by the
government to survey the colony and fix its bound-
ary with Venezuela. He was knighted in 1 845 after
his return to England.

Richard Schomburgk, with the patronage of the
King of Prussia, accompanied his older brother on
the second journey to British Guiana. Plants and
animals collected by the expedition were sent to
the Berlin museum for scientific study where they
were examined by Richard Schomburgk and other
specialists; the mammals were studied by Schom-
bui^ and Cabanis. Richard Schomburgk's three-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

volume account in German of the travels from
1840 to 1844 was published 1847-1848. Roth's
English translation of the first two volumes ap-
peared in 1922-1923.

As a result of the Schomburgk expeditions, Brit-
ish Guiana advanced from a practically unknown
South American country to one of the then best
known for its geography, biota, and ethnology.
Virtually all major physical features of the Guia-
nan region, from the Corentyne River between
British and Dutch Guiana (Suriname), west across
the colony and headwaters of the Rios Branco and
Negro in Brazil to headwaters of the Rio Orinoco
in Venezuela, were traversed, described, and
mapped in detail (fig. 10). The reported observa-
tions on mammals are as good and as welcome
today as when first published.

The following excerpts of observations on mam-
mals taken from the three volumes of the Reise
originated with Richard, Robert, or both. Those
from the earlier published Annals are Robert's.
The mammals were first identified by Cabanis.
Current scientific names, however, are used here
with the Cabanis synonyms in parentheses. None
of the animals were figured in the original works
cited.

MARSUPIALIA

Didelphis marsupialis marsupialis Linnaeus {Di-
delphys cancrivord)

Reise 1:192 (human breast-fed young).

Annals 5:343. ". . . if we could reconcile the
geographical distribution of Z). virginiana over
a space so different in temperature, I should
consider the specimen [of D. marsupialis] I
am now describing a variety of that species;
the circumstance that the ears are of uniform
black would scarcely constitute a specific dif-
ference."

Reise 111:777 (behavior in captivity).
Philander opossum opossum Linnaeus {Didelphys
quica)

Reise 111:777 (distribution).

Annals 5:344 (description; habits).
Caluromys philander philander Linnaeus (^Didel-
phys philander)

Reise 111:777 (distribution).

Annals 5:344 (description; habits).
Marmosa murina murina Linnaeus (Didelphys
dorsiguera. D. musculus Cabanis)

Reise 111:777 (distribution; characters; habitat).

Annals 5:345 (description; habits).
Lutreolina crassicaudata turneri Gunther

Reise 111:777 (predation).

Chironectes minimus minimus Zimmermann
(Chironectes variegatus)
Reise 111:777 (distribution).

CHIROPTERA

Molossus molossus Pallas (Molossus obscurus)
Tonatia bidens Spix (Phyllostoma bidens)
Reise in:772 (habitat; colony size; characters).

EDENTATA

Myrmecophaga tridactyla tridactyla Linnaeus
(Myrmecophaga jubatd)

Reise 11:44, 214, 223, 374 (characters; habits;
defense; chase; flesh).

Reise 111:782 (distribution).

Annals 4:203-207 (characters; habits; capture):
"The young Ant-bear was quite wild at first,
and sought for some dark comer in the room
in which it was confined, in order to hide
itself When we approached it, it put itself
immediately in defense like the adult ones,
and struck out with its right paw, uttering at
the same time a growl like that of an incensed
puppy. After a few days, however, it became
accustomed to its situation, and an Indian
woman took upon her to feed it with milk and
Cassada [cassava] and sometimes White Ants.
It soon showed great attachment to her and
followed her like a dog.

"It appeared to be of a very cold nature;
not only the extremities, but the whole body
felt cold to the touch, although we kept it
wrapped up in a blanket. It preferred, how-
ever, to be nestled, and to be taken up, and
on putting it down it uttered a whining but
not unpleasant sound; when it did not succeed
in attracting attention, and was not taken up
again, the whining sound was raised to a harsh
and grating noise. In following a person, it
directed its course more by the smell than by
sight, and carried its snout close to the ground.
If it found itself at fault, it wheeled round at
right angles upon the hind legs, and snuflTed
the air in all directions, until it found the right
scent again. Of the dimness of its sight we had
various proofs; it hurt itself frequently against
objects that stood in its way, not observing
them until it came in contact with them. Its
power of smelling was exquisite, and it could
discover its nurse, or any person to whom it
had taken a liking, at a considerable distance.
Upon these occasions it would immediately
commence the whining sound so peculiar to

HELDIANA: ZOOLOGY

Fig. 1 0. Map of British Guiana (Guyana) and bordering parts of Venezuela, Dutch Guiana (Suriname), and Brazil.
Robert Herman Schomburgk's routes and surveying areas (1835-1839) shown by large dots. The map (without the
dots) was copied and redrawn by A. Lee Owen for the Roth translation of R. H. Schomburgk (1841).

this animal. It was an expert climber; it hap-
pened that I was one of its favourites, and
whilst writing on my table it used to come
softly behind me, and as soon as it was sure
it had found me out, it climbed up my legs
with great dexterity. It showed its attachment
by licking, and was very gentle and even spor-
tive; we all prized it highly. . . .

"It secretes a liquid substance, transparent
like water, which drops down almost con-
stantly out of its nostrils and mouth; this is
the more remarkable, as it used very little
water. ..."
Tamandua tetradactyla tetradactyla Linnaeus
{Myrmecophaga tamandua)
Reise 111:782 (distribution).

Cyclopes didactylus didactylus Linnaeus

Reise 111:782 (distribution).
Bradypus tridactylus Linnaeus (Bradypus gularis)

Reise 1:142, 258, 455 (capture; swimming;
climbing; mother-infant).

Reise 111:781 (distribution).
Bradypus variegatus Schinz {Bradypus torquatus)
Priodontes giganteus E. Geoffrey

Reise 11:97 (characters; flesh).

Annals 5:32-33 (habits; description).
Dasypus novemcinctus novemcinctus Linnaeus
{Dasypus peba)

Reise 11:24, 29 (excavation).

Annals 5:34 (description; habits; reproduction).
Cabassous unicinctus unicinctus Linnaeus (Dasy-
pus tatouay)

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Reise 111:782 (distribution).

Annals 5:34 (description).
Euphractus sexcinctus sexcinctus Linnaeus (Da-
sypus encoubert)

Reise 111:782 (distribution).
['!\Dasypus sabanicola Mondolfi {Dasypus minu-
tus not Desmarest [= Zaedyus pichiy Des-
marest])

Reise 11:49 1 . "Of the fauna of the Sandhills [for-
mation across Guyana 2 to 40 miles from
coast], the genus Dasypus seemed to be the
most numerous among the mammals and of
the species present in Guiana three are found
on the sandhills alone: Dasypus Peba Desm.
([= D. novemcinctus] lessy of the Arawaks);
D. minutus Desm. (lessy Barakatta of the Ar-
awaks); and D. tatouay Desm. [= Cabassous
unicincius]."
[ly' Dasypus villosus"" {Chaetophractus villosus
Desmarest [not Guianan]

Reise 11:24. "One of the boys brought me an
armadillo {Dasypus villosus Desm.) which he
had surprised on his way across the savannah
[south of the Kanuku Mountains]."

Reise 11:97. "The sharp eyes of a Wapisiana
again noticed something alive moving about
in the savannah below; he quickly ran to the
spot and soon returned carrying another but
smaller [than Priodontes giganteus] armadillo
by the tail. It was Dasypus villosus Desm.
According to the statements of the Indians
this species is particularly distinguished by a
peculiar growth of hair that covers not only
the body but also the plates on the back, is
solely present in the savannahs, and for the
most part lives on carrion ... a characteristic
that is ascribed only to this one species
amongst the seven met with in Guiana."

Reise 111:782 (distribution).

Annals 5:34. "The savanna armadillo is Des-
marest's Dasypus villosus; and, as we were
assured by the Indians, it inhabits only the
plains, and is never to be met in the forest,
the Indians accuse it of feeding occasionally
on carrion."

PRIMATES

Ateles paniscus paniscus Linnaeus

Reise 11:93. "One finds them mostly in com-
panies of 1 6 to 20; often also in lesser number.
I never noticed them on the ground but always
on the highest trees. When exposed to the full
rays of the sun, they lie at full length stretched
out on the branches, to bathe themselves in
iL"

Reise 111:767 (troop size; reproduction; mother-
infant).

Alouatta seniculus seniculus Linnaeus

Reise 1:278, 352 (vocalization; habits; flesh).
Reise 111:768 (characters; distribution; vocal-
ization; social organization; mother-infant).

Cebus apella apella Linnaeus (and other monkey
species)

Reise 1:354 (sociability; troop size, 400-500 in-
dividuals).

Reise 11:247. "It is only in the Canuku Ranges
that I can call to mind having met troops of
monkeys that consisted solely of Cebus apella:
their haunts seem generally limited to partic-
ular localities because except in the Ranges
just mentioned, I have only seen them on the
coast and then always among C capucinus
[C. nigrivittatus] with which the neat little
Callithrix [= Saimiri] sciurea had also often
associated itself. I invariably found Mycetes
[= Alouatta], Ateles, Pithecia and Hapale [=
Saguinus midas] absolutely separate from
one another and even among Pithecia leuco-
cephala [= P. pithecia] never a specimen of
Pithecia [= Chiropotes satanas] chiropotes.""

Reise 111:768-770 (characters; troop size; be-
havior; urine washing; tool use): "I placed
some fruit near the chained monkey out of
arms reach so he tried to sweep it nearer with
his tail. This failing, he searched around as
far as he could and found a stick and with it
managed to roll the fruit to himself."

Cebus nigrivittatus olivaceus Schomburgk (Cebus
capucinus not Linnaeus)

Reise 1:247, 437 (variation; mother-infant; do-
mestication).

Reise 111:770 (most common and widely dis-
tributed Guianan monkey).

Saimiri sciureus sciureus Linnaeus

Reise 1:333 (social relations; mother-infant).
Reise 11:247, 366 (associations).
Reise 111:770 (distribution; not viable in captiv-
ity).

Chiropotes satanas chiropotes Humboldt {Pithecia

chiropotes, P. satanas)
Reise 1:351, 352 (description; social relations;

flesh).
Reise 111:771 (distribution).

Pithecia pithecia pithecia Linnaeus {Pithecia leu-

cocephalus)
Reise 1:352 (social relations).
Reise 111:771 (troop size; distribution).
Royal Geographical Society of London, 6:265

(1836):

FIELDIANA: ZOOLOGY

". . . numerous monkeys jumped from branch
to branch, and, astonished at the uncommon
visit, accompanied us for a considerable dis-
tance [along the banks of the upper Essequibo
River]. Our Caribbees called this species ar-
ieghi, or yahriae; the male has straight long
hair of a shining black, the head rather round,
the forehead and part of the face and neck
covered with short, yellowish hair, part of the
front, the nose, and mouth black, the latter
slightly bearded, hands black, nails claw-like,
except the thumb. The female is different in
colour, and her fur resembles that of the Eu-
ropean hare; her hands are likewise black, and
covered with short yellowish hair, from under
the eyes to the chin extends hair of a similar
colour, but somewhat longer than those of the
front and cheeks, the breast is nearly naked,
and the oshyoides [(sic) oschyoides or scrotal
pad] visible. They jumped with great agility
from tree to tree, the female and sometimes
the male carrying the young ones upon the
back. . . ."

The strongly marked sexual dichromatism
described in 1 836 by Robert Schomburgk was
not discerned by taxonomists until late in the
century. Twelve different names had been be-
stowed on Pithecia pithecia, five of them based
on males, the others on females.

Aotus sp. (Nyctipithecus trivirgatus)

Reise 11:460 (house pet seen at Asacota, Bari-
mani River, NW Br. Guiana).

Saguinns midas midas Linnaeus {Midas rufiman-
us)
Reise 11:366, 367, 505 (distribution; behavior).
Reise 111:772 (distribution; vocalization; captiv-
ity).

CARNIVORA

Nasua nasua vittata Tschudi (Nasua socialis; Na-
sua solitaria)
Reise 11:247-248. "The new Nasua I discovered
here . . . suffered a strange fate in its identi-
fication ... we took it for a new species, but
unfortunately possessing too few natural-his-
tory books to confirm our subspecies, for-
warded it to Berlin with the next assignment
undescribed. I was accordingly all the more
surprised to find that very same Nasua de-
termined as A^. vittata by von Tschudi in his
Untersuchung iiber die Fauna Peruana. The
specimen was shown him on its arrival and
he, recognizing it as new, took the required
notes, and before it was yet described in Ber-

lin, published it in his Fauna Peruana, al-
though it does not occur there."
Annals 5:431-432:

"They live in large societies, and know how
to defend themselves bravely if attacked by
dogs; indeed they fall often en masse upon
them and kill the assailants. They are excel-
lent climbers, and in descending a tree they
always come down head foremost. Their food
consists of insects, fruits, roots and such small
prey as they are able to secure. They are de-
structive to young birds, and expert in digging
after large beetles, for which their claws, which
are very strong, are admirably adapted. They
do not burrow in the ground for a residence."

Procyon cancrivorus cancrivorus Cuvier

Reise 11:443 (behavior).

Annals 4:433-434:

"Although the Racoon [sic] is not an animal
which inhabits the savannahs, its relation to
the preceding genus induces me to give now
the few particulars which I know about its
habits. It frequents the sea coast, and is gen-
erally found in the neighbourhood of inhab-
ited spots, where it is destructive to poultry.
"Among the favourite haunts of these an-
imals are the thickets of Curida bushes {Avi-
cennia tomentosa), which extend along the sea
coast, where they feed upon crabs which they
are expert in killing, first tearing off their claws
or nippers; and being thus disabled from doing
harm, the crab dog or racoon uses its sharp
teeth to break the shell. In their native state
they sleep by day, and issue at dusk in search
of food; birds, insects, roots, and vegetables,
nothing comes amiss; and as they possess a
particular fondness for sweets, I have been
told by practical planters that the injury which
they do to sugar plantations is very consid-
erable.

"They take their food with both paws like
the squirrel, and are fond of dipping it in water.
I have noted with astonishment that they drink
as well by lapping like the dog as by sucking.
I have had several in a domesticated state, all
of which possessed this peculiarity."

Potos flavus flavus Schreber (Cercoleptes caudi-
volvulus)
Reise 11:435 (habits; food; predation).
Annals 5:29 (habits; distribution).

Eira Barbara poliocephala Traill {Galictis barbard)
Reise 11:99 (chase; characters).
Annals 5:30 (habits; distribution): ". . . like the
coati or Nasua, are able to run down a tree

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

. . . head first. They are sometimes tamed and
are thus gentle and playful; but they are easily
excited, and when preparing for defense or
war they erect the hair of their tail."
Galictis vittata vittata Schreber (Galictis allaman-
di)

Reise 11:447 (characters).

Annals 5:31-32 (habits).
['?]Lutreolina crassicaudata Desmarest, or perhaps
Mustela africana Desmarest (Mustela brasi-
liensis?)

Reise 111:775: "I have only a few incomplete
stuffed specimens found among the Arekuna
Indians who wore them as ornaments."
Lutra enudris enudris F. Cu vier (Lutra enydris [sic])

Reise 1:340 (encounter).

Reise 111:775 (distribution).
Pteronura brasiliensis brasiliensis Zimmermann

{Pterura [sic] sambachii)
Otters {Lutra and/or Pteronura)

Reise 1:340; 11:35 (habits).

Annals 5:284-285 (habits):

"We watched a pack of Otters at the Great
Cataracts of the Corentyn, where, at the basin
which one of the cataracts formed, they ap-
peared to carry on their pursuits with great
success. One had secured a Haimura at least
from ten to twelve pounds weight, and carried
it in its mouth to a rock which was partly over
water. Here it began devouring its prey with-
out taking much notice of us, although we
were not twenty yards from it on the opposite
shore. It did not care for our shouting; its
success was however disputed by the Indians,
who got into the canoe and paddled so rapidly
towards the rock, that the Otter saw itself
obliged to retreat and to leave the better half
of the fish to the Indians. Although the Otters
were numerous round the rock, none of them
showed any intention to share the prey with
the successful hunter or to dispute its posses-
sion.

"I have already alluded to their having their
holes on the edge of rivers, sheltered by the
impending bank. Every rock in the vicinity
of their residence bears the mark of their ex-
crements; and their feeding-places are so de-
void of vegetation, if we except the larger
bushes and trees, that they cannot be mistak-
en, even if the number of scales and fish-bones
did not point out the frequency of their visits.
A complete path leads up to these places,
which, in consequences of their ascending and
descending in single file, is hollowed out.

"The young remain for a considerable time
under the protection of their parents, the
mother teaching them to plunge and dive at
approaching danger.

"We had entered the upper Essequibo by
its tributary the Cuyuwini, and passed at the
foot of a ridge of mountains, when we ob-
served on a large ledge of rocks a family of
Otters, consisting of about fifteen, including
old and young. At our approach they broke
out into their peculiar noisy cry, and the par-
ents seizing the young with their mouth they
plunged into the water and disappeared, —and
having placed their young in security, we saw
them shortly after appearing at the head of
our canoe. They raised themselves with half
their body out of the water, snoring for rage
and showing their formidable teeth. At ap-
proaching danger or when apprehensive of it,
they collected in a body, deputing the most
courageous in advance; as our canoe came
nearer, they sank under as if by a precon-
certed sign, and appeared the next moment
within a few yards of it. We saw nothing again
of the young; but the adults and larger-sized
young ones accompanied us, threatening and
snoring, until no doubt we were so far out of
reach of their stronghold that they considered
their progeny now safe. In other instances,
when we attempted to find out their holes,
they became so outrageous that they bit our
paddles and left the print of their teeth. The
Indians know nevertheless how to surprise the
young ones, who are then taken home alive,
and become in a short time so tractable that
they follow their masters like dogs."
Dusicyon thous thous Linnaeus (Canis cancrivo-
rus; Canis azarae)
Reise 11:196, 338 (habits).
Annals 4:430-43 1 (characters; habits):

"They vie in cunning and art with the Eu-
ropean fox, and the depredations which they
commit on the hen-roosts are considerable.
Their favourite haunts are thickets near open
savannahs, and if a pack succeed in entering
the village and in surprising the Indians' poul-
try, few escape, as they completely surround
the roosting-place, and generally carry off their
spoil before the inhabitants have any idea of
their presence. I have been assured by the
Indians that they soon run down deer, and
pursue their game under full cry. They destroy i
in other ways large quantities of game. ... :
They seldom lose, even when domesticated, \

HELDIANA: ZOOLOGY 1

their depredatory habits, and those Indians
who raise them for the sake of procuring a
cross breed with the dog, are obliged to keep
them tied, as otherwise, they would kill all the
fowls and parrots. It is called by the Macusis
Maikang, in Warrau Warityou.

"The variety which has sprung from the
breed between the Indian domestic dog and
the Carasissi more resembles the dog, its body
is however longer in proportion to its size,
and its ears are pricked up. Their progeny
become prolific. They are hardy, and many
of them prove excellent hunters; they are
therefore very much prized by the Indians,
who pay great attention to their training."

Reise 111:775 (distribution).
Felis concolor discolor Schreber

Reise 11:86 (characters; predation).

Annals 4:325-326 (characters; habits; preda-
tion):

"It is very destructive to the cattle farms,
and it is so powerful an animal, that I have
been told by an eye witness, that it killed a
mule and dragged it across a trench to the
opposite side, although the trench was not
quite full of water, and the Puma had to drag
it a few feet up hill, after it landed with its
prey on the other side. My informant, who
had watched its proceedings, had meanwhile
sent for his gun, and shot him while attempt-
ing to pull the mule into the wood. They seem
to be particularly partial to dogs, and a great
number of those which are kept by the settlers
for the purpose of hunting, are killed and eat-
en by them. They follow in the woods the
herds of Peccaries, and watch their motion in
order to seize upon the stragglers, being well
aware that if they attacked the flock, they
would be overpowered and torn to pieces.
They hunt as well by day as in the night, and
feed also on deer and the smaller domestic
animals. They give birth to two young ones,
seldom three, which have spots of a darker
hue, more or less visible, according as the
lights fall upon them, and which I have been
told they lose after the first year. . . ."
Felis onca onca Linnaeus {Felis nigra)

Reise 1:436 (encounter).

Reise 11:34, 85-90, 504 (encounter; characters;
predation; distribution; vocalization; artifacts
of teeth and hides): "Except during the period
when the female has her young, the jaguar
does not seem to possess any particular lair.
... It swims over the widest rivers. . . . When

circling round a camp or cattle-pen, it is al-
ways with a continual purring; not until hunt-
ing at night for its prey does it set up a frightful
roar, that booms through the whole forest."
Annals 4:262-263:

"I consider the number of wild cattle scat-
tered over the savannahs at about 4000, but
I doubt whether they are on the increase, as
man and jaguars commit fearful ravages
among them. . . . Their most deadly enemy is
the greater jaguar, Felis onca, Linn., which
hovers in such quantities about Fort San Joa-
quim, that during the month of June 1838,
twelve individuals were killed by the cattle-
drovers. They are very daring, and sometimes
kill cattle within a few yards of houses that
are inhabited. They care very little for the fires
which are made to prevent their encroach-
ments. If one or a pair of these animals would
take up their quarters in the vicinity of a cattle
farm, scarcely a night passes in which they do
not commit ravages. They do not eat much
of any they kill, perhaps ten or twelve pounds,
and principally of the breast; but they prefer
killing fresh every time they are hungry. When
out of the reach of cattle farms or the wild
herds of the savannahs, they subsist on Pec-
caris, Capybaras, Tapirs, and Deer. . . ."

Felis pardalis melanurus Ball [or 9 Felis onca onca
Linnaeus]

Reise 11:83 (characters; predation).

Annals 4:263:

"Not less destructive is the Turtle-tiger, a
species or variety of the former [Felis onca].
They are of the same strong build as the great-
er jaguar, and very much resemble it both in
form, colour, and disposition of its spots, but
they are about a third less in size. In the vi-
cinity of human habitations they commit great
ravages among domestic animals; Hogs,
Sheep, Goats, &c. are alike exposed to their
attacks, but I never heard of an authenticated
instance of their attacking man, although they
will come boldly to his habitation, and even
enter the houses and carry away the dogs from
the fireside."

Felis tigrina tigrina Schreber

Felis wiedii vigens Thomas (Felis macroura)

Reise 1:85 (characters).
Felis yagouaroundi yagouaroundi E. Geoffrey (Felis
jaguarundi; F. unicolor)

Reise 11:227 (encounter).

Annals 4:327 (description; predation).

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

RODENTIA

Sciurus aestuans aestuans Linnaeus

Reise 11:491 (characters).

Reise 111:778 (distribution).
Echimys armatus armatus I. Geoffroy {Echinomys
hispidus not Desmarest)

Reise 11:498-499: "The strange hedgehog Echi-
nomys hispidus Geoffr. seems most plentiful,
especially in the neighborhood of the small
forest streams. It appears to reside upon the
trees: at least I have never come across it on
the ground. In climbing and springing from
branch to branch it can vie with the smartest
squirrel. The female drops 4 young in the hol-
low limb of a tree, and these soon follow at
their mother's heels: they constitute a special
dainty for the Indians. It seems to be spread
all over British Guiana, because I at least found
it everywhere."
Echimys chrysurus chrysurus Zimmermann

Reise 111:779 (distribution).
Coendou prehensilis prehensilis Linnaeus {Cerco-
labes insidiosus not Kuhl)

Reise 111:779 (habits).
Dasyprocta leporina cayana Lacepede {Dasyprocta
agouti)

Reise 11:80 (food).

Reise 111:779 (distribution; predation; chase).
Myoprocta acouchy acouchy Erxleben (Dasyprocta
acuchy)

Reise 111:779 (distribution).
Agouti paca paca Linnaeus

Reise 11:491, 492 (chase).

Reise 111:780 (distribution; habitat; food).
Hydrochaeris hydrochaeris hydrochaeris Linnaeus
{Hydrochoerus capybara)

Reise 1:418: "Among the many domesticated
animals met with at the settlement [was a] full
grown water-haas. The creature was so tame
that it regularly stuck to the heels of the wom-
en. Although the river Nappi flowed past the
houses not fifty paces away, it never visited
its favorite element otherwise than in com-
pany with the women when they went to draw
water and even then only to drink."

Reise 11:29:

"I often found 6 to 8 of them together [along
the Essequibo River] forming a line in the
middle of which the young were to be seen.
But unless we killed it outright the wounded
animal every time escaped us by immediately
rushing into the water the neighborhood of
which it seldom left."

Reise 111:780 (distribution).

Cavia porcellus guianae (Cavia leucopyga Cabanis
not Brandt)
Reise 11:249:

"Six to eight living specimens would often
be brought to us but without our being able
to keep them alive. The Indians' statement
that they could never by any manner or means
be tamed, was confirmed. Had we ten or twelve
together, none would be alive after the third
day. They live in holes out of which they are
driven by pouring water in, and then easily
caught. ... Its silky fur is attached so deli-
cately to the skin that even the slightest touch
of the hand knocks it off" and leaves a bare
space."

PERISSODACTYLA

Tapirus terrestris terrestris Linnaeus

Reise 11:167 (chase), 169: ". . . since I could not
override the definite instructions given me not
to forward any skins of the larger mammals
to Berlin, I handed the hide over to the In-
dians to make sandals of. I prepared the skel-
eton for the Anatomical museum."

Reise 111:783 (distribution; habitat; forage; jag-
uar; flesh).

ARTIODACTYLA

Tayassu tajacu patira Sonnini (Dicotyles torqua-

tus)
Reise II: 100, 164 (description; habits; chase).
Reise 111:783 (distribution; characters).
Annals 5:401 (description; habits; chase).
Tayassu pecari pecari Link (Dicotyles labiatus)
Reise 11:98, 164 (habits; chase): "June and July

would seem to be the time when they drop

[give birth]."
Reise 111:784 (distribution; herd size).
Annals 5:402 (description; habits; chase).
Mazama americana americana Erxleben (Cervus

rufus)
Reise 11:57 (ectoparasites; habits).
Reise 111:784 (distribution).
Mazama gouazoubira nemorivaga F. Cuvier (Cer-

vus simplicicornis)
Reise 111:785 (distribution; species not seen).
Mazama sp.? [= M. gouazoubira'^] {Cervus humilis

not Bennett [= Pudu puda Molina])
Reise 11:58. "The fourth and smallest species is

known under the name of Wilibisiri {Cervus

humilisl): its home is also in the dense forest."
Reise 11:363. "In the evening the hunters brought

us [in camp at mouth of Aripai, upper Ru-

HELDIANA: ZOOLOGY

pununi, Kanaku Mts.] one of those pretty deer
which the Indians call Walibisiri. It is the
smallest species met with in Guiana, hardly
1 '/3 ft. high."
Odocoileus virginianus gymnotis Wiegmann {Cer-
vus savannarum Cabanis and Schomburgk;
Cervus mangivorus)

Reise 11:57. "The female must throw her young
in March or April because we found amongst
our lot four specimens very advanced in preg-
nancy; but as I have killed deer in a similar
condition during September or October, they
must either throw twice a year, or else they
are not usually limited to any fixed breeding
season. The deer is never present in the for-
ests."

Reise II: 1 57. "In cutting up the venison [secured
in savannas of Rio Cotinga, upper Rio Bran-
co] we found does well advanced in pregnan-
cy, which helped to strengthen my previously
expressed opinion that they either throw twice,
or else have no particular pairing season."

Reise 111:785 (description; distribution).

SIRENIA

Trichechus inunguis Natterer {Manatus australis
not Tilesius)
Reise 11:141, 156:

"The Peixe Boys, as the vaqueiros [cow-
hands] call the Sea cow {Manatus) had already
left the neighborhood of the Fazenda [Rio
Branco above Fort Sao Joaquim] several days
before, the water having commenced falling;
that during high water they usually travel up
as far as the mouth of the Maku which so
many had visited this rainy season, and that
ten had been harpooned. ... As soon as the
Takutu begins to fall a few feet, the Manatis
disappear and make their way back to below
the rapids of the Rio Branco. The search for
more abundant food probably brings them to
the Takutu where their favorite grasses, species
of Panicum and Paspelum, grow in abun-
dance."

[I have observed that when the river drops
a few feet and manatees cannot reach forage
growing on the edge of the embankments, they
move elsewhere, usually downstream.]

CETACEA

Inia geoffrensis Blainville {Delphinus amazonicus)

Reise 11:18: "They would not only raise their

pointed snouts out of the water but mostly a

large portion of their seven to eight foot long
body."
Reise 111:786 (Rio Tacutu, upper Rio Branco,
Brazil, near Guianan border).

IX. Alexander von Humboldt

(1769-1859) and

Aime Bonpland (1773-1858)

Alexander von Humboldt and Aime Bonpland
were rigorously trained scientists highly qualified
to survey the natural resources and native peoples
of a major part of tropical America. Their inves-
tigations and discoveries in the New World from
1799 to 1803 resulted in numerous publications
of primary importance.

Alexander von Humboldt was bom into a
wealthy and distinguished family and could pursue
his cultural interests without stint. His studies in
the arts and sciences prepared him to develop into
one of the most innovative and versatile scientific
investigators of his time, if not all time. He was
at once botanist, zoologist, anthropologist, ecol-
ogist, geologist, cartographer, biogeographer, phy-
sicist, chemist, astronomer, demographer, histo-
rian, mountain climber, poet, artist, and linguist.
He excelled in every field and gained recognition
and prominence in all. Humboldt raised geog-
raphy to a science. Knowledge of the fundamental
principles of climatology is due to him. Last but
not least of his many talents appears in Hum-
boldt's writings, which inspired a generation of
naturalist-travelers, including Charles Darwin.

The young Humboldt's greatest desire was for
an opportunity to apply his skills, knowledge, and
the scientific instrumentation accumulated at his
own expense to the exploration of little-known
lands. After disappointing starts on a number of
prospective expeditions, he visited Spain in June
1799 accompanied by the young French botanist
Aime Bonpland. While in Madrid he had the good
fortune to meet an influential friend who helped
him secure royal orders for travel throughout the
Spanish colonies in America to study natural re-
sources and collect samples of scientific interest.

Humboldt and Bonpland sailed for South
America on 5 June 1799 and landed 16 July 1799
at Cumana, capital of Nueva Andalucia (Vene-
zuela). The remainder of that year and part of the
following were spent in exploration of the coastal
region. Of prime interest, however, was the planned

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

c
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HELDIANA: ZOOLOGY

67°

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ik"^^'! LM

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Q-^ HUMBOLDT AND BONPLAND

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ROUTE IN AMAZONAS, VENEZUELA
1 ON THE Rfo ORINOCO - RIO NEGRO

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Fig. 1 2. Route of Humboldt and Bonpland in Amazonas, Venezuela, from the Rio Orinoco-Atabapo to the Rio
Guainia-Negro via p>ortage between the Rio Temi and Rio Pimichin and the Rio Casiquiare connecting the Negro
and Orinoco.

expedition up the Rio Orinoco for verification of
its reputed connection with the Amazonian Rio
Negro. The exploration began on 27 March 1800
with a three-day inspection of a western tributary,
the Rio Apure. The journey then continued up the
mainstream to the Spanish mission of San Fer-
nando de Atabapo near the confluence of the Rios
Atabapo and Guaviare with the Orinoco. At this
point, the travelers left the Orinoco and continued

up the Atabapo to the tributary Temi, which they
followed to the tiny mission of Yavita, arriving
on 1 May. On 10 May, after portage to the Rio
Pimichin, a tributary of the Guainia, they attained
San Carlos de Rio Negro at the mouth of the Rio
Casiquiare. The next day they headed up the Ca-
siquiare and, after 10 days' travel by water, reen-
tered the Orinoco on 21 May (figs. 1 1-12).
Having confirmed the connection between the

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

waters of the Orinoco and Amazon rivers, the ex-
plorers shipped 750 miles downstream to arrive
at Angostura (Ciudad Bolivar) in mid-June 1800.
After more work on the coast, Humboldt and Bon-
pland departed Venezuela on 24 November 1800
for Havana, Cuba. They remained there until 1 7
March 1 80 1 , then sailed for Colombia with land-
ings along the Rio Sinu on 25 March and Carta-
gena on 30 March. The journey thereafter was
devoted mainly to explorations of the Cordilleras
of Colombia and Ecuador, then through moun-
tains, deserts, and the upper Amazonia of Peru
south to Lima. The few mammals observed or
described during this part of the journey are men-
tioned in Humboldt's (1805-181 1) Recueil.

From Lima, Humboldt and Bonpland em-
barked on 24 December 1 802 for Guayaquil and
left 15 February 1803 for Mexico.

Humboldt's lively Personal Narrative evokes vi-
sions of Venezuelan life and landscapes from
coastal plains to the headwaters of the Rio Ori-
noco. The narrative is replete with descriptions of
geography, ecology, astronomical orientations,
widths, depths, and volumes of rivers, histories,
languages and customs of Indians, Catholic mis-
sions, missionaries, and the human interest trials
and tribulations of the travelers. Information on
mammals, however, is comparatively meager, but
some interesting bits can be quoted or paraphrased
from the Ross translation of the original French
(Humboldt, 1884).

Humboldt and Bonpland found manatees abun-
dant in the Rio Orinoco and tributaries Meta and
Apure, but absent above the cataracts of Mai-
pures. Some of the animals they caught were 1
to 12 feet long and weighed 500 to 800 pounds.
Humboldt's dissection of one (fig. 1 3) revealed "no
vestige of nails on the external surfaces of the fins
which were quite smooth, but little rudiments of
nails appear at the third phalanx when the skin of
the fins is taken off." The lungs, they observed,
consisted of "large cells resembling immense
swimming bladders; they [the lungs] are 3 feet long.
Filled with air they have a bulk of more than a
thousand cubic inches [Humboldt, Ross transla-
tion, 1884, vol. II, p. 169]." Its distinction from
T. manatus was not appreciated, however, until
1 883 when described by Natterer (in Pelzeln, 1 883).
There is also considerable doubt that a clawless
manatee does occur in the Rio Orinoco basin or
anywhere outside the Amazonian watershed.

Dolphins (Sotalia) were seen above and below
the great cataracts of the Orinoco and often swam
alongside the canoe. In the inundated forest of the

divide between the waters of the Orinoco and Ne-
gro, the travelers "were astonished by an extraor-
dinary noise. On beating the bushes a shoal of
toninas (fresh-water dolphins) four feet long sur-
rounded our boat. They fled across the forest,
throwing out those spouts of compressed air and
water. . . ."

Other Venezuelan mammals mentioned in the
narrative include the expected jaguar, otter, deer,
peccaries, capybara, and vampire bats.

Monkeys, however, absorbed more of Hum-
boldt's attention than other animals. He carried
with him a number of live simians captured in the
upper Rio Orinoco region for shipment to the Jar-
din des Plantes in Paris via the Antillean island
of Guadeloupe. The newly discovered bearded saki
{Chiropotes satanas chiropotes Humboldt; fig. 14)
died before transshipment, but its skin was saved
and arrived in Paris. The type specimen of red
howler, Simla urslna Humboldt (= Alouatta se-
nlculus arctoides Cabrera) survived the journey,
whereas the first-known douroucouli or night
monkey {Aotus trivlrgatus Humboldt; fig. 14) suc-
cumbed in Guadeloupe.

Humboldt often mentioned the ubiquitous,
highly visible howler or araguato {Alouatta seni-
culus). At one time he saw from the road below
troops of 30 to 40 individuals crossing through the
trees. In a carefully deployed experiment in Ara-
gua, he calculated the distance the howler's vo-
calization could be heard as 800 toises (6 ft 4.73
inches x 800 = 5,1 15 ft) or nearly 1 mile (5,280
ft).

Humboldt (Ross translation, 1884, vol. II, p.
453) recounts the Indian tale of bearded sakis
(Chiropotes) and uacaries (Cacajao) of the Orinoco
"placing themselves in a circle and, by striking the
shell [of the Brazil nut pericarp] with a stone, suc-
ceed in opening it so as to take out the triangular
nuts." Although Humboldt dismissed the story as
fabulous, he did believe that the monkeys cracked
the shell of the Bertholletia nut with their teeth to
obtain the meat which they devoured with gusto.

Belief in the existence of a hairy man of the
woods was practically universal. The missionary
Father Gili gravely related to Humboldt the tale
of a woman "in the town of San Carlos in the
Llanos of Venezuela who much praised the gentle
character and attentions of the man of the woods.
She is stated to have lived several years with one
in great domestic harmony, and only requested
some hunters to take her back because she and the
children (a little hairy also) were weary of living
so far from the church and the sacraments." Hum-

FIELDIANA: ZOOLOGY

IkbU.

^ %

Fig. 1 3. The Orinoco clawless manatee, supposedly Trichechus inunguis Natterer left, lateral (1) and ventral (2)
views; right, head from above (1), mouth, upper inner view (2), mouth, lower inner view (3), mouth, side view (4),
and trunk, sagittal section (5); original illustrations by Humboldt; from Humboldt (1838).

boldt resented that he and Bonpland "were every-
where blamed, in the most cultivated class of so-
ciety, for being the only persons to doubt the reality
of the great anthropomorphic monkey of Ameri-
ca."

Humboldt's Recueil d 'Observations de Zoologie
et d 'Anatomic Comparee, a collection of memoires
published as a volume in 181 1-1812, deals with
many species of invertebrates and vertebrates, but
a large share of the text is about monkeys. One
memoir with excellent illustrations by Humboldt
is on the comparative anatomy of the hyoid bone
and larynx of the cotton-top tamarin (Saguinus
oedipus oedipus Linnaeus; fig. 1 4), and that of the
red howler {Alouatta seniculus seniculus Lin-
naeus), the Colombian squirrel {Sciurus granaten-
sis granatensis Humboldt; fig. 14), birds, and croc-

odiles, all from the Rio Magdalena region. Another
memoir on the carnivores includes descriptions
of Gulo quitensis (= Conepatus chinga quitensis
Humboldt) from Quito, Ecuador, Mustela sinuen-
sis (= Eira barbara sinuensis Humboldt), from the
Rio Sinu, Colombia, and a discourse on other
mustelids and the kinkajou {Potos Jlavus Schre-
ber). The memoir on monkeys of the upper Rio
Orinoco and connecting Rios Casiquiare and Ne-
gro includes the original descriptions o^ Aotus tri-
virgatus, Chiropotes satanas chiropotes, Cacajao
melanocephalus, Callicebus torquatus lugens, La-
gothrix lagothricha, and Cebus albifrons. A chap-
ter on the monkeys of Colombia and the upper
Amazonian region includes the description of a
representative each of Cebus capucinus Linnaeus
from the Rio Sinu, A teles belzebuth marginatus E.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

V M. PSIT TAl- 1 S AR Al' RAN A.

N° Ml. SI n HIS (.RANATF.NSIS.

_;?•«•

V. vm. SIMIA OKDIHl S.

-. •»«

Fig. 1 4. Monkeys and anatomical dissections from Humboldt (1811): upper left, Simia melanocephala Humboldt
(= Cacajao melanocephalus), holotype; lower left, two views of Simia trivirgata Humboldt (= Aotus trivirgaius),
holotype; upper right, Simia satanas Hoffmannsegg (= Chiropotes satanas satanas), lectotype; lower right, throat
cartilages of Psiltacus araurana Linnaeus (= Ara araurana), Sciurus granatensis Humboldt, and Simia oedipus
Linnaeus (= Saguinus oedipus oedipus).

HELDIANA: ZOOLOGY

Table 8. New World monkeys (Platyrrhini) recorded by Humboldt (1812); the arrangement is phylogenetic.

Current name

Humboldt synonym

Figure

Callitrichidae

Callithrix jacchus jacchus Linnaeus, 1758
Callithrix jacchus penicillata E. Geoffroy, 1812
Callithrix jacchus geoffroyi Humboldt, 1812
Callithrix jacchus aurita E. Geoffroy, 1812
Callithrix humeralifer humeralifer E. Geoffroy,

1812
Callithrix argentata melanura E. Geoffroy, 1812
Callithrix argentata argentata Linnaeus, 1771
Saguinus fuscicollis fuscus Lesson, 1840
Saguinus labiatus labiatus E. Geoffroy, 1812
Saguinus midas niger E. Geoffroy, 1 803
Saguinus midas midas Linnaeus, 1758
Saguinus oedipus oedipus Linnaeus, 1758
Leontopithecus rosalia rosalia Linnaeus, 1 766

Cebidae

Saimiri sciureus cassiquiarensis Lesson, 1 840

Callicebus moloch moloch Hoffmannsegg, 1 808

Callicebus torquatus lugens Humboldt, 1811

Callicebus torquatus torquatus Hoffmannsegg, 1 808

Callicebus personatus personatus E. Geoffroy, 1812

Actus trivirgatus Humboldt, 1811

Actus azarae azarae Humboldt, 1811

Pithecia mcnachus monachus E. Geoffroy, 1812

Pithecia pithecia pithecia Linnaeus, 1 766

Chiropotes satanas satanas Hoffmannsegg, 1 808
Cacajac melanocephalus Humboldt, 1811
Alouatta caraya Humboldt, 1812
Alouatta seniculus arctoidea Cabrera, 1 940
Alouatta seniculus straminea Humboldt, 1812
Cebus capucinus capucinus Linnaeus, 1758
Cebus nigrivittatus nigrivittatus Wagner, 1 848

Cebus apella apella Linnaeus, 1 758

Cebus apella xanthosternos Wied-Neuwied, 1 820
Cebus apella nigritus Goldfuss, 1810

Lagcthrix lagcthricha lagothricha Humboldt, 1812
Lagothrix lagcthricha cana E. Geoffroy, 1812
Lagcthrix flavicauda Humboldt, 1811
Atetes paniscus chamek Humboldt, 1812
Ateles paniscus paniscus Linnaeus, 1 766
Ateles belzebuth belzebuth E. Geoffroy, 1 806
Ateles belzebuth marginatus E. Geoffroy, 1809

Brachyteles arachnoides E. Geoffroy, 1 806

Jacchus leucccephalus Geoffroy, 1812

Simla leonina Humboldt, 1805, not Shaw, 1800
Simla Ursula Hoffmannsegg, 1808

Not Simla sciurea Linnaeus

Simla amicta Humboldt, 1811

Pithecia rufiventer E. Geoffroy, 1812; Simla leuco-
cephala E. Geoffroy, 1812

Simla ursina Humboldt, 1805, not Bechstein, 1800

Simla hypoleuca Humboldt, 1811

Simla capucina Humboldt, 1812, not Linnaeus,

1758
Cebus barbatus Humboldt, 1812, attributed to E.

Geoffroy
Simla variegata Humboldt, 1812, not Kerr, 1 792
Simla cirrifera Humboldt, 1812; Cebus niger E.

Geoffroy, 1812

Simla chuva Humboldt, 181 1, p. 340; 1812, p. 362,
footnote 2

Geoffroy from lower Amazonia, Alouatta senicu-
lus Linnaeus from the Rio Magdalena, and La-
gothrix flavicauda Humboldt from northern Peru.
In an addendum, Humboldt listed all platyrrhine
monkeys known to 1812. They are arranged in
Table 8 by current scientific names with Hum-
boldt's synonyms.

X. PARAGUAY

The Paraguayan province, claimed by Spain,
was first visited in 1526 by Sebastian Cabot and
then explored by Cabeza Alvarez Nunez de Vaca
in 1541. For the next two centuries, waves of mis-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Fig. 1 5. Map of Azara's Paraguay and adjacent parts of Brazil and Argentina; from Azara (1809).

FIELDIANA: ZOOLOGY

sionaries and colonists penetrated to the remotest
comers of the province in quest of climates or
environments that resembled or could be trans-
formed into the familiar ones of Spain.

The monumental Histoire du Paraguay by the
Jesuit missionary Pierre Francois Xavier de Char-
levoix (1682-1761), published in 1757, describes
the land that extended from the Atlantic to the
eastern base of the Andes between latitudes 1 5°
and 35° in the drainage basin of the Rio Parana-
Paraguay. It relates the history of the province
from the time of the conquest, describes native
customs, conversions to Christianity, and estab-
lishment of missions. The little of natural history
in the text adds nothing about wild mammals not
already recorded by others. Two decades later Fe-
lix de Azara wrote the most complete natural his-
tory account of the mammalian fauna of Paraguay
for its time and ever since.

Felix de Azara (1746-1811)

The Spaniard Don Felix de Azara (1 746-1 811),
an army engineer, was commissioned in 1781 to
assist in defining the boundaries between Spanish
and Portuguese territories. Unmapped territories
between Brazil and Paraguay were assigned to
Azara, but the Portuguese showed no interest in
their share of the work. With time on his hands
and a disposition toward the natural sciences, Azara
devoted nearly the full 20 years, from 1781 to
1800, of his American residence to the study of
geography, Guarani Indians, and the birds and
mammals of Paraguay and northeastern Argen-
tina between 24° and 36°S and about 54°3r to
56°W (or 60°W of Greenwich) (fig. 15).

With no schooling in the natural sciences and
no books for reference or guidance, Azara de-
pended on his own resources. They proved ade-
quate. Azara recorded his observations with care,
precision, meticulous attention to detail, and rig-
orous exclusion of speculation and fantasy. His
anatomical descriptions, measurements, and ac-
counts of behavior were based on animals ob-
served in the wild or in captivity, usually in his
own home or garden. Useful information received
from others was credited to the informants. Pop-
ular beliefs and hearsay were labeled as such.
Without other sources of information, Azara used
the Guarani names for most of the amimals he
described and Spanish epithets for the remainder.

The manuscript of the mammals or quadrupe-
dos of Paraguay contained accounts of 66 species.

Shortly after its completion, the author received
a shipment of several volumes of a Spanish trans-
lation of Buffon's Histoire Naturelle. Not surpris-
ingly, Azara found in them much with which to
disagree, but some of his adverse criticism was
unfair. Azara knew Paraguayan mammals better
than anyone else, but only a minority of the species
were the same as the Neotropical species described
in the Histoire Naturelle, and those that were the
same did not always behave in the same way at
different times or in different places.

Azara sent a copy of the manuscript of the quad-
rupedos to his brother, Jose Nicolas, then Spanish
ambassador to Paris, who arranged for publication
in that city after translation into French by M.-L.-E.
Moreau de Saint-Mery. A year after his return to
Spain in 1801, Azara secured publication in Ma-
drid of the original Spanish manuscript with
emendations and addition of 1 1 species, for a total
of 77.

Azara may not have been aware that as many
as 62 of the 77 species he described were still un-
known to science. His clear and precise charac-
terization of each of the species or subspecies,
however, provided contemporary cataloguers and
systematists with the bases for the descriptions of
50 new species, many with their vernacular ap-
pellations in the binomial. Actual specimens served
as types for the remaining 1 2 species.

The mammals described by Azara are listed be-
low, with the scientific name of each given first
followed by its local name(s). The page references
are to Azara's works in French (Essais, 1801),
Spanish (Apuntamientos, 1802), and the Voyage
(1809). The last is a French translation in four
volumes of Azara's travels in Paraguay with sep-
arate atlas, but only the first volume and atlas
contain information on mammals.

Tapirus terrestris Linnaeus, 1758
Mborebi, Essais I, p. 1; Mborebi, Apunt., I, p.
1; Mborebi ou tapir. Voyage, p. 246.
Tayassu G. Fischer, 1814
Coure ou Tayazou, Essais, I, p. 18; Cures o
Tayaziis, Apunt., I, p. 14; Cure ou tayazii.
Voyage, p. 248.
Tayassu pecari albirostris Illiger, 1815
Tagnicati, Essais, I, pp. 2 1 , 25; Taiiicati, Apunt.,
p. 19; Tanicati, Voyage, p. 249.
Bibliographic type of the subspecies.
Tayassu tajacu Linnaeus, 1758
Taytetou, Essais, I, pp. 21,31; Taytetvi, Apunt.,
I, p. 23; Taytetu, Voyage, p. 249.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

CERVIDAE

Gazou, Essais. I, p. 43; Venados, Apunt., I, p.
29; Guazu, Voyage, p. 250.
Blastocenis dichotomus Illiger, 1815
Gouazoupoucou, Essais, I, p. 70; Guazu-pucu,
Apunt., I, p. 33; guazu-pucu. Voyage, p. 250.
Bibliographic type of the species.
Blastoceros bezoarticus leucogaster Goldfuss, 1817
Gouazouti, Essais, I, p. 77; Giiazu-ti, Apunt., I,
p. 41; Guazu-ti, Voyage, p. 251.
Bibliographic type of the subspecies.
Mazama americana gouazoupita Fischer, 1814
Gouazoupita, Essais, I, p. 82; GUazu-pita,
Apunt., I, p. 5 1 ; Guazu-pita, Voyage, p. 252.
Bibliographic type of the subspecies.
Mazama gouazoubira gouazoubira Fischer, 1814
Gouazoubira, Essais, I, p. 86; Giiazu-bira,
Apunt., I, p. 57; Guazu-bira, Voyage, p. 252.
Bibliographic type of the species.

DIDELPHIDAE

Micoures, Essais, I, p. 240; Fecundos, Apunt.,
I, p. 204; Feconds, Voyage, p. 281.
Didelphis albiventris Lund, 1 840
Micoure premier, ou micoure propement dit,
Essais, I, p. 244; Micure, Apunt., I, p. 209;
Micure, Voyage, p. 283.
Caluromys lanatus Olfers, 1818
Micoure second, ou Micoure laineux, Essais, I,
p. 175; Lanoso, Apunt., I, p. 221; Lanoso,
Voyage, p. 287.
Holotype in alcohol, no. 528, Museo de Cien-
cias Naturales, Madrid, captured 22 July
1789, by Felix d'Azara (Cabrera, 1916,
Bol. Real Soc. espanola Hist. Nat., 16,

p. 1).

Lutreolina crassicaudata Desmarest, 1 804

Micoure troisieme, ou micoure a queue grosse,
Essais, I, p. 284; Coligrueso, Apunt., I, p.
229; Coligrueso, Voyage, p. 290.
Bibliographic type of the species.
Marmosa pusilla Desmarest, 1 804

Micoure quatrieme, ou micoure a queue longue,
Essais, I, p. 290; Colilargo, Apunt., I, p.
251; Colilargo, Voyage, p. 291.
Bibliographic type of Marmosa macrura Ol-
fers, 1818 (= M. pusilla Desmarest).
Micoure sixieme, ou micoure nain, Essais, I, p.
304; Enano, Apunt., I, p. 262; Enano, Voy-
age, p. 284.
Bibliographic type of Marmosa pusilla Des-
marest, 1804.
Monodelphis brevicaudis Olfers, 1818

Micoure cinquieme, ou micoure a queue courte,
Essais, I, p. 295; Colicorto, Apunt., I, p.
258; Colicorto, Voyage, p. 293.
Bibliographic type of the species.

MYRMECOPHAGIDAE

Hormigueros, Apunt., I, p. 61.
Myrmecophaga tridactyla Linnaeus, 1758

Gnouroumi, ou Yoquoui, Essais, \, p. 89; Nu-
rumi o Yoqui, Apunt., I, p. 66; Nurumi ou
tamandua. Voyage, pp. 253, 255.
Tamandua tetradactyla Linnaeus, 1758 (fig. 16)

Cagouare, Essais, \, p. 103; Cagiiare, Apunt., \,
p. 74; Cagiiare, Voyage, pp. 253, 256; Atlas,
pi. VII (tamandua noir), pi. VIII (Cag-
uouare).

FELIDAE

Gatos, Apunt., I, p. 85.
Felis onca Linnaeus, 1758
Yagouarete, £'55a/5, 1, p. 1 14; Yaguarete,^;?Mm.,
I, p. 91; Yaguarete, Voyage, p. 258; Atlas,
pi. IX.
Yagiiarete negro, Apunt., I, p. 114; Yaguarete
noir. Voyage, p. 267.
Felis concolor Linnaeus, 1771
Gouazouara, Essais, I, p. 133; Giiazuara, Apunt.,
I, p. 120; Guazuara, Voyage, p. 268.
Felis geoffroyi D'Orbigny and Gervais, 1 844
Mbaracaya, ^/7Mn/., I, p. 147; Baracaya, Voyage,
p. 271.
Note: Said not to exist in Paraguay.
Felis species?

Negro, Apunt., I, p. 154; Chat noir. Voyage, p.
273.
Felis pardalis Linnaeus, 1758
Chibigouazou, Essais, I, p. 152; Chibi-giiazii,
Apunt., I, p. 132; Chibi-guazu, Voyage, p.
269.
Herpailurus yagouaroundi eyra Fischer, 1814 (fig.
16)
Yagouaroundi, Essais, I, p. 171; Yaguarundi,
Apunt., I, p. 156; Yaguarundi, Voyage, p.
273, Atlas, pi. X (Yagouarondi, black
phase); Eyra, Essais, I, p. 177; Eyra, Apunt.,
I, p. 159; Eyra, Voyage, p. 274 (red phase).
Bibliographic type of the subspecies.
Felis colocolo pajeros Desmarest, 1816
Chat pampa, Essais, I, p. 1 79; Pajero, Apunt.,
I, p. 160; Pajero, Voyage, p. 274.
Bibliographic type of the species.
Note: Said not to exist in Paraguay.

FIELDIANA: ZOOLOGY

.(■ ').i.'..M.ir<imll /' / /./,,• ).,.„..„.,■'. I

l.r (' .1M(. 11.11 .■ /' ' oil 'r.llll.lllllll.l I'm.II I/^/ ,l.;y; /.„.,„„./«., < .,,

Fig. 1 6. Two of Azara's Paraguayan animals: top, le yagouarondi (= Eira yagouarondi eyra Fischer); bottom, le
cagouare or cagiiare (= Tamandua tetradactyla Linnaeus); from Azara ( 1 809).

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

MUSTELIDAE

Furets, Essais, I, p. 185; Hurones, Apunt.. I, p.
167.
Galictis cujafurax Thomas, 1907
Petit furet, Essais, I, p. 190; Huron menor,
Apunt., I, p. 182; Huron minor. Voyage, p.
276.
Eira barbara Linnaeus, 1758
Grand furet, Essais, I, p. 197; Huron mayor,
Apunt., I, p. 172; Huron major, Voyage, p.
275, Atlas, pi. XI.
Conepatus chinga suffocans Illiger, 1815
Yagoure, Essais, I, p. 211; Yagiiare, Apunt.. I,
p. 187; Yaguare, Voyage, p. 277.
Bibliographic type of the subspecies.
Pteronura brasiliensis paranensis Rengger, 1830
Loutre, Essais. I, p. 348; Nutria, Apunt., I, p.
304; Loutre, Voyage, p. 304.

CANIDAE

ZoTTO, Apunt., I, p. 264; Renard, Voyage, p. 295.
Chrysocyon brachyurus Illiger, 1815
Agouara-gouazou, Essais, I, p. 307; Agiiara-
guazvj, Apunt., I, p. 266; Aguara-guazii,
Voyage, p. 296.
Bibliographic type of the species.
Dusicyon gymnocercus Fischer, 1814
Agouarachay, Essais, I, p. 317; Aguarachai,
Apunt., I, p. 271; Aguarachay, Voyage, p.
298, Atlas, pi. XII.
Bibliographic type of the species.

PROCYONIDAE

Procyon cancrivorus nigripes Mivart, 1886
Agouarapope, Essais, I, p. 324; Pope, Apunt.. I,
p. 278; Pope, Voyage, p. 299.
Nasua nasua spadicea Olfers, 1818
Couati, Essais, I, p. 334; Cuati, Apunt.. I, p.
293; Cuati, Voyage, p. 301.
Bibliographic type of the subspecies.

LEPORIDAE

Sylvilagus brasiliensis paraguensis Thomas, 1 90 1
Tapiti, Essais. II, p. 57; tapiti, Apunt.. II, p. 32;
Tapity, Voyage, p. 313.

RODENTIA

Roedores, Apunt., II, p. 68.
Myocastor coypus bonariensis Commerson, 1805
Quouiya, Essais, II, p. 5; Quiya, Apunt., II, p.
1; Quiya, Voyage, p. 308.

Hydrochaeris hydrochaeris dabbenei Rovereto,
1913
Capiygoua, Essais, II, p. 12; Capibara, Apunt..
II, p. 8; Capibara, Voyage, p. 309.
Agouti paca Linnaeus, 1758

Pay, Essais, II, p. 20; Pai, Apunt., II, p. 14; Pay,
Voyage, p. 310.
Dasyprocta azarae paraguayensis Liais, 1872
Acouti, Essais. II, p. 26; Acuti, Apunt.. II, p. 21;
Acuty, Voyage, p. 312.
Vizcacia maximus Desmarest, 1817
Vizcache, Essais. II, p. 41; Vizcacha, Apunt.. II,
p. 45; Vizcacha, Voyage, p. 316.
Bibliographic type of the species.
Note: Vizcacia Schinz, 1 824 (Naturg. Abbild.
Saugeth., p. 243) antedates Viscaccia
Schinz, 1825 and Lagostomus Brookes,
1825.
Dolichotis patagonum Zimmermann, 1780
Lievre pampa, Essais. II, p. 5 1 ; Liebre patagona,
Apunt., I, p. 51; Lievre patagon, Voyage, p.
318.
Note: Said not to exist in Paraguay.
Cavia aperea hypoleuca Cabrera, 1953
Aperea, Essais, II, p. 65; Aperea, Apunt., II, p.
37; Aperea, Voyage, p. 314.
Euryzygomatomys spinosus Fischer, 1814

Rat premier, ou rat epineus, Essais, II, p. 73;
Espinoso, Apunt., II, p. 76; Epineuse, Voy-
age, p. 326, Atlas, pi. XIII.
Bibliographic type of the species.
Oryzomys megacephalus Fischer, 1814
Rat second, ou rat a grosse tete, Essais, II, p.
82; Cola igual al cuerpo, Apunt., II, p. 87;
Cola igual al cuerpo. Voyage, p. 330.
Bibliographic type of the species.
Note: Antedates Oryzomys capito Olfers.
Oryzomys angouya Fischer, 1814
Rat troisieme, ou rat angouya, Essais, II, p. 86;
Anguya, Apunt.. II, p. 89; Anguya, Voyage,
p. 331.
Bibliographic type of the species.
Note: Antedates Oryzomys buccinatus Olfers.
Reithrodon auritus Fischer, 1814

Rat quatrieme, ou rat oreillard, Essais, II, p. 9 1 ;
Orejon, Apunt., II, p. 83; Orejon, Voyage,
p. 329.
Bibliographic type of the species.
Oxymycterus rufus Fischer, 1814
Rat cinquieme, ou rat roux, Essais, II, p. 94;
Hocicudo,/ipM/7/., II, p. 80; Hocicudo, Voy-
age, p. 328.
Bibliographic type of the species.

HELDIANA: ZOOLOGY

Oryzomys nigripes Olfers, 1818
Rat sixieme, ou rat a tarse noir, Essais, II, p.
98; Colilargo, Apunt.. II, p. 91; Colilargo,
Voyage, p. 331.
Bibliographic type of the species.
Calomys laucha Olfers, 1818
Rat septieme, ou rat laucha, Essais, II, p. 102;
Laucha, Apunt., II, p. 96; Laucha, Voyage,
p. 333.
Bibliographic type of the species.
Coendou insidiosus Olfers, 1818
Couiy, Essais, II, p. 105; Cuiy, Apunt., II, p. 55;
Cuiy, Voyage, p. 320.
Bibliographic type of the species.
Akodon colibreve Brants, 1827

Colibreve, /i/7M/i/., II, p. 86; Colibreve, Voyage,
p. 329.
Bibliographic type of the species.
Note: Akodon obscurus Waterhouse, 1837, is
probably a junior synonym, but see Lang-
guth(1978).
Ctenomys tucotuco Brants, 1827
Tucutuco, Apunt., II, p. 89; Tucutuco, Voyage,
p. 324.
Bibliographic type of the species.
Akodon agreste Brants, 1827

Agreste, Apunt., II, p. 94; Agreste, Voyage, p.
332.
Bibliographic type of the species.
Note: Antedates Akodon azarae Fischer.
"Musdubius", Fischer, 1829 [= ?]
Blanco debaxo, Apunt., II, p. 97; Blanco-de-
baxo, Voyage, p. 333.
Bibliographic type of the species.

DASYPODIDAE

Tatous, Essais, II, p. 122; Tatiis, Apunt., II, p.
99; Tatous, Voyage, p. 334.
Priodontes maximus giganteus, E. Geoffroy, 1 803
Tatou premier, ou grand tatou, Essais, II, p.
132; Maximo, Apunt., II, p. 110; Grand
tatou ou geant. Voyage, p. 336.
Bibliographic type of the subspecies.
Euphractus sexcinctusjlavimanus Desmarest, 1804
Tatou second, tatou poyou, ou tatou a main
jaune, Essais, II, p. 142; Poyu, Apunt., II,
p. 118; Tatu-poyu, Voyage, p. 338.
Bibliographic type of the subspecies.
Cabassous tatouay Desmarest, 1 804
Tatou troisieme, ou tatou tatouay, Essais, II, p.
155; Tatuai, Apunt., II, p. 131; Tatuai, Voy-
age, p. 341.
Bibliographic type of the species.

Chaetophractus villosus Desmarest, 1 804
Tatou quatrieme, ou tatou velu, Essais, II, p.
164; Peludo, Apunt., II, p. 140; Tatou velu,
Voyage, p. 343.
Bibliographic type of the species.
Dasypus novemcinctus niger Desmarest, 1 804
Tatou cinqui^me, ou tatou noir, Essais, II, p.
175; Negro, Apunt., II, p. 144; Tatou noir,
Voyage, p. 346.
Bibliographic type of the subspecies.
Dasypus hybridus Desmarest, 1 804
Tatou sixieme, ou tatou mulct, Essais, II, p. 1 86;
Mulita, Apunt., II, p. 156; Tatou mulita,
Voyage, p. 348.
Bibliographic type of the species.
Zaedyus pichiy Desmarest, 1 804
Tatou septieme, ou tatou pichiy, Essais, II, p.
192; Pichiy, Apunt., II, p. 158; Tatu-pichy,
Voyage, p. 345.
Bibliographic type of the species.
Tolypeutes matacus Desmarest, 1 804
Tatou huitieme, ou tatou mataco, Essais, II, p.
197; Mataco, Apunt., II, p. 161; tatou-ma-
taco. Voyage, p. 350.
Bibliographic type of the species.

PLATYRRHINI

Singes, Essais, II, p. 206; Micos, Apunt., II, p.
167; Singes, Voyage, p. 351.
Alouatta caraya Humboldt, 1812

Caraya, Essais, II, p. 208; Caraya, Apunt., II, p.
169; Caraya, Voyage, p. 351.
Bibliographic type of the species.
Cebus apella cay, Illiger, 1815
Cay, Essais, II, p. 230; Cay, Apunt., II, p. 182;
Cay, Voyage, p. 354.
Bibliographic type of the subspecies.
Aotus azarae azarae Humboldt, 1812

Miriquouina, Essais, II, p. 243; Miriquina,
Apunt., II, p. 195; Miriquina, Voyage, p.
356.
Bibliographic type of the species.
Callithrix jacchus penicillatus E. Geoffroy, 1812
Titi, Essais. II, p. 254; Titi, Apunt., II, p. 200;
Titi, Voyage, p. 359. "N'est pas du Para-
guay, mais du Bresil." [Description is of a
captive pair seen in the province of Buenos
Aires.]

MICROCHIROPTERA

Chauve-souris, Essais, II, p. 264; Murcielagos,
Apunt., II, p. 288; chauve-souris. Voyage,
p. 382.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Artibem lituratus Olfers, 1818

Chauve-souris premiere, ou chauve-souris ob-
scure et rayee, Essais, II, p. 269; Obscuro
listado, Apunt., II, p. 291.
Bibliographic type of the species.
Vampyrops lineatus E. Geoffroy, 1810
Chauve-souris seconde, ou chauve-souris brune
et rayee, Essais, II, p. 271; Pardo listado,
Apunt., p. 292
Bibliographic type of the species.
Desmodus rotundus E. Geoffroy, 1810
Chauve-souris troisieme, ou chauve-souris brune,
Essais, II, p. 273; Mordedor, Apunt., II, p.
293.
Bibliographic type of the species.
[Azara was first to distinguish true vampires
from other bats, particularly Vampyrum
spectrum.]
Sturnira lilium E. Geoffroy, 1810

Chauve-souris quatrieme, ou chauve-souris
brun-rougeatre, Essais, II, p. 277; Pardo
roxizo, Apunt., II, p. 299.
Bibliographic type of the species.
Noctilio leporinus rufescens Olfers, 1818

Chauve-souris cinquieme, ou chauve-souris
rougeatre, Essais, II, p. 280; Roxizo, Apunt.,
II, p. 301.
Bibliographic type of the subspecies.
Molossus ater castaneus E. Geoffroy, 1805
Chauve-souris sixieme, ou chauve-souris cha-
taine, Essais, II, p. 282; Castano, Apunt.,
II, p. 302.
Bibliographic type of the subspecies.
Lasiurus cinereus villosissimus E. Geoffroy, 1806
Chauve-souris septieme, ou chauve-souris brun-
blanchatre, Essais, II, p. 284; Pardo blan-
quizco, Apunt., II, p. 303.
Bibliographic type of the subspecies.
Histiotus velatus I. Geoffroy, 1824

Orejon, Apunt., II, p. 304.
Tadarida laticaudata E. Geoffroy, 1 805
Chauve-souris huitieme, ou chauve-souris ob-
scure, Essais, II, p. 286; Obscuro, Apunt.,
II, p. 305.
Bibliographic type of the species.
Molossus molossus Pallas, 1 766

Chauve-souris neuvieme, ou petit chauve-sou-
ris obscure, Essais, II, p. 288; Obscuro me-
nor, Apunt., II, p. 306.
Molossus crassicaudatus E. Geoffroy, 1 805
Chauve-souris dixieme, ou chauve-souris bnin-
cannelle, Essais, II, p. 290; Pardo acane-
lado, Apunt., II, p. 307.
Bibliographic type of the species.

Myotis ruber E. Geoffroy, 1 806

Chauve-souris onzieme, ou chauve-souris can-
nelle, Essais, II, p. 292; Acanelado, Apunt.,
II, p. 308.
Bibliographic typ>e of the species.
Myotis albescens E. Geoffroy, 1 806

Chauve-souris douzieme, ou chauve-souris
brun-obscur, Essais, II, p. 294; Pardo ob-
scuro, Apunt., II, p. 309.
Bibliographic type of the species.

Johann Rudolph Rengger (1795-1832)

Azara was followed by Johann Rudolph Reng-
ger, a Swiss pharmacist and naturalist, who arrived
in Paraguay in 1819 and devoted himself to the
study of its mammals. His six-year study culmi-
nated in the Naturgeschichte der Saeugethiere von
Paraguay, published 1830. A total of 59 species
was described, including four as new of which only
Calomys callosus and Proechimys longicaudatus
survived revisions. Azara distinguished 77 species,
or 1 8 more, but several are not strictly Paraguayan.
Among the Paraguayan forms missed by Rengger
but recognized by Azara are the murine opossum
(Marmosa), hairy armadillo (Chaetophractus),
three-banded armadillo (Tolypeutes), skunk (Co-
nepatus), tucotuco (Ctenomys), four cricetine ro-
dents, and two bats. Well over 100 species are
presently known from Paraguay.

No doubt Azara set standards for the high qual-
ity and accuracy of Rengger's descriptions and be-
havioral accounts. The wealth of information in
the Naturgeschichte has hardly been tapped by
modem mammalogists.

XI. Chile

Giovanni Ignazio Molina (1737-1829)

Knowledge of Chilean land mammals as a re-
gional fauna begins with publication of the Saggio
in 1782 by the Jesuit priest Don Giovanni (Juan)
Ignazio Molina, who lived in Chile the first 30
years of his life. Expulsion of the Jesuits from the
country obliged Molina to emigrate in 1768 and
settle in his ancestral Italy. What Molina knew
about Chilean mammals he learned before 1768;
much of what he wrote about them thereafter suf-
fered from a decayed memory.

Molina was a naturalist in the broadest sense

HELDIANA: ZOOLOGY

and was familiar with the Systemce of Linnaeus.
He was not, however, particularly dedicated to any
one branch of science, and his descriptions of the
Chilean mammals are, for the most part, vague,
inaccurate, and sometimes composite. A few of
his subjects were fanciful, and none of the re-
mainder were closely examined. Nevertheless, by
dint of elimination and stretches of the imagina-
tion, modem mammalogists have come to agree-
ment on the application of most of the Linnaean
names proposed by Molina for the likeliest species
he may have had in mind.

Thirty kinds of mammals were described in the
Saggio. According to Osgood (1943, p. 15), five
of them are unidentifiable, four (armadillos) are
extraterritorial, two are but one and the same, and
one is duplicated. The 14 still valid, with names
dating from Molina, 1782, are Lutra felina, My-
ocastor coypus, Conepatus chinga, Galictis cuja,
Dusicyon culpaeus, Felis guigna, Felis colocolo,
Felis concolor puma, Spalacopus cyanus, Octodon
degus, Vizcacia vizcacia, Pudu puda, Vicugna vi-
cugna, and Hippocamelus bisulcus. Remaining
species, notably the larger mammals, recorded by
Molina were well known to early voyagers, chron-
iclers, and naturalists and had already received
Linnaean names.

first volume (1847) of eight on zoology contains
virtually all Chilean mammals known at the time.
Fifty-four species are described, with accounts of
habits, habitat, and geographic distribution of each.
For the most part. Gay worked from actual spec-
imens brought to him by natives or observed by
him on his travels throughout the country. On his
return to France, Gay included in his studies the
Chilean material preserved in the Paris Natural
History Museum.

The species recorded by Gay include Marsupi-
alia, 2 (4% of the total); Chiroptera, 7 (13%); Gar-
ni vora, 12 (22%); Pinnipedia, 6(11%); Rodentia,
23 (43%; myomorphs, 24%, caviomorphs, 18%);
Artiodactyla, 3 (5%). Among the 30 species re-
corded by Molina, only 3 or 10% are rodents. Of
the 20 Chilean species collected by Darwin, 12 or
60% are rodents. In this volume Patterson and
Feigl recognize 93 living Chilean sF)ecies, of which
53 or 57% are rodents (33% myomorphs, 24%
caviomorphs), and 1 or 1 1 % are bats.

XII. Peru

Johann Jacob von Tschudi (1818-1889)

Eduard Friedrich Poeppig (1798-1868)

The German naturalist Eduard Poeppig is known
for his Reise in Chile, Peru, and on the Rio Ama-
zonas during the years 1827-1832. The account
of his travels, in two volumes, was published
1835-1836. The Chilean mammals recorded in-
clude seals, sea lions, and elephant seals, the degu,
Spalacopus cyanus Molina (Psammomys nocti-
vagus Poeppig, a synonym), the coypu, and a small
canid, probably Dusicyon griseus Gray. In Antuco,
Province of Bio Bio, he encountered the pudu,
huemul, and two species of bats, one described as
Nyticyus varius (= Lasiurus borealis bonariensis
Lesson & Gamot, 1827), the other as Nycticyus
macrotus (currently Histiotus macrotis Poeppig,
1835).

Claudio Gay (1800-1873)

Between the years 1 844 and 1871, Claudio Gay,
French naturalist and longtime resident of Chile,
produced 25 volumes, including two of plates, on
the history, geography, and biota of Chile. The

The Swiss biologist Johann von Tschudi was
bom in the town of Glarus and studied the sciences
at Swiss, French, and German universities. In-
spired by the accounts of the travels of Humboldt
and Darwin in South America, Tschudi sailed on
27 February 1838 from Le Havre for Peru. The
first landing on the continent was made 5 June
1 838 on the Chilean island of Chiloe. After a delay
of about three weeks and many observations of
the natural history of the island, von Tschudi
reembarked for Callao, Pern, with short stopovers
in Valdivia and Juan Femandez.

From August 1838 through most of 1843, von
Tschudi traveled over much of Peru. Of particular
interest to him were the higher vertebrates and the
physical factors controlling their geographic dis-
tribution. He distinguished faunal zones based on
mling ecological features. The major zones were
Pacific coast, Andean altitudinal zones of westem
and eastem slopes, and the tropical Amazonian
selva. Apparently, no one had preceded von
Tschudi in the recognition of definable biogeo-
graphic areas in the New World.

The narrative of von Tschudi's travels in Pern
was published in 1846 in German, followed in
1847 by Thomasina Ross's English translation.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

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HELDIANA: ZOOLOGY

The scientific accounts of the mammals are found
in a preliminary report (1844a) and first part of
the Untersuchungen liber die Fauna Peruana, pub-
lished later the same year ( 1 844b).

Although von Tschudi attempted to provide the
fullest account possible of Peruvian mammals, it
appears he had little or no contact with the ma-
jority of them. Most of his characterizations and
life history accounts are taken from Humboldt,
Spix, Wied-Neuwied, other European travelers and
natives. Camelids, the dominant animals of the
Peruvian landscape fascinated von Tschudi, and
he wrote more about them than of other animals.
His description of a vicuria hunt is quoted below
from the Ross translation (Tschudi, 1 847, pp. 2 1 9-
220).

The Indians seldom employ fire-arms in
hunting the vicunas. They catch them by
what they term the chacu. In this curious
hunt, one man at least belonging to each
family in the Puna villages takes a part, and
women accompany the train, to officiate as
cooks to the hunters. The whole company,
frequently amounting to seventy or eighty
individuals, proceeds to the Altos (the most
secluded parts of the Puna), which are the
haunts of the vicuiias. They take with them
stakes, and a great quantity of rope and cord.
A spacious open plain is selected, and the
stakes are driven into the ground in a circle,
at intervals of from twelve to fifteen feet
apart, and are connected together by ropes
fastened to them at the height of two or two
and a half feet from the ground. The circular
space within the stakes is about half a league
in circumference, and an opening of about
two hundred paces in width is left for en-
trance. On the ropes by which the stakes are
fastened together the women hang pieces of
colored rags, which flutter about in the wind.
The chacu being fully prepared, the men,
some of whom are mounted on horseback,
range about within a circuit of several miles,
driving before them all the herds of vicurias
they meet with, and forcing them into the
chacu. When a sufficient number of vicunas
is collected, the entrance is closed. The timid
animals do not attempt to leap over the ropes,
being frightened by the fluttering rags sus-
pended from them, and, when thus secured,
the Indians easily kill them by the tolas.
These bolas consist of three balls, composed
either of lead or stone; two of them heavy,

and the third rather lighter. They are fas-
tened to long, elastic strings, made of twisted
sinews of the vicuria, and the opposite ends
of the strings are all tied together. The Indian
holds the lightest of the three balls in his
hand, and swings the two others in a wide
circle above his head; then taking his aim at
the distance of about fifteen or twenty paces,
he lets go the hand-ball, upon which all the
three balls whirl in a circle, and twine round
the object aimed at. The aim is usually taken
at the hind legs of the animals, and the cords
twisting round them they become firmly
bound. It requires great skill and long prac-
tice to throw the bolas dexterously, espe-
cially when on horseback: a novice in the
art incurs the risk of dangerously hurting
either himself or his horse, by not giving the
balls the proper swing, or by letting go the
hand-ball too soon.

The vicuiias, after being secured by the
bolas, are killed, and the flesh is distributed
in equal portions among the hunters. The
skins belong to the Church. The price of a
vicuna skin is four reals. When all the ani-
mals are killed, the stakes, ropes, &c., are
packed up carefully, and conveyed to another
spot, some miles distant, where the chacu is
again fixed up. The hunting is continued in
this manner for the space of a week. The
number of animals killed during that inter-
val varies according to circumstances, being
sometimes fifty or sixty, and at other times
several hundred. During five days I took part
in a chacu hunt in the Altos of Huayhuay,
and in that space of time 122 vicurias were
caught. With the money obtained by the sale
of the skins a new altar was erected in the
church of the district. The flesh of the vicuiia
is more tender and better flavored than that
of the llama. Fine cloth and hats are made
of the wool. When taken young, the vicurias
are easily tamed, and become very docile;
but when old, they are intractable and ma-
licious. At Tarma I possessed a large and
very fine vicuria. It used to follow me like a
dog whenever I went out, whether on foot
or on horseback.

The frequent hunting seems not to have
the effect of diminishing the numbers of these
animals. If in the vicinity of the villages
where chacus are frequently established, they
are less numerous than in other parts, it is
because, to elude the pursuit of the hunters,

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 9. Peruvian mammals according to Tschudi (1844a,b); current scientific names are used followed by
Tschudi's synonym or misidentification, local names, and figure in this text; extralimital species are bracketed;
arrangement of taxa follows Tschudi.

Current wune

Tschudi synonym or
misidentification

Local name

Figure

Aieles paniscus chamek Hum-
boldt

LagothrLx lagothricha poeppigi

Schinz
LagolhrLx flavicauda Humboldt
Alotmtta seniculus Linnaeus
[Mycetes rufimanus Kuhl]
Cebus apella Linnaeus
Cebus albifrons Humboldt
[Cebus capucinus Linnaeus]
Saimiri boliviensis peruviensis

Hershkovitz
Callicebus torquatus Hofimann-

segg (subsp.?)
[Callicebus personatus E. Geof-

froyl
Aotus nigriceps E)ollman
[Chiropotes]

Saguinus mystax mystax Spix
Saguinus nigricollis Spix
Saguinus fuscicollis Spix
[Saguinus midas midas Lin-
naeus]
[Leontopithecus rosalia chryso-
melas Kuhl]

Chiroptera

Phyllostomus elongatus E. Geof-

fh)y
Phyllostomus hastatus Pallas
Phvlloslomus discolor Wagner,

i843
Artibeus cinereus Gervais

Stumira erythromos Tschudi
Sturnira oporophilum Tschudi

Glossophaga soricina Pallas
Anoura geoffroyi peruana

Tschudi
Eptesicus innoxius Gervais
Histiotus macrotus Poeppig
Noctilio leporinus Linnaeus
Noctilio albiventris Desmarest
Tadarida brasiliensis I. Geoffroy
Xfolossus molossus Pallas?
Eumops auripendulus Shaw

Molossus ater E. Geofiroy

[Promops nasutus Spix]

Carnivora

Tremarctos omatus F. Cuvicr

Nasua nasua montana Tschudi

Potosflavus Schreber
Eira barbara Linnaeus

Ateles marginatus; Aieles ater;
Ateles pentadactylus

Lagothrix humboldti; Lagothrix

canus
Mycetes flavicaudatus (sic)
Mycetes stramineus
Alouatta belzebul
Cebus robustus

Chrysothrix sciureus

Callithrix amictus

Callithrix personatus

Nyctipithecus trivirgatus
IPithecia*
[Midas labiatus]
[Midas labiatus]
[Midas labiatus]
[Midas rufimanus]

[Midas chrysomelas]

Chuva; maquisapa; chamek;

mahmonda; machucusillo;

supaya
Mono oki; choko

Coro [= coto?]
Macaquito

Tocon

Hatmnmasu

Phyllostomus innominatum

Tschudi
Phyllostomus (Artibeus) pusil-

lum

Phyllostomus (Sturnira) oporo-
philum Tschudi

Glossophaga amplexicauda

Glossophaga (Choeronycteris)
peruana Tschudi

Vespertilio innoxius

Vespertilio ( Vesperugo) velatus

Noctilio unicolor

Noctilio affinis

Molossus (Dysopes) naso

Molossus (Dysopes) velox

Molossus (Dysopes) ferox; Dy-
sopes longimanus

Molossus (Dysopes) myosuros
Tschudi; Molossus anonymus
Tschudi

Dysopes fitmarius

Ursus fivgilegus Tschudi
Nasua socially, Nasua solitaria,
Nasua leucorhynchos Tschudi
Cercoleptes caudivolvidus
Galictis barbara

Hucamari
Achuna, mishash

Cushumbi
Omeyro

HELDIANA: ZOOLCXJY

Table 9. Continued.

Current name

Tschudi synonym or
misidentification

L4>cal name

Figure

Carnivora {continued)
Mustela frenata agilis Tschudi
Conepatus chinga Molina

Lutra felina Molina
Lutra montana Tschudif
Dusicyon thous Linnaeus
Felis concolor Linnaeus
Felis onca Linnaeus

Felis pardalis Linnaeus
Feiis wiedii Schinz

Felis yagouaroundi E. Geoffroy

PlNNIPEDIA

Otaria flavescens Shaw

Marsupialia

Didelphis marsupialis Linnaeus

Metachirus nudicaudatus E.

Geoffroy
Philander opossum Linnaeus
Marmosa noctivaga Tschudi
Marmosa impavida Tschudi
Marmosa murina Linnaeus
Caluromys lanatus ornatus

Tschudi

RODENTIA

Sciurus aestuans Linnaeus
Sciurus pyrrhinus Thomas
Sciurus stramineus Eydoux and

Souleyet
Sciurus spadiceus tricolor

Tschudi
Proechimys sp.?
Chinchilla brevicaudata Water-
house
Lagidium peruanum Meyen
Lagidium viscacia Molina^
[Octodon degus Molina]
[Myocastor coypus Molina]
Coendou bicolor Tschudi
Dasyprocta leporina Linnaeus
Dasyprocta variegata Tschudi
Akodon boliviensis Meyen
Phyllotis darwini Waterhouse
Oryzomys longicaudatus de-
structor Tschudi
Oryzomys melanostoma Tschu-
di
Rhipidomys leucodactylus

Tschudi
Agouti paca Linnaeus
Hydrochaeris hydrochaeris Lin-
naeus
Cavia porcellus Linnaeus

Molina (Thiosmus) mapurita;

Mephitis furcata; Mephitis

amazonica
Lutra chilensis

Canis azarae

Felis onza

Felis macrura (sic = Felis ma-

croura); Felis celidogaster
Felis yaguaruruii

Otaria jubata; Otaria ulloae
Tschudi; Otaria aurita Hum-
boldt (in Tschudi)

Didelphys azarae
Didelphys myosuros

Poma, leon

Choque china, yana cheque,

tigre
Uturunco

Mucamuca, jarachupa

\Sciurus variabilis]

[Echinomys leptosoma]
Eriomys chinchilla

Lagidium peruvianum {sic)
Lagidium pallipes
[Octodon cummingii]
[Myopotomus coypus]
Sphingurus {sic) bicolor
Dasyprocta aguti Linnaeus

Acodon boliviense
Hesperomys darwini
Hesperomys destructor

Hesperomys melanostoma

Hesperomys {Rhipidomys) leu-
codactylus
Coelogenys fulvus
Hydrochoerus capybara

Cavia cutleri

Cutspi or cushpi

Cuy del monte

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 9. Continued.

Current name

Tschudi synonym or
misidentification

Local name

Figure

Lagomorpha

Sylvilagus brasiliensis Linnaeus

Edentata

Bradypus vahegatus Schinz
[Bradypus torquatus lUiger]
Dasypus novemcinctus Linnaeus
Cabassous unicinctus Linnaeus
Tamandua tetradactyla Lin-
naeus
Cyclopes didactylus Linnaeus

Perissodactyla

Tapirus terrestris Linnaeus

Tapirus pinchaque Roulin

Artiodactyla
Tayassu tajacu Linnaeus
Tayassu pecari Link
Lama glama Linnaeus
Lama pacos Linnaeus
Lama guanicoe Miiller
Vicugna vicugna Molina
Mazama americana Erxleben
Mazama gouazoubira peruana

Tschudi
Hippocamelus antisensis d'Or-

bigny

Lepus brasiliensis

Bradypus infuscatus

Dasypus 9-cinctus {sic)
Dasypus tatuay {sic = tatouay)
Myrmecophaga tamandua

Myrmecophaga didactyla

Quirquincho

Tapirus americanus

Tapirus villosus

Dicotyles torquatus

Dicotyles labiatus

Auchenia lama

Llama

Auchenia paco

Alpaca

Auchenia huanaco

Auchenia vicuna

Vicuna

Cervus rufus

Cervus nemorivagus var. per-

Liucho, venado

uana

Cervus antisiensis

Tarush, taruga

* Sakis {Pithecia) evidently not seen by von Tschudi. His descriptions are of bearded sakis {Chiropotes) after
Humboldt (1811), which do not occur in Peru,
t May not be an otter, according to Thomas (1908, p. 393).
% The species was known to occur in parts of formerly southwestern Peru now in Chile.

they seek refuge in the Altos, where they are
found in vast numbers. Several modem
travelers have lamented the diminution of
the vicunas, but without reason. In fonner
times those animals were hunted more ac-
tively than at present.

Von Tschudi's journeys in the puna inspired
him to poetic descriptions of the habits, particu-
larly the visual propensities, of its denizens.

Herds of vicuiias approached me with cu-
rious gaze, and then on a sudden fled with
the swiftness of the wind. In the distance I
observed stately groups of huanacos turning
cautiously to look at me, and then passing
on. The Puna stag (tarush) slowly advanced
from his lair in the mountain recesses, and
fixed on me his large, black, wondering eyes,
whilst the nimble rock rabbits (viscachas)
playfully disported and nibbled the scanty

herbage growing in the mountain crevices.
(Tschudi, Ross translation, 1 847, p. 249)

On descending the eastern slope of the Cordi-
lleras to the subtropical zone inhabited by a greater
variety of different kinds of mammals, von Tschu-
di (Ross translation, 1847, p. 275) romanticized
that:

. . . the swift-footed roe [Mazama sp.] of the
Cordillera roams here and dwells in the
thickets, avoiding the warm forest. The dark
brown coati {Nasua montana, Tsch.) howls
and digs at the root of trees in search of food,
the shy opossum crawls fearfully under the
foliage; the lazy armadillo creeps into his
hole, but the ounce [Felis onca] and the lion
[Felis concolor] seldom stray hither to con-
test with the black bear {Ursus frugilegus
Tsch.) the possession of his territory. The

FIELDIANA: ZOOLOGY

little hairy tapir (Tapirus villosus, Wagn.)
ventures only at twilight out of his close am-
bush to forage in the long grass.

The systematic arrangement in the Untersu-
chungen is said to include all mammals known at
the time to occur in Peru. By von Tschudi's count,
the fauna consists of 1 1 9 species in 48 genera.
These totals include domestic animals, the intro-
duced house mouse, some duplicated names of
native species, and a number of others not known
to occur in Peru. In terms of currently recognized
species found in Peru, von Tschudi's combined
lists (1844a, pp. 244-255; 1844b, pp. 6-20; 21-
264) consist of 87 species in 58 genera. The species
are listed in Table 9 with von Tschudi's synonyms
or misidentifications. Author attributions of the
synonyms are omitted unless they are to von
Tschudi himself Vernacular names, if given, are
included. Extralimital species are shown in brack-
ets. In the case of unrevised groups or where two
or more subspecies occur in Peru without possi-
bility of determining which were described by von
Tschudi, only the specific names are given.

XIII. Patagonia

Alcide Charles Victor d'Orbigny (1802-1857)

The French-bom Alcide d'Orbigny was educat-
ed by his country's leading naturalists. His apti-
tudes were recognized by authorities of the Mu-
seum National d'Histoire Naturelle, and with that
institution's financial and material assistance, he
sailed for South America charged with making a
scientific survey of the southern half of the con-
tinent. Circumstances restricted his studies and
collections of mammals almost entirely to Argen-
tina and Bolivia.

D'Orbigny left France 3 1 July 1 826 and arrived
in Rio de Janeiro 24 September 1826 on his way
to Montevideo where he landed on 29 September.
The natural history of the region between Mal-
donado east of Montevideo and Buenos Aires en-
gaged his attention for several months.

On 14 February 1827, d'Orbigny ascended the
Rio Parana and arrived 1 5 March at the important
fluvial port of Corrientes, capital of the province
of the same name. With the town as base, d'Or-
bigny explored the province throughout much of
one year.

On his return to Buenos Aires in April 1828, he
made stops in Entre Rios and Santa Fe. Beginning

June 1828 and continuing through 1829, his at-
tentions were devoted to faunal studies in the
provinces of Buenos Aires and Rio Negro. The
chronology of the early part of 1829, as given by
d'Orbigny (1835-1847) in the Voyage, confuses
time spent in the two provinces with that spent in
Corrientes. In any event, d'Orbigny was clearly in
Buenos Aires and Rio Negro during the last half
of 1 829. He returned to Montevideo in December
1 829 and on 29 December sailed on to Patagonia
and Chile.

Cape Horn was rounded on 19 January 1830
and Valparaiso, Chile, was reached 16 February.
Because of the political unrest in the country, d'Or-
bigny sailed to the then Bolivian port of Cobija,
where he landed on 8 April; 20 April found him
in Arica and Tacna, both ports then in Peru's pos-
session. After some investigation of the coast,
d'Orbigny left Tacna on 19 May for La Paz, the
mountain capital of Bolivia, arriving there 28 or
29 May.

For the next three years, d'Orbigny explored,
mapped, and sampled the natural resources of the
country. He crisscrossed Bolivia from La Paz east
to the Paraguayan border and from Potosi in the
south to the lower Rio Mamore in the north. D'Or-
bigny's actual itinerary is almost impossible to
track because of the inaccuracies of the then avail-
able maps. Modem maps aided Pilleri and Arvy
(1977) in their reconstmction of the itinerary in
chronological sequence (fig. 1 8).

A complete account of d'Orbigny's South Amer-
ican joumey with observations on and descrip-
tions of the geology, paleontology, living plants,
animals, and Indians is contained in seven huge
volumes published serially from 1 835 through 1847
in Paris under the title Voyage dans I'Amehque
Meridionale. A full report on the mammals was
reserved for the last, or perhaps a separate pub-
lication, but a turn in d'Orbigny's fortunes inter-
rupted the work. A number of colored plates of
mammals believed new to science and a few short
articles on others had already been published. So
that all would not be lost, a synoptic systematic
report on the mammals collected was published
in 1 847 jointly with the distinguished mammal-
ogist Paul Gervais, as number 2 of volume 4 of
the Voyage. Brief notes on distribution and be-
havior accompany the abbreviated descriptions of
each species. The species are listed in Table 10
with abstracted locality data. Scientific names used
are current with synonyms and misidentifications
added. The specimens are deposited in the Mu-
seum National d'Histoire Naturelle in Paris.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

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FIELDIANA: ZOOLOGY

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Fig. 1 9. Animals of the d'Orbigny Bolivian Expedition: upper left, Callithrix entomophagus d'Orbigny (= Saimiri
boliviensis boliviensis I. Geoffroy and Blainville); upper right, Callithrix donacophilus d'Orbigny (= Callicebus do-
nacophilus donacophilus); lower left, Felis geoffroyi d'Orbigny and Gervais (= Felis colocolo geoffroyi); lower right.
Mephitis humboldtii (= Conepatus chinga suffocans Illiger); from d'Orbigny and Gervais (1847).

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 10. Mammals of the southern half of South America, mostly Bolivia and Argentina, recorded by d'Orbigny
and Gervais (1847); the arrangement is phylogenetic.

Current name

d'Orbigny and Gervais synonym

Locality

Figure

Chiroptera

Noctilio albiventris Desmarest,
1818

Noctilio leporinus rufipes d'Or-
bigny, 1835

Tonatia sylvicola d'Orbigny,
1835

Artibeus planirostris Spix, 1 823

Desmodus rotundus E. Geoffroy,
1810

Myotis nigricans Wied-Neu-
wied, 1821

Eptesiciis furinalis d'Orbigny
and Gervais, 1 847

Myotis albescens E. Geoffroy,
1806

Myotis ruber E. Geoffroy, 1 806

Histiotus velatus I. Geoffroy,
1824

Tadarida brasiliensis I. Geof-
froy, 1824

Molossus crassicaudatus E. Geof-
froy, 1805

Primates

Saimiri boliviensis boliviensis I.

Geoffroy and Blainville, 1 834
Callicebus donacophilus donaco-

philus d^Orhigny, 1835
Alouatta seniculus sara Elliot,

1910
Cebi4s apella paraguayanus

Fischer, 1829

Carnivora

Dusicyon gymnocercus Fischer,

1814
Chrysocyon brachyurus Illiger,

1815
Tremarctos ornatus F. Cuvier,

1825
Procyon cancrivorus nigripes

Mivart, 1886
Nasua nasua solitaria Wied-

Neuwied, 1821
PotosflavusSchxc\xT, 111 A

Lyncodon patagonicus Blain-
ville, 1842

Galictis cujafurax Thomas,
1907

Conepatus chinga suffocans Illi-
ger, 1815

Lutra platensis Waterhouse,
1838

Noctilio affinis d'Orbigny, 1835 BOLIVIA: Moxos Province

BOLIVIA: Chiquitos and

Moxos provinces
BOLIVIA: Yuracare territory,

base of eastern Cordillera
BOLIVIA: Chiquitos Province

Not Vespertilio perspicillatus
Linnaeus, 1758

Desmodus rufus Wied-Neuwied,
1824; Edostoma cinerea d'Or-
bigny, 1835

Vespertilio hypothrix d'Orbigny
and Gervais, 1847

Vespertilio isidori d'Orbigny and
Gervais, 1847

Molossus rugosus d'Orbigny,

1835, not Molossus nasutus

Spix, 1823
Molossus moxensis d'Orbigny,

1835; Molossus velox Tem-

minck, 1827

Calithrix (sic) entomophagus
d'Orbigny, 1835

Not Stentor stramineus E. Geof-
froy
Cebus fulvus var.

Not Canis cancrivorus Desma-
rest, 1820
Canis jubatids Desmarest, 1820

Not Procyon cancrivorus Cuvier,

1798
Nasua fusca Desmarest, part

Cercoleptes caudivolvulus Schre-
ber, 1774

Not Mustela brasiliensis Gme-
lin, 1788

Mephitis castaneus d'Orbigny
and Gervais, 1 847, not Me-
phitis humboldtii Gray, 1837

BOLIVIA: Chiquitos

BOLIVIA: Moxos

ARGENTINA: Corrientes

ARGENTINA: Corrientes
BOLIVIA: Chuquisaca

ARGENTINA: Corrientes

BOLIVIA: Moxos and Chiqui-
tos provinces

BOLIVIA: Chiquitos; Moxos;

Santa Cruz
BOLIVIA: Moxos Province

BOLIVIA: Santa Cruz; Chiqui-
tos; Moxos

BOLIVIA: near Santa Cruz de
la Sierra

BOLIVIA: Chiquitos

Tropical South America to 41°S

BOLIVIA: Cochabamba; Chu-
quisaca

BOLIVIA: Chiquitos; ARGEN-
TINA: Corrientes

BOLIVIA: tropics to 30'^

BOLIVIA: foot of eastern Cordi-
llera
ARGENTINA: Rio Negro

19
19

ARGENTINA: Rio Parana in
Provinces Buenos Aires and
Corrientes

FIELDIANA: ZOOLOGY

Table 10. Continued.

Current name

d'Orbigny and Gervais synonym

Locality

Figure

Carnivora

Felis colocolo pajeros Desma-

rest, 1816
Felis geoffroyi d'Orbigny and

Gervais, 1847
Felis concolor Linnaeus, 1771

Felis onca Linnaeus, 1758

PiNNIPEDIA

OtariaflavescensShs^N, 1800

Arctocephalus australis Zimmer-

mann, 1782
Mirounga leonina Linnaeus,

1758

Artiodactyla

Mazama gouazoubira Fischer,
1814

Blastoceros bezoarticus Lin-
naeus, 1758

Hippocamelus antisensis d'Or-
bigny, 1834

Blastocerus dichotomus Illiger,
1815

RODENTIA

Sciurus spadiceus Olfers, 1818

Eligmodontia typus F. Cuvier,
1837

Octodon degus Molina, 1782

Octodontomys gliroides, Gervais
and d'Orbigny, 1 844

Ctenomys boliviensis Water-
house, 1848

Ctenomys magellanicus Bennett,

1835
Microcavia australis Gervais

and d'Orbigny, 1833
Galea flavidens Brandt, 1835

Dolichotis patagonum Zimmer-
man, 1780

Dasyprocta azarae Lichtenstein,
1827

Cetacea

Inia boliviensis d'Orbigny, 1834

[Pontoporia blainvillei Gervais
and d'Orbigny, 1 844; not part
of d'Orbigny collection]

Lagenorhynchus cruciger Quoy
and Gaimard, 1 824

Lissodelphis peroni Lacepede,
1804

Otaria jubata Schreher, 1776
Otaria porcina Molina, 1782
Phoca proboscidea Peron, 1817

Cervus simplicicornis Illiger,
1815

Not Cervus campestris F. Cu-
vier, 1817

Cervus paludosus Desmarest,
1822

Not Sciurus igniventris Wagner,
1842

Not Ctenomys brasiliensis
Blainville, 1826

Dasyprocta patachonica Desma-
rest, 1820

Not Dasyprocta nigricans Wag-
ner, 1842

ARGENTINA: from 35°-45'S
ARGENTINA: Pampas to 44'S 19

BOLIVIA; ARGENTINA: to
Straits of Magellan

Tropical South America not be-
yond 40^; ARGENTINA:
Pampas; Serrania de Tandil

ARGENTINA: S mouth Rio

Negro
ARGENTINA: coast; PERU:

coast
ARGENTINA: Rio Negro, near

mouth

Tropical South America to 28°S

Lowland savannas to northern
Patagonia

BOLIVIA: La Paz; Cochabam- 20

ba; Chuquisaca; rarely below
3500 m

ARGENTINA: Corrientes; BO-
LIVIA: Chiquitos

BOLIVIA: Chiquitos

ARGENTINA: Corrientes

CHILE: Santiago de Chile
BOLIVIA: La Paz

ARGENTINA: Corrientes; BO-
LIVIA: Santa Cruz de la Sie-
rra

ARGENTINA: northern Pata-
gonia

ARGENTINA: Rio Negro

BOLIVIA: Cochabamba; Chu-
quisaca; La Paz

ARGENTINA: northern Pata-
gonia; Corrientes

Tropical South America

BOLIVIA: rivers of Moxos and 20

Chiquitos
URUGUAY: Montevideo

Atlantic Ocean (57»-76'«, E and
S of Cape Horn)

Atlantic Ocean (48°-64'«); At-
lantic-Pacific Oceans around
Cape Horn

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

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<N 53

FIELDIANA: ZOOLOGY

Charles Robert Darwin (1809-1882)

Charles Robert Darwin was bom in Shrews-
bury, England, to a wealthy and distinguished fam-
ily. Although his early schooling emphasized the
classics, Darwin's interests since boyhood were in
natural history, particularly of the insects he col-
lected, and in hunting as a sport. As a university
student, he dropF)ed out of medical school after
two years, then took up theology, and abandoned
that after three years. Nevertheless, through the
influence of his teachers, he developed and sharp-
ened his interests in biology and geology, and his
reading of Humboldt's Personal Narrative of
Travels to the Equinoctial Regions of America fired
him with a zeal for travel and discoveries in distant
and unexplored lands.

The opportunity for travel in exotic parts soon
came. At age 22, with his mostly self-acquired
knowledge of geology and systematic biology and
exjjerience as a collector and hunter, Charles Dar-
win accepted the unsalaried post of Naturalist on
H.M.S. Beagle for a five-year cruise of chrono-
metrical explorations round the world. The ex-
periences on the voyage, which began 27 Decem-
ber 1831 (fig. 21), transformed Darwin into the
leading naturalist of his time and were the prime
source of inspiration for Darwin's theory of or-
ganic evolution by natural selection.

The Beagle touched the South American main-
land at Bahia (now Salvador), Brazil, on 29 Feb-
ruary 1 832 for a short stay. Before the ship left for
Rio de Janeiro in March, Darwin captured and
prepared for study a specimen of the very common
phyllostomid bat, Carollia perspicillata Linnaeus,
his first mammalian sp)ecimen of the exp)edition.

In Rio de Janeiro, Darwin was taken on a hunt
by an old Portuguese priest. Two howler monkeys
{Alouattafusca E. Geoffroy), described by Darwin
( 1 839, p. 32) as "two large bearded monkeys," had
been shot the day before by his companion. Dar-
win wrote:

These animals have prehensile tails, the ex-
tremity of which, even after death, can sup-
port the whole weight of the body. One of
them thus remained fast to a branch, and it
was necessary to cut down a large tree to
procure it. This was soon effected and down
came tree and monkey with an awful crash.

The priest later presented Darwin with an eyra cat
(Herpailurus yagouaroundi eyra Fischer) that had
just been killed in the Gavea mountain.

In July 1832 the Beagle left Brazil for the Pa-
tagonian subregion. Up to this time, Darwin's zoo-
logical collections consisted mainly of insects and
mollusks. Because only negligible contact with the
rich mammalian fauna of Brazil had been made,
Darwin was deprived of a basis for direct com-
parisons with the comparatively poor but largely
unique mammalian fauna of the Patagonian
subregion, which he studied zealously. As a result,
his attention focused on morphological and eco-
logical differences between the individual species
(or subspecies) he collected or observed in La Pla-
ta, Bahia Blanca, Patagonia, the Falklands, Chile,
and the Galapagos and the same or nearly related
species of Paraguay and Chile described by Azara
and Molina. How much would Darwin's concept
of the origin of life been affected if his thoughts
had been directed primarily to faunas and faunal
regions rather than to species and their geographic
variation?

The Beagle remained in the area of La Plata
from July 1832 to July 1833, affording Darwin
opportunities to collect near Maidonado, a short
distance up the coast from Montevideo. The Bea-
gle then sailed south to the mouth of the Rio Ne-
gro. While the vessel's crew mapped and took
soundings up and down the coast between the Rio
Negro and Rio Plata, Darwin made a number of
excursions into the Pampas, Bahia Blanca, Sierra
de la Ventana, Rio Colorado, Rio Parana, and Rio
Uruguay. Many observations were made on the
behavior and habitat of mammals characteristic
of the region, but few animals were actually col-
lected. Among the species mentioned are arma-
dillos (known as pichiy, peludo, apar, and mulita),
the Patagonian hare or mara (misnamed "agou-
ti"), the capybara, cavia, skunk, puma, jaguar,
guanaco, and pampas deer. Darwin (1839, p. 144)
was fascinated by the viscacha's packrat-like hab-
its such as:

dragging every hard object to the mouth of
its burrow; around each group of holes
many bones of cattle, stones, thistle stalks,
hard lumps of earth, dry dung, etc., are col-
lected into an irregular heap, which fre-
quently amounts to as much as a wheelbar-
row would contain. I was credibly informed
that a gentleman, when riding on a dark night,
dropped his watch; he returned in the morn-
ing, and by searching the neighborhood of
every bizcacha hole on the line of the road,
as he exp>ected, soon found it.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

lOV

Coplapo'U

Coqulabo pLl-
Coocepcirfn J9 \ l Buenos Aires

^-xJUjtontevldeo
Y (SIERRA DF, ^fo Plata

CHARLES DARUIN

TOTAGE OF B.M.S. BEACLE

SOUTH AMERICAN LOCALITIES

(1832-1835)

CBOHOS*-

AXCRIPELACO^^ ,'

-■w

Good Success

(■snia dc Good Uickf

80*

Hp-y^^ ^/San Julian

FALKLAND ISLANDS

tllUA DEL ruioo

Canai fagla

J 1 L.

60*

50*

30*

Fig. 2 ! . Map showing principal South American stations visited by Charles Darwin ( 1 832-1 835) on world cruise
of H.M.S. Beagle (1832-1836).

HELDIANA: ZOOLOGY

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 1 1 . Mammals collected or observed by Oanvin in the Maldonado Region, Uruguay and parts of Argentina,
and those recorded by Waterhouse (1838-1839); the arrangement is phylogenetic.

Current name

Waterhouse synonym or
misidentification

Locality

Figure

Marsupialia

Didelphis albiventris Lund

Lutreolina crassicaudata Des-

marest
Monodelphis dimidiata Wagner

Chiroptera

Tadarida brasiliensis I. Geoffroy

Edentata

Dasypus hybridus Desmarest

Zaedyus pichiy Desmarest

Chaetophractus villosus Desma-
rest
Tolypeutes matacus Desmarest

Carnivora

Dusicyon gymnocercus Fischer

Felis colocolo pajeros Desmarest
Galictus cujafurax Thomas
Lutra platensis Waterhouse
Conepatus chinga gibsoni

Thomas
Felis concolor acrocodia Gold-
man
Felis onca palustris Ameghino

Artiodactyia
Blastoceros bezoartiats Lin-
naeus

Lama guanicoe Muller

RODENTIA

Myomorpiia

Oryzomys flavescens Waterhouse
CaJomys laucha Olfers

Eligmodontia typus Cuvier

Holochilus brasiliensis darwini
Thomas

Reithrodon physodes typicus
Waterhouse

Akodon azarae Fischer

Akodon colibreve Brants

Scapteromys tumidus Water-
house

Oxymycierus rufus nasutus
Waterhouse

Caviomorpiia

Cavia porcellus Linnaeus
Hydrochaeris hydrochaeris Lin-
naeus

Didelphis azarae AucL

Didelphis brachyura Auct.
Not Dysopes nasutus Spix

Dasypus minutus AucL

Not Canis azarae Wied-Neu-
wied

Not Galictis vittata Schreber

Not Cervus campestris Cuvier

Mus bimaculatus Waterhouse;

Mus gracilipes Waterhouse
Mus elegans Waterhouse

Mus arenicola Waterhouse
Mus obscurus Waterhouse

Cavia cobaia Auct.
Hydrochoerus capybara Auct.

URUGUAY: Maldonado
URUGUAY: Maldonado

URUGUAY: Maldonado

URUGUAY: Maldonado
ARGENTINA: Banda Oriental,

Entre Rios
ARGENTINA: Bahia Blanca

(observed)
ARGENTINA: Bahia Blanca

(observed)

ARGENTINA: La Plata (ob-
served)
ARGENTINA: Bahia Blanca
URUGUAY: Maldonado
URUGUAY: Maldonado
ARGENTINA: Bahia Blanca

(observed)
ARGENTINA: the pampas (ob-
served)
ARGENTINA: in the Rio Pa-
rana (observed)

URUGUAY: Maldonado; AR-
GENTINA: Bahia Blanca;
Rio Negro

ARGENTINA: Rio Negro (ob-
served)

URUGUAY: Maldonado
ARGENTINA: Bahia Blanca

ARGENTINA: Bahia Blanca
ARGENTINA: Bahia Blanca

URUGUAY: Maldonado

URUGUAY: Maldonado
URUGUAY: Maldonado
URUGUAY: Maldonado

URUGUAY: Maldonado

URUGUAY: Maldonado
URUGUAY: Maldonado

HELDIANA: ZOOLOGY

Table 1 1 . Continued.

Current name

Waterhouse synonym or
misidentification

Locality

Figure

Caviomorpha (continued)

Dolichotis patagonum Zimmer-
man
Vizcacia maximus Desmarest

Ctenomys brasiliensis Blainville

Lagostomus trichodactylus
Brookes

ARGENTINA: Rio Negro (ob-
served)
URUGUAY: Maldonado

URUGUAY: Maldonado

Darwin could not explain the viscacha's behavior.
Pampas deer (Blastoceros bezoarticus) were
abundant throughout the La Plata region. Darwin
(1839, p. 55) saw

very many small herds, containing from five
to seven animals each, near the Sierra Ven-
tana, and among the hills north of Maldo-
nado. If a person crawling close along the
ground, slowly advances toward a herd, the
deer frequently, out of curiosity, approach
to reconnoitre him. I have by this means
killed, from one spot, three out of the same
herd. Although so tame and inquisitive, yet
when approached on horseback, they are ex-
ceedingly wary. In this country nobody goes
on foot, and the deer knows man as its en-
emy only when he is mounted and armed
with the bolas.

The jaguar by some accounts is a man-killer, by
others, fears man. Darwin (1839, p. 159) records
several instances reported to him of man-killing
jaguars of the Rio Parana region.

The Beagle left the Rio Plata on December 1 833
for Puerto Deseado, or Port Desire, on the Pata-
gonian coast. The mammals collected by Darwin
and reported by Waterhouse (1838-1839), with
descriptions and supplementary notes by Darwin,
are listed in Table 1 1, with the Waterhouse syn-
onyms (misidentifications included). Added are the
few species Darwin mentioned in his Journal but
did not collect. Unless otherwise indicated, all
species are from the neighborhood of Maldonado,
Uruguay.

The geology and natural history of Patagonia
investigated by Darwin included those of the Straits
of Magellan and Tierra del Fuego (December 1 832-
January 1833; May-June 1834), Puerto Deseado
(Port Desire) (December 1833-January 1834),
Santa Cruz (April-May 1834), and the Falkland
Islands (March 1834). The Beagle itself (fig. 22)
sailed up the Rio Santa Cruz to a point 1 40 miles

from its mouth in the Atlantic Ocean to about 60
miles from the nearest arm of the Pacific Ocean
on the opposite side of the cordillera.

Darwin was greatly impressed by the number,
variety, and great size of fossil mammals, mostly
Pleistocene, exposed on the Patagonian plains.
These, he (1839, p. 209) believed, were confir-
mation of the "law" that existing animals in an
area have a close relation in form with extinct
species in the same area. The natural causes for
extinction, however, eluded Darwin. After pro-
posing and rejecting a number of explanations, the
nonevolutionist Darwin (1839, p. 212) concluded
that

the whole series of animals, which have been
created with f>eculiar kinds of organization,
are confined to certain areas; and we can
hardly suppose these structures are only ad-
aptations to peculiarities of climate or coun-
try; for otherwise, animals belonging to a
distinct type, and introduced by man, would
not succeed so admirably even to the exter-
mination of the aborigines. On such grounds
it does not seem a necessary conclusion that
the extinction of species, more than their
creation, should exclusively depend on the
nature (altered by physical change) of their
country. All that at present can be said with
certainty, is that, as with the individual, so
with the species, the hour of life has run its
course, and is spent.

The small number of extant large mammals and
great number and variety of small mammals, also
impressed Darwin (1839, p. 215).

Patagonia, poor as she is in some resj)ects,
can, however, boast of a greater stock of
small rodents than p)erhaps, any other coun-
try in the world. Several sjjecies of mice are
externally characterized by large thin ears
and a very fine fur. These little animals

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

swarm amongst the thickets in the valleys,
where they cannot for months together taste
a drop of water. They all seem to be can-
nibals, for no sooner was a mouse caught in
one of my traps than it was devoured by
others. A small and delicately-shaped fox
which is likewise very abundant, probably
derives its entire support from these small
animals.

The guanaco was regarded as the characteristic
quadruped of the Patagonian plains (Darwin, 1 839,
p. 215).

Herds of fifty or a hundred were common,
and, as I have said, we saw one which must
have contained at least five hundred. The
puma with the condor in its train, follows
and preys upon these animals. The footsteps
of the former were to be seen almost every-
where on the banks of the river [Santa Cruz];
and the remains of several guanaco, with
their necks dislocated, and bones broken,
showed how they had met their death.

In March 1834, Darwin visited the Falkland
Islands. On a tour he encountered large numbers
of horses, cattle, swine, and rabbits {Oryctolagus
cuniculus Linnaeus [= Lepus magellanicus Lesson
and Gamot] in domestic and feral states. The an-
imals had been brought by French colonists in
1 764. Darwin wrote (p. 249),

The only quadruped native to the island, is
a large wolf-like fox [Dusicyon (Iculpaeus)
australis Kerr] which is common to both
East and West Falkland. I have no doubt it
is a peculiar species and confined to this
archijjelago. . . . These wolves are well known
. . . [for] their lameness and curiosity. ... To
this day their manners remain the same. . . .
As far as I am aware, there is no other in-
stance in any part of the world, of so small
a mass of broken land, distant from a con-
tinent, possessing so large a quadruped pe-
culiar to itself. Their numbers have rapidly
decreased; they are already banished from
that half of the island which lies to the east-
ward of the neck of land between St. Sal-
vador Bay and Berkeley Sound. Within a
very few years after these islands shall have
become regularly settled, in all probability
this fox will be classed with the dodo, as an
animal which has perished from the face of
the earth.

The mammals collected by Darwin in the Ar-
gentine Patagonia (including Falkland Islands) and
bordering parts of the Chilean Straits of Magellan
are listed in Table 12.

The Chilean leg of the cruise began in May 1 834
with the passage of the Beagle into the eastern
mouth of the Straits of Magellan and ended July
1835 with departure from Copiapo in northern
Chile. While the Beagle sailed up and down the
Chilean coast, Darwin explored the coast, islands,
archipelagos, and cordillera. He crossed the Andes
on 21 March 1835 through the Portillo Pass south
of Santiago, and proceeded to the town of Men-
doza in Argentina. Differences observed between
the biota of eastern and western versants of the
cordillera impressed Darwin. The mountains, he
(1839, p. 399) reasoned,

have existed as a great barrier, since a period
so remote that whole races of animals must
subsequently have perished from the face of
the earth. Therefore, unless we suppose the
same species to have been created in two
different countries, we ought not to expect
any closer similarity between the organic
beings on opposite sides of the Andes, than
on shores separated by a broad strait of the
sea.

The correlation between geographic isolation and
faunal peculiarity was noted in other circum-
stances. Darwin (1839, p. 439) observed that

next to lizards, mice appear to be able to
support existence on the smallest and driest
portions of the earth— even on islets in the
midst of great oceans. I believe it will be
found, that several islands, which possess no
other warm-blooded quadruped, have small
rodents peculiar to themselves.

Ratadas or rat plagues in Chile also caught Dar-
win's attention. One of the earliest recorded for
Oryzomys longicaudatus longicaudatus, viewed
through the eyes of Darwin (in Waterhouse, 1 838,
p. 40), "overran the wooded country south of Con-
cepcion, in swarms of infinite numbers."

The mammals of Tierra del Fuego tallied by
Darwin (1839, p. 300) included, besides cetaceans
and phocids,

one bat [not named but likely Histiotus
montanus magellanicus Philippi], a mouse
with grooved front teeth {Reithrodon of
Waterhouse) and two other species, the tu-

HELDIANA: ZOOLOGY

Table 12. Patagonian mammals collected by Darwin and recorded by Waterhouse (1838-1839); arrangement is
phylogenetic.

Current name

Waterhouse synonym or
misidentification

Locality

Figure

Carnivora

Dusicyon australis Kerr

Dusicyon griseus Gray

Felis colocolo pajeros Desmarest

Artiodactyla

Lama guanicoe Miiller

RODENTIA

Oryzomys longicaudatus magel-
lanicus Bennett

Akodon xanthorhinus Water-
house
Akodon canescens Waterhouse

Auliscomys micropus Water-
house

Graomys griseojlavus Water-
house

Phyllotis xanthopygus Water-
house

Reithrodon physodes cunicu-
loides Waterhouse

Euneomys chinchilloides Water-
house

Myocastor coypus Molina

Microcavia australis I. Geoffroy
and d'Orbigny

Dolichotis patagonum Zimmer-
man

Cetacea

Lagenorhynchus cruciger Quoy
and Gaimard

Canis antarcticus Shaw
Not Canis azarae Wied-Neu-
wied

Auchenia llama Desmarest

Cavia patachonica Shaw

Delphinus fitzroyi Waterhouse

Falkland Islands
ARGENTINA: PaUgonia

ARGENTINA: Santa Cruz

ARGENTINA: Patagonia

ARGENTINA: Puerto de
Hambre (Port Famine);
CHILE: Straits of Magellan

CHILE: Peninsula de Hardy

ARGENTINA: Puerto Deseado
(Port Desire); Santa Cruz

ARGENTINA: Rio Santa Cruz,
Santa Cruz

ARGENTINA: Rio Negro

ARGENTINA: Puerto Deseado
(Port Desire); Santa Cruz

ARGENTINA: Puerto Deseado
(Port Desire); San Julian; Rio
Santa Cruz, Santa Cruz

ARGENTINA: Eastern entrance
to Straits of Magellan

ARGENTINA: Rio Chubut

ARGENTINA: 41°S to Straits
of Magellan

ARGENTINA: Patagonia

ARGENTINA: Golfo San Jose,
42°30'S, Chubut

cotuco (the greater number of these rodents
are confined to the eastern and dry part), a
fox, sea-otter, guanaco, and one deer [un-
named but likely Hippocamelus bisulcus].
The latter animal is rare, and is not, I be-
lieve, to be found south of the Straits of
Magellan, as happens with the others.

With respect to geographic distribution, Darwin
(1839, p. 300),

observing the general correspondence of the
cliffs of soft sandstone, mud, and shingle, on
the opposite side of the Strait, together with
those on some intervening islands [was]
strongly tempted to believe that the land was
once joined and thus allowed animals so del-
icate and helpless as the tucotuco, and
Reithrodon to pass over.

The tucotuco in question is Ctenomys magellan-

icusfueginus Philippi (Osgood, 1943, p. 1 19). Dar-
win (1839, p. 327) also mentioned the occurrence
of the puma {Felis concolor) in Tierra del Fuego,
and related something of its habits in other parts
of Chile and Argentina.

The type specimen of Darwin's zorro {Dusicyon
fulvipes Martin), peculiar to the island of Chiloe,
was discovered by Darwin (p. 34 1 ) on 6 December
1834 sitting on the rocks and so intently absorbed
in watching the maneuvers of two ship's officers
engaged in surveying,

that I was able, by quietly walking up be-
hind, to knock him on the head with my
geological hammer. This fox, more curious
or more scientific, but less wise, than the
generality of his brethren, is now mounted
in the museum of the Zoological Society.

Sea otters {Lutrafelina Molina) were described
by Darwin (in Waterhouse, 1838, p. 24) as ex-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

ceedingly common amongst the innumerable
channels and bays which form the Chonos Ar-
chipelago.

. . . they may generally be seen quietly swim-
ming with their heads just out of water amidst
the great entangled beds of kelp, which
abounds on this coast. They burrow in the
ground, within the forest, just above the
rocky shore, and I was told, that they some-
times roam about through the woods. This
otter does not, by any means, live exclu-
sively on fish. One was shot whilst running
to its hole with a large volute-shell in its
mouth; another (I believe the same species)
was seen in Tierra del Fuego devouring a
cuttle fish. But in the Chonos Archipelago
perhaps the chief food of this animal, as well
as that of the immense herds of great seals,
and flocks of terns and cormorants, is a red-
coloured crab (belonging to the family Mar-
rouri) of the size of a prawn, which swims
near the surface in such dense bodies, that
the water appears of a red colour. This spec-
imen weighed nine pounds and a half

The vampire bat which Darwin (1839, p. 25)
recognized as a species of d'Orbigny's genus Edos-
toma (= Des modus) was singled out as

often the cause of much trouble, by biting
the horses on their withers. The injury is
generally not so much owing to the loss of
blood, as to the inflammation which the
pressure of the saddle afterwards produces.
The whole circumstance has lately been
doubted in England; I was therefore fortun-
ate in being present when one was actually
caught on a horse's back. We were bivouack-
ing late one evening near Coquimbo, in Chile,
when my servant, noticing that one of the
horses was very restive, went to see what
was the matter, and fancying he could dis-
tinguish something, suddenly put his hand
on the beast's withers, and secured the vam-
pire. In the morning, the spot, where the bite
had been inflicted, was easily distinguished
from being slightly swollen and bloody. The
third day afterwards we rode the horse, with-
out any ill effects.

The Chilean mammals collected and others, only
observed by Darwin, are listed in Table 13.

Departing Chile on 12 July 1835, the Beagle
sailed north along the Peruvian coast before turn-
ing west to the Galapagos Islands. Darwin's ac-

counts of the stopovers in Iquique, Callao. and
Lima make no mention of indigenous mammals.

The Galapagos Archipelago, it seemed to Dar-
win ( 1 839. p. 454), was "a little world within itself;
the greater number of its inhabitants both vege-
table and animal being found nowhere else." Dar-
win (p. 464) "endeavoured to make as nearly a
p)erfect collection in every branch as time permit-
ted" but the only land mammals he found were
the Chatham Island rice rats described by Water-
house as Oryzomys galapagoensis (fig. 23), and the
introduced Rattus on James Island.

Darwin's investigations of the Galapagos fauna,
Ijarticularly the birds, lizards, tortoises, and cer-
tain plants stirred old beliefs and generated new
and conflicting thoughts. However, at the time he
wrote his journal in October 1835, Darwin (1839,
p. 474) made no

attempt to come to any definite conclusions,
as the sp)ecies have not [as yet] been accu-
rately examined; but we may infer, that, with
the exception of a few wanderers, the organic
beings found on this archipelago are peculiar
to it; and yet their general form strongly par-
takes of an American character. . . . This
similarity in type between distant islands and
continents, while the sp)ecies are distinct, has
scarcely been noticed. The circumstances
would be explained according to the views
of some authors, by saying that the creative
power had acted according to the same law
over a wide area.

Writers on Darwin, quoting from his revised
(1845) edition of the Journal, attribute to Darwin
more foresight on the origin of species than is ap-
parent in the first (1839) edition quoted here. At
the time of its publication, two years delayed, Dar-
win, still a creationist and believer in the immut-
ability of species, had yet to know the identity or
specific aflinities of the vast majority of the plants
and animals he had collected. This knowledge
served him later for definition and elaboration of
thoughts expressed in the second and other revised
editions of the Journal, but not in the first.

The following impressions of the biota of the
Galapagos Islands in the second edition (p. 372 of
an 1 899 "authorized edition") and oft quoted in
whole or in part by various authors, are absent in
the first.

The natural history of the islands is emi-
nently curious and well deserved attention.
Most of the organic productions are aborig-
inal creations, found nowhere else; there is

FIELDIANA: ZOOLOGY

Table 13. Chilean mammals collected or only observed by Darwin, and those identified by Waterhouse (1838-
1839); the arrangement is phylogenetic.

Current name

Waterhouse synonym or
misidentification

L4>cality

Figure

Marsupialia

Marmosa elegans Waterhouse

Chiroptera

Histiotus montanus magellani-

cus Philippi
Myotis chiloensis Waterhouse
Tadarida brasiliensis I. Geoffroy
Desmodiis rotundus dorbignyi

Waterhouse

Carnivora

Dusicyon culpaeus magellanicus

Gray
Dusicyon fulvipes Martin
Dusicyon griseus Gray

Lutra felina Molina
Felis concolor Linnaeus

Artiodactyla
Hippocamelus bisulcus Molina

RODENTIA

Oryzomys longicaudatus longi-
caudatus Bennett

Oryzomys longicaudatus magel-
lanicus Bennett

Akodon olivaceus olivaceus
Waterhouse

Akodon olivaceus brachiotus
Waterhouse

Akodon xanthorhinus xantho-
rhinus Waterhouse

Abrothrix longipilis longipilis
Waterhouse

Phyllotis darwini darwini Water-
house

Reithrodon chinchilloides
Waterhouse

Abrocoma bennetti Waterhouse

Spalacopus cyanus Molina

Myocastor coypus Molina

Octodon degus Molina

Ctenomys magellanicus fueginus
Philippi

Not Dysopes nasutus Spix

Not Canis azarae Wied-

Neuwied
Lutra chilensis Bennett

Mus renggeri Waterhouse

Abrocoma cuvieri Waterhouse
Poephagomys ater Cuvier

Octodon cummingii Bennett

Valparaiso

Tierra del Fuego (observed)

Chiloe

Valparaiso

Coquimbo

Copiapo; Straits of Magellan

Chiloe 22

Copiapo; Straits of Magellan

Chonos Archipelago
Tierra del Fuego and central

Chile to 10,000 ft elevation

(observed)

Tierra del Fuego (observed)

Concepcion

Puerto de Hambre, Straits of

Magellan
Valparaiso; Coquimbo

Chonos; Chiloe

Hardy Peninsula, Tierra del

Fuego
Coquimbo

Coquimbo 22

Straits of Magellan

Valparaiso; Aconcagua

Valparaiso

Chonos Archipelago

Valparaiso

Tierra del Fuego (observed)

even a difference between the inhabitants of
the different islands; yet all show a marked
relationship with those of America, though
separated from that continent by an open
space of ocean, between 500 and 600 miles
in width. The archijjelago is a little world
within itself, or rather a satellite attached to
America, whence it has derived a few stray
colonists, and has received the general char-
acter of its indigenous productions. Consid-
ering the small size of these islands, we feel

the more astonished at the number of their
aboriginal beings, and at their confined range.
Seeing every height crowned with its crater,
and the boundaries of most of the lava-
streams still distinct, we are led to believe
that within a period, geologically recent, the
unbroken ocean was here spread out. Hence,
both in space and time, we seem to be
brought somewhat near to that great fact—
that mystery of mysteries— the first appear-
ance of new beings on this earth.

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

"-(SI

HELDIANA: ZOOLOGY

XIV. Georges Louis Leclerc de Buffon
(1707-1788)

Georges Louis Leclerc de Buffon was bom into
wealth and devoted his life to scientific labors; he
won recognition as the leading naturalist of his
time. In 1739 he was appointed keeper of the Jar-
din du Roi in Paris (now the Jardin des Plantes),
which he turned into one of the most important
centers of biological research during the 18th cen-
tury. Buffon's lifetime work was a general natural
history in 36 volumes. The first volume dealt with
science in general, the second with man, the next
1 3 with nonhuman mammals ( 1 750-1 767). These
were followed by nine volumes on birds, seven
volumes (1789) supplementary to the preceding,
and the last five on minerals, including fossils.

Treatment of most species in the Histoire Na-
turelle is usually monographic. Gross descriptions,
including measurements and weights, are based
on individuals received in the Jardin du Roi. Geo-
graphic distribution of the species is included with
the description. Habits observed in captivity and
mentioned in the literature are recorded. Anatom-
ical descriptions by Daubenton, Buffon's collab-
orator, are of the skeleton, with soft parts and
tegumentary structures of particular interest.
Complete bibliographic references and synony-
mies, including those to the 10th edition of the
Linnaean Systema Natura, accompany each spe-
cies account.

Buffon drew together much if not everything
known of a species, often an indiscriminate com-
posite of species. Most of the information was
compiled, some of it original. Many life history
notes were received from correspondents, partic-
ularly M. de la Borde, the royal physician resident
in Cayenne, French Guiana. Another correspon-
dent, M. Saint Lurrent of Trinidad, believed he
had solved the mystery of marsupial birth (cf p.
40). At a certain stage of development, he in-
formed Buffon, the embryonic op>ossum crawled
from the uterus through a tube at the end of which
it found a long teat to which it remained attached
until fully developed. An easily verifiable discov-
ery by Daubenton (in Buffon) was that tapirs have
simple stomachs, not the complex ruminant type
claimed by Bajon (above). Buffon reported that
domestic cats kill but do not eat shrew- or short-
tail opossums of the genus Monodelphis. House
cats do indeed kill these animals and usually de-
posit them whole in the middle of the path leading
from the house to the garden.

Most of the illustrations of mammals and all

anatomical drawings of the Histoire Naturelle are
original. A small sampling is reproduced here (figs.
24-25).

Buffon was the first naturalist to recognize re-
gional faunas as such and to discriminate between
Old World species and different but similar ap-
pearing or like-named species of the New World.
He perceived the platyrrhine-catarrhine dichoto-
my of primates, and the phylogenetic distance be-
tween the groups. He further distinguished pre-
hensile-tailed monkeys from non-prehensile-tailed
species, and cebids from callitrichids by their un-
gues and teeth.

Buffon's sense of rivalry with the contemporary
Linnaeus led him to find fault with and cast scorn
on the binomial system used in the Systema Na-
turae. Buffon argued for retention of vernacular
names for species as well as a makeshift vernacular
terminology for generic or supergeneric groups.

Lack of a scientific system of nomenclature in
Buffon's work, and the almost universal adoption
of the Linnaean binomial system by contemporary
and later authors caused the Histoire Naturelle to
be regarded as no better than a layman's encyclo-
pedia of science. It has been republished with many
revisions in many editions and languages. It is
unfortunate that Buffon's important contributions
to life histories, morphology, and evolutionary bi-
ology were largely ignored by Darwin and are little
appreciated today. It seems that the greater luster
credited to Darwin owes much to the dimming of
Buffon's because of his lack of organization and
consistency in his writings.

XV. Faunal Origins and Distribution

Early attempts to explain observed similarities
and differences between Old and New World
mammals all supposedly descended from occu-
pants of Noah's ark, began with the 1 6th century
philosopher and chronicler Acosla and in some
quarters continues to this day.

Jose de Acosta (1539-1600)

Jose de Acosta argued that the animals of the
New World had not been carried there by man.
His evidence indicated that New World man
brought nothing but himself over a land route. The
possibility that animals migrating from the ark
might have crossed the Atlantic Ocean by swim-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

IIJI.IXI /W ,.♦ j^^ ,„

J"/ X-W /-.ff //y

I.B 5.VOOI IV \ ri.(;.\|HK.\IKNl

AvPBi.r. Msnt, i>K :«i»iT.

/■//. 11// /..y

i.A URAXDK riiAivr-soiRis mn- im- i.ajht h" h iu'vaxnb

IK c- \i;; \i

Fig. 24. Mammals figured in the Histoire Naturelle of Buffon: upper left, le saki (= Pithecia pithecia Linnaeus,
male; from Buffon, 1767); upper right, le sagouin singe de nuit (= Pithecia pithecia Linnaeus, female; from Buffon,
1 789); lower left, la grande chauve-souris fer-de-lance de la Guyanne (= Phyllostomus hastatus Pallas; from Buffon,
1 789); lower right, le cabiai (= Hydrochaeris hydrochaeris Linnaeus; from Buffon, 1 764).

FIELDIANA: ZOOLOGY

rtijii j-^ lit

I'fFK l"|-K 111' fllll.l

l>K. MIR IK llf, l.A <.l'^.^NNK

/■/ Win ..^ f-

Xm >U

'CXXl/.fja td>

1,/

1i- B^^Bl

^i»'

Via '■

\*«.ll m

)<': fwfl

■^/

i '^fjS

'' ■'u t

"*■■ ■ r

-,-:5V

^>^^%P

vi.: ' 'utA m

■^3^5^

»«yr'-^->—

:^;

i^?5??S»«—

-ttt;

IK I \1l\RIN NK«i«l

L II* ossr.rsE i>K 1.A noxvi! iir. i.'Ar.ot'Arrr.

Fig. 25. Mammals figured in the Histoire Naturelle of BufTon: upper left, la mouffette du Chili (= Conepatus
chinga Molina); upper right, la grande marte de la Guyanne (= Eira barbara Linnaeus); lower left, le tamarin n^e
(= Saguinus midas Linnaeus); lower right, hyoid apparatus and thyroid cartilage of the throat ofAlouatta seniculus
Linnaeus); from Buffon (1789).

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

ming or island hopping was also dismissed be-
cause, as Acosta pointed out, none of the animals
was known to occur on oceanic islands. The leg-
endary island of Atlantis, which might have been
part of a former transatlantic archipelago, was
treated as fable. Other conjectures discarded,
Acosta resolved that New and Old World northern
continents were or had been connected, or very
nearly approximated, at their polar extremities.
Differences between New and Old World species,
he affirmed, could be explained by the disappear-
ance of connecting Old World populations, mu-
tations among the New World sijecies engendered
by their isolation in different environments, or by
degeneration. No accounting or explanations were
needed for "imperfect" organisms such as rats,
frogs, insects, or vermin in general. These, it was
commonly held, arose spontaneously from decay-
ing matter.

Antonio Vazquez de Espinosa (1560/1575-1630)

Antonio Vazquez de Espinosa agreed with the
explanation of a northern migratory or connecting
route but added (1948, chap. 36), in the awkward
phrasing of Clark's translation, that

near the Straits of Magellan in what is called
Tierra del Fuego, which is still not well
known or explored, and there are numerous
other quarters where the mainland of the
New World could have communicated with
that of the Old, or at least have lain so close
as to afford passage not merely for the peo-
ples who settled the New World, but the
various kinds of animals which live in
them— many of species well known in Eu-
rope and elsewhere, and others peculiar and
unique in the New World, like the Peruvian
sheep [llamas], the guanacos [regarded as the
wild form of llamas], vicuiias and tarugas
[Hippocamelus bisulcus].

Carolus Linnaeus (1707-1778)

Linnaeus played no direct role in the develop-
ment of Neotropical mammalogy apart from pro-
viding scientific names for some species discov-
ered or described by others. His impact on the
scientific world, however, was enormous. Like the

philosophers before him, he believed all species
were created as they are now in one place from
which they spread in search of the habitats for
which they were specially created. This idea of a
staging area as the center of origin and dispersal,
still dominant today in the minds of some students
of Neotropical mammalogy, was critically reex-
amined by Buffon.

Georges Louis Leclerc de Buffon (1707-1788)

Buffon, on comparing faunas of New and Old
Worlds, was impressed by similarities between
some of the species and differences between others.
His explanation for similar-apjjearing sp>ecies, like
that of the chroniclers, was that a land connection
permitted passage of animals from Old to New
World. Differences between New and Old World
species, he suggested, in agreement with the chron-
icler Jose Acosta, could have resulted from de-
generation, environmental pressures, or isolation
of the New World derivatives. On the other hand,
Buffon argued, species peculiar to the New World
or without Old World analogs must have arisen
in situ, an opinion already intimated by Vazquez
de Espinosa.

Buffon was the first naturalist to envision the
mammals of the region as a community or fauna
that might well have originated indeF>endently of
other regional faunas. Noah's ark had no place in
his concept of faunal origins, and he rejected as
too short the scripturally based 6,000-year esti-
mate of the earth's age. Buffon's ideas of organic
evolution and multiple centers of origin were nov-
el and prepared the minds of his and succeeding
generations for the acceptance of Darwinian evo-
lution.

Linnaeus, the arch exponent of the fixity of
species and their origin and dispersal from a single
center, conceived the elements of his binomial sys-
tem as symbols for nailing down his credo. The
system was so good it proved to be the best yet
devised for the expression of genetic relationships
between species and the surest base for the con-
struction of evolutionary sequences in nominate
terms. On the other hand, Buffon, independent of
the religious constraints of his time and evolu-
tionist in thought if not always in words, never
attained the stature of his contemporary for lack
of a competing system, key, code, or standard that
would bring cohesion to his rambling philoso-
phies.

FIELDIANA: ZOOLOGY

Johann Andreas Wagner (1797-1861)

Johann Andreas Wagner, the foremost masto-
zoologist of his generation and author of a mono-
graph on the geographic distribution of mammals,
summarized (1844, p. 13) the three current but
disparate opinions on mammalian origin and dis-
persal. First, all species were created in one and
the same region and spread from there to all cor-
ners of the earth. Second, the species could have
been created in separate localities in the same or
different regions. Finally, each species could have
arisen spontaneously anywhere and developed ac-
cording to its peculiar constraints.

Zoogeographers of the early half of the 19th cen-
tury divided the world into major faunal regions
correlated primarily with climate. Wagner (1844)
separated the earth into four provinces: the Nord-
liche north of 30°N, the Mittlere between BCN
and 30°S, excluding the Australische roughly be-
tween 0°S-55°S and 1 30°W-200''W, and the South
American Magellanische, south of 30°S. The South
American portion of the pantropical Mittlere
Province extended from Mexico southward to
southern Brazil and central Chile. Wagner's de-
scriptions of the provincial faunas included tab-
ulations of their resfjective genera and included
species.

Maximilian Prinz von Wied-Neuwied (1782-1867)

Scriptural constraints were not evident in the
thinking of the field naturalists. Maximilian Prinz
von Wied-Neuwied recognized the limitations of
geographic range as a property of a species.

Johann Jacob von Tschudi (1818-1889)

Tschudi attempted to follow Wied-Neuwied in
defining specific ranges, but nearly all were based
on the presumption that the geographic range of
the species coincided with ecological life zones.
The ecological life zones of Peru described by von
Tschudi on the basis of fauna, flora, and climate,
are the first of their kind for any Neotropical re-
gion.

Charles Robert I>arwin (1809-1882)

The young Darwin also recognized geographic
limitations of distribution in the light of physical

barriers such as mountains and large bodies of
water.

XVI. Inventories to Middle of
19th Century

Systema Naturm of Linnaeus, 1758, 1766

The 1 0th edition of the Systema Naturce pub-
lished in 1758 by the Swedish naturalist Carolus
Linnaeus ( 1 707-1 778), marks the beginning of the
consistent application of his binomial system of
zoological nomenclature. According to the uni-
versally accepted International Code of Zoological
Nomenclature, names for animals published be-
fore 1 758 are not available, no matter how clearly
defined the species. Likewise, zoological names for
species published after 1757 that are not binomial
or do not satisfy all provisions of the Code are not
available. The effect of the Code in practice is that
species without Linnaean names are treated as un-
known to science.

The 1 0th edition of the Systema Naturce lists a
total of 1 72 species of mammals, exclusive of ma-
rine cetaceans, each with its binomen consisting
of a defined generic and defined specific name.
Subsequent revisions of the bases for the names
revealed that some represented more than a single
species, others were duplicates or synonyms, and
a few were equivocal or belonged to unidentifiable
animals. The revisions, however, made no signif-
icant change in the total number of real mam-
malian species known to Linnaeus in 1758.

The 1 2th and last revised edition of the Systema
Naturae by Linnaeus himself, published in 1766,
lists a world total of 208 mammalian species. Ta-
ble 14 compares the relative numbers of world.
Neotropical, and Nearctic genera and species in
the Linnaean 10th and 12th editions of the Sys-
tema Naturce with the totals in Buffbn's Histoire
Naturelle. Cetaceans are omitted because they are
oceanic species known before the discovery of
America.

Primary sources for the definition and naming
of the Linnaean New World species were speci-
mens preserved in the Swedish museums, partic-
ularly the Adolphi Friderici Regis Museum, and
primary bibliographic references. Such references
for the Neotropical mammals were the works of
Marcgraf (1648), Anson (1748), Browne (1756),
and Seba (1 734-1 765). For both Neotropical and

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

Table 14. Number of world and New World mammals known to Linnaeus (1758, 1766) and to Buffon (1750-
1 789) and their percentage of world total.

Author

Date

World total

Neotropica

Nearctica

Linnaeus

1758

35 genera

18(51%)

15(43%)

1 72 species

45 (26%)

23(13%)

Linnaeus

1766

36 genera

20 (55%)

15(42%)

208 species

55 (26%)

28(13%)

BulTon

1750-1789

251 species

78(31%)

47(19%)

Nearctic species, Linnaeus cited Hernandez (1651)
and Edwards ( 1 743- 1751), and for Nearctic species
alone, Catesby (1731) and Kalm (1753).

Histoire Naturelle of Buffon, 1750-1789

In volume 9 of his Histoire Naturelle, published
in 1 76 1 , Buffon estimated a world total of ap-
proximately 200 known mammalian species of
which 1 30 were Old World, 70 New World. When
the sp)ecies of all 1 3 volumes on mammals and the
supplementary volumes are counted, the total is
251, of which 78 are Neotropical, 47 Nearctic.

The greater number of species recognized by
Buffon as compared with those of Linnaeus re-
flects his better knowledge of mammals and wider
use of the available literature. Neither authority
searched the works of the chroniclers for descrip-
tions or figures of New World mammals.

Synopsis Mammalium of Schinz, 1844

The Schinz catalogue of Recent mammals of the
world, published 1844, brings the inventory of
mammalian species to near the cutoff date of this
part of the history of Neotropical mammalogy.
The totals indicate that about 50% of all New World
mammalian species now known had been de-
scribed. The vast majority of the remaining 50%
described since the middle of the 19th century are
as small as or smaller than common tree squirrels.
With respect to Neotropical mammals, by mid-
1 9th century about 90% of known sp)ecies larger
than common tree squirrels had been described.
In contrast, no more than about 1 0% of the smaller
forms, mainly marsupials, bats, and rodents, had
been named.

The Schinz catalogue is summarized in Table
1 5. A first glance at the figures of the first order,
the Marsupialia, may suggest the list is skewed.
Only one marsupial species, Didelphis virginiana,
is known to occur in Nearctica. Schinz recorded

three because the species had been ovemamed at
the time. It is believed, however, that "under-
named" or composite species, as well as over-
named identical species, are more or less evenly
distributed in all three columns. As a result, the
bottom line totals, particularly the percentages, are
fair estimates of the real number of species known
to science at the time of Schinz's compilation. The
percentages have not changed significantly since.

XVII. Summary

Knowledge of Neotropical mammals from
1492 to the mid- 17th century was mainly an ag-
gregation of anecdotes often riddled with myth,
folklore, and untested generalizations. Eurojjeans
identified New World species with similar-ap-
pearing or similar-behaving Old World species and
used the same names for them. Descriptions of
mammals were usually comparisons with familiar
European animals; measurements, rarely given,
were rough estimates. Habitat when mentioned
was usually on the order of "forest," "plain," or
"river."

Descriptions of animals often included use as
food or pets, medicinal merits, or value of rawhide
or bones in the manufacture of artifacts. Habits
were usually described in terms of reactions to
man when hunted or in captivity, or as harmful
or benign to his person or property. Mammals—
the term had not yet been concocted— were the
hairy beasts of the earth. Whales and manatees
were fish and could be eaten on Fridays. Bats were
something else, mostly vampires; mice and other
small mammals were vermin, in a class with frogs
and cockroaches.

Mammal collecting during this period was gen-
erally limited to capture of live individuals for
domestication, as pets, or for exhibition in zoos,
circuses, or fairs. Dead animals were sometimes
skinned and stuffed or bottled in brandy. The
crudely prepared or pickled specimens, if not live

HELDIANA: ZOOLOGY

animals, often served as models for the woodcut
drawings of early treatises on natural history. Some
specimens were purchased for museums or cabi-
nets of collectors, including those of Linnaeus,
King Frederick Adolph of Sweden, Reaumur of
Paris, the King of France (Jardin du Roi), or the
shelves of the Dutch pharmacist, Albert Seba. Most
of the Neotropical specimens probably originated
in the South American possessions of Holland and
France.

The few crude attempts at classification of mam-
mals during the 16th and 17th centuries were hard-
ly more than random arrangements equivalent to
shopping lists. Species, being individually created
kinds, were unrelated to other created kinds, or
simply arose spontaneously from putrefying mat-
ter.

The scientific study of mammals, or mammal-
ogy, of the Neotropical Region began with the ex-
plorations of northeastern Brazil by Georg Marc-
graf and culminated with the publication in 1648
of his Historic Rerum Naturalium Brasilia. His
accounts of the included 32 sp)ecies of mammals
reveal the glimmer of an attempt at natural group-
ings of kinds or the beginnings of a classification
of Neotropical mammals. Insofar as is known,
none of MarcgraPs animals were preserved. Lin-
naean names for the species of the Historice were
based on bibliographic references to their descrip-
tions and figures (cf. fig. 2, table 1).

The first expedition to the Neotropical Region
actually committed to the collection and perma-
nent preservation of mammals (and other objects)
for scientific study was the Brazilian Viagem Fi-
losofica, 1783 to 1792, conceived by the Portu-
guese government and conducted by the Brazilian-
bom naturalist Alexandre Rodrigues Ferreira. The
large number of specimens gathered by the ex-
pedition was deposited in Lisbon's Museu d'A-
juda. The specimens of monkeys that had been
carried away to the Paris Natural History Museum
were studied by the French scientist Etienne Geof-
froy St.-Hilaire. His descriptions were published
without reference to source of material.

Alexander von Humboldt followed on the heels
of the Viagem Filosofica with his explorations of
northwestern South America from 1799 through
1802. His expedition was highly successful and in
scope has rarely been equaled by other "one-man"
surveys of a large portion of the South American
continent. The personal narrative of his travels
inspired successive naturalist-travelers, most no-
tably the explorers of Brazil, Spix and Martius,
Maximilian Wied-Neuwied, and Johann Natterer.

Table 1 5. Number of world, Neolropic, and Nearc-
tic species (subspecies) of mammals known to Schinz
( 1 844); species common to both regions are included in
both.

Order

World

Neotropica

Nearctica

Marsupialia

138

31(22%)

4(3%)

Insectivora

114

2(2%)

21 (18%)

Chiroptera

326

110(34%)

21 (6%)

Primates

281

73 (26%)

0(0%)

Edentata

24 (77%)

0(0%)

Camivora

303

58(19%)

41 (13%)

Pinnipedia

1 1 (28%)

2(5%)

Sirenia

2 (40%)

1 (20%)

Perissodactyla

2(9%)

0(0%)

Artiodactyla

186

1 1 (6%)

12(6%)

Lagomorpha

4(8%)

14(27%)

Rodentia

563

152(27%)

104(18%)

Cetacea

2 (50%)

0(0%)

Proboscidea

0(0%)

Hyracoidea

0(0%)

Totals

2,072*

482 (23%)

220(11%)

* The estimated number of species in 1972 (Hersh-
kovitz, p. 332, table 3) is Neotropica 810, Nearctica 442
or an approximate doubling since 1 844 in both regions,
with a slightly greater increase (less than 2%) in Nearctica
relative to Neotropica. Increase since then has been al-
most exclusively Neotropical.

Later there was von Tschudi, who traveled in Peru;
d'Orbigny, who journeyed in Patagonia but did
his finest and most lasting work in Bolivia on a
scale almost equal to that of Humboldt's; and
Darwin, who voyaged around the southern half of
South America and the Galapagos Islands in
H.M.S. Beagle.

The brothers Schomburgk, motivated by Hum-
boldt's trip up the Rio Orinoco to its connection
via the Casiquiare Canal with the Rio Negro trib-
utary of the Rio Amazonas, completed the trajec-
tory by plotting the course of the upper Rio Negro
to its connection with the Casiquiare. Their ex-
plorations of the British Guianas and bordering
parts of Brazil and Venezuela yielded the first
large collection of Guianan mammals, all depos-
ited in the Berlin Natural History Museum.

Chilean mammals became fairly well known
through the reports of Molina (1782), Poeppig
(1836), and Gay (1847). The mammals of Para-
guay, their distribution, habits, or biology in gen-
eral, became better known through the efforts of
Felix de Azara than those of other Neotropical
countries.

The 200-year period from MarcgraPs ( 1 648) re-
port to the last of those of Schomburgk (1 848) was
one of survey and inventory of South American
mammals. The published reports and personal

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

narratives of travel provided much reliable data
on geographic distribution, habitat, life histories,
ecological backgrounds, itineraries and maps of
the expeditionary routes, and stopping and col-
lecting localities. Descriptions of the collected
mammals, most of them by the naturalist-trav-
elers themselves, were often based on skeletal,
dental, and soft parts in addition to purely tegu-
mentary characters. Their classifications were pu-
tatively natural groupings on the ordinal, family,
and, as a rule, the generic levels. The prevailing
belief in the biblical version of creation and fixity
of species, not confessed in writing, did not blind
systematists of the period to evident relationships
between species and their clusterings into supra-
specific groups. Descriptions of species were,
nevertheless, typological. Subspecies or geograph-
ic races were, at best, vaguely conceived but de-
scribed as species. The infrequent or rare use of
trinomials was accidental or equivocal and not
certainly intended for a clearly defined geographic
race. The term usually used for deviates firom
"types" was "variety."

Controversies regarding origin of species or fau-
nas centered on where, not how. Philosopher-
chroniclers of the first era accepted Noah's ark
literally as the one place of origin and disjiersal of
the Recent fauna. Acosta may have been the first
to suggest the former existence of intercontinental
connections for passage of Old World animals into
the New World.

More and better knowledge of the world's fauna
during the second era revealed the weaknesses or
fallacy of the ark dogma. Staunch creationists such
as Linnaeus pointed instead to a vaguely located
region as the place from which all species dispersed
to occupy predestined habitats for which they had
been created. Other authorities like Buffon argued
for multiple centers of origin, with sp)ecies origi-
nating in the habitats for which they were adapted.
Darwin also believed in multiple places of origin,
or faunal regions separated by geographic barriers
but with some trepidation. The belief in multiple
creations was heretical.

Inconsistencies between religious dogma and
realities did not prevent Wied-Neuwied from rec-
ognizing the geographic range of a species (or sub-
species) as a property of that species. Another
advance beyond scriptures was the concept of eco-
logical life zones contributed by von Tschudi, who
plotted them for Peru on the basis of plants, an-
imals, and climate.

The total of named Neotropical sjjecies of mam-
mals counted ft-om 1758, the year of publication

of the 10th edition of the Linnaean Systema Na-
tura and starting date of zoological nomenclature,
to mid- 1 9th century, exceeded by far that of the
Nearctic region and any other equivalent area of
the world. Neotropical mammals were also better
known than those of other continents except west-
em Europe.

By mid- 1 9th century, about 90% of currently
known Neotropical mammalian species larger than
common tree squirrels had already been described,
but no more than about 10% of the smaller forms.

The great number and variety of Neotropical
mammals (and animals generally) known to sci-
ence by mid- 19th century and the accumulated
knowledge gained from study of living and pre-
served specimens in field and laboratory, much of
it contributed by Charles Darwin, helped pscvt the
way to the Darwinian revolution of the next half
century.

XYIII. Acknowledgments

I am indebted to Benjamin W. Williams, As-
sociate Librarian and Librarian of Rare Books,
Field Museum of Natural History, for p)ermission
to consult at pleasure in the Museum's Mary W.
Runnells Rare Book Room the books needed for
writing this article; and to Bruce D. Patterson,
Robert M. Timm, Ronald H. Pine, Debra Mos-
kovitz, and J. A. Gagliano for reviewing the manu-
script. Map>s shown in Figures 11, 12, and 2 1 were
prepared by the author with assistance of Mary
Anne Rogers from accounts of the travelers cited
and other sources. Photographic reproductions of
the figures are by Field Museum of Natural His-
tory Staff Photographer Ron Testa. Technical
Assistants Barbara Brown and Mary Anne Rogers
typed the manuscript and contributed in other ways
toward its completion.

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368.

HELDIANA: ZOOLOGY

A New Superfamily in the Extensive Radiation
of South American Paleogene Marsupials

Rosendo Pascual and Alfredo A. Carlini

ABSTRACTS

Significant new mammals have been recovered from the Colhuehuapian mammal-bearing
beds (latest Oligocene) exposed in the Gaiman region of Patagonia (Chubut Province, Argentina).
Some fragmentary mandibles and isolated teeth belong to a new genus and species, Patagonia
peregrina. The mandibular and dental specializations of this form are so distinctly convergent
on those of some fossorial rodents that it is regarded as a distinct clade of South American
marsupials. It represents the type of a new family, Patagoniidae, which is assigned to a new
superfamily, Patagonioidea, which represents a natural evolutionary group in the same sense
as other superfamilies of marsupials recognized by Simpson. Its systematic position within the
superorder Marsupialia awaits comprehensive analysis of those enigmatic marsupials (Groe-
berioidea and Argyrolagoidea) apparently most closely related to it.

Varios nuevos y significativos mamiferos han sido recogidos de capas mamaliferas del Col-
huehuapense (Oligoceno tardio) expuestas en la region de Gaiman, Patagonia (Chubut, Argen-
tina). Algunos fragmentos mandibulares y dientes aislados pertenecen a un nuevo genero y
especie, Patagonia peregrina. Esta forma presenta especializaciones mandibulares y dentarias
tan distintamente convergentes hacia las de albunos roedores cavadores que es considerada
como un distinto clado de marsupiales sudamericanos. Representa el tipo de una nueva Familia,
Patagoniidae, que es asignada a la nueva Superfamilia Patagonioidea, porque representa un
grupo evolutivo natural como los de otras Superfamilias de marsupiales reconocidas por Simp-
son. Su posicion sistematica dentro del Superorden Marsupialia depende del analisis integrado
de aquellos marsupiales enigmaticos (Groeberioidea y Argyrolagoidea) aparentemente mas
estrechamente relacionados a el.

Novos mamiferos foram recuperados dos leitos de Colhuehuapian (do alto Oligoceno), ex-
postos na regiao de Gaiman, Patagonia (Provincia de Chubut, Argentina). Fragmentos man-
dibulares e dentes isolados pertencem a um novo genero e especie, Patagonia peregrina. As
especializa9oes mandibulares e dentais encontradas sao tao claramente convergentes as de alguns
roedores fossorios, que esta forma e considerada uma classe distinta de marsupiais sulameri-
canos. A especie representa o tipo de uma nova familia, Patagoniidae, a qual e designada a
uma nova superfamilia, Patagonioidea, por formar um grupo evolutivo bem definido, como o
formam as outras familias de marsupiais, reconhecidas por Simpson. A posi9ao sistematica
dos Patagonioidea, dentro da superordem Marsupialia, aguarda uma analise mais compreensiva
dos marsupiais ainda enigmaticos (como Groeberioidea e Argyrolagoidea) aparentemente e seus
relativos mais proximos.

From the Division Paleontologia Vertebrados, Museo
de La Plata, Paseo del Bosque, 1900 La Plata, Argentina;
and CONICET, Argentina.

PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 99

Introduction

The taxon described in this paper is yet another
example of the great adaptive radiation and dis-
persal of marsupials in South America. It repre-
sents a second line of marsupials that is convergent
on the rodent adaptive zone (cf Groeberioidea—
Patterson, 1952; Simpson, 1970c; Clemens &
Marshall, 1976). However, it is distinct from pre-
viously named forms, not only phylogenetically
but also ecologically.

The new form does not suggest that marsupials
attained the breadth and diversity of rodent ad-
aptations, but it does show that marsupials oc-
cupied the rodent adaptive zone in previously un-
imagined ways. This new marsupial indicates that
marsupial radiations in South America were al-
most as broad and reached as great extremes as
those in Australia. The find is consistent with the
view that "A complete record of South American
marsupials would certainly include a large number
of taxa, probably some of high categorical rank,
now unknown" (Simpson, 1970a, p. 59). This and
other forms recently found in northwestern Ar-
gentina (Pascual, 1980a, b, 1981, 1983) validate
Simpson's prophetic suggestion that ". . . major
parts of marsupial evolution were occurring in areas
and facies inadequately sampled, if at all, by the
known fossil deposits and the collections so far
made" (Simpson, 1970a, p. 58). These deposits
indicate the value of applying new sample-col-
lecting techniques at mammal-bearing localities
that are supposedly well known; it is only neces-
sary to find new, appropriate facies.

The new ecological type from the Paleogene pro-
vides evidence to support Gould's (1983) view of
"early experimentation, later standardization,"
with a consequent reduction in diversity. As in
therians (Pascual et al., 1985) the diversification
of South American marsupials took place princi-
pally in the Paleogene.

Measurements reported in Table 1 are depicted
in Figure 3 and are given in millimeters. The ab-
breviation MACN CH is used for the Museo Ar-
gentino de Ciencias Naturales "Bernardino Ri-
vadavia" (Buenos Aires), Coleccion Chubut.

Classification

Superfamily PATAGONIOIDEA nov.

The only known family of this taxon is the Pat-
agoniidae. The superfamily is sufficiently charac-

terized by the diagnoses of that family and its only
known species. Justification for its superfamilial
rank is given in a later section on affinities.

Family Patagoniidae nov.

Type— Patagonia gen. nov. The only known ge-
nus.

Known Distribution— Late Oligocene. Col-
huehuapian from Central Patagonia (Chubut
Province, Argentina).

Diagnosis.— Small marsupials with the same
reduced number of lower teeth as the Groeberi-
idae, but with a different dental formula: 1.1.0.3.
Open-rooted and rodent-like lower incisor, oval
in cross section, strongly curved, although not as
much as in the Groeberiidae, and with the intra-
alveolar portion differently arranged. The incisor
extends lingually along the ventral border of the
mandible to the root of the inflected crest beneath
the last molar, where it forms a prominence sim-
ilar to that of hystricognathous rodents, but ven-
trally. Lower canine smaller, procumbent, appar-
ently incisor-like and closed-rooted, separated from
the cheekteeth by a short, crested diastema at al-
veolar level. Lower cheekteeth rectangular in cross
section, decreasing in size posteriorly, hypselo-
dont, rootless, wholly surrounded by enamel, and
slightly curved, with the concavity forward. Hor-
izontal ramus of the mandible short and deep, with
the highest part posterior, beneath the masseteric
fossa, where the body of the mandible becomes
strongly convex and inflected; deep pterygoid fos-
sa, limited ventrally by a flange like that found in
Argyrolagidae and in some Australian marsupials
(e.g., Macropodidae); strong, salient coronoid pro-
cess; masseteric fossa relatively deep but reduced,
dorsally situated with a prominent masseteric crest;
subvertical symphysis unfused, with nearly smooth
symphyseal surfaces.

PATAGONIA gen. nov.

Etymology- From Patagonia, its geographical
record.

Type— Patagonia peregrina sp. nov.

Known Range and Diagnosis— Same as that
of the family.

Patagonia peregrina sp. nov. Figures 1-3

Etymology— From Latin peregr inns, strange or
rare.

100

FIELDIANA: ZOOLOGY

«f *^ «

Fig. 1 . Patagonia peregrina gen. et sp. nov. A-B, Stereopairs of MACN CH-865, a fragment of a right mandibular
ramus with m,.,: A, occlusal view; B, posterior view; C-D, X-ray of fragments of two right mandibular rami with
i,, alveolus of c,, and m,., complete (C, holotype; MACN CH-869) and with alveoli of i,, and c,, and m,., complete
(D, MACN CH-865). Graphic scale = 2 mm.

HoLOTYPE-MACN CH-869 (fig. 2A-B). Frag-
ment of right mandibular ramus with three cheek-
teeth, intra-alveolar portion of the incisor, and
alveolus of the canine.

Hypodigm — Holotype and the following:

MACN CH-864, part of right mandibular ramus
with first and second cheekteeth, part of alveolus
of the third, and part of alveoli of incisor and
canine; MACN CH-865, part of right ramus with
three cheekteeth and alveoli of the incisor and

PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS

101

'C,

alveolus
alveolus

alveolus

Fig. 2. Patagonia peregrina gen. et sp. nov. A-B, Stereopairs of MACN CH-869 (holotype), a fragment of a right
mandibular ramus, with i,, alveolus of c,, and m,.,: A, labial view; B, lingual view. Graphic scale = 2 mm. C,
Stereopairs of MACN CH-867, a fragment of a left mandibular ramus, with alveoh of i,, c, and mj, and m,_2; occlusal
view.

canine; MACN CH-866, part of left ramus with
the second and third cheekteeth, and part of al-
veoli of the first cheektooth and the incisor, MACN
CH-867, part of the left ramus with the first and
second cheekteeth, and part of alveoli of the third
cheektooth, incisor, and canine; MACN CH-868,
part of the right ramus with three cheekteeth and

alveolus of the incisor; MACN CH-870, part of
left ramus with first and second cheekteeth and
part of the alveolus of the third; MACN CH-874,
part of right ramus with the second cheektooth,
alveoli of the first and third cheekteeth, and part
of the alveolus of the incisor; MACN CH-875,
part of right ramus with the second and third

102

HELDIANA: ZOOLOGY

cheekteeth and part of the alveolus of the incisor;
and MACN CH-876, three isolated upper(?)
cheekteeth.

Horizon and Locality— Both the holotype and
the hypodigm come from the Trelew Member of
the Sarmiento Formation (see Mendia & Bayar-
sky, 1981) and are Colhuehuapian (Late Oligo-
cene) in age. Apparently they were found in the
upper unit, exposed on the south side of the Chu-
but River valley, Chubut Province, Argentina
(Central Patagonia; see Fleagle & Bown, 1983, pp.
242-244). Quite probably this corresponds to
Simpson's "stratum F of Fig. 1," which is part of
his "Trelew beds" (= "Trelewense"). The material
was recovered by O. E. Donadio, M. Soria, J. G.
Fleagle, and T. M. Bown (see Fleagle & Bown,
1 983) through dry-screening local deflation lag de-
posits.

Diagnosis— The only known species of the fam-
ily.

Description— Dentition— See Figures 1 A,C-D;
2-3. Each side of the lower jaw has one fully ro-
dent-like gnawing incisor, only incompletely pre-
served in the holotype; it is posteriorly bordered
by a relatively shallow and conical alveolus (the
tooth being absent in all specimens at hand) sep-
arated from the medial one by bone and set at a
relatively oblique angle (figs. IC-D; 2C). Homol-
ogies of these teeth are uncertain, but the rodent-
like medial tooth is surely an incisor, designated
for description as i,. The shape and disposition of
the second alveolus agrees with the procumbent
canine of Polydolopidae (Epidolopinae; cf Paula
Couto, 1952, 1961; Pascual & Bond, 1981) and
Prepidolopidae(Pascual, 1980b, fig. 2D-E); it thus
appears that this tooth is c,. This alveolus is fol-
lowed by a short diastema at alveolar level, then
three cheekteeth, all rectangular in cross section
(with some differences among them) and in close
approximation, forming a molariform series. They
are surrounded by enamel on all sides and are not
strictly lobate, nor are the trigonid, talonid, or
original cusps clearly indicated, as occlusion with
the uppers was mediated through practically flat
areas. The dentine forms a shallow basin sur-
rounded by the highest enamel layer, which is
slightly higher on the lingual side. There is a slight-
ly deeper anteroposterior wear groove, extending
from the anterolabial comer to the posterolingual
one (fig. lA). Grinding involved a longer propal-
inal movement and a shorter ectental stroke. The
homologies of these teeth with the more numerous
ancestral series cannot be determined. Plausibly

/r-

^[ii^dEJE).--]^

Fig. 3. Patagonia peregrina gen. et sp. nov. Outline
of a right mandibular ramus fragment, with alveoli of i,
and c,, and m,., complete (MACN CH-865), showing
the measurements of Table 1 . A, Labial view; B, occlusal
view; C, cheekteeth series (m ,.3); D, lingual view. Graph-
ic scale = 2 mm.

they are homologous with those typically desig-
nated m,_3 in marsupials and are so designated
here, yielding the lower dental formula 1.1.0.3.
which is provisionally homologized as i,, c,, m,.,.
However, many specialized marsupials from the
South American fossil record show tendencies (1)

PASCUAL &. CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS

103

Table 1 . Dimensions of specimens of Patagonia peregrina gen. et sp. nov. (see fig. 3 for measurement references).

Dimensions

Specimen

MACN CH-864
MACN CH-865
MACN CH-866
MACN CH-867
MACN CH-868
MACN CH-869
MACN CH-870
MACN CH-874
MACN CH-875

3.68 4.08 1.40

1.12

4.08

3.60

1.00

0.96

1.40

1.20

1.08

1.12

0.96

5.40

1.00

1.08

1.00

0.92

1.20

1.00

0.96

1.00

1.04

1.00

0.92

1.36

1.08

1.00

0.96

0.92

5.12

1.40

1.00
1.08

1.00

1.00
0.96

4.80

1.28

1.16

1.24

1.08

6.76

2.80

3.00 4.60

2.20 4.40

6.20

1.80
1.60

1.80

2.40

to elongate or modify either the pj (Polydolopidae
and Parabderitini caenolestids; see Marshall, 1980)
or the m, (Abderitinae and Palaeothentinae caen-
olestids), and (2) to reduce (e.g., Caenolestidae) or
lose (Polydolopidae) the m4. All teeth except the
c, are completely hypselodont and rootless. The
cheekteeth are slightly curved, with the concavity
forward (fig. IC-D).

Incisor ("/J— Incompletely preserved in the ho-
lotype (fig. 2A-B). The anterior end is broken, but
the posterior end is unaltered, showing an open
pulp cavity with no sign of root formation. It is
as elongate and curved as in the most specialized
caviomorphs (e.g., Ctenomyidae), although not
upcurved posteriorly. The extra-alveolar part
would have been nearly vertical, although not so
much as in the Groeberiidae (cf. fig. 4B,D). The
tooth extends along the ventral border lingually,
first beneath the m, and then lingually to far below
the mj, terminating where the ventral border be-
comes an inflected flange (MACN CH-865, fig. 1 B)
reminiscent of kangaroos; the base of the alveolus
shapes a superficial prominence similar to that of
hystricognathous rodents, although enveloped by
the inflected ventral border. It is approximately
oval in cross section, with the long axis oriented
dorsoventrally and with a flatter medial surface.
Apparently enamel covers the entire tooth, but a
noticeably heavier layer extends as a ventral band.

Canine fie J— None of the specimens in the hy-
podigm include this tooth, but its alveolus (figs.
1 C-D; 2C) is oblique, tapered, and relatively shal-
low, indicating a closed-rooted tooth; its oblique
orientation suggests that the occlusal apex was ap-
pressed against the occlusal tip of i,.

First Molar (m,)—The first molar is separated
from the c, by a crested diastema (fig. 1 A) as long
as m,. It is the largest cheektooth, almost rect-
angular in cross section, with the longer lateral

faces slightly concave; the lingual face sometimes
bears a very shallow groove along the intra-alveo-
lar portion. The anterior face is convex, occasion-
ally somewhat pointed; the posterior face is almost
flat, forming angles with the lateral faces slightly
greater than 90°.

Second Molar (^wj— The second molar is irreg-
ularly quadrate, with the lateral faces slightly con-
vex and the anterior and posterior ones flatter. The
posterolingual angle is less than 90°, whereas the
others are almost 90°. Its width is similar to that
of m, (fig. lA).

Third Molar (m^J—The third molar is the small-
est cheektooth, being subtriangular in cross section
rather than square. The anterior face is slightly
convex, lingually flatter, and labially more strongly
curved; the labial face converges posterolingually
with the lingual face, forming a rounded pillar
rather than a well-defined posterior face (fig. lA).

Mandible— No nearly complete mandible is
known, but parts of the horizontal ramus and base
of the coronoid process are known. These parts
indicate the mandible is extremely short and deep,
like that of Groeberia minoprioi, although very
different in other respects (cf. fig. 4B,D). The sym-
physis is subvertical and unfused, with a nearly
smooth symphyseal surface (i.e., normal in struc-
ture instead of fused and forming the odd medial
posterior projection peculiar to Groeberiidae). The
depth of the mandible increases abruptly toward
the mj. The deep masseteric fossa appears to be
F)eculiarly confined to a dorsal position, as the
masseteric crest is situated at a level between the
alveolar rim and the lowest level of the rounded
and inflected ventral border (figs. 1 B; 2 A). A sim-
ilar condition is found in some Abderitinae caen-
olestids (e.g., Parabderites bicrispatus; Marshall,
1976, fig. 8a), although in P. bicrispatus the man-
dibular body is not as deep and the alveolar border

104

FIELDIANA: ZOOLOGY

not as extensively inflected. The coronoid process
has its root beneath the m,, forming a strong, sa-
lient lamina (known only by its root), so that a
conspicuous diagonal valley is formed between the
coronoid and the alveolar border behind m, (fig.
lA); a similar structure is present in Groeberia
minoprioi (see Patterson, 1952, p. 41); the valley
is open labially and lingually limited by a prom-
inence similar to that present in Australasian Po-
toroinae.

In many respects this strong, salient, ascending
ramus and correlated features are reminiscent of
highly fossorial caviomorphs, such as burrow-in-
habiting Ctenomyidae. Although the mandibular
angle is not preserved in any of the specimens, it
probably was inflected, as suggested by the inflec-
tion of the ventral border, beginning at the level
of mj, which defines a lingual flanged crest (figs.
IB; 2B) similar to that producing the extremely
inflected angle in the Macropodidae. This lingual
ventral flanged crest seems to be the lingual border
of an expanded and relatively deep pterygoid fos-
sa, resembling that of argyrolagids (see Simpson,
1 970a). There is a relatively large alveolar foramen
within the pterygoid fossa, level with the alveolar
border and within a pit (fig. 3D), and a mental
foramen beneath the anterior face of m, at the
level of the alveolus of i, (fig. 2 A).

Affinities

As in the case of Groeberia (see Simpson, 1 970c),
the conclusion that Patagonia is a marsupial rests
on a combination of definite, negative, and indi-
rect evidence. The most definite evidence for its
being a marsupial is the inflected ventral border
of the mandible and probably the related inflected
angle. This evidence alone is inconclusive, as a
few marsupials lack an inflected angle and a few
placentals have one. However, no known placental
has such an extended and upturned flange-shaped
inflection, and even in marsupials it is rarely so
well developed (e.g., Groeberiidae [Patterson,
1952]. Argyrolagidae [Simpson, 1970a,b], and the
Australasian Macropodidae). Unlike Groeberia,
Patagonia has other characters supporting its mar-
supial affinities, namely the lower procumbent in-
cisor-like canine. In the Epidolopinae (Pascual &.
Bond, 1981) there are three procumbent lower
teeth, the third being unquestionably the canine.
Within the more advanced Polydolopidae (Poly-
dolopinae), there are one or three procumbent
lower teeth; in the latter case, evidence suggests

they consist of two incisors and a canine, the me-
dial incisor being quite reduced and the canine
well developed, single, and closed-rooted.

As in Groeberia the negative evidence is that
Patagonia has no features precluding its reference
to the Marsupialia. It does exhibit characters mak-
ing reference to any Eutheria highly improbable.
Its habitus is rodent-like, but its two differentially
procumbent lower teeth rule out reference to the
Rodentia. While the incisor is rodent-like in shape,
it is oriented differently than that in rodents, ex-
tending along the ventral border of the horizontal
ramus, first below the m,, then lingually to other
molars, without curving upward. It apparently
shapes the ventral border of the mandible. In ad-
dition, the short diastema extends at the level of
alveoli. Among known rodents, only Paramyidae
and Ischyromyidae developed diastemas at the al-
veolar level, but even in these groups, the incisor
extends as in other rodents, not as in Patagonia.
A more-or-less rodent-like habitus was also char-
acteristic of some notoungulates, especially among
Typotheria and Hegetotheria, but insofar as known
not so extreme in development as in Patagonia.
Neither the enlargement of the incisor nor the re-
duction of the cheekteeth is known in any prim-
itive Paleocene notoungulates or in other South
American "ungulates." Even later rodent-like no-
toungulates were much less specialized than the
Oligocene Patagonia. South American marsupials
diverged very early into unique evolutionary lin-
eages (see Simpson, 1970a-c, 1971, 1 980; Pascual,
1980a,b, 1981; Paula Couto, 1979; Reig, 1981).

Patagonia peregrina is unquestionably a mar-
supial because its unique and diagnostic combi-
nation of characters are unknown in any eutherian.
Nevertheless, it could be regarded as another of
the extinct South American mammals considered
by some as incertae sedis and by others as a tertium
quid with regard to the eutherian-marsupial di-
chotomy (McKenna, 1980; Reig, 1981). However,
the marsupial affinities of other peculiar fossil
mammals from South America remain unques-
tioned, despite weaker support than that offered
here for Patagonia. For example, the basis for con-
sidering the Polydolopidae as marsupials is the
combination of an inflected mandibular ramus,
palatal vacuities, and a cheektooth formula of

1-3 1-4

P— -- and ^-r~z- These characters were formerly

used to exclude the polydolopids from the Allothe-
ria. But, as these characters are present in prim-
itive therians outside South America, their diag-
nosis of marsupials can be considered an "act of

PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS

105

Fig. 4. Labial (1) and occlusal (2) outlines of mandibles, showing the diflFerent development of incisor. A, Ar-
gyrolagus parodii Rusconi; B, Groeberia minoprioi Patterson; C, Proargyrolagus bolivianus Wolff, D, Patagonia
peregrina gen. et sp. nov. Graphic scale = 2 mm.

faith based on . . . geography and stratigraphic po-
sition rather than on . . . biology" (McKenna, 1 980,
pp. 58-59). We beheve that assignment of poly-
dolopids to marsupials represents the most par-
simonious conclusion.

Like the newly described Proargyrolagus boli-
vianus (Wolff, 1984), Patagonia peregrina is
another peculiar marsupial that appears in the fos-

sil record without known ancestors (see Simpson,
1970c, p. 16) only to vanish again soon afterward:
Groeberiidae (Divisaderan Age, Late Eocene); Pat-
agoniidae (Colhuehuapian Age, Late Oligocene);
Necrolesiidae (Santacrucian Age, Early Miocene);
Argyrolagidae (Huayquerian to Uquian Ages, Late
Miocene to Early Pleistocene). We believe there
are cedent reasons to think of Proargyrolagus bo-

106

nELDL\NA: ZOOLOGY

livianus Wolff, 1984, described as a Deseadan ar-
gyrolagid, as possibly representing a distinct fam-
ily of Argyrolagoidea.

This raises the question of the position of Pat-
agonia among the varied ranks of South American
marsupials. The previous descriptions and illus-
trations demonstrate that Patagonia peregrina has
many peculiarities that are rare, differently devel-
oped, or completely absent in other marsupials (cf
fig. 4). The most striking of these are:

1 . Mandible extremely short and deep, with un-
fused subvertical symphysis, dorsally posi-
tioned masseteric fossa, ventral border in-
flected at level of the mj, enveloping there
the alveolus of the incisor.

2. Presence in each ramus of mandible of one
rodent-like rootless incisor that extends lin-
gually along ventral border of mandible to
below the m,.

3. Presence in each ramus of one procumbent
canine, single- and closed-rooted, scarcely
separated from the incisor and with the oc-
clusal apex probably appressed to the inci-
sive apex.

4. Three rectangular and continuously growing
cheekteeth arranged in close sequence.

These and other less striking characters under-
score the unique specializations of Patagonia per-
egrina, leading to its assignment to a new family,
Patagoniidae. But the distinctive combination of
characters in the Patagoniidae identify it as a dis-
tinct evolutionary group, that is, a different clado-
genetic unit. Simpson (1945, 1970a, 1980) des-
ignated natural evolutionary groups of marsupials
as suF>erfamilies. Following this line of reasoning,
Patagoniidae should be allocated to a new super-
family, the Patagonioidea.

What are the affinities of this new superfamily
to other superfamilies within the superorder Mar-
supialia? Any discussion of its affinities depends
on the systematics of other taxa, many of which
are problematic. The systematics of fossil and ex-
tant South American marsupials, including the
merits of recognizing Marsupialia as a superorder,
are discussed by Simpson (1970a, 1971) and Pas-
cual (1980b).

The majority of South American marsupials
represent the order Polyprotodonta; this is roughly
equivalent to Ride's ( 1 964) Marsupicamivora, but
also includes Ameghino's Paucituberculata (see
Pascual, 1980b; contra Kirsch, 1977a,b; Reig,
1981). There is as yet no compelling argument to

include any South American families within the
Australasian order Diprotodonta (Reig, 1981), de-
spite some suggestions to the contrary (e.g., Pas-
cual & Herrera, 1 973, 1 975). While the allocations
of these groups seems unambiguous, the positions
of most remaining groups (e.g., Argyrolagidae,
Necrolestidae, and Groeberiidae) remain uncer-
tain. With some reservation, Kirsch (1977b) in-
cluded the Necrolestidae in the polyprotodont
Borhyaenoidea (as did Patterson, 1958), and the
Groeberioidea and Argyrolagoidea within the
Paucituberculata. Independently, Clemens and
Marshall ( 1 976) also treated these animals as mar-
supials, recognizing each as superfamilies: Argy-
rolagoidea, Necrolestoidea, and Groeberioidea.
Like Simpson, they made these assignments with
disclaimers that the interrelationships of these
groups were far from clear.

Reig (1981, p. 60) not only questioned whether
the Argyrolagidae (his Microtragulidae) were mar-
supials, as none of its known characters are ty-
pologically diagnostic, but conjectured probable
affinities to the Anagalida. Further, without rig-
orous analysis, he suggested that the Argyrolagidae
could be treated as an independent order, pro-
posing the name Argyrolagida. He concluded that
only more intensive study or additional records
could substantiate allocation of this order to the
Metatheria or the Eutheria.

Remains of Patagoniidae exhibit a unique mo-
saic of characters, some of which are absent or
differently developed in Groeberiidae and Argy-
rolagidae. Despite their similarities, each of these
taxa apF)ears prima facie to represent indep)endent
evolutionary trends. To assess their interrelation-
ships, common and distinctive characters of each
must be carefully weighed. Remains of Argyro-
lagoidea obtained in the same horizon and locality
as the hypodigm of Patagonia peregrina should
be particularly useful in this regard and are now
under study. Ordinal and subordinal allocation of
the Patagonioidea await this more comprehensive
analysis. Known representatives of this taxon are
so highly derived, as is the case with other peculiar
marsupials, that their relationships to other mar-
supial groups are obscure and can only be clarified
by an expanded record of earlier forms.

Ecology and Historical Biogeography

Biological inferences of Patagonia are necessar-
ily limited to the mandibular fragments thus far

PASCUAL & CARLINL NEW SUPERFAMILY OF PALEOGENE MARSUPIALS

107

known. These demonstrate unique characters
among marsupials, Hving or extinct, which are ob-
viously related to a particular mode of life. No
known eutherian possesses such mandibular fea-
tures. Superficially it is similar to Groeberia, both
being rodent-like marsupials: each has a short and
deep mandible with a single enlarged, open-rooted
incisor, deeply extended along the mandible, with
the extra-alveolar part apparently nearly vertical.
These represent functional not phylogenetic sim-
ilarities, as similar states were attained by different
routes: in Groeberia this tooth extended within an
odd medial posterior projection of the symphysis,
whereas in Patagonia the intra-alveolar portion is
truly rodent-like, in being extended along the hor-
izontal ramus (cf fig. 4B,D). No doubt both were
powerfial gnawers as the lower incisor worked al-
most vertically, much more so than in most ro-
dents. The unknown face and snout of Patagonia
was probably short and deep; whether it had two
pairs of lagomorph-like upper incisors like Groe-
beria remains unknown. Related to this gnawing
specialization, both Groeberia and Patagonia show
a short diastema near the alveolar level and a re-
duced number of postincisive teeth, four in both;
however, Patagonia has three cheekteeth, whereas
Groeberia has four. The rodent-like habitus of Pat-
agonia is especially advanced, because the three
cheekteeth are truly hypselodont, rectangular-
shaped in cross section, with at most only shallow
lateral grooves representing the remnants of an-
cestral bilobate cheekteeth.

This combination of features suggests food was
obtained by gnawing and prepared for swallowing
by grinding. It represents extraordinary conver-
gence on some desert-adapted and fossorial forms,
such as the Octodontidae. The evolution of cheek-
teeth toward a rectangular shape and numerically
reduced sequence has been recognized as occurring
within the Octodontoidea (from the Octodontidae
to the Ctenomyidae; Pascual et al., 1965). The
dental features of Patagonia are also convergent
on those of the desert-dwelling African Bathyer-
gidae, particularly to the sand rat Heterocephalus
glaber, and to the North American Geomyidae.
These convergent anatomical features suggest that
Patagoniidae were probably fossorial marsupials.

Anatomical convergence of Patagonia on des-
ert-dwelling fossorial rodents is curious, because
prevailing conditions in central Patagonia during
the Colhuehuapian Age were not highly favorable
to desert dwellers. The first record of platyrrhine
monkeys in Patagonia occurs at the same locality
and level (Fleagle & Sown, 1983) as Patagonia.

Many other vertebrate remains recovered at this
site (see Bordas, 1939; Donadio, 1983) suggest an
environment of well-watered tropical woodlands.
Conversely, however, both Argyrolagoidea and
very advanced Cephalomyidae rodents from this
site (currently under study) show dental features
reminiscent of desert or at least drier environ-
ments. Generally, the Colhuehuapian vertebrate
fauna from central Patagonia (see Pascual, 1970;
Pascual & Odreman Rivas, 1971; Marshall et al.,
1983) is composed of both forest and open-coun-
try types, presumably brought together in a sub-
tropical savanna. Thus, the Patagoniidae, Ceph-
alomyidae, and Argyrolagoidea occurred in
apparently inappropriate environments, probably
restricted to xeric patches in the subtropical sa-
vanna mosaic. Because the Colhuehuapian Pata-
gonioidea were already highly specialized for xeric
habitats, they probably evolved earlier in the Pa-
leogene. It therefore seems likely that ancestral
forms existed in the Deseadan (Early Oligocene).
Another highly specialized group of marsupials,
the Argyrolagoidea, suggests this hypothesis. For-
merly believed present in the record fi-om the
Huayquerian (Late Miocene) to the Uquian (Early
Pleistocene; see Marshall et al., 1983), argyrola-
goids have now been reported from the Deseadan
of Bolivia (Wolff, 1984), and here from the Col-
huehuapian beds of central Patagonia.

The pre-Deseadan record contains no potential
ancestor for either Argyrolagoidea or Patagonioi-
dea. Simpson (1970c, p. 17) proposed that "these
groups (including Groeberioidea) evolved in what
are now (and quite likely were then) the tropics
and are picked up in our record only when they
spread rather briefly to what was for them a mar-
ginal area." It seems quite probable that the en-
vironments responsible for their initial divergence
were poorly or not represented in the known fossil
record.

Global diastrophic movements in the Late
Eocene, and apparently related climatic and en-
vironmental changes, are thought to be responsi-
ble for the cosmopolitan turnover in Early Oh-
gocene mammal communities (Kurten, 1971). This
turnover also occurred in South America (Pascual,
1984). Mammal communities in the Deseadan
(Early Oligocene) are substantially different from
Eocene communities in composition (see Pascual
et al., 1 985), apparently reflecting Stehlin's '^^grande
coupure."" The apparently sudden occurrence of
the Argyrolagoidea. and probably the Patagonioi-
dea, in the Deseadan Age is probably another ex-
ample of this global turnover.

108

nELDL\NA: ZOOLOGY

It is remarkable that, to the numerous succes-
sive parallel trends ("successive trends" or "iter-
ation"; Simpson, 1953, pp. 248-259; 1961, p. 127)
in the evolutionary history of South American
mammals, especially from the Deseadan on, can
be added the convergence of Oligocene patagoniid
marsupials and Pliocene to Recent ctenomyid ro-
dents on a common morphology. These conver-
gences are products of similar responses to re-
peated environmental conditions. The anatomical
and functional similarities of Patagonia peregrina
with the extant Ctenomys are so striking that we
are tempted to call the former the "marsupial tuco-
tuco."

Acknowledgments

All of the material studied here was discovered
by 1983 and 1984 expeditions of the Museo Ar-
gentino de Ciencias Naturales "Bernardino Ri-
vadavia" (MACN), in which Lie. Oscar E. Don-
adio and Lie. Miguel Soria (both of MACN) and
the American paleontologists John G. Fleagle and
Thomas M. Bown participated. Dr. Jose F. Bo-
naparte, Chief of the Seccion Paleontologia Ver-
tebrados, MACN, and responsible for these ex-
peditions, generously put this and other marsupial
material at our disposal. The x-ray plates were
made by Dr. Roberto Guevara, Profesor de Odon-
tologia, Universidad Nacional de La Plata, by the
authority of the Dean, Dr. Oscar Barletta. We thank
all of them very much.

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110

FIELDIANA: ZOOLOGY

An Additional 14-Chromosome Karyotype and
Sex-Chromosome Mosaicism in
South American Marsupials

Milton H. Gallardo and Bruce D. Patterson

ABSTRACTS

The karyotype of Rhyncholestes Osgood is described for the first time. The karyotype has
2n = 14 and is similar in most respects to karyotypes of similar number found in other American
and Australasian genera in several families. The karyotype of somatic (bone marrow) tissues
from male Dromiciops Thomas is presented for the first time; surprisingly, it differs from the
2n = 14 complement previously reported from female bone marrow and male gonads. The
2n = 1 3 karyotype found in bone marrow of male Dromiciops lacks a minute element thought
to be the Y chromosome. This instance of somatic chromosome elimination represents the first
case reported for American marsupials and presents an interesting parallel to sex-chromosome
mosaicism among Australasian Peramelidae and Petauridae.

El cariotipo de Rhyncholestes Osgood es descrito por primera vez. El cariotipo consta de
2n = 14 y es muy similar a cariotipos de igual mimero encontrados en otros generos americanos
y australoasiaticos de varias familias. El cariotipo de tejidos somaticos (medula osea) de un
Dromiciops Thomas macho es presentado por primera vez; sorprendentemente, difiere del
complemento 2n = 14 reportado previamente de medula osea femenina y gonadas masculinas.
El cariotipo 2n = 1 3 encontrado en medula osea del Dromiciops macho carece de un diminuto
elemento que supuestamente corresponde al cromosoma Y. Este ejemplo de eliminacion so-
matica de cromosomas representa el primer caso reportado en marsupiales americanos y pre-
senta un interesante paralelo con el extenso mosaicismo de los cromosomas sexuales descrito
entre las formas australoasiaticas.

Descreve-se pela primeira vez, o cariotipo de Rhyncholestes Osgood. O cariotipo e de 2n =
1 4, e, na maioria de seus aspectos, assemelha-se aos cariotipos de numeros similares encontrados
em outros generos americanos e austral^sios. O cariotipo de tecidos somdticos (da medula ossea)
de Dromiciops Thomas machos e descrito pela primeira vez. Supreendentemente, este cari-
otipo difere do complemento de 2n = 1 4, previamente descrito para a medula ossea das femeas
e para as gonadas dos machos. No cariotipo de 2n = 13, encontrado na medula ossea de
Dromiciops machos, falta um elemento miudo, possivelmente o cromossomo Y. Este e o
primeiro exemplo documentado da elimina9ao somatica de um cromossomo em marsupiais
americanos, e apresenta um paralelo interessante ao mosaico frequentemente encontrado nos
cromossomos sexuais de outras formas austral^sias.

From the Institute de Ecologia y Evolucion, Univer-
sidad Austral de Chile, Casilla 567, Valdivia, Chile; and
Division of Mammals, Field Museum of Natural His-
tory, Chicago, IL 60605-2496.

GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS 1 1 1

Introduction

Several unusual cytological features, including
low diploid number (Hayman & Martin, 1969;
Reig et al., 1977), paternally derived X inactiva-
tion (Lyon, 1974a,b), multiple sex-chromosome
systems (Hayman & Martin, 1969; Schneider,
1977), somatic elimination of sex-chromosomes
(Schneider, 1977; Close, 1984), and sperm con-
jugation (Biggers & Creed, 1962; Biggers & De-
Lamater, 1965), have made marsupials interesting
subjects of cytological research. These studies have
clarified fundamental cytological mechanisms.
Additionally, results of the research have shed Ught
upon directions of chromosomal evolution and
upon interrelationships of lineages (Hayman &
Martin, 1969; Reig et al., 1977; Sharman, 1982).

A 14-chromosome karyotype occurs in several
distinct lineages in all living American families:
Didelphidae (Reig et al., 1 977), Microbiotheriidae
(Spotomo & Fernandez, 1971; Reig et al., 1972),
and Caenolestidae (Hayman et al., 1971). This
karyotype also occurs in several Australian mar-
supial lineages (Hayman & Martin, 1969) and is
therefore considered the primitive chromosome
number for Metatheria (Reig et al., 1977). Direc-
tion of chromosome evolution in Metatheria has
proceeded via centromeric dissociations— with
pericentric inversions superimposed on the basic
Robertsonian mechanism— to give rise to the re-
maining 2n = 1 8 and 2n = 22 karyotypes known
for American forms (Hayman & Martin, 1969;
Reig et al., 1977). Extremes of karyotypic varia-
tion in Australasian marsupials are 2n = 10 to 32
(Schneider, 1977).

Two autochthonous and endemic South Amer-
ican genera, Rhyncholestes and Dromiciops, are
especially interesting from an evolutionary view-
point. Both are represented by a single species and
occur only in the temperate Valdivian rainforests
of southern Chile and Argentina. Rhyncholestes.
one of three extant genera of Caenolestidae, is
widely isolated from its relatives in the northern
Andes and presents some striking morphological
specializations. Dromiciops. thought by some to
have special affinities with Australasian lineages
(Sharman, 1982; Szalay, 1982), is the only hving
genus of the otherwise extinct Microbiotheriidae
(Marshall, 1982). Its affinities with other marsu-
pial genera are currently uncertain. In this note we
present the first somatic karyotypes of male Rhyn-
cholestes raphanurus and Dromiciops australis.
Additionally, we document the first instance of

somatic sex-chromosome mosaicism in South
American marsupials.

Materials and Methods

Seven specimens of D. australis (five males and
two females) from Valdivia (39°32'S, 72°52'W),
Osorno (4r06'S, 72°30'W), and Concepcion
(37»26'S, 73°19'W) provinces, Chile, were ana-
lyzed by the in vivo colchicine-hypotonic citrate
technique using bone marrow as a source of mi-
toses (Patton, 1967). Modifications of the same
procedure were used for the one R. raphanurus
collected at La Picada, Volcan Osorno (41*^6'S,
72''30'W); incubation with colchicine lasted 2.5
hours and a slightly more hypotonic solution of
sodium citrate was used. A total of 4 1 9 mitotic
plates was examined: 29 1 from male and 1 20 from
female D. australis and 10 from R. raphanurus.
Museimi specimens were deposited in the Collec-
tion of Mammals, Instituto de Ecologia y Evolu-
cion, Universidad Austral de Chile, and Field Mu-
seum of Natural History.

Results and Discussion

Rhyncholestes raphanurus presents a 2n = 14
complement, consisting of three pairs of large
metacentric, one pair of medium-sized metacen-
tric, and two pairs of small metacentric autosomes.
The sex-chromosomes are an acrocentric X and a
minute Y (fig. 1). This karyotype differs morpho-
logically from the didelphid 2n = 1 4 in not show-
ing a clear break between chromosome groups A
and B. It also differs in arm ratios (table 1) from
the other living caenolestids, Lestoros and Caeno-
lestes (see Hayman et al., 1971). Moreover, the
interstitial region of the short arm of pair two shows
an achromatic area, resembling a secondary con-
striction, not described in other caenolestids (but
see discussion in Sharman, 1982). Nevertheless, a
2n = 14 karyotype characterizes all three genera
of Caenolestidae, which supports previous claims
that this karyotype is primitive for Metatheria
(Hayman & Martin, 1969; Hayman et al., 1971;
Reig et al., 1 977) and reinforces the pattern of low
karyotypic variation within marsupial families.

Secondary constrictions can serve as chromo-
some markers and are thus useful, in the absence

112

FIELDIANA: ZOOLOGY

of banding data, for phylogenetic reconstruction.
However, the secondary constriction evident in
the karyotype of Rhyncholestes is unreported in
other South American marsupials, although sec-
ondary constrictions are widespread among Aus-
tralasian marsupials (Hayman &. Martin, 1969).
Considering commonality and in-group and out-
group comparisons, we regard the secondary con-
striction of Rhyncholestes as apomorphic. Thus,
the similar structures of Australasian marsupials
were apparently independently derived and can-
not be traced back to some marsupicamivorous
or other common ancestor.

Chromosome counts from all four male D. aus-
tralis consistently indicated 2n = 13 chromo-
somes. The diploid number for females was 2n =
14 as was previously reported (Spotomo & Fer-
nandez, 1971; Reig et al., 1972). No differences
among our specimens from geographically isolated
localities were detected, nor were secondary con-
strictions evident.

Electron microscope studies of sex-chromo-
somes in spermatocytes of D. australis and the
didelphid Marmosa elegans demonstrate striking
similarities (Fernandez et al., 1979). These simi-
larities suggest that a 2n = 14 karyotype should
be present in D. australis, its Y chromosome should
resemble that of M elegans, and both genera should
exhibit an XX/XY sex-chromosome system.

We have consistently found 2n = 13 chromo-
somes in somatic tissues of male Dromiciops and
2n = 14 in female somatic tissue. The missing
chromosome in males is dotlike and probably the
Y chromosome (fig. 1). Translocation of the Y to
an autosome is an unlikely mechanism for the dif-
ferences between sexes because males have 2n =
14 in germinal cells and because the sex vesicle
appears normal (Fernandez et al., 1979). While it
is possible that such a small chromosome might
be overlooked in one or a few chromosomal
spreads, its universal absence in all counted plates
makes this alternative highly unlikely. Available
data favor a somatic elimination of the Y chro-
mosome.

Previous studies have shown that both consti-
tutive and facultative heterochromatin can be de-
leted from marsupial cells in vivo without appar-
ent deleterious effects on cell replication and
survival (Hayman & Martin, 1969). Most exam-
ples of somatic elimination of sex-chromosomes
in marsupials involve the X chromosome in dos-
age compensation (e.g., perameUds and petaurids;
Close, 1984). Mitotic figures from the testes of

(( a (»

A-1

B-1

C-1

B-1

A-2 A-3

II •«

C-2 XY

D& Si! \l

A-1 A-2 A-3

lOjLL

»'' A« ^^

c-1 c-2 XX

iiii !3 W

A-1 A-2 A-3

B-1

Ift 6A

c-1 c-2 X

Fig. 1 . Karyotypes from bone marrow cells of A,
Rhyncholestes raphanurus, male; B, Dromiciops aus-
tralis (2n = 1 4), female; C, Dromiciops australis (2n =
13), male.

GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS

113

Table 1 . Arm ratios Gong arm/short arm) of Rhyn-
cholestes autosomes (ratios are based on 1 counted plates;
sex chromosomes are acrocentric).

Pair 1:

1.43

Pair 2:

1.53

Pair 3:

1.39

Pair 4:

1.44

Pair 5:

1.42

Pair 6:

1.46

Dromiciops do show the XY constitution. There-
fore, male zygotes begin development as XY, and
the Y is retained in the germinal cell line, but is
lost in at least some somatic tissues. More studies
will be needed to determine the extent of this mo-
saicism in other tissues.

We believe this instance of sex-chromosome
mosaicism probably represents a parallel, inde-
pendently derived case from that in Australasian
forms. However, it could be used to support Sza-
lay's (1982) assertion that Dromiciops is more
closely related to Australasian lineages than any
other American form, belonging in the Australa-
sian cohort Australidelphia. In this regard it is
noteworthy that Sharman's (1982) analysis of gross
chromosomal morphology suggested that the 2n =
1 4 karyotype of Dromiciops (virtually identical to
those of some burramyids, peramelids, and Vom-
batus ursinns) might be highly similar to that of
the common ancestor of Australian marsupials.
This instance of sex-chromosome mosaicism also
bears on Archer's (1976) contention that pera-
melids, which also exhibit sex-chromosome mo-
saicism, appear to be derivatives of didelphids in
basicranial anatomy. Banding studies of chro-
mosomal morphology in these groups are needed
to help resolve these various suggestions.

A "ratchet" model for the evolution of the Y
chromosome and dosage compensation has been
suggested (Charles worth, 1978). Initially an active
chromosome, the Y is homologous to the X, but
chiasmata formation (and thus recombination be-
tween the two) is suppressed (e.g., Ohno, 1967).
A gradual accumulation of deleterious mutations
could account for its erosion over time, leading to
minute size. In the didelphid Monodelphis dimid-
iata. the synaptonemal complex is absent in the
X-Y pairing region. Structural elements of the
complex are present, but their assembly seems in-
hibited by the shortness of the Y chromosome. It
could be argued that, in Monodelphis and other
metatherians with dotiike Y chromosomes, Y
function is apparently reduced to sex determina-

tion, unnecessary in at least some somatic tissues.
We favor the evolutionary erosion of the Y chro-
mosome and its lack of function in the bone mar-
row tissue of Dromiciops australis as ultimate
causes for this sex-chromosome mosaicism. Late
replication of highly heterochromatinized DNA,
a proximate mechanism for sex -chromosome mo-
saicism suggested in dosage compensation (Hay-
man &. Martin, 1974), may account for the acci-
dental loss of the minute Y chromosome during
mitotic divisions of somatic cells of Dromiciops.

Acknowledgments

We thank Brian K. Lang and Peter L. Meserve
for assistance in obtaining specimens at La Picada.
We received financial support from the Direccion
de Investigacion, Universidad Austral de Chile (S-
83-03), Field Museum of Natural History, Amer-
ican Philosophical Society (Johnson Fund # 1 646),
and National Geographic Society (#2582-82). Dr.
R. Fernandez kindly facilitated study of Dromi-
ciops material. The constructive criticisms of J. A.
W. Kirsch, P. Myers, and J. L. Patton on an earlier
draft are thankfiiUy acknowledged.

Literature Cited

Archer, M. 1976. The basicranial region of marsupi-
camivores (Marsupialia), inter-relationships of car-
nivorous marsupials, and affinities of the insectivo-
rous marsupial peramelids. Zoological Journal of the
Linnean Society, 59: 2 1 7-322.

BiGGERS, J. D., AND R. F. S. Creed. 1962. Conjugate
spermatozoa of the North American opossum. Nature
(London), 196: 1112-1113.

BiGGERS, J. D., AND E. D. DeLamater. 1965. Mar-
supial spermatozoa pairing in the epididymis of
American forms. Nature (London). 208: 402—404.

Charlesworth, B. 1978. Model for evolution of Y
chromosomes and dosage compensation. Proceedings
of the National Academy of Science, USA, 75: 5618-
5622.

Close, R. L. 1984. Rates of sex chromosome loss dur-
ing development in different tissues of the bandicoots
Perameles nasuta and Isoodon macrourus (Marsupi-
alia: Peramelidae). Australian Journal of Biological
Science, 37: 53-61.

Fernandez-D., R., S. Berrios, and J. Pincheira. 1979.
Position of the nucleolus within the nuclei of pachy-
tene spermatocytes of Dromiciops australis and Mar-
mosa elegans (EHdelphoidea-Marsupialia). Experien-
tia (Basel), 35: 1021-1023.

Havman, D. L., and p. G. Martin. 1969. Cytogenetics

114

FIELDIANA: ZOOLOGY

of marsupials, pp. 191-217. In Benirschke, K., ed..
Comparative Mammalian Cytogenetics. Springer, New
York.
. 1974. Mammalia. I: Monotremata and Mar-

supialia, pp. 1-110. In John, B., ed.. Animal Cyto-
genetics. Vol. 4: Chordata 4. Gebriider Bomtraeger,
Berlin.

Hayman, D. L., J. A. W. KiRscH, P. G. Martin, and
P.F.Waller. 1971. Chromosomal and serological
studies of the Caenolestidae and their implications for
marsupial evolution. Nature (London), 231: 194-195.

Lyon, M. F. 1974a. Evolution of X-chromosome in-
activation in mammals. Nature (London), 250: 651-
653.

. 1974b. Mechanisms and evolutionary origins

of variable X-chromosome activity in mammals. Pro-
ceedings of the Royal Society of London, Series B,
187: 243-268.

Marshall, L. G. 1982. Systematics of the South
American marsupial family Microbiotheriidae. Field-
iana: Geology, n.s., 10: 1-75.

Ohno, S. 1967. Sex Chromosomes and Sex-Linked
Cells. Springer- Verlag, Berlin, 192 pp.

Patton, J. L. 1967. Chromosome studies of certain
pocket mice, genus Perognathus (Rodentia: Hetero-
myidae). Journal of Mammalogy, 48: ll-'il.

Reig, O. a., R. Fernandez, and O. A. Spotorno. 1 972.
Further occurrence of a karyotype of 2n = 14 chro-
mosomes in two species of Chilean didelphoid mar-
supials. Zeitschrift fur Saugetierkunde, 37: 37—42.

Reig, O. A., A. L. Gardner, N. O. Bianchi, and J. L.
Patton. 1977. The chromosomes of the Didelphi-
dae (Marsupialia) and their evolutionary significance.
Biological Journal of the Linnean Society, 9: 191-216.

Schneider, L. K. 1977. Marsupial chromosomes, cell
cycles, and cytogenetics, pp. 51-93. In Hunsaker II,
D. D., ed.. The Biology of Marsupials. Academic Press,
New York.

Sharman, G. B. 1 982. Karyotypic similarities between
Dromiciops australis (Microbiotheriidae, Marsupi-
alia) and some Australian marsupials, pp. 711-714.
In Archer, M., ed.. Carnivorous Marsupials, Vol. IL
Royal 2toological Society of New South Wales, Sydney,
804 pp.

Spotorno, O. A., and D. R. Fernandez. 1971. The
chromosomes of the "monito del monte" Dromiciops
australis Philippi. Mammalian Chromosomes News-
letter 12(2): 40-41.

SzALAY, F. S. 1982. A new appraisal of marsupial phy-
logeny and classification, pp. 621-640. In Archer, M.,
ed.. Carnivorous Marsupials, Vol. II. Royal Zoological
Society of New South Wales, Sydney, 804 pp.

GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS

115

Notes on the Black-Shouldered Opossum,
Caluromysiops irrupta

Robert J. Izor and Ronald H. Pine

ABSTRACTS

Caluromysiops is distinct from the three species of Caluromys in external, cranial, dental,
skeletal, and phallic characters, although the two genera are certainly more closely related to
each other than to any other extant genus. Much uncertainty remains regarding the ecology and
distribution of this rare opossum.

Caluromysiops es distinto de las tres especies de Caluromys en caracteres extemos, craneales,
dentales, esqueletales y falicos, aunque los dos generos son por cierto mas cercanamente rela-
cionados entre si que lo es ningun otro genero existente. Todavia hay mucha incertidumbre en
relacion a la ecologia y distribucion de este rara raposa.

Caluromysiops difere das tres especies de Caluromys en carateres extemos, craniais, dentais,
esqueletais e falicos, embora sejam os dois generos claramente mais proximos entre si do que
entre qualquer outro genero atualmente existente. A ecologia e a distribuifao desta rara especie
continuam muito pouco conhecidas.

Introduction

The black-shouldered opossum, Caluromysiops
irrupta Sanborn, is the rarest of the larger didel-
phids. Its history as a subject of scientific study is
peculiar, beginning with a very late discovery
(195 1); also, many more specimens have been dis-
played in zoos (15) than have been collected for
museums directly from the wild (2). Despite the
paucity of associated data and other shortcomings,
zoo animals have been the source of some valuable
information during this study.

Materials and Methods

All zoos known or suspected to have kept Cal-
uromysiops were contacted for information on the

From the Division of Mammals, Field Museum of
Natural History, Chicago, Illinois 60605-2496. Dr. Pine's
present address is Illinois Mathematics and Science
Academy, Aurora, Illinois 60506-1039.

acquisition, history in captivity, and eventual dis-
position of animals. All known specimens pre-
served in collections were examined by one or both
of us, and all tag data were recorded.

Results

Small sample sizes have hampered previous
work on this species, and have affected this study
to some extent. Most published information is
based on single specimens. Some of the characters
described by Sanborn (1951) as diagnostic are in-
dividually or ontogenetically variable and not re-
liable for identification in all cases.

For example, the extent of the hair on the dor-
sum of the tail is distinctive in Caluromysiops,
although not as extreme as originally described.
On immature animals such as the holotype, the
furred area reaches nearly to the end of the tail.
Adults, however, lack fur on the distal 1 5-20 mm.

IZOR & PINE: CALUROMYSIOPS IRRUPTA

117

A^ ■ -i;

Fig. 1. Ventral and lateral views of the cranium and lateral view of the mandible of adult male Caluromysiops
irrupta. fmnh 60698. Certain incisors have fallen out and have been lost.

118

FIELDIANA: ZOOLOGY

Fig. 2. Adult female Caluromysiops irrupta with two young (one is specimen no. cvg M-17 BE 173). Photo
courtesy of Edward T. Maruska and the Cincinnati Zoo.

The portion of the tail covered dorsally with fur
is still much more extensive than even in Calu-
romys lanatus, in which only the proximal 50%-
70% is covered. Except perhaps for some Glironia,
Caluromysiops is unique among didelphids in that
the fur extends onto a distal unpigmented portion
of the tail. The distal one-quarter to one-third of
the tail fur is also white. In other genera of didel-
phids, individuals with some distal portion of the
tail skin unpigmented have fur of the tail confined
to the proximal pigmented area of the tail skin.

The most striking external feature of Caluro-
mysiops is probably the pair of dark lateral and
dorsal stripes. These typically arise on the back of
the hand and run up the inner side of the forelimb
onto the shoulder, where they reach their greatest
width of 15-30 mm. They approach each other
middorsally but usually do not merge, and run in
narrowing parallel bands to the rump. In one old
individual, cvg M-30 BE 95, which had been dis-
played for six years and eight months at the Cin-

cinnati Zoo, the pattern is obscured by a general
grizzling. A common variant of the pattern has
the back of the hand white, with the dark stripe
beginning as a sharply delineated black band
around the wrist. This feature may occur on one
or both forefeet.

As Sanborn and others have noted, some in-
dividuals of the woolly opossums Caluromys der-
bianus and C. lanatus have coloration suggesting
the characteristic dorsal markings of Caluromy-
siops. In the species of Caluromys, there is typi-
cally a darker brown or reddish dorsal area which
grades into the paler, grayer sides of the body. In
some individuals, this darker region is bisected on
the back of the head, neck, and shoulders by a
middorsal gray streak. The supposed similarity to
Caluromysiops, however, is not at all close. The
darker dorsal areas in Caluromys are most sepa-
rated in the place where in Caluromysiops they
are closest to merging. Moreover, the individuals
of Caluromys having the gray middorsal stripe

IZOR & PINE: CALUROMYSIOPS IRRUPTA

119

Table 1 . Caluromysiops irrupta formerly exhibited in zoos.

Sex

Date arrived

Date died

Disposition of remains

Acquisition data

Bronx 2too (New York 2^ological Society)

F 10 Sept. 1962 28 July 1969 Incinerated?
F 20 Nov. 1963 26 Dec. 1964 amnh 208101

National Zoo (Smithsonian Institution, Washington, D.C.)

'Iquitos, Ecuador" (presumably Peru)

M 4 Nov. 1969
M 31 Mar. 1971

12 Apr. 1971

(Sent to Lincoln Park
Zoo, 11 Oct. 1972)
usNM 396160

C. Chase, Miami

From Oklahoma City Zoo

Oklahoma City Zoo

F 23 Nov. 1965
M 28 Jan. 1967

F 19 Dec. 1965

20 Oct. 1970
5 Aug. 1967

Discarded?

(Sent to National Zoo,

31 Mar. 1971)
Discarded?

C. Chase, Miami

Lincoln Park Zoo, Chicago

F 18 Aug. 1972
M 18 Aug. 1972

(pouch young)
F 18 Aug. 1972

(pouch young)
M 11 Oct. 1972

4 Apr. 1973

25 Mar. 1973

16 Sept. 1974

Discarded?

FMNH 121522
FMNH 60154
FMNH 60398

...

Brookfield Zoo (Chicago Zoological Society)
M 21 Apr. 1970 19 Aug. 1971

FMNH 60698

R. Baudy, Center Hill, Fla.

Tarpon Springs Zoo

M (1 Aug. 1972

"rec'd in lab")

USNM 397626

...

Cincinnati Zoo

F 25 July 1965
F 25 July 1965

(young w/F)
F 25 July 1965

(young w/F)
M 1 July 1967

8 Mar. 1967
6 Nov. 1965

11 Dec. 1967

27 Feb. 1974

Discarded?
CVGM-17BE 173

Discarded?

cvG M-30 BE 95

Peru, via Animated Shippers, Miami
Peru, via Animated Shippers, Miami

Peru, via Animated Shippers, Miami

Cuxio (= Cuzco?), Peru, via C. Chase,

Miami

Data from E. Maruska, M. Jones, J. Eisenberg, A. Dittmar, A. Hamer, C. Chase (all in litt.), and Collins (1973).
AMNH = American Museum of Natural History, New York; usnm = National Museum of Natural History, Washing-
ton, D.C; FMNH = Field Museum of Natural History, Chicago; cvg = personal collection of E. Maruska, Director,
Cincinnati Txio.

Table 2. Measurements of Caluromysiops irrupta.

Sex

No.

Total Hind

length Tail length foot

Greatest Medial

skull Basal palatal

length Condylo- length length

(incl. incisive (incl. (incl.

Ear incisors) length incisors) incisors)

CVG M-30 BE 95

617

333.5

63.7

61.0

57.3

30.0

USNM 397626

630+

330 +

64.5

63.4

59.1

32.3

USNM 396 1 60

590

340

63.6

62.5

57.3

30.5

FMNH 60698

63.4

60.6

56.5

30.5

AMNH 208101

570

310

62.6

60.7

56.4

29.7

120

FIELDIANA: ZOOLOGY

also strongly tend to have the palest extremities,
whereas Caluromysiops has extremities with broad
blackish bands (on the inner side of the forelimbs
and outer side of the hind limbs).

Other differences in pelage include the Mar-
mosa-like eye rings and the median facial stripe
of all Caluromys, which are completely lacking in
most Caluromysiops and only faintly suggested in
a few. There is no feature of the color pattern
indicating that Caluromys and Caluromysiops
represent simple variants of a single evolutionary
trend.

Cranially, the extant didelphids present a rather
restricted array of morphologies. All have the same
dental formula. The skulls differ primarily in size,
in the presence and arrangement of palatal vacu-
ities, and in details of the masticatory apparatus
such as sagittal crests, shape of the zygomata, and
the postorbital processes. To our knowledge, a key
to the skulls of the genera has never been con-
structed. It is not surprising, therefore, that it is
difficult to find trenchant cranial characters sup-
porting the distinctiveness of Caluromysiops as a
genus. In the context of the family's relative uni-
formity, this does not necessarily argue against
generic distinction. Pine, however, indicated in
Honacki et al. (1982) that he prefers to regard
Caluromysiops as a subgenus o^ Caluromys, most-
ly because of similarity in skull shape.

The dentition of Caluromysiops irrupta was de-
scribed by Sanborn (1951) as having larger M'--
and m,., than Caluromys. He noted the absence
in the holotype of M^, M^*, and m4 and attributed
the lack of an M^ to its probable loss in the cleaning
of the skull, but did not discuss the absence of the
other molars. The holotype is a juvenile and the
developing alveolus of the m4 is quite evident, so
the tooth is probably unerupted. The larger size
of the molars is generally a valid character distin-
guishing Caluromysiops from Caluromys. Some
individuals oi Caluromysiops may never have had

the minute P', which is frequently lost in adults,
but otherwise the dental formula conforms to that
of the other didelphids. The single root of the usu-
ally spicule-like P' differs from the condition in
Caluromys, in which the tooth is double rooted,
or at least very broad with an incipient division.
There is a strong tendency in Caluromys for the
small cusps on the labial stylar shelf to be subdi-
vided into as many as nine small, low cusps. Cal-
uromysiops typically has five such cusps, each being
higher and more distinct than in Caluromys.

Caluromys and Caluromysiops are united by the
apparently derived character (Archer, 1 982) of clo-
sure of the maxillary palatal fenestrae. This feature
alone is sufficient to distinguish them from all oth-
er living New World marsupials, with the possible
exception of some Marmosa. Archer apparently
erred in attributing such closure to Glironia. Cal-
uromysiops is slightly farther along in the process
than Caluromys, with only small, round, paired
foramina remaining at the maxillopalatal suture.
Species of Caluromys have more or less elongate
foramina.

Several cranial features of Caluromysiops sug-
gest adaptations for strong biting forces. The sag-
ittal crest in adults is very pronounced, and the
zygomatic arches are robust and widely bowed
outward. Rostral length is relatively shorter than
in Caluromys, and the mandible is deeper, with
the ascending ramus broader and more upright.
This seems incongruous in view of the description
by Janson et al. (1981) of nectarivorous behavior.
Zoo animals, however, have readily accepted a
varied diet including animal products (Collins,
1973), and the species probably only exploits nec-
tar and pollen opportunistically.

Cranial asymmetry is prevalent in our sample.
About half of the skulls examined had some sort
of deviation of the rostral axis relative to that of
the braincase, or deflection of the sagittal crest
from the midline.

Table 2. Continued.

Post-

Breadth

zygo-

Depth

Inter-

post-

Post-

matic

brain-

Maxil-

Man-

orbital

Zygo-

brain-

Length

case

Length

lary

dibular

con-

pro-

con-

matic

case

longer

(incl.

mand.

tooth-

M'-

tooth-

striction

cesses

striction

breadth

width

nasal

bullae)

mandible

ramus

row

12.7

22.1

8.2

38.2

23.4

25.1

22.2

45.8

47.2

22.8

9.5

28.3

13.9

21.1

36.9

23.2

24.4

20.6

46.8

48.6

23.2

8.7

28.8

12.4

21.4

7.8

39.2

22.9

25.1

22.1

47.5

23.1

29.4

14.0

20.1

9.5

37.0

23.6

23.9

9.3

11.3

18.3

9.2

38.0

23.3

25.8

22.3

9.1

IZOR & PINE: CALUROMYSIOPS IRRUPTA

121

Postcranial anatomy of the black-shouldered
opossum displays some interesting but as yet
inexplicable differences from that of woolly opos-
sums. The hind limbs of Caluromysiops are rel-
atively much shorter than the forelimbs. The fore-
arm is especially long. In addition, all of the skeletal
elements are more heavily built than in Caluro-
mys, with larger articular surfaces. Both genera
exhibit a slightly offset articulation of the second
metacarpal, which allows the animals to spread
the second and third digits and grasp small branch-
es between them. This schizodactylous grip, also
found in phalangeroids, is useful for slow, delib-
erate climbers which may back up along a branch
rather than turn around to proceed headfirst. The
tail has 30-31 vertebrae, compared to 36-38 in
Caluromys, and has well-developed chevron bones
throughout its length.

Rosenthal ( 1 972, 1 975) noted that a female Cal-
uromysiops was received at the Lincoln Park Zoo
with pouch young, which 40 days later still lacked
markings and body hair. Details of pouch anatomy
were not provided.

All of the didelphids examined to date have a
more or less cleft glans penis. Biggers ( 1 966) noted
that Caluromys derbianus differed from other
species he examined in the greater extent of the
cleft (half the length of the penis), in the contin-
uation of medial urethral grooves to the apices,
and in the rounded, slightly bulbous ends of the
glans. The single available dissected-out specimen
of a Caluromysiops penis (fmnh 60698) suffered
some postmortem deterioration and may not be
completely representative, but still shows clearly
a very deeply split glans (ca. 4 cm) with distinctly
enlarged, rounded tips. The urethral grooves also
seem to extend nearly to the ends.

These characters of the genitalia would seem to
ally Caluromys and Caluromysiops. However,
Caenolestes also has a deeply cleft glans penis (Os-
good, 1921), and many Australian marsupials ex-
hibit some version of the same phenomenon, so
it may represent a shared primitive character.
Moreover, a large majority of didelphid species
have not been evaluated in this regard, and the
significance cannot be properly assessed. Genitalia
of mammals lacking bacula generally have been
less studied, even though soft tissue structure can
be equally informative (Woolley, 1982), and our
cursory survey of preserved material indicates
considerable undocumented variety.

A remarkable feature, poorly preserved on fmnh
60698, but manifest on the protruding penial apex
of FMNH 60398, is a dense covering of small (ca.

1 mm), comified, recurved spines. These are dis-
tributed primarily on the rather rugose tip and
medial sides of the glans, along the urethral groove. |
Osgood (1921) described the glans of Caenolestes
as rugose proximally and covered distally by small i
circular papillae, but Biggers ( 1 966) noted no such
structures on Caluromys or other didelphids ex- j
amined.

The taxonomic affinities of Caluromysiops ir-
rupta have been controversial at both the generic
and suprageneric levels. Cabrera (1958), Hersh-
kovitz (in Marshall, 1982), and Pine (in Honacki
et al., 1982) have suggested that its evident rela-
tionship to Caluromys might be better expressed
by including it in the latter genus. The present
authors are divided on the question of whether
this change would improve the current arrange-
ment.

Reig's (1955) assertion that this species belongs
in the Microbiotheriidae has received adequate
refutation (Segall, 1969; Szalay, 1982). Kirsch's
(1977) attempt to subdivide the Didelphidae is
undermined by the fact that his subfamily names
Caluromyinae and Dactylopsilinae, as proposed,
are nomina nuda. Given his uncertainty about the
contents of the supposed subfamilies of didel-
phids, this fact could spare future workers consid-
erable confusion, although the names may have
since become available inadvertantly in subse-
quent publications.

As most zoo animals have changed hands sev-
eral times before reaching their final destinations,
there is little likelihood of accurate field data ac-
companying them. Among dubious origins re-
ported for zoo-held Caluromysiops are Sao Paulo,
Brazil, and Iquitos, Ecuador (sic). According to J.
A. Davis, Jr. (in litt.), the latter animal "was said
by the dealer to have been captured in a backyard
on the outskirts of Iquitos, Peru"; see also Bridges
(1968) and Davis (1965). Another purported lo-
cality, Cuxio, Peru, has not been located and may
represent a transcription error for Cuzco.

There are only three unquestioned locality rec-
ords, all from southern Amazonian Peru, as fol-
lows:

Peru: Depto. Cuzco; Prov. Quispicanchis,
Quince Mil (13°16'S, 70°38'W), 680 m, fmnh
68336 (the holotype).

Peru: Depto. Madre de Dios; Itahuania (12°47'S,
7 1°1 3'W), skull is fmnh 84426, skin is in the
Museo Nacional de Historia Natural "Javier
Prado", Lima.

Peru: Depto. Madre de Dios; Manu National

122

FIELDIANA: ZOOLOGY

Park, Cocha Cashu Biological Station ( 1 1°55'S,
7 1°1 8'W) (Janson et al., 1981; Terborgh et al.,
1984; Emmons, 1984).

These three localities are within 1 50 km of each
other, along the western margin of the Amazon
basin, between 400-700 m elevation. The only
sympatric species of Caluromys recorded is C la-
natus.

Simonetta's (1979) report of a Caluromysiops
near Leticia, Colombia, is a problem. Although
we are unable to locate the original account, it is
our opinion that this record is best discounted.
The photograph appears to have been staged with
a captive specimen, since the species is nocturnal
(Collins, 1973; Janson et al., 1981; Terborgh et
al., 1984). Leticia is at least 900 km from the three
well-documented localities, and one of the mu-
seum specimens we examined (usnm 397626) is
known to have passed through Leticia from an
unknown source en route to a zoo in Florida. Le-
ticia is the location of a major animal dealership,
and the point of exportation of many Amazonian
species to the U.S. The dusky brown color on the
crown of the head, which Simonetta suggests may
differentiate his Colombian specimen subspecifi-
cally, is variable in the material we examined, and
is probably of no taxonomic importance.

Conclusions

Caluromysiops irrupta is a species which has
often been erroneously or incompletely character-
ized in the scientific literature. There are now
enough specimens in collections to allow reason-
ably complete treatments of its morphology, al-
though it remains an almost complete ecological
and behavioral enigma.

Acknowledgments

The authors thank those individuals and insti-
tutions listed in Table 1 for their invaluable as-
sistance in compiling these data, and for loans of
specimens in their care. Anita McQuaig, Linda E.
Pine, Nobuko Etoh Pine, Joyce Shaw, and Mary
Reed helped with the manuscript. Joseph A. Davis
and the editors and reviewers made many helpful
suggestions.

Literature Cited

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(Thylacinidae, Marsupialia), the phylogenetic position
of the Thylacinidae and the problem of apriorisms in
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BiGGERS, J. D. 1 966. Reproduction in male marsupials,
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the Zoological Society of London, 15: 1-559.

Bridges, W. 1968. The Bronx Zoo Book of Wild An-
imals. New York Zoological Society and Golden Press,
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Cabrera, A. 1958. Catalogo de los mamiferos de
America del Sur. I. (Metatheria-Unguiculata-Camiv-
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turales "Bernardino Rivadavia": 2k>ologia (1957), 4:
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Collins, L. R. 1973. Monotremes and Marsupials.
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tion, Washington, D.C., 323 pp.

Davis, J. A., Jr. 1965. Agreat year for rarities. Animal
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Emmons, L. H. 1 984. Geographic variation in densities
and diversities of non-flying mammals in Amazonia.
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HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds.
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Janson, C, J. Terborgh, and L. H. Emmons. 1981.
Non-flying mammals as pollinating agents in the Am-
azonian forest. Biotropica, 12(Suppl.): 1-6.

KiRSCH, J. A. W. 1977. The comparative serology of
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tralian Journal of Zoology, supp. sen, 52: 1-152.

Marshall, L. G. 1982. Evolution of South American
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H. Genoways, eds.. Mammalian Biology in South
America. Special Publication Series, F*ymatuning Lab-
oratory of Ecology, University of Pittsburgh, 6: 1-539.

Osgood, W. H. 1921. A monographic study of the
American marsupial, Caenolestes. Field Museum of
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Reig, O. A. 1955. Noticiapreliminarsobrelapresencia
de microbiotherinos vivientes en la fauna sudameri-
cana. Investigaciones Zool6gicas Chilenas, 2: 121-130.

Rosenthal, M. A. 1972. Observations on the water
opossum or yapok (Chironectes minimus). Proceed-
ings 48th Annual Conference American Association
of Zoological Parks and Aquariums held in Portland,
Oregon, Oct. 1-5, 1972: 95-98.

. 1975. Observations on the water opossum or

yapok Chironectes minimus in captivity. International
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Sanborn, C. C 1951. Two new mammals from south-
em Peru. Fieldiana: Zoology. 31: 473-477.

Segall, W. 1 969. The middle ear region of Dromi-
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IZOR & PINE: CALUROMYSIOPS IRRUPTA

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SiMONETTA, A. M. 1979. First record of Ca/Mrow>'5;op5 cies of Cocha Cashu Biological Station, Manu Na-

from Colombia. Mammalia, 43: 247-248. tional Park, Peru. Fieldiana: Zoology, n.s., 21: 1-29.

SzALAY, P. S. 1982. A new appraisal of marsupial phy- Woolley, P. A. 1982. Phallic morphology of the Aus-

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ed., Carnivorous Marsupials. Royal Society of New alia): A new taxonomic tool?, pp. 767-781. /« Archer,

South Wales. M., ed.. Carnivorous Marsupials. Royal Society of New

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124 FIELDIANA: ZOOLOGY

Feeding Habits of the Opossum

{Didelphis marsupialis) in Northern Venezuela

Gerardo A. Cordero R. and Ruben A. Nicolas B.

ABSTRACTS

The food items in the annual diet of the opossum {Didelphis marsupialis) in northern Ven-
ezuela are reported by season, sex, and dental age. One hundred eight opossums were sampled
in 21 different sites on a monthly basis from March 1983 to March 1984. The number of food
items recorded varies seasonally. By volume, animal foods (63.5%) are more important than
plant foods (22.9%) throughout the year. Birds (2 1 .5%), mammals (1 5.3%), insects (14.8%), and
fruits (12.8%) are the most prominent foods, by volume. Feeding habits of males and females
do not differ significantly. However, diets of young and old animals are different.

Se seiialan los componentes de la dieta anual del rabopelado {Didelphis marsupialis) en el
norte de Venezuela por epoca del ano, sexo y edad. El muestreo se hizo mensualmente colec-
tandose 108 animales desde Marzo 1983 a Marzo 1984 de 21 localidades diferentes. El numero
de componentes de la dieta varia estacionalmente. En terminos de volumen, los alimentos de
origen animal (63.5%) son mas importantes que los de origen vegetal (22.9%) a traves del aiio.
Las aves (21.5%), los mamiferos (15.3%), los insectos (14.8%) y las frutas (12.8%) son las
alimentos mas sobresalientes, en terminos de volumen. Los habitos alimentarios de los machos
y las hembras no difieren significativamente. Sin embargo, las dietas de los animales jovenes
y viejos son diferentes.

Relata-se os componentes da dieta anual do gamba {Didelphis marsupialis) no norte da
Venezuela, por epoca, sexo e idade. Amostras foram coletadas mensalmente de marfo de 1983
a marfo de 1 984, e de 2 1 locais diferentes, para um total de 108 animais examinados. O nvimero
dos componentes da dieta varia sasonalmente. Em termos de volume, os alimentos de origem
animal (63,5%) sao mais importantes do que os de origem vegetal (22,9%) atraves do ano. Aves
(21,5%), mamiferos (15,3%), insetos (14,8%), e frutos (12,8%) foram os alimentos mais abun-
dantes por volume. Apesar dos habitos alimentares nao diferirem entre machos e femeas, a
dieta dos animais jovens difere da dieta dos adultos.

Introduction of at least seven of 70 species are known (Fleming,

1972; Hunsaker, 1977; Atramentowicz, 1982;
Feeding habits of neotropical didelphid mar- Streilein, 1982; Charles-Dominique, 1983; cf.
supials are poorly known, in spite of their high Kirsch & Calaby, 1977). However, the informa-
diversity and broad geographical distribution. Diets tion reported for most species is based on quali-
tative data. This paper reports the food items in-
gested by opossums {Didelphis marsupialis) in

From the Facultad Ciencias Institute de Zoologia northern Venezuela throughout the year by sea-
Tropical, Apartado 47058, Caracas 1041 -A, Venezuela. son, sex, and dental age.

CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS 125

Study Area

Fieldwork was conducted mainly in the Barlo-
vento region of the State of Miranda and within
the city of Caracas and its surroundings in north-
em Venezuela (10°00'-l(y30'N, 66°00'-67°00'E).
The climate is highly seasonal, with a humid pe-
riod of nine months (May-January) and a dry pe-
riod of three months (February-April) in Barlov-
ento and seven months of rainfall (May-
November) and five months of drought (Decem-
ber-April) in Caracas. Annual mean temperature
of Barlovento is 26° C versus 20.6° C in Caracas
and its surroxmdings. Rainfall is 2,053 mm at Bar-
lovento and 1,011 mm at Caracas. Elevations
sampled range from 40 m to more than 1 ,000 m
above sea level. According to the Holdridge Life
Zones (Ewel et al., 1976), the vegetation of Bar-
lovento is primarily a humid tropical forest,
whereas that of Caracas is mostly in premontane
humid forest.

Materials and Methods

The sample of 108 opossums was assembled
fi"om March 1 983 to March 1 984, either from road-
kills or hunting. Fifty-two animals were taken from
nine localities at Barlovento, whereas 56 speci-
mens were taken from 1 2 localities in or near Ca-
racas. Body measurements, sex, and dental age of
each animal were recorded. Age determination was
based on tooth eruption and wear (Petrides, 1 949;
Tyndale-Biscoe & MacKenzie, 1976), permitting
their grouping into seven age classes (Cordero, un-
publ. data, see Appendix 1). Stomach contents were
analyzed according to Korschgen's (1980) rec-
ommendations. Each stomach and its contents were
placed in a fine sieve ( 1 -mm diameter mesh screen)
and thoroughly washed under running water in
order to separate fine from coarse material. After
measuring the entire volume of the contents, each
item was separated under a dissecting microscope
and its volume recorded. A reference collection
was used for the identification of insects.

Results

Opossum Foods and Seasonal Variation

Six (5.6%) of the 108 stomachs we examined
were empty. Numbers of stomachs with items were:

dry season, 1 6(15.7%) and wet season, 86 (84.3%).
Data for these 102 stomachs appear in Table 1
and Figure 1 . Percentage of volume and frequency
of occurrence are shown for each class of items in
Table 1.

Considerable seasonal variation exists in the
number of food items recorded. During the dry
season, the most important food items are mam-
mals, birds, and insects. In the wet season, birds
are more important by volume than mammals or
insects, and fruits seem to be of greater impor-
tance. Gastropods are ingested in a higher pro-
portion during dry season than wet season. Snakes,
toads, and earthworms are consumed only in the
latter period.

Food of animal origin is more important (63.5%
by volume) than plant food (22.9%) in the diet of
opossums throughout the year. By volume, birds
(21.5%), mammals (15.3%), insects (14.8%), and
fi-uits (12.8%) are the principal foods ingested by
opossums. In terms of frequency, insects (49.1%),
fiiiits( 18.6%), birds (12.7%), and mammals (8.8%)
contribute to the annual diet.

Domestic cats {Felis catiis) and rats (Rattus rat-
tus) were considered as prey items of opossums
because no dipteran carrion larvae were observed
in stomach contents. However, unidentified mam-
malian remains are more important than those of
cat and rat by both volume and frequency. Birds
ingested by opossums were either chickens (Gallus
sp.) or young birds which were more numerous in
the wet season; during the dry season, chickens
were recorded as carrion. Avian material account-
ed for 12.7% of the stomach contents and 21.4%
by volunie. Snakes and toads are consumed at low
levels in relation to their abundance in study sites,
suggesting that these food items are of little im-
portance for opossums in northern Venezuela.

Insects of at least nine families occurred in 49.0%
of the stomachs, with an annual volume of 14.8%.
Beetles and grasshoppers accounted for the ma-
jority of insects consumed.

Slugs (Veronicellidae) were recorded in the rainy
season (1.7% by volume), whereas Vulimulidae
are important in the dry season (6.3% by volume).
Centipedes and earthworms were poorly repre-
sented in the stomachs.

Fruits such as Psidium guajava and Giiazuma
ulmifolia are very important in the diet of opos-
sums. By both volume and frequency of occur-
rence, ftiiits are more important in the rainy sea-
son.

Miscellaneous foods such as garbage (paper,
plastic bags, felt, thread filaments), particulate ma-

126

FIELDIANA: ZOOLOGY

Table 1. Percentages of volume (V) and frequency (F) of food items of opossums in northern Venezuela in 1983
and 1984, by season and for the year.

Wet

season

Dry

season

Annual

Food items

% V

Animals

65.95

90.65

51.22

114.40

63.48

97.03

Mammalia

15.42

5.81

14.75

20.19

15.31

8.82

Felis catus

4.14

2.32

3.44

1.96

Rattus rattus

0.41

7.69

0.07

0.98

Mammal remains

11.28

3.49

14.34

12.50

11.80

5.88

Aves

23.52

13.95

11.27

7.69

21.46

12.74

Gallus sp.

6.42

2.32

5.34

1.96

Young birds

10.04

3.49

8.3S

4.90

Bird remains

7.06

8.14

11.27

7.69

7.77

5.88

Reptilia

0.41

1.16

0.34

0.98

Snake remains

0.41

1.16

0.34

0.98

Amphibia

1.97

1.16

1.64

0.98

Bufo sp.

1.97

1.16

1.64

0.98

Insecta

14.82

47.65

14.70

63.45

14.81

49.01

Coleoptera

7.36

23.24

3.69

23.07

6.76

21.56

Passalidae

0.21

1.16

1.20

7.69

0.28

1.96

Scarabaeidae

5.07

11.63

1.54

7.69

4.48

9.80

Coccinelidae

1.54

7.69

0.26

0.98

Curculionidae

0.31

2.32

0.26

1.96

Meloidae

0.31

1.16

0.26

0.98

Carabidae

0.12

1.16

0.10

0.98

Remains

1.34

5.81

1.12

4.90

Orthoptera

5.12

16.28

0.87

7.69

4.41

14.71

Acrididae

5.12

16.28

0.87

7.69

4.41

14.71

Cursores

0.64

2.32

8.09

25.00

1.89

5.88

Blattaria

0.43

1.16

8.09

25.00

1.72

4.90

Phasmida

0.21

1.16

0.17

0.98

Lepidoptera

1.66

4.65

2.05

7.69

1.72

5.88

Larvae

1.66

4.65

2.05

7.69

1.72

5.88

Homoptera

0.04

1.16

0.03

0.98

Cicadidae

0.04

1.16

0.03

0.98

Mollusca

1.71

8.14

6.25

7.69

2.47

5.88

Veronicellidae

1.71

8.14

1.42

4.90

Vulimulidae

6.25

7.69

1.05

0.98

Chilopoda

1.73

5.81

0.15

7.69

1.46

4.90

■ Annelida

1.20

5.81

1.00

4.90

Lumbricidae

1.20

5.81

1.00

4.90

Carrion

5.17

1.16

4.10

7.69

4.99

3.92

Dendrophidion parcarinatum

5.17

1.16

4.30

0.98

Gallus sp.

4.10

7.69

0.69

2.94

Plants

22.05

43.01

27.26

32.69

22.92

39.22

Fruits

14.19

20.92

6.15

7.69

12.84

18.63

Psidium gtmjava

5.70

13.95

6.15

7.69

5.77

13.73

Guazuma ulmifolia

6.21

2.32

5.17

1.96

Passiflora sp.

1.91

3.49

1.59

1.96

Mangifera sp.

0.37

1.16

0.31

0.98

Grass remains

0.83

2.32

0.69

1.96

Plant remains

7.03

19.77

21.11

25.00

9.39

18.63

Miscellaneous

2.38

19.76

8.71

12.40

1.98

18.62

Paper

trace

11.63

trace

9.80

Plastic bags

trace

4.65

trace

6.20

trace

4.90

Felt

trace

1.16

trace

0.98

Thread filaments

trace

2.32

trace

6.20

trace

2.94

Particulate Material

9.62

12.79

21.52

18.75

11.62

11.76

CORDERO &. NICOLAS: FEEDING HABITS OF OPOSSUMS

127

GRASS 0.7 %
SNAKES 0.3%
CENTIPEDES 1.5%

EARTHWORMS 1.0%

Fig. 1. Proportionate annual volumes of major groups of items from 102 stomach contents of opposums from
northern Venezuela between March 1983 and March 1984.

terial, and plant remains comprised 2.0%, 1 1.6%,
and 9.4% by volume, respectively. Garbage items
were only recorded for those animals collected in
or near Caracas.

Variation of Food Items by Sex

Feeding habits of male and female opossums
are compared in Table 2. By volume, males con-
sume mainly fruits (22.8%), birds (17.1%), plant
remains (15.4%), and insects (14.5%), whereas fe-
males consume mammals (31.4%), birds (14.5%),
insects (11.6%), and fruits (8.8%). However, by
frequency of occurrence, males consume primarily
insects (30%), fruits (19.2%), and plant remains
(15.6%); females consume insects (28.6%), plant
remains (12.2%), mammals (10.2%), and fruits
(10.2%). Both comparisons by means of a Mann-
Whitney U test indicate no significant differences
between the sexes.

Table 2. Food items, by sex, in terms of volume (V)
and frequency (F).

Males

Females

(N =

= 53)

(N =

31)

Food items

% V

Mammalia

9.6

3.6

31.4

10.2

Aves

17.1

6.0

14.5

8.1

Reptilia

0.7

1.2

Amphibia

3.3

1.2

Insecta

14.5

30.0

11.6

28.6

Mollusca

2.3

6.0

4.0

6.1

Chilopoda

1.2

3.6

0.08

2.0

Annelida

0.6

2.4

1.8

4.1

Carrion

2.0

Fruits

22.8

19.2

8.8

10.2

Plant remains

15.4

15.6

7.6

12.2

Miscellaneous

0.7

3.6

5.2

6.1

Particulate material

12.4

7.2

13.0

10.2

N = Sample size.

128

FIELDIANA: ZOOLOGY

A Mann-Whitney U test was also used to com-
pare volumes of principal food groups (mammals,
birds, amphibians, fruits, insects, and plant re-
mains) in the diets of male and female opossums;
no significant differences were detected. Despite
this, the composition of the diet suggests that males
are more arboreal than females. However, a / test
comparing the capture frequencies of both sexes
on the ground in a 26-hectare grid indicates no
significant differences {P > 0.05; Cordero, unpubl.
data). The grid contained 18x18 National live-
traps, with a distance of 30 m between stations
and rows, and was run from December 1981 to
May 1984.

Variation of Food Items by Age

Food of opossums by age classes appears in Ta-
ble 3. Note that the number of food items increases
as animals become older. By volume and fre-
quency of occurrence, animals of younger ages (I,
II, III, and IV) consume mainly invertebrates,
fruits, and plant remains, while older animals (ages
V, VI, and VII) take those items plus mammals
and birds, which become more important as the
animal ages.

Diets of opossums were compared by successive
ages, that is, II with III, III with IV, and so on, by
Mann-Whitney U tests. No significant differences
were detected. However, when diets of young and
old animals were compared, significant differences
were demonstrated. Nothing has been published
on age-related diet variation for D. virginiana, D.
albiventris, or any other marsupial.

Discussion

These results provide a preliminary view of the
annual diet of Didelphis marsupialis in north-
western Venezuela. This study shows that opos-
sums, while omnivorous, are more carnivorous
and insectivorous than herbivorous or frugivo-
rous. However, we accept these patterns guardedly
because they may represent methodological arti-
facts: (1) most of our specimens (84.3%) were tak-
en in the wet season, so that trophic habits during
the dry season are imprecisely known; and (2) the
rinsing step in processing stomach contents may
have inadvertently washed away traces of fruit pulp
that might have been studied using other methods.

Our results indicate that insects, fruits, birds,
and mammals figure prominently in the annual
diet. These figures contrast with those reported by

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• 00 r^

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Ov 00 00
vO PO rn

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r-' — — • vd r~-' o

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d — ■

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■O

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00
iri

■<)...«-)

— Tf

<t • • • 00

- (N

•*

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■n • • • Os

■* ■*

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«N

>r) o\ O

. . . . O
; ; ; ; O

: : : : 8

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Ji = 3

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cu S cu

CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS

129

Molins de la Sema and Lorenzo (1 982) in a study
of stomach contents of 47 Didelphis marsupialis
sampled from February 1981 to May 1982 in the
lowlands of Sierra de Perija in the State of Zulia,
northwestern Venezuela. In their study, the order
of importance of food items, by frequency, is as
follows: plant leaves (68.3%), fruits (56.2%), rep-
tiles (42.6%), insects (29.2%), amphibians (28.8%),
birds (14.3%), mammals (15.1%), mollusks
( 1 2.2%), and seeds ( 1 1 .4%). The effects of seasonal
and habitat differences in the two studies may ex-
plain these differences, since the main vegetation
types of the lowlands of Perija are dry and humid
tropical forests, with eight months of rainfall
(April-November) and four months of drought
(December-March).

Other studies have also shown that opossums
feed on vertebrates. The volume we recorded for
mammalian prey (15.3%) is low in comparison
with diets determined for the Virginia opossum
{Didelphis virginiana), except for Lay's ( 1 942) 7%
value. Hopkins and Forbes (1980) also recorded
cats and rats in low frequencies and volumes in
the diets of opossums in Oregon. Similarly, do-
mestic chickens figured prominently in the diet of
our specimens and have been reported as prey or
carrion of D. virginiana in New York (Hamilton,
1951, 1958), Missouri (Reynolds, 1945), Iowa
(Wiseman & Hendrickson, 1950), Michigan
(Taube, 1947), and Kansas (Sandidge, 1953). In
contrast, snakes and toads were taken infrequent-
ly, paralleling the results of Blumenthal and Kirk-
land (1976), who reported traces of amphibians in
the diets of Pennsylvania Didelphis, and of Wise-
man and Hendrickson (1950), who showed rep-
tiles have a frequency of 1% in the diet of Iowa
opossums. The importance of insects in the diet
of our animals is somewhat lower than that pre-
viously reported for opossums in Michigan (30.4%;
Gardner, 1982, citing Dearborn, 1932), Missouri
(34.2%; Reynolds, 1 945), and Kansas (42.2%; San-
didge, 1953). However, the volumes we report are
higher than those in literature records for New
York (Hamilton, 1951, 1958), Oregon (Hopkins
& Forbes, 1980), and Pennsylvania (Blumenthal
& Kirkland, 1976). Records for other inverte-
brates are also similar to those in existing literature
reports (e.g., Taube, 1947; Hamilton, 1951, 1958;
Reynolds, 1945; Sandidge, 1953).

Our data and literature records indicate that Di-
delphis species have similar diets, embracing a wide
range of food items. More detailed studies, espe-
cially of food-use in relation to availability, will
be needed to establish the degree of euryphagy.

Acknowledgments

This study was partly granted by CONICET
Project SI- 1158. We thank J. Ojasti for sugges-
tions and review of the manuscript. We greatly
appreciate the editorial assistance of B. Patterson.
The staff members of the Estacion Experimental
Rio Negro, Universidad Simon Rodriguez pro-
vided logistical support during fieldwork. L. Du-
que and R. Martinez helped us in the identification
of snakes and slugs, and E. Pannier provided some
stomach contents. To all of them, our thanks.

Literature Cited

Atramentowicz, M. 1982. Influence du milieu sur
I'activite locomotrice et la reproduction de Caluromys
philander (L). Revue d'Ecologie Appliquee (Terre Vie),
36: 373-395.

Blumenthal, E. M., and G. L. Kirkland. 1976. The
biology of the opossum, Didelphis virginiana in south-
central Pennsylvania. Proceedings of the Pennsylvania
Academy of Science, 50: 81-85.

Charles-Dominique, P. 1983. Ecology and social ad-
aptations in didelphid marsupials: Comparison with
eutherians of similar ecology, pp. 395-422. In Eisen-
berg, J. F., and D. G. Kleiman, eds., Advances in the
Study of Mammalian Behavior. Special Publication
of the American Society of Mammalogy, no. 7.

EwEL, J. J., A. Madriz, and J. A. Tosi. 1976. Zonas
de vida de Venezuela. Fondo Nacional de Investiga-
ciones Agropecuarias, Caracas, 265 pp.

Fleming, T. H. 1972. Aspects of the population dy-
namics of three species of opossums in the Panama
Canal Zone. Journal of Mammalogy, 53: 619-623.

Gardner, A. L. 1982. Virginia opossum {Didelphis

virginiana), pp. 3-36. In Chapman, J. A., and G. A.

Feldhamer, eds., Wild Mammals of North America.

Johns Hopkins University Press, Baltimore.
Hamilton, W. J. 1951. The food of the op>ossum in

New York State. Journal of Wildlife Management, 15:

258-264.

. 1958. Life history and economic relations of

the opossum {Didelphis marsupialis virginiana) in New
York Slate. Cornell University Agricultural Station
Memoir, 354: 1-48.

Hopkins, D. D., AND R. B. Forbes. 1980. Dietary pat-
terns of the Virginia oix)ssum in an urban environ-
ment. The Murrelet, 61: 20-30.

HuNSAKER, D. 1977. Ecology of New World marsu-
pials, pp. 95-156. In Hunsaker II, D., ed.. Academic
Press, New York.

KiRSCH, J. A. W., andJ. H. Calaby. 1977. The species
of living marsupials: An annotated list, pp. 9-26. In
Stonehouse, B., and D. Gilmore, eds.. The Biology of
Marsupials. The Macmillan Press Ltd., London and
Basingstoke.

130

FIELDIANA: ZOOLOGY

KoRSCHGEN, L. J. 1980. Procedures for food-habits
analyses, pp. 1 13-127. /n Schemnitz, S. D., ed.. Wild-
life Management Techniques. The Wildlife Society,
Washington, D.C.

Lay, D. W. 1 942. Ecology of the opossum in eastern
Texas. Journal of Mammalogy, 23: 147-159.

MOLINS DE LA SeRNA, M., AND J. LORENZO PrIETO. 1982.

Alimentacion del rabipelado {Didelphis marsupialis)
de la Sierra de Perija. Acta Cientifica Venezolana, 33:
410.

Petrides, G. a. 1949. Sex and age determination in
the opossum. Journal of Mammalogy, 30: 364-378.

Reynolds, H. C. 1 945. Some aspects of the life history
and ecology of the opossum in central Missouri. Jour-
nal of Mammalogy, 26: 361-379.

Sandidge, L. L. 1953. Food and dens of the opossum
{Didelphis virginiana) in northeastern Kansas. Kansas
Academy of Science, 56: 97-106.

Streilein, K. E. 1982. Ecology of small mammals in
the semiarid Brazilian Caatinga. I. Climate and faunal
composition. Annals of Carnegie Museum, 51: 79-
107.

Taube, C. M. 1947. Food habits of Michigan opos-
sums. Journal of Wildlife Management, 11: 97-103.

Tyndale-Biscoe, C. H., and R. B. Mackenzie. 1976.
Reproduction in Didelphis marsupialis and D. albi-
ventris in Colombia. Journal of Mammalogy, 57: 249-
265.

Wiseman, G. L., and G. D. Hendrickson. 1 950. Notes
on the life history and ecology of the oi>ossum in south-
east Iowa. Journal of Mammalogy, 31: 331-337.

Appendix 1.
supialis.

E>ental age classes for Didelphis mar-

Tooth

Age

eruption

Wear

class

(months)

dP' M'

3.0-3.5

dP' M^

4.5-5.0

dP' M'

III

6.2-6.7

P' M'

7.9-8.7

P' M*

10.9-11.7

P' M-

M'-2

12.8-14.1

P' M*

VII

> 16.1

Source: G. A. Cordero (unpublished data).

CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS

131

Notes on Distribution of Some Bats
from Southwestern Colombia

Michael S. Alberico

ABSTRACTS

Noteworthy range extensions are presented for Noctilio albiventris, Rhinophylla alethina,
Sturnira aratathomasi, and Lonchophylla handleyi, including the second Colombian report for
the last. A previous report of Molossops brachymeles is clarified as representing M. abrasus.

Se presentan algunas notables extensiones del rango de distribucion para las especies Noctilio
albiventris, Rhinophylla alethina, Sturnira aratathomasi y Lonchophylla handleyi, este ultimo
siendo el segundo reporte para Colombia. Un reporte anterior de Molossops brachymeles se
clarifica como representativo de M. abrasus.

Apresentam-se notaveis exten^oes mas distribui96es das especies Noctilio albiventris, Rhin-
ophylla alethina, Sturnira aratathomasi, e Lonchophylla handleyi, esta ultima sendo apenas o
segundo registro para a Colombia. Clarifica-se o registro anterior de Molossops brachymeles
como representativo de M. abrasus.

Introduction

Despite considerable interest in Neotropical
mammals, southwestern Colombia remains poor-
ly understood in this respect. This is mainly a
result of a lack of adequate collections caused by
the inaccessible nature of much of the zone. Early
collecting expeditions to which we owe much of
our knowledge were undertaken around the turn
of the century by personnel of the American Mu-
seum of Natural History and summarized by Allen
(1916). Bats were typically underrepresented in
these early collections because of inadequate col-
lecting techniques in use at the time. Now, with
the aid of Japanese mist nets, we are able to obtain
more complete samples of bat communities. In
this report I present results of a continuing col-
lecting effort during the past five years in this poor-
ly known region, extending the known distribution

From the Departamento de Biologia, Universidad del
Valle, Call, Colombia.

of Noctilio albiventris, Lonchophylla handleyi,
Rhinophylla alethina, Sturnira aratathomasi, and
Molossops abrasus.

All specimens mentioned were collected in mist
nets, prepared as standard study skins with skulls,
and deposited in the mammal collection of the
Departamento de Biologia, Universidad del Valle,
Cali, Colombia (UV).

Distribution

Noctilio albiventris

The lesser bulldog bat was recently reviewed by
Davis (1976) and by Hood and Pitocchelli (1983).
Both mapped the distribution as including eastern
Colombia across the Llanos and Amazonas and
the northern Caribbean coast. Davis (1976) re-
ported the altitudinal range of the species as ex-
tending up to 1 , 1 00 m. We have found this species
to be common in the upper Cauca valley, between

ALBERICO: DISTRIBUTION OF COLOMBIAN BATS

133

the Cordillera Central and the Cordillera Occi-
dental of the Andes, where the elevation reaches
this approximate limit. Fifteen specimens from
the Departamento (= state) del Valle del Cauca
and adjacent Departamento del Cauca were com-
pared with the descriptions and measurements of
all subspecies recognized by Davis (1976). This
population is indistinguishable from N. a. minor
in all characters examined and undoubtedly fol-
lows the Rio Cauca south from the Caribbean low-
lands. A similar southern extension is most prob-
able in the valley of the Rio Magdalena to the
Departamento de Huila, but has yet to be con-
firmed by collections.

Specimens Examined— Cauca: Rio Palo, 18 km
S, 5 km E Puerto Tejada, 3°04'N, 76°22'W, 1,050
m (3 92, UV3 1 3, 324, 325); Valle del Cauca: 2 km
S, 4 km W Candelaria, 3°23'N, 76°23'W, 1,000 m
(1 (5, UV676); Universidad del Valle (Melendez
Campus), 8 km S Cali, 3°22'N, 76°32'W, 1,000 m
(5 S6, UV2602, 2603, 2604, 2608, 2609; 2 99,
UV2605, 2607); 13 km S, 1 km E Cali, 3°22'N,
76°32'W, 1,000 m (2 66, UV2620, 2611; 1 9,
UV2612).

Lonchophylla handleyi

This species was described on the basis of spec-
imens from Peru and southern Ecuador by Hill
(1980), who suggested that some individuals in
existing collections might be misidentified as L.
robusta. Lonchophylla handleyi was first reported
for Colombia by Alberico and Orejuela (1982),
who collected a single individual from near the
Ecuadorian border at 850 m. A specimen recently
collected from the Departamento del Valle del
Cauca at 480 m provides the second record for
Colombia. Both specimens are larger (greatest
length of skull, 28.4 and 28.6 mm, respectively)
than the largest L. robusta reported by Hill (1980)
for Peru and Ecuador and are larger than any L.
robusta in our collections from western Colombia.
Both Colombian specimens of L. handleyi are from
the lower slope Andean forests, probably one of
the last habitats to be intensively sampled for
mammals in this country. The presence of this
species in a relatively narrow elevational band
within this habitat type attests to the importance
of continued collecting in the Pacific slope of the
Andes in southwestern Colombia.

Specimens Examined— Nariilo: 5 km E Junin,
l''20'N, 78°08'W, 850 m (1 6, UV3007); Valle del

Cauca: Rio Cajambre, approx. 60 km S Buena-
ventura, 3°20'N, 77°00'W, 480 m (1 9, UV3694).

Rhinophylla alethina

This species was described based on specimens
from western Colombia in the Departamento del
Valle del Cauca (Handley, 1 966) and until recently
was known only from the type locality. Albenco
and Orejuela (1982) reported it from Narifio near
the Ecuadorian border and suggested that it might
have a broader geographic range than previously
thought, which was confirmed by Baud (1982) who
reported the species for Ecuador. Our collections
show R. alethina to be relatively common in the
Pacific lowlands and the adjacent lower slopes of
the western Andes up to 850 m. That this species
was only recently described and remains poorly
known is undoubtedly due to insufficient collect-
ing in the forests of this zone.

Specimens Examined— Nariflo: 5 km E Junin,
1°20'N, 78°08'W, 850 m (3 66, UV3029, 3033,
3036; 5 99, UV3030, 3031, 3032, 3034, 3035).
Valle del Cauca: Alto Anchicaya, 35 km S, 20 km
E Buenaventura, 3°34'N, 76°54'W, 400 m (2 66,
UV3166, 3167); Rio Azul, 5 km N, 25 km W
Darien, 3°59'N, 76°44'W, 560 m (1 9, UV3391);
Rio Cajambre, approx. 60 km S Buenaventura,
3°20'N, 77°00'W, 480-520 m (l 6, UV3702; 1 9,
UV3703); Rio Cahma, 13 km N, 14 km E Bue-
naventura, 4°00'N, 76°59'W, 40 m (1 9, UV2809).

Stumira aratathomasi

In their description of this species, Peterson and
Tamsitt (1968) reported three specimens, the ho-
lotype from the Departamento del Valle del Cauca
in western Colombia and two from an unknown
locality in Ecuador. They stated that it might be
restricted to the Pacific side of the Andes. Thomas
and McMurray ( 1 974) provided measurements for
the holotype and six individuals collected near the
type locality and suggested that this species may
be common at high elevations in the western An-
des of Colombia. Our recent collections extend the
known range some 150 km to the north in the
Cordillera Occidental and, more importantly, re-
cord the presence of S. aratathomasi in the Cor-
dillera Central, where it was previously unknown.
This species appears to inhabit medium to high
elevation forests which are relatively continuous

134

FIELDIANA: ZOOLOGY

in Colombia, and its occurrence both farther to
the north and in the Cordillera Oriental is likely.
Specimens Examined— Valle del Cauca: Cor-
dillera Central: Hacienda "Los Alpes," 6 km S, 1 1
km E Florida, 3°16'N, 76°09'W, 2,400 m (1 9,
UV3482); Cordillera Occidental: Betania, 10 km
N, 15 km W Bolivar, 4°26'N, 76''19'W, 1,800 m
(1 9, UV3876); Parque Nacional "Los Farallones
de Cali," 1 km S, 1 6 km W Cali, 3°22'N, 76°4 1 'W,
2,600 m (1 9, UV3373); Paso de Galapagos, 8 km
N, 4 km E El Cairo, 4°50'N, 76°12'W, 1,800 m
{2 66, UV4131,4133).

Molossops abrasus

This species was reported for Colombia by Al-
berico and Naranjo-H. (1982) as M. brachymeles,
based on specimens from the Cauca valley in
northern Valle del Cauca. Although often referred
to by this latter specific epithet (see Cabrera, 1958;
Freeman, 1981), the holotype of Dysopes abrasus
from Brazil has been shown to represent this species
(Husson, 1962; Carter & Dolan, 1978). The Co-
lombian record extends the known distribution of
M. abrasus in western South America from An-
dean Peru some 1 ,600 km to the north.

Specimens Examined— Valle del Cauca: 1 1 km
S, 2 km W Cartago, 4''39'N, 75°56'W, 930 m (2
66, UV2451, 2452; 1 9, 2453).

Acknowledgments

This report is the result of the combined efforts
of many friends and students, too numerous to
mention by name, who have collaborated either
by accompanying the author in the field, by sharing
specimens collected during other activities, or both.
However, a few individuals have contributed more
than could be expected in the normal turn of events,
and their support in the field and out has been
especially important in the present study: Eduardo
Velasco, Gloria Giral, Alonso Gonzalez, Guiller-
mo Cantillo, and Luz Marina Alberico. To these,
the author is most appreciative.

Literature Cited

Alberico, M., and L. G. Naranjo-H. 1982. Primer
registro de Molossops brachymeles (Chiroptera: Mo-
lossidae) para Colombia. Cespedesia, II: 141-143.

Alberico, M., AND J. E. Orejuela. 1982. Diversidad
especifica de dos comunidades de murcielagos en Na-
rino, Colombia. Cespedesia, Suplemento no. 3(4 1-42):
31-40.

Allen, J. A. 1916. List of mammals collected in Co-
lombia by the American Museum of Natural History
expeditions, 1910-1915. Bulletin of the American
Museum of Natural History, 35: 191-238.

Baud, F. J. 1982. Presence de Rhinophylla alethina
(Mammalia, Chiroptera) en Equateur et repartition
actuelle du genre en Amerique du Sud. Revue Suisse
de Zoologie, 89: 8 1 5-82 1 .

Cabrera, A. 1958. Catalogo de los mamiferos de
America del Sur. Revista del Museo Argentino de
Ciencias Naturales "Bernardino Rivadavia", Ciencias
Zoologicas, 4: 1-307.

Carter, D. C, and P. G. Dolan. 1978. Catalogue of
type specimens of neotropical bats in selected Euro-
pean museums. Special Publications, The Museum,
Texas Tech University, 15: 1-136.

Davis, W. B. 1976. Geographic variation in the lesser
noctilio, Noctilio albiventris (Chiroptera). Journal of
Mammalogy, 57: 681-107.

Freeman, P. W. 1981. A multivariate study of the
family Molossidae (Mammalia, Chiroptera): Mor-
phology, ecology, evolution. Fieldiana: Zoology, n.s.,
7: 1-173.

Handley, C. O., Jr. 1966. Descriptions of new bats
{Choeroniscus and Rhinophylla) from Colombia. Pro-
ceedings of the Biological Society of Washington, 79:
83-88.

Hill, J. E. 1 980. A note on Lonchophylla (Chiroptera:
Phyllostomatidae) from Ecuador and Peru, with the
description of a new species. Bulletin of the British
Museum (Natural History), Zoology Series, 38: 233-
236.

Hood, C. S., and J. Pitocchelll 1983. Noctilio al-
biventris. Mammalian Species, 197: 1-5.

HussoN, A. M. 1962. The bats of Suriname. Rijks-
museum van Natuurlijke Historic, Leiden, 58: 1-282.

Peterson, R. L., and J. R. Tamsitt. 1968. A new
species of bat of the genus Sturnira (family Phyllosto-
matidae) from northwestern South America. Life Sci-
ences Occasional Papers, Royal Ontario Museum, 12:
1-8.

Thomas, M. E., and D. N. McMurray. 1974. Ob-
servations on Sturnira aratathomasi from Colombia.
Journal of Mammalogy, 55: 834-836.

ALBERICO: DISTRIBUTION OF COLOMBIAN BATS

135

Distributional Records of Bats

from the Caribbean Lowlands of Belize

and Adjacent Guatemala and Mexico

Timothy J. McCarthy

ABSTRACTS

Thirty new species records are presented for the bat fauna of Belize, along with secondary
records for eight bats that had been recorded previously from that country. Contiguous lowland
localities in Guatemala provided new department records: nine for El Peten, five for Izabal,
and two for Alta Verapaz. The El Peten records include the first confirmation of Vampyrum
spectrum in Guatemala. One state record for Quintana Roo, Mexico, is reported. These species
represent the genera Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops,
Micronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma,
Trachops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vam-
pyressa, Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bau-
erus, Eumops, and Molossus. Range extensions are acknowledged for Saccopteryx leptura,
Diclidurus virgo, Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyl-
lostomus discolor, Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vam-
pyrodes caraccioli, Artibeus toltecus, and Bauerus dubiaquercus. A checklist of the bat fauna of
Belize, which stands at 66 species, is presented.

Se registran 30 especies que no habian sido citadas antes para la fauna de murcielago de
Belice, con registros secundarios para ocho murcielagos ya conocidos de ese pais. En ciertas
localidades contiguas de las tierras bajas de Guatemala, se obtuvieron nuevos registros depar-
tamentales: nueve de El Peten, cinco de Izabal, y dos de Alta Verapaz. Los registros de El Peten
incluyen la primera confirmacion de Vampyrum spectrum en Guatemala. Ademas, se presenta
un nuevo registro estatal para Quintana Roo, Mexico. Las especies obtenidas estan segregadas
en los generos Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Mi-
cronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trach-
ops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa,
Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bauerus, Eu-
mops, y Molossus. Para cada una de las siguientes especies de murcielagos se anota el alcance
geografico de su distribucion conocida: Saccopteryx leptura, Diclidurus virgo, Noctilio leporinus,
Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor, Vampyrum spec-
trum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli, Artibeus tolte-
cus, y Bauerus dubiaquercus. Se incluye una lista de 66 especies que representan la fauna de
murcielagos de Belice.

Apresenta-se records de 30 novas especies de morcegos para Belice, e de oito especies pouco
conhecidas no pais. Areas adjacentes, na Guatemala, providenciaram novos records para: El

From the Department of Mammalogy, American Mu-
seum of Natural History, Central Park West at 79th
Street, New York, NY 10024.

MCCARTHY: DISTRIBUTION OF BATS 137

Peten (nove especies), Izabal (cinco especies), e Alta Verapaz (dois especies). Os records de El
Peten incluem as primeiras confirma^oes de Vampyrum spectrum na Guatemala. Um novo
record para Quintana Roo, Mexico, e incluido. Estas especies representam os generos Saccop-
teryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Micronycteris, Lonchorhina,
Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trachops, Chrotopterus. Vam-
pyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa, Chiroderma, Artibeus,
Centurio. Diphylla, Natalus, Myotis, Eptesici4s, Lasiurus. Bauerus, Eumops, e Molossus. Re-
conhece-se extensoes nas areas onde sao encontrados Saccopteryx leptura, Diclidurus virgo,
Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor,
Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli,
Artibeus toltecus. e Bauerus dubiaquercus. Apresenta-se uma lista da fauna de morcegos em
Belice, que agora conta com 66 especies.

Introduction

Inventories of bat communities in Mexico and
Central America have increased significantly dur-
ing the last twenty-five years (Jones et al., 1977).
Although the resulting data have enhanced our
knowledge of the distributions and the zoogeo-
graphical relationships of species, incomplete sur-
veys exist for certain regions. The northern low-
lands along the Caribbean coast of Honduras,
Guatemala, Belize, and Quintana Roo, in Mexico,
is one such region. Travel within this coastal ver-
sant has improved with agricultural and settle-
ment expansion. The isolation of Belize from its
neighbors has been reduced with the construction
of roads in Guatemala's frontier of El Peten and
Mexico's former territory of Quintana Roo. A
paved road from Izabal now connects El Peten
and Belize with the Pan-American Highway in
western Guatemala. Road development continues
within Belize for all-weather travel.

Belize is situated within the Caribbean lowland
drainage of northern Central America. Contiguous
with Belize on this eastern slope is the eastern
portion of the department of El Peten to the west
and, to the south, the department of Izabal, both
of Guatemala. Southern Quintana Roo of penin-
sular Mexico borders to the north (see fig. 1 and
Gazetteer). The topography of these Caribbean
lowlands extends from the lower ranges (600 m
and below) of the eastern Sierra de Chama, the
Sierras de las Minas, the Sierra de Santa Cruz, the
Sierra del Meredon, and the Montarias del Mico
in Alta Verapaz and Izabal, and the Maya Moun-
tains of southern Belize and southeastern El Peten
to the low undulating relief of southern Quintana
Roo. The Maya Mountains represent a heavily
eroded Paleozoic formation that now ranges at the
top from 671 to 853 m in elevation, with the high-

est peak at 1113m (Wright et al., 1959). Annual
rainfall in portions of Izabal averages from 3,000
to nearly 5,000 mm (Portig, 1976). Over 4,500
mm of rainfall was reported from the most south-
em coastal area of Belize. North and northwest-
ward of the Maya Mountains, rainfall decreases
appreciably to less than 1 ,500 mm in north-central
El Peten and northern Belize, where less than 1 ,400
mm was recorded near the Quintana Roo border
(Walker, 1973). The severity of this northward
reduction of rainfall is intensified by the increased
lack of surface drainage into the Yucatan Penin-
sula of Mexico. Because the limestone shelf of
northern Belize has geological affinities with the
Yucatan Peninsula (Wright et al., 1 959), the south-
em limit of this peninsula can be considered the
fault line extending from north of the Maya Moun-
tains westward through the northem shore of Lake
Peten-Itza, El Peten (Wadell, 1938; West, 1964).
Effectively, the northem plain of Belize and north-
em El Peten are portions of the Yucatan Peninsula.
The northward shift from alluvial soils to shallow
calcareous soils, along with the mentioned cli-
matic changes, create edaphic conditions that af-
fect the composition and the structure of the vege-
tation that can be supported (Lundell, 1 934, 1 937;
Standley & Record, 1936; Wright el al., 1959; Pen-
nington & Samkhan, 1968). The potential effect
of this transitional physiography on the distribu-
tion and relative abundance of bats in this Carib-
bean lowland region will require further inventory
studies.

This paper documents 30 new records for Belize.
A checklist of the known bat fauna for this country
is annotated in the Appendix. Sixty-six species are
cited. Included here are also records from nearby
localities for El Peten, Izabal, and Alta Verapaz,
Guatemala, and Quintana Roo, Mexico. Nine
species records from EI Peten, five records from

138

HELDIANA: ZOOLOGY

Izabal, and two records from Alta Verapaz in-
crease the number of reported species for these
departments to 35, 31, and 40, respectively (see
Jones, 1966; Carter etal., 1966; Rick, 1968; Smith,
1972; LaVal, 1973a; Martinez R., 1980; Mc-
Carthy, 1982). Jones et al. (1973) and Bimey et
al. (1974) summarized the records for 31 species
from Quintana Roo, and this paper provides one
additional record.

Materials and Methods

The bats that I collected during the years 1974-
1984 in Belize and El Peten (Parque Nacional Ti-
kal), Guatemala, were obtained principally with
mist nets set at ground level; aerial netting and the
use of a bat trap were limited. Unless otherwise
stated, mist netting was carried out during the first
half of the night. A limited number of specimens
were obtained with hand nets or plastic funnel
traps at roost sites. Specimens were prepared as
standard museum skins with skulls and/or skele-
tons, or as fluid-preserved specimens. These
vouchers are housed in Field Museum of Natural
History, Chicago (FMNH); The Museum, Mich-
igan State University, East Lansing (MSU); Car-
negie Museum of Natural History, Pittsburgh
(CM); and American Museum of Natural History,
New York (AMNH).

A survey of 45 museum collections in the United
States, Canada, Mexico, Guatemala, and England
resulted in additional specimens from Belize, El
Peten, Izabal, Alta Verapaz, and Quintana Roo.
Pertinent specimens (147) have been included in
this report from the following institutions [collec-
tors in brackets]: American Museum of Natural
History, New York [N. Sullivan]; British Museum
(Natural History), London, England (BM) [R. H.
L. Disney; P. Williams; A. M. Hutson; R. E. Steb-
bings]; Carnegie Museum of Natural History [N.
A. Bitarj; Field Museum of Natural History [L.
de la Torre]; Florida State Museum, University of
Florida, Gainesville (FSM) [F. J. Bonaccorso];
Museum of Zoology, Louisiana State University,
Baton Rouge (LSUMZ) [D. M. Uy]; Royal On-
tario Museum, Toronto, Canada (ROM) [R. L.
Peterson; J. Kamstra; J. Fragoso]; Texas Coop-
erative Wildlife Collections, Texas A«&,M Uni-
versity, College Station (TCWC) [D. C. Carter; M.
D. Engstrom]; Texas Tech University, Lubbock
(TTU) [P. Diamond]; and United States National

Museum of Natural History, Washington, D.C.
(USNM) [E. L. Tyson].

Systematic arrangement of species accounts and
nomenclature, unless otherwise indicated, follow
Jones et al. (1977) and Handley (1980). Disney
(1968) did not provide data for the first records
of Pteronotus davyi, Tonatia minuta, and Eptesi-
cusfurinalis from Belize. Those data are presented
in the respective accounts of this report, with ad-
ditional records. Further secondary records from
Belize of Mimon crenulatum, Trachops cirrhosus,
Glossophaga commissarisi, Vampyressa pusilla,
and Eumops auripendulus are also included. All
of the species accounts are discussed in the context
of their range and elevational distributions in
Mexico and Central America. Hall (1981) was the
primary reference for this unless cited otherwise.

Forest types in Belize follow Wright et al. ( 1 959),
whose classification was partially based on the sea-
sonal formation series (Beard, 1 944), which refers
to structural appearance. The correct political
alignments between the states of the Yucatan Pen-
insula are inconsistent among a number of pub-
lished maps. The state boundary between Quin-
tana Roo and Campeche on the map in Figure 1
(see also Gazetteer) is based on a number of Gov-
ernment of Mexico (Secretaria de Programacion y
Presupuesto) maps, including "Carta Topografica,
Merida" (1:1 ,000,000; 1 979 and 1 983) and "Mapa
Geografica" (1:5,000,000; 1980).

Species Accounts
Family EMBALLONURIDAE
Subfamily EMBALLONURINAE
Saccopteryx leptura (Schreber, 1 774)

Specimen Examined— BELIZE. Toledo: 2. 1 km
NNE Salamanca Camp, Columbia Forest, 1 9 (cm).

The known distribution of this small sac-winged
bat north of Panama extends through Costa Rica
and Nicaragua to Chiapas along the Pacific ver-
sant. The presence of predominantly lowland Sac-
copteryx leptura in southern Belize represents a
country record and an extension of its distribution
along the Caribbean side from southeastern Nic-
aragua.

Small bats were observed foraging up to heights
of 13-13.5 m during the twilight period of the
evening. Flight appeared to be concentrated within

MCCARTHY: DISTRIBUTION OF BATS

139

a small, open area below the lower canopy of ev-
ergreen seasonal forest. A short mist net was hand-
hoisted to capture (24 March) this adult specimen.
Saccopteryx bilineata was collected shortly after
the capture of S. leptura.

Balantiopteryx io Thomas, 1 904

Specimens Examined— GUATEMALA. EI Pe-
ten: Poptun, Finca Ixobel, \S 66, 1 8 99 (cm).

The restricted distribution of Balantiopteryx io
ranges from the Gulf lowlands of Veracruz, Oa-
xaca, and Tabasco to the lowlands of Belize and
eastern Guatemala. Kirkpatrick et al. ( 1 975), Cart-
wright and Kirkpatrick (1977), and Sanborn (1936)
represent the previous records for Belize and Iza-
bal. The Poptun locality represents the first record
for El Peten.

The specimens reported here were collected ( 1 2
June) by N. A. Bitar as they exited from a cave
surrounded by secondary forest. The distribution
of this colonial species may be restricted in part
by the availability of adequate cave habitats as
roosting sites.

Subfamily DICLIDURINAE

Diclidurus virgo Thomas, 1903

Specimen Examined— BELIZE. Cayo: 1.5 km
SSW Roaring Creek, 1 6 (fmnh).

The white bat is represented by relatively few
localities in Middle America, which extend from
western (Nayarit) and eastern (southern Veracruz)
Mexico through Central America. Specimens from
southwestern El Peten were reported by Jones
(1966). The single specimen from Belize repre-
sents a northward range extension in the Carib-
bean lowlands from northwest Honduras (Carter
& Dolan, 1978) and a record for the country.

The single bat apparently was roosting on the
trunk of a fig tree (Ficus insipida) overhanging a
pool along the Roaring Creek River. It was cap-
tured (May) by C. Tzul after being observed on a
number of occasions roosting near, but not among,
a group of Rhynchonycteris naso. Jones ( 1 966),
Starrett and Casebeer ( 1 968), and Handley ( 1 976)
commented on the high foraging habits of Dicli-
durus. Similar to the molossid bats, Diclidurus
probably concentrates its foraging efforts at levels
well above the tree canopy and beyond the reach
of conventional collecting techniques, except fire-

arms. This may explain why there are few speci-
mens available in collections.

Goodwin ( 1 969) considered Diclidurus virgo at
best not more than subspecifically different from
D. albus. Both species were recognized by Ojasti
and Linares (1971), who questioned Goodwin
( 1 969) because they believed that his South Amer-
ican comparative material represented D. virgo and
not D. albus.

Family NOCTILIONIDAE

Noctilio leporinus mastivus (Vahl, 1797)

Specimens Examined— BELIZE. Cayo: Banana
Bank, 1 <5, 1 9 (fmnh); Barton Creek, at Western
Hwy., 1 9 (fmnh). Stann Creek: Melinda, Stann
Creek River, 1 6 (fmnh). Toledo: 1.2 km E Agua-
cate, Aguacate River, 2 66 (cm), 1 9 (bm); Big Fall,
vicinity Rio Grande Bridge, 1 9 (fmnh); Salamanca
Camp, 1 9 (bm).

The fishing bat occurs along the riparian habi-
tats of river systems, inland lakes, and coastlines
in primarily lowland regions from northwestern
(northern Sinaloa), eastern (southern Veracruz),
and peninsular (Yucatan) Mexico throughout Cen-
tral America (Davis, 1973;Hellebuycketal., 1985).
Dickerman et al. (1981) reported a locality for
Noctilio leporinus from Alta Verapaz as in the Ca-
ribbean drainage when it was clearly in the Rio
Usamacinta drainage of the Gulf lowlands. The
Belizean localities extend northward the recorded
occurrence of A^. leporinus from Izabal and north-
western Honduras (Carter et al., 1966).

All of the specimens were obtained (March,
April, May, July, August) over rivers and a pond
except for one individual, which was mist netted
(28 August) low over a pasture adjacent to a flood-
ed river. This bat was foraging primarily for insects
since its feces contained the chitinous remains of
these prey. Additional fishing bats from the lo-
calities in Cayo and Stann Creek districts were
captured, banded, and released. This bat was com-
mon along the South Stann Creek drainage. Cocks-
comb Basin. A specimen belonging to M. Craig,
Belize Audubon Society, was collected at Indian
Church (Lamanai), New River Lagoon, Orange
Walk District. An old specimen of M leporinus in
the collections of British Museum (Natural His-
tory) was registered in 1 909 without pertinent field
data. The two peninsular records from Campeche
(Jones et al., 1973) and Yucatan (Bimey et al.,
1974) were obtained in coastline habitat along the

140

FIELDIANA: ZOOLOGY

Gulf of Mexico. Although subsurface drainage
predominates north of Belize into Quintana Roo,
shallow inland "lagunas" are fairly common and
probably support Noctilio populations.

Family MORMOOPIDAE
Pteronotus davyi fulvus (Thomas, 1 892)

Specimens Examined— BELIZE. Cayo: Central
Farm, 1 <5 (cm), 1 $ (fmnh); Ontario, 5.5 km W
Teakettle, 1 S (fmnh); Unitedville, 9 km WSW
Teakettle, 1 <5 (fmnh). Orange Walk: Tower Hill,
B.S.I, compound, 3 99 (fmnh). Toledo: Aguacate,
1 (3 (cm); 1.2 km E Aguacate, 1 $ (bm), 1 $ (cm);
Rice Station, 2 6$ (fmnh); 0.4 km W Rice Station,
1 $ (fmnh); San Antonio, 1 $ (fmnh). GUATE-
MALA. El Peten: Parque Nacional Tikal, 1 $ (msu).

Smith (1972) summarized the majority of the
capture localities for this subspecies of naked-
backed bat, which ranges from northwestern (So-
nora), northeastern (Tamaulipas), and peninsular
(Yucatan) Mexico southeastward to Honduras and
El Salvador, but omitted the only record for Belize
(Disney, 1968). Parque Nacional Tikal is the first
record for El Peten, and the Belizean specimens
provide additional records for Belize.

Disney ( 1 968) did not present data for his single
Pteronotus davyi specimen. This male was ob-
tained ( 1 November) in Cayo District, at Listowel
along the Belize River, and is housed in British
Museum (Natural History). The subsequent spec-
imens reported here were collected (October-De-
cember, May, July, August) in open areas, bor-
dering on vegetation and buildings, and over water.
The specimen from El Peten was captured (25
March) along a trail in upland deciduous forest.
An additional P. davyi from Tikal was captured,
banded, and released.

Pteronotus personatus psilotis (Dobson, 1878)

Specimens Examined— BELIZE. Toledo: 1 .2 km

E Aguacate, Aguacate River, 1 9 (bm), 4 33, 1 9
(CM); Big Fall, 1.5 km WSW Rio Grande Bridge,
1 $ (fmnh); 0.8 km NW Blue Creek, 1 9 (fsm);
Crique Jute, 1 $ (fmnh); Crique Lagarto, 1 km
NW San Antonio, 1 9 (fmnh); Jacinto Creek, at
Punta Gorda Road, 1 3 (msu); 0.4 km W Rice
Station, 1 9 (fmnh); Salamanca Camp, 1 9 (usnm);
San Antonio, 1 9 (fmnh); 0.9 km WNW San Pedro
Columbia, 1 9 (fmnh).

The distribution of Pteronotus personatus psi-
lotis extends from western (southern Sinaloa) and
eastern (Tamaulipas) Mexico southeastward to
Honduras and El Salvador (Smith, 1972), with
Caribbean lowland localities in Campeche (Jones
et al., 1973), El Peten (Jones, 1966), and Alta Ver-
apaz (Jones, 1966). Elevations range from 123 to
984 m. These localities from southern Belize are
the first records for the country.

Fifty-three percent of the small moustache bats
were collected (March, May, July) over open water;
the remainder were foraging (January, April, Au-
gust, December) in open areas adjacent to build-
ings or corralled cattle.

Mormoops megalophylla megalophylla

Peters, 1864

Specimens Examined— BELIZE. Belize: 6.6 km

N Churchyard, 1 9 (cm). Cayo: 1.6 km NW Au-
gustine, Rio Frio, 1 3 (ttu). Stann Creek: Melinda,
1 3 (fmnh). Toledo: Forest Home, 1 3 (msu); Pueb-
lo Viejo, 1 3 (fmnh). GUATEMALA. Izabal: 25
km SSW Puerto Barrios, 1 3 (tcwc).

The leaf-chinned bat has been reported through-
out Mexico, Guatemala, El Salvador, and Hon-
duras (Smith, 1972). Davis and Carter (1962),
Jones (1966), and Taibel (1977) provided lowland
records for El Peten and Alta Verapaz. Elevations
range from near sea level to 2270 m. These lo-
calities are the first records for Belize and Izabal.

Except for one Belizean specimen, which was
captured (9 June) in a cave, these leaf-chinned bats
were associated (March, April, December) with
open areas bordering on forest or orchard edges,
including pine savanna. One Mormoops specimen,
which was registered into the British Museum
(Natural History) collections in 1892, may have
been obtained in the vicinity of Belize City. The
Guatemalan specimen was collected by D. C. Car-
ter.

Family PHYLLOSTOMIDAE
Subfamily PHYLLOSTOMINAE

Micronycteris brachyotis (Dobson, 1878)

Specimens Examined— BELIZE. Cayo: 1 km

NW Augustine, 2 $S (fmnh). Toledo: Crique Ne-
gro, Columbia Forest, 1 3 (bm).

The first Middle American specimen of Micro-

McCARTHY: DISTRIBUTION OF BATS

141

nycteris brachyotis was initially reported from Nic-
aragua as M. syhestris by Goodwin (1946), but
was correctly identified by Sanborn (1949). Sub-
sequent records are from the Gulf-Caribbean low-
lands of southern Veracruz (Medellin L. et al.,
1983), Oaxaca (Schaldach, 1964), Chiapas (Davis
et al., 1964), and El Peten (Jones, 1966; Rick,
1 968; McCarthy, 1 982); and the Pacific-Caribbean
versants of Costa Rica (Howell & Burch, 1974;
Starrett, 1976; LaVal & Fitch, 1977) and Panama
(Handley, 1966; Fleming etal., 1972; Bonaccorso,
1979). Reported elevations range from 40 to 594
m. The present specimens are the first records of
the yellow-throated bat for Belize.

Two specimens were captured (29 July) as they
exited from a cave into a low deciduous seasonal
forest, and a third bat was taken (28 May) along
a path in an evergreen seasonal forest. The two
specimens from Cayo, which I obtained while as-
sisting a histoplasmosis survey, were listed as "M.
bardyoiis" in a preliminary report (Quinones et
al., 1978, p. 559) and no specific locality data were
provided.

Micronycteris megalotis mexicana Miller, 1898

Specimens Examined— BELIZE. Corozal: San

Antonio, 2 km NW Corozal, 1 6 (fmnh). Orange
Walk: San Antonio, Rio Hondo, 2 55, 1 2 (fmnh).
Toledo: Aguacate, 1 6 (cm); Big Fall, 2 km E Rio
Grande Bridge, 1 2 (bm); Cuevas Creek Bridge, 10
km NW Punta Gorda, 1 5, 1 2 (bm), 1 S (amnh),
1 2 (msu); Nimli Punit, 1 2 (cm); Rocky Run Ranch,
4.8 km NW Punta Gorda, 1 3, 1 2 (bm); Union
Camp, 2 22 (bm); Vista Hermosa Ranch, 3.7 km
WNW Punta Gorda, 1 2 (cm). GUATEMALA. El
Peten: Parque Nacional Tikal, 1 <5 (fmnh).

The distribution of this subspecies of big-eared
bat extends from western (Jalisco), eastern (south-
em Tamaulipas), and peninsular (Yucatan) Mex-
ico, along the Pacific coastal and highland regions,
to Costa Rica. Gardner et al. ( 1 970) suggested that
the southern extent of Micronycteris megalotis
mexicana is in the Cordillera Talamanca of Costa
Rica. This species has been recorded most often
at lowland-moderate elevations, up to 2870 m.
Specimens from Isla Cozumel, Quintana Roo, rep-
resent the only record for Quintana Roo (Jones et
al., 1973). The records of A/, m. mexicana which
are reported here are the first for Belize and El
Peten.

Belizean specimens were obtained (May, July,
August, November) in diurnal roost sites (shallow

caves and limestone chambers, bridge approach-
ments, abandoned rum factory boiler) and col-
lected in forest habitats (riparian marsh, evergreen
and semi -evergreen, deciduous semi -evergreen,
and deciduous seasonal). The Tikal specimen was
captured (6 June) roosting in a passageway of an
excavation tunnel within a ruin complex. A second
juvenile male was captured, banded, and released
(29 July) in escobal palm (Cryosophila argentea)
forest, 1.9 km SE Tikal Reservoir.

Micronycteris nicefori Sanborn, 1 949

Specimen Examined— BELIZE. Toledo: 0.4 km

NE Aguacate, 1 2 (fmnh).

Handley ( 1 966) documented the first specimens
of Micronycteris nicefori north of South America,
from Panama. Subsequently, it has been reported
from southeastern Nicaragua (Baker & Jones, 1975)
and both the dry Pacific (Starrett, 1976) and wet
Caribbean (LaVal, 1977) lowlands of Costa Rica.
These Central American localities range from near
sea level to over 100 m. This first record from
Belize also represents a significant Central Amer-
ican range extension along the Caribbean versant.

The M. nicefori specimen reported here was mist
netted on 1 5 December along a track in hilltop,
evergreen seasonal forest.

Micronycteris schmidtorum Sanborn, 1935

Specimens Examined— BELIZE. Corozal: Pat-
chakan, 2 22 (fmnh). Orange Walk: 1 .3 km W San
Antonio, Rio Hondo, 1 6 (fmnh). Toledo: Big Fall,
1 km E Rio Grande Bridge, 1 6 (cm).

Micronycteris schmidtorum was described (San-
bom, 1935) from specimens collected in the Ca-
ribbean lowlands of Izabal. An additional Gua-
temalan specimen was recorded in the Pacific
piedmont (Dickerman et al., 1981). The remaining
Central American records represent both the Pa-
cific and Caribbean lowland slopes from Honduras
(Sanbom, 1941), Nicaragua (Davis et al., 1964;
Baker & Jones, 1 975), Costa Rica (Starrett & Case-
beer, 1968; Fleming et al., 1972; Howell & Burch,
1974; LaVal & Fitch, 1977), and Panama (Han-
dley, 1 966). Specimens from Yucatan assigned to
M. schmidtorum by Villa-R. (1966) were reiden-
tified as M. megalotis by Jones et al. (1973). An
identification of M. schmidtorum (Jones et al.,
1 973) for a specimen from Isla Cozumel, Quintana
Roo, was questioned by Hall (1981) because this

142

FIELDIANA: ZOOLOGY

specimen previously was identified as M. mega-
lot is (Jones & Lawlor, 1 965). I examined this spec-
imen (University of Kansas 9 1 539) and agree that
it is M. schmidtorum. The northern distribution
of this big-eared bat extends to the Caribbean coast
of the Yucatan Peninsula. The specimens reported
here are the first records for Belize.

At Parque Nacional Tikal, one juvenile and two
adult females, which were captured (30 July) in a
hollow tree (Bursera semirouba) of an upland de-
ciduous seasonal forest, were photographed, band-
ed, and released. This site was revisited during the
following March, but no Micronycteris were found.
These individuals of M. schmidtorum were the
first seen in El Peten. Similarly, Sanborn (1935)
and Starrett and Casebeer (1968) reported indi-
viduals from tree hollows. The Belizean specimens
were captured (February, September, November)
in the orchard vegetation of a village, along a sec-
ondary forest edge, and in riparian secondary
vegetation.

Lonchorhina aurita aurita Tomes, 1863

Macrophyllum macrophyllum (Shinz, 1821)

Specimens Examined— BELIZE. Cayo: Sibun
River at Indian Creek, 1 $ (fmnh). Toledo: Big
Fall, 1.7 km NE Rio Grande Bridge, 1 $ (cm).

Tabasco, Mexico, represents the northernmost
occurrence for the long-legged bat, which is known
from both the Caribbean and Pacific regions of
Central America. Primarily a lowland inhabitant,
Macrophyllum macrophyllum ranges from 40 to
almost 600 m. These specimens represent a Ca-
ribbean lowlands range extension from north-
western Honduras (Valdez «& LaVal, 1971) and
the first records for Belize.

Harrison and Pendleton (1974), Gardner (1977),
and Dickerman et al. (1981) indicated that long-
legged bats may be closely associated with aquatic
habitats. Similarly, the Belizean specimens were
obtained ( 1 7 March, 1 April) from along the Sibun
River, although not directly above water, and over
the surface of the Rio Grande. The first bat was
taken at approximately 0340 in a stand of shade
trees, dominated by cohune palms (Orbignya co-
hune), at the edge of an open pasture.

Specimens Examined— BELIZE. Stann Creek:

5.3 km WNW Quam Bank, Cockscomb Basin, 1
9 (CM). Toledo: 0.8 km NW Blue Creek, 1 3, 1 2
(amnh); Crique Jute Village, 1 9 (cm); Crique Ne-
gro, Columbia Forest, 1 S (bm), 1 $ (usnm); 2. 1 km
NNE Salamanca Camp, Columbia Forest, 3 66
(CM). GUATEMALA. El Peten: Poptun, Finca
Ixobel, 2 66 (cm).

Lonchorhina aurita was first recorded in Middle
America from Panama (Miller, 1912). Subsequent
collecting has found this cave-dwelling bat north-
ward through Central America to southeastern
(southern Veracruz, Oaxaca, Tabasco) and pen-
insular (Quintana Roo) Mexico. Predominately
lowland, this distinctive leaf-nosed bat extends up
to more than 1500 m in representative habitats.
Jones et al. (1973) reported the only record from
Quintana Roo, while specimens from Izabal (San-
bom, 1936) are apparently the next Caribbean ver-
sant record north of eastern Costa Rica (Nelson,
1965); records from Nicaragua and Honduras are
lacking. The specimens examined for this account
are the first records from Belize and El Peten.

All specimens from Belize were captured (March,
April, May, August) in deciduous seasonal and
evergreen seasonal forests. The Guatemalan bats
were captured by N. A. Bitar as they exited from
the cave discussed in the Balantiopteryx io ac-
count.

Tonatia bidens bidens (Spix, 1823)

Specimens Examined— BELIZE. Cayo: Rio

Frio, 1 .6 km W Augustine, 1 9 (cm). Toledo: Nimli
Punit, 1 6 (cm); Orange Creek, 1.5 km SW Punta
Gorda, 1 6 (msu); 2. 1 km NNE Salamanca Camp,
Columbia Forest, 1 6 (cm); 2.2 km NNE Sala-
manca Camp, Columbia Forest, 1 9 (cm).

Goodwin (1946) first recorded Tonatia bidens
in Central America from the Pacific lowlands of
Costa Rica. Other humid lowland records include
both the Caribbean and Pacific versants of Pan-
ama, continuing along the Caribbean corridor of
Nicaragua, Honduras, and Guatemala. The north-
ernmost record is from eastern Chiapas (Medellin
L., 1983). The Guatemalan records are from the
Caribbean lowlands of El Peten (McCarthy, 1982)
and Izabal (Carter et al., 1966). Elevations range
from near sea level to around 660 m. The present
specimens constitute the first records from Belize.

Four adult males were taken (March, April) over
a creek in a low transitional forest, in a high ev-
ergreen seasonal forest, and in a deciduous sea-
sonal forest. A subadult male was captured (24
September) in the courtyard of a Mayan archae-
ological site located in a high deciduous seasonal
forest.

MCCARTHY: DISTRIBUTION OF BATS

143

Tonatia evotis Davis and Carter, 1978

Specimen Examined- GUATEMALA. El Pe-
ten: Parque Nacional Tikal, 1 6 (fmnh).

Davis and Carter ( 1 978) described Tonatia evo-
tis on the basis of its smaller size in comparison
to T. sylvicola; a female from Izabal was designated
as the holotype. El Peten is part of a Gulf-Carib-
bean distribution which extends from southern
Veracruz, Tabasco, Chiapas, and Campeche to Be-
lize, and continues along northern Honduras (Da-
vis & Carter, 1978). Martinez R. (1980) recorded
an additional eastern Guatemalan locality in Aha
Verapaz. All recorded elevations are less than 100
m. The T. evotis from Tikal represents the first
record for El Peten.

Two adult males and one pregnant female were
mist netted (20 February, 29 and 25 March) in
Tikal along the Uaxactun Road, at a permanent
water pool in escobal palm forest, and in an upland
deciduous seasonal forest. One male and the fe-
male were banded and released.

Tonatia minuta Goodwin, 1 942

Specimens Examined— BELIZE. Cayo: 1.1 km

W Augustine, 1 2 (fmnh); Central Farm, at Belize
River, 1 9 (fmnh); 1.2 km E Macaw Bank, 1 2
(fmnh). Toledo: Big Fall, 1.7 km NE Rio Grande
Bridge, 1 2 (msu); San Lucas, 1 2 (msu).

This small Tonatia was originally described from
the Caribbean coast of Nicaragua as T. nicaraguae
(Goodwin, 1942a). Its Middle American distri-
bution is lowland ( 1 5 to 6 1 m) along Caribbean
and Pacific versants, from southern Veracruz
(Lackey, 1 970) to El Peten, Guatemala (McCarthy,
1982) and Belize (Disney, 1968), continuing
through Honduras (LaVal, 1969; Valdez & LaVal,
1971; Greenbaum & Jones, 1978), Nicaragua
(Jones et al., 1971; Greenbaum & Jones, 1978),
and Costa Rica (Gardner et al., 1 970; LaVal, 1 977),
to Panama (Davis et al., 1964; Handley, 1966).
This account represents additional records for the
small round-eared bat in Belize.

Disney (1968) reported no data for the first To-
natia minuta specimen from Belize, which was a
female collected (25 November) in Cayo District,
at Listowel, along the Belize River. This specimen
was deposited in British Museum (Natural His-
tory). The additional specimens reported here were
captured (November, January, February, April,
May) over rivers or in a deciduous seasonal forest.

The name minuta is applied in accordance with

the discussion by McCarthy ( 1 982). Gardner ( 1 976)
referred to a personal communication with C. O.
Handley, Jr., who suggested that all small Tonatia
(including minuta) represent a single species, T.
brasiliense. Because the taxonomy is poorly under-
stood, a systematic review of this group would be
useful.

Mimon cozumelae Goldman, 1914

Specimens Examined— BELIZE. Belize:

Churchyard, Sibun River, 1 2 (fmnh). Cayo:
"Mountain Pine Ridge", 2 33, 1 2 (bm); 0.8 km W
Augustine, 1 6 (cm); 1 km NW Augustine, 2 $6
(fmnh); Barton Creek, at Western Hwy., 2 $S, 3
22 (fmnh). Toledo: vicinity Aguacate, 2 $S, 2 22
(cm), 1 (5 (fmnh); Crique Negro, Columbia Forest,
1 2 (bm); Pueblo Viejo, 1 3, 1 2 (fmnh); 2. 1 km
NNE Salamanca Camp, Columbia Forest, 2 6S
(cm); 2.2 km NNE Salamanca Camp, Columbia
Forest, 1 <5 (cm); vicinity Union Camp, 2 5(5, 1 2
(bm), 2 22 (cm).

This spear-nosed bat ranges from southeastern
(northern Oaxaca, southern Veracruz) and pen-
insular (Yucatan, Quintana Roo) Mexico south-
eastward along the humid Caribbean side of Cen-
tral America. Specimens from Isla Cozumel,
Quintana Roo, provided the original description
for this species (Goldman, 1914). Recorded ele-
vations extend to 495 m. The Belizean localities
reported here are the first records for the country.

Mimon cozumelae were collected (January,
March, May, July, August, September, December)
along the edge of deciduous and semi-evergreen
seasonal forests bordered with pasture, on riparian
flood plains, over rivers, along paths in high de-
ciduous, semi-evergreen seasonal forests, and in
caves.

Schaldach (1964), Villa-R. (1966), and Hall
(1981) considered cozumelae a subspecies of ben-
nettii. I tentatively accept cozumelae at the specific
level.

Minion crenulatum keenani Handley, 1 960

Specimens Examined— BELIZE. Cayo: Listow-
el, Baking Pot, 1 S (fmnh). Toledo: Crique Negro,
Columbia Forest, 1 6 (usnm).

There are few records for Mimon crenulatum
keenani from Middle America. The distribution
of this distinctive spear-nosed bat extends along
the Caribbean versant, from Panama (Handley,

144

FIELDIANA: ZOOLOGY

1966; Bonaccorso, 1979), Costa Rica (Gardner et
al., 1970; LaVal, 1977), Nicaragua (Greenbaum &
Jones, 1978), Belize (Ruiz, 1983), El Peten
(McCarthy, 1982), and Campeche (Jones, 1964)
to the Gulf lowlands of eastern Chiapas (Medellin
L., 1 983). All recorded elevations range below 265
m. These specimens are the second and third rec-
ords from Belize. The first record (Ruiz, 1 983) was
obtained near Blue Hole, 14 km SE Belmopan,
Cayo District.

One Mimon crenulatum was captured (8 Oc-
tober) in a house after it flew through an open
window. The house was situated along the Belize
River in an agricultural area. The second specimen
was netted (29 March) along a path in evergreen
seasonal forest. E. L. Tyson collected the specimen
from Toledo District.

Phyllostomus discolor verrucosus Elliot, 1905

Specimens Examined— BELIZE. Toledo: Cri-
que Lagarto, 1 km NW San Antonio, 1 S (fmnh);
1 km NNE Salamanca Camp, Columbia Forest, 1
(3 (cm). GUATEMALA. Alta Verapaz: Lanquin,
Lanquin Cave, approx. 1 49 km WSW Puerto Bar-
rios, 1 (5, 1 5 (fmnh).

Records of Phyllostomus discolor extend from
southern (Oaxaca, Veracruz) Mexico along both
the Pacific and Caribbean corridors of Central
America. Records are more common at lower el-
evations, less than 600 m. The new records from
southern Belize provide a limited range extension
northward from eastern Izabal (Sanborn, 1936).

An adult from Crique Lagarto was captured ( 1
January) along the edge of low secondary forest
bordering this settlement. The head of the bat was
covered with yellow pollen. The second specimen
was netted (21 March) in secondary vegetation,
which resulted from slash-bum agriculture. Whit-
ish pollen dusted the face, chest, and ventral wing
surfaces. A male subadult Phyllostomus discolor
that was taken ( 1 3 July) along a fenceline of sec-
ondary vegetation between two pastures, 1 .9 km
ENE Rio Grande Bridge, Big Fall, Toledo District,
was photographed, banded, and released. L. de la
Torre apparently captured (3 June) the two Phyl-
lostomus from Alta Verapaz inside the entrance
of Lanquin Cave.

I tentatively follow Jones et al. (1977) in as-
signing the specimens of Phyllostomus discolor
from the Caribbean lowlands to the subspecies
verrucosus. Sanborn (1936, p. 98) recognized ver-
rucosus subspecifically, stating the "available mea-

surements of rf/5co/or would place them much clos-
er to verrucosus."'' He suggested the Panamanian
P. d. discolor are assignable to verrucosus based on
larger size. Felten (1956) and ^urt and Stirton
(1961) concurred with his statement by referring
a large series from El Salvador to verrucosus; with
the availability of greater series of specimens, Da-
vis and Carter (1962) indicated they could not
recognize two subspecies of P. discolor in Central
America and northern South America, acknowl-
edging only P. d. verrucosus. Handley ( 1 966) ap-
parently disagreed as he recognized the subspecies
discolor in Panama. Multivariate analysis of mor-
phological data (Power & Tamsitt, 1 973) suggested
this species might be monotypic.

Phylloderma stenops septentrionalis

Goodwin, 1940

Specimens Examined— BELIZE. Toledo: Cri-
que Negro, Columbia Forest, 1 2 (usnm); 2. 1 km
NNE Salamanca Camp, Columbia Forest, 2 $S

(CM).

This rarely encountered species has been re-
corded north of Panama from the Caribbean coast
of Costa Rica (LaVal, 1977), the highlands of Hon-
duras (Goodwin, 1940), the Caribbean lowlands
of Guatemala (McCarthy, 1982), and the Gulf
lowlands of Chiapas (Carter et al., 1966). Limited
elevational data are from lowland to approxi-
mately 1320 m. The specimens oi Phylloderma
stenops from Belize represent the eighth, ninth,
and tenth specimens north of Panama and the first
records from Belize.

All specimens were mist netted (March, Decem-
ber) in similar evergreen seasonal forest habitats.
E. L. Tyson collected the specimen from Crique
Negro.

Handley ( 1 966) regarded the Panamanian spec-
imens to be Phylloderma stenops stenops, and those
from northward into Middle America were thought
to be subspecifically different from the nominal
species. LaVal (1977) did not designate a subspe-
cies for his Costa Rican specimen.

Trachops cirrhosus coflini Goldman, 1925

Specimens Examined— BELIZE. Orange Walk:

Richmond Hill (Goat Hill), 8.9 km SSW Orange
Walk Town, 1 3, 1 $ (cm). Toledo: 2.2 km NNE
Salamanca Camp, Columbia Forest, 1 $ (cm).

MCCARTHY: DISTRIBUTION OF BATS

145

GUATEMALA. Izabal: 25 km SSW Puerto Bar-
rios, 1 $ (tcwc).

This lowland subspecies of the fringe-lipped bat
is recognized from eastern (southern Veracruz) and
southeastern (eastern Oaxaca) Mexico southeast-
ward to Nicaragua. Recorded elevations are from
near sea level to approximately 330 m. Jones
(1966), Rick (1968), and McCarthy (1982) pro-
vided records for El Peten. The description of this
subspecies was based on specimens from eastern
El Peten (Goldman, 1925). The first Belizean rec-
ords were reported from Belize District in the vi-
cinity of Belize City (Sanborn, 1941) and Rock-
stone Pond (Pendergast, 1979). The specimen from
Izabal is the first record for that Guatemalan de-
partment.

D. C. Carter obtained the single specimen from
Izabal on 19 February. The additional Belizean
specimens were mist netted (March, April) in de-
ciduous marsh and evergreen forests.

Chrotopterus auritus (Peters, 1856)

Specimens Examined— BELIZE. Toledo: vicin-
ity Crique Negro, Columbia Forest, 1 9 (fmnh);
1.6 km NNE Salamanca Camp, Columbia Forest,
1 9 (fmnh).

Chrotopterus was first reported in Central Amer-
ica from El Salvador (Burt & Stirton, 1961). Sub-
sequently, this carnivorous bat has been recorded
from southern (southern Veracruz, northern Oa-
xaca, Chiapas) and peninsular (Yucatan, Quintana
Roo) Mexico southeastward throughout Central
America at lowland and upland elevations (40 to
over 1 880 m). Chrotopterus auritus has been re-
ported from Quintana Roo (Jones et al., 1 973) and
El Peten (Rick, 1968; McCarthy, 1982). These
specimens from southern Belize provide the first
records for the country.

The Belizean specimens were netted (10 April,
28 July) in an evergreen seasonal forest at ground
level along a path and at a height of about 13.7m
over an intermittent stream bed. Both were active
during the morning hours, 0418 and 0330, re-
spectively.

The subspecific name Chrotopterus auritus au-
ritus has been applied to Middle American pop-
ulations (Jones et al., 1971). Carter and Dolan
(1978) stated the type specimen for Vampyrus au-
ritus Peters, 1856, actually was collected in Santa
Catarina, Brazil, not in Mexico. The discussion by
Carter and Dolan (1978, p. 37) suggested that Pe-

ters based his description on one or more speci-
mens from Brazil and compared these with a spec-
imen from an unrecorded locality in Mexico as
the "verwandten Art aus Mexico." Handley ( 1 966)
doubted that subspecies were recognizable.

Vampyrum spectrum (Linnaeus, 1758)

Specimen Examined— BELIZE. Toledo: Santa
Elena, 1 S (fmnh).

Two localities in southern Veracruz, Mexico
(Goldman, 1917; Navarro L., 1 979) are the north-
westernmost records of the false vampire bat's
Middle American distribution, which continues in
Nicaragua (Dobson, 1 878; Allen, 1910), Costa Rica
(Casebeer et al., 1963; Armstrong, 1969; Gardner
et al., 1970; Howell &. Burch, 1974; Vehrencamp
et al., 1977; LaVal & Fitch, 1977), and Panama
(Handley, 1966; Peterson & Kirmse, 1969; Bo-
naccorso, 1979). Although primarily lowland in
distribution, its highest recorded elevation was
about 1815m. The occurrence of Vampyrum spec-
trum in the Caribbean lowlands of Belize is doc-
umented by this specimen.

There appears to be no definite record of this
carnivorous bat from Guatemala (Jones, 1966).
Dobson (1878, p. 471) recorded "Guatemala" as
part of the Central American range for Vampyrum,
but did not list any examined specimens. Alston
(1879-1882, p. 39) stated Dobson (pers. comm.)
saw specimens from Guatemala, although Alston
realized the collector, O. Salvin, had not obtained
specimens of Vampyrum; hence, the identification
of this species is doubtful. Five false vampire bats
were mist netted on three separate dates in Parque
Nacional Tikal, El Peten. Two females were cap-
tured during the dry season (22 and 24 March) in
an upland deciduous seasonal forest, in the vicin-
ity of Central Plaza of the archaeological site, and
at a permanent water pool in escobal palm forest,
2.6 km SE Central Plaza. Two females and one
male were netted during the wet season (22 July)
at a location along an archaeological transect in
escobal palm forest, 1 km SE Tikal Reservoir. All
of these bats were released after being observed,
measured, and/or photographed. These individ-
uals provide the first record for Guatemala and,
along with the specimen from Belize, bridge an
intermittent distribution that now extends north-
ward toward peninsular Mexico.

The Vampyrum spectrum from Belize was cap-
tured (8 April) during the early morning (0300) in

146

FIELDIANA: ZOOLOGY

an open field. We were "trapping" Desmodus ro-
tundus during a vampire bat control effort in the
village. This large bat was captured after it made
a number of low passes over horses and mules,
which were encircled by mist nets. The bat died
while enroute to captivity via an assistant.

The Central American population of Vampy-
rum was described as a distinct subspecies, V. s.
nelsoni (Goldman, 1914), but Handley (1966) ar-
gued that the species was monotypic.

Subfamily GLOSSOPHAGINAE

Glossophaga commissarisi commissarisi

Gardner, 1962

Specimens Examined— BELIZE. Belize: Rock-
stone Pond, 2 SS, 3 99 (rom). Toledo: Aguacate, 1
9 (fmnh), 1 9 (cm); Big Fall, 1 km SE Rio Grande
Bridge, 2 $S (cm); Forest Home, 1 9 (fmnh); 2.8
km NNW Punta Gorda, 1 9 (fmnh). GUATE-
MALA. Izabal: 25 km SSW Puerto Barrios, 7 SS,
6 99 (tcwc).

Webster and Jones (1982) summarized the dis-
tribution for this subspecies of nectivorous bat,
which was documented from eastern (Veracruz)
and southern (Oaxaca, Chiapas) Mexico and
southern Belize southeastward throughout Central
America. Hellebuyck et al. (1985) recently re-
ported records from El Salvador. The specimens
from Izabal are the first records from this Gua-
temalan department. The specimens from Belize
District extend northward the distribution of Glos-
sophaga commissarisi along the Caribbean low-
lands.

According to D. C. Carter's field notes, the ma-
jority of the Guatemalan Glossophaga commis-
sarisi were mist netted (February, March) over a
stream and in the adjacent undisturbed forest.
Many of these nectivorous bats were captured in
association with night-blooming "bat flowers"
bordering on a stream. The Belizean specimens
reported (Webster & Jones, 1982) from Lubaan-
tun, Toledo District, were collected ( 1 8 April) in
a disturbed semi-evergreen seasonal forest. Ad-
ditional specimens were secured (January, July,
September, December) in secondary and orchard
vegetation of villages, in riparian secondary vege-
tation, and from the hollow of a mamey tree (Pou-
teria mammosa).

Subfamily STENODERMATINAE
Uroderma bilobatum molaris Davis, 1968

Specimen Examined— MEXICO. Quintana Roo:

2 km N, 8 km W Bacalar, 1 $ (tcwc).

Davis (1968) recognized this subspecies of the
tent-making bat from the Gulf-Caribbean versant
of southern Veracruz, Tabasco, northeast Oaxaca,
northern Chiapas, Belize, Honduras, Nicaragua,
Costa Rica, and northwest Panama. Disney (1968)
and Pendergast (1979) also reported the occur-
rence of Uroderma bilobatum from Belize. The
specimen reported here represents the first record
for Quintana Roo and a marginal range extension
into the Mexican peninsula of Yucatan.

The above specimen was taken in a net on 6
August by M. D. Engstrom along a path leading
to an inland lagoon.

Vampyrops helleri helleri Peters, 1866

Specimens Examined— BELIZE. Cayo: Banana
Bank, 5 99 (fmnh); 0.8 km W Macaw Bank, 1 6
(fmnh). Toledo: Big Fall, 1.9 km ENE Rio Grande
Bridge, 1 9 (amnh), 1 9 (cm), 1 $ (msu); Crique
Negro, Columbia Forest, 1 $ (bm); Forest Home,
1 (5 (fmnh), 1 (5 (msu); Salamanca Camp, 1 S (bm),
1 (5 (fmnh), 1 9 (usnm); 1.8 km NNE Salamanca
Camp, Columbia Forest, 1 9 (fmnh); vicinity Union
Camp, 1 9 (bm), 2 99 (cm).

The Middle American records of this fruit bat
indicate a distribution from sea level to elevations
of over 1 300 m and a range from southeastern
Mexico (southern Veracruz, Oaxaca, Tabasco)
throughout Central America. Lowland records
have been reported from El Peten (Rick, 1968)
and Izabal (Carter et al., 1966). This account con-
stitutes the first records from Belize.

Eighty-seven percent of the Vampyrops helleri
specimens were captured along or in proximity to
waterways. Eleven additional individuals were re-
leased at Banana Bank, where a concentration of
stenodermatines (Sturnira, Uroderma, Vampyres-
sa, Chiroderma, Artibeus, and Vampyrops) was
observed. The remaining localities were in upland
evergreen seasonal forest and in disturbed village
vegetation. A specimen in the collection of St.
John's College, Belize City, was collected by E. L.
Tyson in Columbia Forest.

I follow Dickerman et al. (1981) for the taxo-
nomic assignment of the subspecific epithet.

MCCARTHY: DISTRIBUTION OF BATS

147

Vampyrodes caraccioli major G. M. Allen, 1908

Specimens Examined— BELIZE. Toledo: Agua-
cate, 1 (5 (CM); Big Fall, 1 .9 km ENE Rio Grande
Bridge, 1 S (cm), 1 6 (fmnh); Big Fall, 2.1 km E
Rio Grande Bridge, 1 S (bm); Crique Negro, Co-
lumbia Forest, 1 S (bm), 1 5, 1 9 (msu); Salamanca
Camp, 1 S (usnm); 1.6 km N Salamanca Camp,
Columbia Forest, 1 S (fmnh); 2. 1 km NNE Sala-
manca Camp, Columbia Forest, 4 66, I 9 (cm); San
Antonio, 1 9 (fmnh).

The published distribution of Vampyrodes car-
accioli major northwestward of Costa Rica and
Panama is confined to the Gulf-Caribbean low-
lands as far as southern Mexico (Oaxaca, southern
Veracruz, Chiapas); elevational data are less than
300 m. The records from Belize extend the range
of this stenodermatine north of Izabal (Sanborn,
1936).

The Belizean localities represent habitats of ri-
parian lowland and upland evergreen seasonal for-
ests and village secondary vegetation. The capture
dates cover both the dry and wet seasons (March,
April, May, July-September, December).

I follow Carter and Dolan (1978) for the correct
spelling of Vampyrodes caraccioli.

Vampyressa pusilla thyone Thomas, 1 909

Specimens Examined— BELIZE. Cayo: 1.6 km

NW Augustine, 3 66, I 9 (cm); Banana Bank, 1 9
(fmnh); Blancaneaux, 8.3 km NNE Augustine, 1
9 (fsm). Toledo: vicinity Aguacate, 1 9 (bm), 3 99
(cm); 1.2 km E Aguacate, 1 3, 1 9 (cm); Big Fall,
1 km E Rio Grande Bridge, 1 9 (cm); Big Fall, 2. 1
km E Rio Grande Bridge, 1 6 (cm); Big Fall, 1 .9
km ENE Rio Grande Bridge, 1 5, 1 9 (cm), 1 9
(fmnh); Crique Negro, Columbia Forest, 1 6 (msu),
1 6 (usnm); Forest Home, 1 6 (msu); Pueblo Viejo,
1 9 (fmnh); 1 .6 km NNE Salamanca Camp, Co-
lumbia Forest, 1 3, 2 99 (fmnh).

The general distribution of the little yellow-eared
bat extends from southern (Oaxaca, southern Ve-
racruz, Chiapas) and peninsular (Campeche) Mex-
ico and continues southeastward along the Carib-
bean slope to both the Pacific and Caribbean
corridors of southern Nicaragua, Costa Rica, and
Panama, into South America. Elevational data are
primarily lowland, from sea level up to a recorded
2200 m. Peterson (1966) reported the only record
of Vampyressa pusilla in Belize, from Rockstone
Pond, Belize District. There are also previous rec-
ords from El Peten (Rick, 1 968) and southeastern

Campeche (Jones et al., 1973). This account pro-
vides additional records of this species.

These specimens of Vampyressa pusilla were
collected (February-May, July-September, De-
cember) in moist habitats, the majority of which
were associated directly with riparian vegetation
or in village and pasture-edge vegetation situated
near rivers. Evergreen seasonal forest provided an
upland habitat.

Chiroderma villosum jesupi J. A. Allen, 1900

Specimens Examined— BELIZE. Cayo: Banana

Bank, 1 3, 5 99 (fmnh). Corozal: Chan Chen, 1 6
(fmnh). Toledo: Big Fall, vicinity Rio Grande
Bridge, 1 6 (fmnh); Big Fall, 1 .7 km NE Rio Grande
Bridge, 1 9 (msu); Big Fall, 1 .9 km ENE Rio Grande
Bridge, 1 3, 1 9 (cm); San Antonio, 1 6 (fmnh); 1
km WNW San Pedro Columbia, 1 9 (fmnh). GUA-
TEMALA. EI Peten: Parque Nacional Tikal, 1 6
(fmnh).

The Middle American occurrence of Chiroder-
ma villosum has been documented in southern
(Oaxaca, southern Veracruz, Chiapas) and pen-
insular (Campeche, Quintana Roo) Mexico, Gua-
temala, Nicaragua, Costa Rica, and Panama. Hel-
lebuyck et al. (1985) recently reported this fruit
bat from El Salvador. Locality records reach from
the coastal lowlands to upland habitats at 1 300 m.
Southeastern Campeche (Jones et al., 1973) and
northern Quintana Roo (Bimey et al., 1974) are
previous Caribbean lowland localities, in addition
to these first records from Belize and El Peten.

All but one of the Belizean Chiroderma were
associated either directly with or in the vicinity of
riparian evergreen or semi-evergreen seasonal for-
ests (April, May, August, September, December).
One individual was captured (15 November) in
village orchard vegetation. Five additional indi-
viduals were released at Banana Bank. The Tikal
specimen was captured (24 March) along the per-
manent water pool mentioned in the Tonatia ev-
otis account.

Artibeus toltecus toltecus (Saussure, 1 860)

Specimens Examined— BELIZE. Cayo: vicinity
Augustine, 2 66, 4 99 (fsm); 1 .6 km NW Augustine,
Rio Frio, 1 3, 1 9 (fmnh), 5 66 (ttu), 4 66 (cm);
"Rio On," ? km N Augustine, 1 9 (ttu); 1.1 km
S Baldy Beacon, Bald Hills, 3 99 (cm); vicinity San
Luis, 7.1 km SSW Augustine, 1 9 (ttu). Toledo:

148

FIELDIANA: ZOOLOGY

Orange Point, 1 2 (fmnh); Pueblo Viejo, 3 9$
(fmnh); Union Camp, 5 S6, 4 92 (cm).

In his revision of the small Artibeus of Middle
America, Davis (1969) recognized the range of
Artibeus toltecus toltecus from southern Tamau-
lipas, Mexico, southeastward along the mountain-
ous region of the Gulf versant, upland of southern
Mexico, Guatemala, Honduras, Nicaragua, and
Costa Rica. He did not examine Panamanian spec-
imens. Handley (1966) summarized the Pana-
manian localities for /I. toltecus. This bat primarily
occurs at elevations between 328 and 1640 m,
although elevations near sea level were recorded
(Davis, 1969). Consequently, the occurrence of ^.
toltecus in the Maya Mountain range of southern
Belize and southeastern El Peten was not unex-
pected. These Belizean localities represent the first
northern Caribbean lowland records.

The Belizean localities range in elevation from
near sea level to approximately 720 m. Artibeus
toltecus is more common at the higher elevations.
These dark-colored Artibeus were captured (De-
cember-February, April, June, September) in hab-
itats of deciduous seasonal forest, semi-evergreen
seasonal forest, transitional forest, and pine forest-
savanna.

The subspecies toltecus is applied, based on the
proximity of Belize to its distribution as defined
by Davis (1969).

Centurio senex senex Gray, 1 842

Specimens Examined— BELIZE. Belize: 1 .4 km
S San Pedro, Ambergris Caye, 1 3, 1 9 (fmnh).
Cayo: 1.6 km NW Augustine, Rio Frio, 1 <5 (ttu);
vicinity Augustine, Rio On, 1 9 (ttu); Blanca-
neaux, 8.3 km NNE Augustine, 1 9 (fsm); Central
Farm, 1 3, 1 9 (fmnh); Teakettle, Young Gal Road
at Belize River, 1 3, 1 9 (fmnh); Xunantunich, 1
$ (fmnh). Corozal: 1.2 km E, 1.6 km N Corozal,
1 (5 (LSUMZ). Orange Walk: 1.6 km NW San An-
tonio, Rio Hondo, 1 9 (fmnh). Toledo: Big Fall,
1 .9 km ENE Rio Grande Bridge, 1 S (cm); Crique
Negro, Columbia Forest, 1 5, 1 9 (usnm); Forest
Home, 1 9 (amnh); vicinity Union Camp, 2 99
(BM), 1 9 (CM). GUATEMALA. Alta Verapaz: Lan-
quin, vicinity Lanquin Cave, approx. 149 km WSW
Puerto Barrios, 1 $ (amnh). Izabal: 25 km SSW
Puerto Barrios, 1 3, 5 99 (tcwc).

The recorded distribution of the wrinkle-faced
bat extends from western (southern Sinaloa),
northeastern (southern Tamaulipas), and penin-
sular (Campeche and Quintana Roo) Mexico and

continues southeastward through Central America
at principally lower to upland elevations (sea level
to 1882 m). The records given here are the first
for Belize, Alta Verapaz, and Izabal.

The distribution of this unusual bat in Belize
reflects apparent ecological flexibility. Centurio se-
nex has been captured in low littoral forest and
mangrove swamp edge on the coastal sand strip
of Ambergris Caye, to about 720 m in evergreen
and semi-evergreen seasonal forest on the south-
ern slope of the Maya Mountains. Evergreen sea-
sonal and transitional forests, secondary forest, and
agriculturally disturbed areas provide additional
habitats. This bat was captured throughout the
year. Two males and one female were mist netted
and released at Orange Point, Toledo District.
Brother N. Sullivan collected (15-17 January) the
specimen from Alta Verapaz, but I assume the bat
was captured outside of Lanquin Cave. The spec-
imens from Izabal were obtained (February,
March) by D. C. Carter and field party. Field data
are limited, but four Centurio were captured over
a stream.

Diphylla ecaudata Spix, 1823

Specimens Examined— BELIZE. Cayo: vicinity
Augustine, 1 S (rom); San Antonio, 1 6 (fmnh).
Toledo: Crique Jute, 1 $ (amnh); San Antonio, 1
9 (fmnh); Santa Elena, 1 9 (fmnh).

The distribution of Diphylla ecaudata appears
primarily restricted along the Gulf side and in the
Yucatan Peninsula of Mexico southeastward
throughout Central America, where this bat occurs
from the coastal lowlands up into the mountainous
highlands (1880 m). The hairy-legged vampire bat
has been recorded from El Peten (McCarthy, 1 982)
and Quintana Roo (Jones et al., 1973). The spec-
imens reported here are the first records from Be-
lize.

Four of the localities represent village environ-
ments where Diphylla was captured (April, July,
August, December) along with Desmodus rotundus
during vampire bat control activities. Mist netting
was carried out in direct immediacy to domestic
livestock and homes. The feeding activities of £>/-
phylla in these villages were not documented, al-
though one blood meal was obtained for analysis.
P. Boreham, Imperial College Field Station, En-
gland, reported (in litt.) a weak precipitin reaction
for a mammal host from the blood meal sample
without a response for bird or reptile. It is not
known if this blood meal was obtained in the vil-

McCARTHY: DISTRIBUTION OF BATS

149

lage (Santa Elena). Gardner (1977) summarized
the sanguivorous preference ofDiphylla as for pri-
marily avian hosts. The hairy-legged vampire from
Augustine was apparently taken (22 February) in
a deciduous seasonal forest.

Family NATALIDAE

Natalus stramineus saturatus

Dalquestand Hall, 1949

Specimens Examined— BELIZE. Cayo: 1.6 km

NW Augustine, Rio Frio, 2S6,2 9i (fsm); 0.8 km
W Augustine, 2 66, 3 22 (cm); 1.5 km N Augustine,
5 22 (cm); Sibun Camp, Hummingbird Hwy. at
Silver Creek, 1 2 (fmnh). Orange Walk: Richmond
Hill (Goat Hill), 8.9 km SSW Orange Walk Town,
1 2 (cm). Stann Creek: Kendal, 1 6 (fmnh). Toledo:
vicinity Aguacate, 1 3, 3 22 (cm); 1.2 km E Agua-
cate, 1 2 (cm); Vista Hermosa, 3.7 km WNW Punta
Gorda, 8 66, 6 22 (fmnh).

The northern range of Natalus stramineus sa-
turatus extends from both northwestern (Sinaloa)
and northeastern (Nuevo Leon) Mexico, including
the Yucatan Peninsula, southeastward through
Central America where the number of records for
this species is noticeably reduced beyond Guate-
mala to Panama. Although predominately a low-
land species, elevations were recorded as high as
2400 m. The presence of the funnel-eared bat in
Belize was anticipated, as it appears to be well
reported throughout the Gulf-Caribbean versant.

Those specimens obtained (April, August, Sep-
tember) at roost sites in Belize were from caves.
Other capture localities include low riparian forest
and open areas bordering on forest, in orchard
habitats, and alongside a building.

Family VESPERTILIONIDAE

Subfamily VESPERTILIONINAE

Myotis elegans Hall, 1962

Specimens Examined— BELIZE. Belize: Belize
City, Landivar, 1 2 (amnh), 1 3, 1 2 (fmnh), 1 6
(msu); Mussel Creek, 7.5 km W Burrell Boom, 1

6, 1 2 (fmnh).

LaVal (1973a) summarized the lowland distri-
bution of Myotis elegans. ranging from the Gulf
(eastern San Luis Potosi, Veracruz), Pacific coastal
(Chiapas), and peninsular (southeastern Cam-

peche) regions of Mexico to Honduras, Nicaragua,
and northeastern Costa Rica. Subsequent records
were reported from the Pacific side of Costa Rica
and the Caribbean lowlands of El Peten (LaVal,
1977; McCarthy, 1982). The majority of eleva-
tions are less than 1 20 m, ranging to 750 m. These
additional Caribbean lowland localities are the first
records from Belize.

Two elegant Myotis were netted (1 July) along
a tractor track, in low riparian vegetation domi-
nated by bamboo and thistle palms. Four indi-
viduals were obtained (January, February, May,
December) at a coastal locality in low vegetation
bordering on disturbed mangrove (Rhizophora
mangle, Avicennia germinans) habitat.

Eptesicus furinalis gaumeri

(J. A. Allen, 1897)

Specimens Examined— BELIZE. Belize: Belize
City, Landivar, 1 2 (cm). Cayo: Central Farm, 2
22 (cm), 5 66, 16 22 (fmnh), 2 66,19 (TTu); Little
Vaquero Creek, 9.3 km NNW Augustine, 1 5, 1 2
(fsm); Ontario, 5.5 km W Teakettle, 1 2 (fmnh);
Teakettle, 1 6 (fmnh). Corozal: Estero Lagoon, 4
km W Patchakan, 1 3, 1 2 (fmnh); Santa Clara, 1
2 (fmnh). Orange Walk: Honey Camp Lagoon, 1
6, 2 22 (fmnh); Tower Hill, B.S.I, compound, 3 66
(CM), 1 5, 4 22 (fmnh); 2 km SSW Tower Hill
Bridge, 1 2 (cm). Stann Creek: Melinda, 3 22 (fmnh);
Dangriga (Stann Creek), 1 <5 (usnm). Toledo: Or-
ange Creek, 1.5 km S Punta Gorda, 1 2 (msu);
Punta Gorda, 1 2 (msu).

The Mexican distribution of Eptesicus furinalis
gaumeri ranges from the western (Jalisco) and the
eastern (San Luis Potosi) versants southeastward
to South America. Davis (1965), Disney (1968),
and Starrett and Casebeer (1968) reported records
from all of the Central American countries except
El Salvador. Lowland elevations range from near
sea level to 1800 m, the majority being below 500
m. This tropical brown bat has been reported from
El Peten (Rick, 1968; McCarthy, 1982) and Quin-
tana Roo (Jones et al., 1973). The localities here
are additional records for Belize.

Disney ( 1 968) did not present locality data for
his two specimens of Eptesicus furinalis. Both were
males, captured (16 November, 29 December) in
Cayo District, near Central Farm and Esperanza
(4.5 km W Central Farm). These are located in
British Museum (Natural History). An additional
1 96 individuals were captured from three of the
localities reported here; the majority of these were

ISO

FIELDIANA: ZOOLOGY

banded and released during a behavioral study.
The majority was found in direct association with
buildings, utilizing the infrastructure of the walls
or floors and the space behind window shutters as
roost sites. Individuals have been taken over water
(creeks and a swimming pool) at three localities
and in riparian vegetation along two lagoons.

Lasiurus borealis (Muller, 1776)

Speomens ExAMnsfED— BELIZE. Orange Walk:
Tower Hill, B.S.I, compound, 1 9 (fmnh). Stann
Creek: 5.3 km NNW Quam Bank, Cockscomb
Basin, 1 9 (cm). GUATEMALA. El Peten: Parque
Nacional Tikal, 1 9 (fmnh).

The subspecies teliotis ranges southward from
both the western and eastern regions of Mexico to
Oaxaca and the northern Yucatan Peninsula.
Specimens of Lasiurus borealis from the Guate-
malan central highlands were assigned by Jones
(1966) to the Central American subspeciesyra«/z/7,
based on Handley (1960). Carter et al. (1966) as-
signed specimens from both lowland and highland
localities in Chiapas Xofrantzii, suggesting that the
region of the Isthmus of Tehuantepec represents
the break heXwecn front zii and teliotis. Hall (1981)
concurred with this arrangement. Similarly, Jones
et al. (1973) suggested that southern Mexico, in-
cluding the Yucatan Peninsula, may represent a
zone of intergradation between frantzii and teli-
otis. Few specimens of L. borealis are available
from El Salvador (Burt & Stirton, 1961), Honduras
(Goodwin, 1942b), Nicaragua (Davis & Carter,
1962— as L. b. teliotis), Costa Rica (Goodwin,
1 946; Gardner et al., 1 970) and Panama (Handley,
1 966). Recorded elevations (near sea level to about
2540 m) are primarily low or moderate (< 1 155
m). Koopman (1959) reported the only record from
Quintana Roo. This account represents the first
records for Belize and eastern Guatemala from El
Peten.

The red bats captured in Belize (April, May)
were netted over a stream and a swimming pool.
The Tikal specimen was taken (30 July) while it
was flying in an open area near a large man-made
reservoir.

I hesitate to assign a subspecific designation be-
cause I see no practical purpose in doing so until
adequate series of specimens from throughout the
range of Lasiurus borealis become available. Han-
dley (1960) had fewer specimens of L. borealis at
hand for a proper evaluation of subspecific vari-

ation. Consequently, the limits of the distributions
for the recognized subspecies remain unresolved.

Lasiurus ega (Gervais, 1855)

Specimens Examined— BELIZE. Belize: Trop-
ical Park, Mi. 14.5 Western Hwy., 1 S (fmnh).
Orange Walk: Tower Hill, B.S.I, compound, 2 6i
(FMNH), 1 (3 (cm). Stann Creek: 5.3 km WNW Quam
Bank, Cockscomb Basin, 2 66, I 9 (cm). Toledo:
Big Fall, 1.7 km NE Rio Grande Bridge, 1 9 (cm);
Orange Creek, 1 .5 km SW Punta Gorda, 1 6 (msu).

Similar to Lasiurus borealis, the distribution for
the two recognized subspecies of the yellowish bat
is not well understood. While L. e. panamensis
was recognized along the Pacific versant of Chia-
pas (Baker &. Patton, 1 967) and Guatemala (Dolan
& Carter, 1979;Dickermanetal., 1981), Goodwin
( 1 969) identified panamensis from the moderate
elevations of the Gulf drainage in northern Oaxaca
and suspected L. e. xanthinus may occur in the
drier Pacific portion of that state. Baker et al. ( 1 97 1 )
determined the variation in karyotypes and pelage
color of L. ega from near Brownsville, Texas, re-
sembled those from eastern coastal and southern
Mexico and referred the Texas specimens to L. e.
panamensis. Meanwhile, L. e. xanthinus was rec-
ognized in the Yucatan Peninsula (Jones et al.,
1973; Bimey et al., 1 974). The yellow bat is poorly
represented from the remainder of Central Amer-
ica, which includes Honduras (Goodwin, 1942b;
LaVal, 1969; Greenbaum &. Jones, 1978), Costa
Rica (Goodwin, 1946; Starrett & Casebeer, 1968;
Gardner et al., 1970; LaVal & Fitch, 1977), and
Panama (Handley, 1966). Where designated, the
subspecies panamensis has been applied to these
preceding Central American localities, although
Hall (1981) did not acknowledge panamensis north
of Costa Rica. Elevational data are similar to those
for L. borealis. Ingles (1958) reported two L. ega
from Quintana Roo. Jones et al. (1973, p. 23)
translated Ingles's locality from Spanish as "Puer-
to Morelos" when it was actually a collection site
only 16 km east of the state border with Yucatan,
along the highway from Valladolid (Yucatan) to
Puerto Morelos (Quintana Roo). Alvarez and Ra-
mirez-P. (1972) cited an additional Caribbean
lowland record from southeastern Campeche. This
account provides the first L. ega records from Be-
lize.

Eight yellowish bats were captured (April, May)
over streams, a river, and a swimming pool.
Another was netted ( 1 8 August) at about 5 m above

MCCARTHY: DISTRIBUTION OF BATS

151

the ground while circhng a building located in grass-
sedge savanna.

Lasiurus intermedius intermedius

(H. Allen, 1862)

Specimen Examined— BELIZE. Toledo: Crique
Jute, 1 9 (cm).

The range of this subspecies of the large yellow
bat extends southeastward from Mexico to Hon-
duras (Handley, 1960; Carter et al., 1966), El Sal-
vador (Hellebuyck et al., 1985), and Guatemala
(Carter et al., 1 966). Lasiurus intermedius has been
recorded in Mexico from the northern Yucatan
Peninsula and Chiapas northwestward to Texas
along the eastern coast and to Sinaloa on the Pa-
cific side. Recorded elevations range from lowland
to highland (1620 m) habitats. A single specimen
of L. intermedius from northern Quintana Roo
(Bimey et al., 1974) provided the only record for
that Mexican state. This Belizean specimen rep-
resents the first record for the country.

The above specimen was obtained on 30 March
over the stream Crique Jute surrounded by sec-
ondary vegetation.

Bauerus dubiaquercus (Van Gelder, 1959)

Specimens Examined— BELIZE. Cayo: 1.6 km

NW Augustine, Rio Frio, 1 2 (rom). Toledo: 2. 1
km NNE Salamanca Camp, Columbia Forest, 1 6

(CM).

The published localities of the rarely encoun-
tered Bauerus dubiaquercus are scattered from the
Islas Tres Marias (Nayarit), Jalisco, and southern
Veracruz in Mexico to eastern Honduras and Cos-
ta Rica (Engstrom & Wilson, 1981; Dinerstein,
1985). Mainland elevations range from approxi-
mately 460 to 1450 m and appear to represent
mid-elevation and montane forest habitats (Pine,
1 966; Pine et al., 1 97 1 ; Engstrom & Wilson, 1981;
Dinerstein, 1985). These first occurrences of Bau-
erus in Belize extend northward a scattered dis-
tribution along the northern Caribbean lowlands
in Central America.

J. Kamslra and J. Fragoso collected (8 July) one
specimen inside the main Rio Frio cave, located
in a deciduous seasonal forest at approximately
410m. The second Bauerus was netted (26 March)
along an open forestry track in an evergreen forest
at about 180 m.

Engstrom and Wilson (1981) and Martin and
Schmidly (1982) evaluated the taxonomic status
of Antrozous (Bauerus) dubiaquercus and con-
cluded the chromosomal, cranial, postcranial, and
phallic differences between this bat and Antrozous
(Antrozous) pallidus were sufficient to recognize
Bauerus as a distinct genus. I follow their conclu-
sions and agree that the species is monotypic since
the mainland sample size that previously was as-
signed to A. d. meyeri Pine, 1971, was limited to
a total of five specimens representing both sexes.

Family MOLOSSIDAE

Eumops auripendulus auripendulus (Shaw, 1 800)

Specimens Examined— BELIZE. Orange Walk:

Orange Walk Town, 1 ? (cm); Tower Hill, B.S.I,
compound, 1 9 (fmnh).

The recorded distribution of Eumops auripen-
dulus auripendulus includes both moist uplands
and drier lowland coastal and plateau areas, rang-
ing from eastern Oaxaca, Tabasco, Quintana Roo,
and Belize, through Guatemala, western Hondu-
ras, El Salvador, western Nicaragua, Costa Rica,
and Panama, into South America (Eger, 1974;
Greenbaum & Jones, 1978). Villa-R. (1956) and
Eger (1974), respectively, reported this free-tailed
bat from Quintana Roo and Belize (Belize District:
Rockstone Pond). This account provides the sec-
ond and third records for Belize.

The Orange Walk specimen consists of a man-
dible and partial skull, which were recovered from
an owl {Tyto alba) roost in a church tower. The
second specimen was discovered (July) alive by L.
G. Hoevers, after it apparently was attacked by a
bird.

Eumops bonariensis nanus (Miller, 1900)

Specimens Examined— BELIZE. Orange Walk:

Orange Walk Town, 2 ?? (cm).

Eger (1977) summarized the few available Mid-
dle American localities for this small mastiff bat,
which are limited to southeastern Mexico (south-
em Veracruz, Tabasco, Yucatan), eastern Hon-
duras, and Panama. These and additional locali-
ties in Panama (Dolan & Carter, 1979) and
Nicaragua (Hall, 1981) are restricted to coastal
lowland environments. This is the first recording
of Eumops bonariensis for Belize.

152

HELDIANA: ZOOLOGY

Entire specimens oi Eumops bonariensis as yet
are unavailable from Belize. Documentation is
based on two fragmented sets of maxillary tooth-
rows, which were sifted from regurgitated rubble
beneath an owl {Tyto alba) roost in a church tower.

Moiossus ater nigricans (Miller, 1 902)

Specimens Examined— GUATEMALA. Izabal:

25 km SSW Puerto Barrios, 2 33, 4 99 (tcwc).

The black mastiff bat is a common inhabitant
of roof spaces throughout its lowland Middle
American range, from western (Sinaloa) and east-
em (Tamaulipas) Mexico southeastward into South
America. This species has been reported from the
Caribbean lowlands of Quintana Roo (Jones et al.,
1973) and Belize (Murie, 1935; Pendergast, 1979).
These Moiossus ater from Izabal are the first rec-
ord for that department. Apparently, all of these
specimens were collected on 1 5 February over a
stream by D. C. Carter and his field party.

Addendum

While this paper was in press, other papers and
additional information concerning bats in Belize
came to my attention. Two recent papers provide
new country records of the glossophagines Li-
chonycteris obscura (Hill, 1985) and Hylonycteris
underwoodi (McCarthy & Blake, 1987), which
increase the known bat fauna to 68 species. Both
records are from Toledo District. McCarthy and
Blake (1987) also reported the occurrence of the
following bats: Rhynchonycteris naso, Saccopteryx
bilineata, Balantiopteryx io, Noctilio leporinus,
Pteronotus parnellii, Micronyctehs megalotis, M.
nicefori, M. schmidtorum, Tonatia evotis, Mimon
cozumelae, Phyllostomus discolor, Trachops cir-
rhosus, Chrotopterus auritus, Vampyrum spec-
trum, Glossophaga soricina, Carollia brevicauda,
C. perspicillata, Sturnira lilium, Uroderma bilo-
batum, Vampyressa pusilla, Artibeus jamaicensis,
A. lituratus, A. phaeotis, A. watsoni, Centurio se-
nex, Natalus stramineus, Rhogeessa tumida, Bau-
erus dubiaquercus, and Eumops underwoodi.

Hill, J. E. 1 985. The status of Lichonycteris degener
Miller, 1931 (Chiroptera: Phyllostomidae). Mam-
malia, 49(4): 579-582.

McCarthy, T. J., and M. Blake. 1 987. Noteworthy

bat records from the Maya Mountains Forest Re-
serve, Belize. Mammalia, 51(1): 161-164.

A bat specimen from Stann Creek was reported
(Miller & Allen, 1928, p. 180) as Myotis nigricans
nigricans. I examined this specimen at USNM and
found it to be an immature Eptesicus furinalis. It
is included in this paper under the species account
for the latter species.

Miller, G. S., Jr., and G. M. Allen. 1928. The
American bats of the genera Myotis and Pizonyx.
Bulletin of the United States National Museum, 144:
1-218.

Silva-Taboada and Koopman (1964, p. 3) re-
ported specimens of Tadarida laticaudata (= Nyc-
tinomops laticaudatus) from Corozal District. Most
of the bat species discussed in an unpublished dis-
sertation by A. M. Cartwright were also reported
by Kirkpatrick et al. (1975) and Cartwright and
Kirkpatrick (1977). The remaining identifications
(Cartwright, 1977, pp. 240, 242-246, 250, 251),
which were from Belize District, included Rhyn-
chonycteris naso, Saccopteryx bilineata, Carollia
brevicauda, C. perspicillata, Sturnira lilium, Arti-
beus lituratus, A. phaeotis, Des modus rotundus,
Eptesicus furinalis, Rhogeessa tumida, and Mo-
iossus moiossus. Baker et al. (1985, p. 236) re-
ported cytogenetic data from specimens of Rho-
geessa tumida that I collected in Belize District.

Baker, R. J., J. W. Bickham, and M. L. Arnold.
1985. Chromosomal evolution in Rhogeessa (Chi-
roptera: Vespertilionidae): Possible speciation by
centric fusions. Evolution, 39(2): 233-243.

Cartwright, A. M. F. 1977. Patterns of Neotrop-
ical chiropteran reproduction including histological
and ecological aspects of bats collected in Belize.
Ed.D. diss.. Ball State University, Muncie, Ind., 278
pp.

Silva-Taboada, G., and K. F. Koopman. 1964.
Notes on the occurrence and ecology of Tadarida
laticaudata yucatanica in eastern Cuba. American
Museum Novitates, 2175: 1-6.

A mammal checklist was included in a resource
paper on Belize (Hartshorn et al., 1984). The list
of bats supposedly was a compilation of known
and expected species. The result is inaccurate and
undocumented. The reader is referred to the
checklist in the Appendix as correct.

Hartshorn, G., et al. 1984. Belize. Country En-
vironmental Profile. A Field Study. United States

MCCARTHY: DISTRIBUTION OF BATS

153

Agency for International I>evelopment, San Jose,
Costa Rica. 151 pp.

Additional specimens of Micronycteris mega-
lotis (2: Belize District, Cayo District), Artibeiis
toltecus (11: Cayo District), Centurio senex (2: Cayo
District), and Diphylla ecaudata (3: Cayo District)
were found in the mammal collection of Royal
Ontario Museum. D. J. Tallman collected speci-
mens of M. megalotis ( 1 ) and Mimon cozumelae
(2) from Orange Walk District, which were de-
posited in Bell Museum of Natural History, Uni-
versity of Minnesota. I secured further voucher
specimens (AMNH) of Mimon cozumelae (1),
Vampyrodes caraccioli ( 1 ), Vampyressa pusilla ( 1 ),
and Bauerus dubiaquercus (1) from Toledo Dis-
trict.

Certain specimens (Sturnira lilium and Rho-
geessa tumidd) that were catalogued by Dobson
(1878, pp. 540, 246) were listed as collected in
"Honduras." These were obtained by D. Dyson
and H. Cuming between November 1844 and late
1845. During that time, "Honduras" correspond-
ed to the present region that extends from southern
Quintana Roo, Mexico, southeastward to northern
Honduras. Many early collectors did not differ-
entiate between the area of Belize ("British settle-
ment in Honduras") and that of the Republic of
Honduras ("Spanish Honduras"), but recorded
only "Honduras" or "Bay of Honduras" without
further locality data. The above specimens did not
originate from present day Honduras, but were
collected in Belize. Additional specimens of Mi-
cronycteris megalotis (Dobson, 1 878, p. 479) from
the "Bay of Honduras" and Rhynchonycteris naso
(Dobson, 1878, p. 368) from "Honduras" remain
orphaned records of the historical literature.

Uroderma bilobatum was reported (Sanchez-H.
et al., 1986) from southern Quintana Roo while
this volume was delayed. Ten specimens were col-
lected at Ruinas de Kohunlich (18°23'N; 88''42'W),
about 16 km W Estevez on the Belizean border.
Four other species {Pteronotus davyi, Mormoops
megalophylla, Tonatia evotis, T. minuta) were
documented for the first time from Quintana Roo,
from localities within 35 km of the northern bor-
der of Belize. The known bat fauna of Quintana
Roo is now represented by 36 species.

Sanchez-H., O., G. Tell£z-G., R. A. Medelun, and
G. Urbano-V. 1986. New records of mammals
from Quintana Roo. Mexico. Mammalia, 50(2): 275-
278.

Acknowledgments

Grateful acknowledgments are due H. Flowers,
O. Rosado, R. Belisle, and E. O. Bradley, E>e-
partment of Forestry, Belize, and H. Topsey, W.
Branche, E. A. Graham, and J. Palacio, Depart-
ment of Archaeology, Belize, for the necessary per-
mission to conduct fieldwork. The Department of
Agriculture, Belize, provided opportunities and
support while I headed the Vampire Bat Education
and Control Program (1975-1978). Division of
Mammals, Field Museum of Natural History, pro-
vided partial equipment needs (1974) and partial
funding ( 1979). Mellon North American Mammal
Research Institute, O'Neil Fund, and Section of
Mammals, Carnegie Museum of Natural History,
supported further fieldwork (1982, 1984). Nu-
merous colleagues and friends have been of direct
assistance to my needs in Belize, and the following
are but a few: E. Awe, P. Boreham, H. Bums, W.
D. Burton, J. Cab, G. Cal, G. C. Canto, J. Chun,
M. Craig, L. de la Torre, L. Dieckman, R. Foster,
P. W. Freeman, P. Gamble, E. Garcia, J. P. and
M. Garcia, H. H. Genoways, E. Gomez, L. Hen-
derson, R. W. Henderson, L. G. and E. Hoevers,
A. M. Hutson, R. J. Izor, T. Juring, E. King, K.
Leacock, D. Owen-Lewis, N. MacKenzie, G. T.
McCarthy, H. Pastor, R. H. Pine, A. Rabinowitz,
M. L. Reed, C. J. Rushin, K. and T. Salisbury, R.
Schmidt, B. M. Silva, M. F. Valentine, L. Waight,
L. Wilkins, and A. C. S. Wright. Various regiments
and the Royal Air Force of Her Majesty's British
Forces provided certain logistical support over the
years.

My fieldwork in Parque Nacional Tikal, El Pe-
ten, Guatemala, was made possible while assisting
D. J. Howell and J. G. Cant during their respective
visits to the park. M. Dary-R., Universidad de
San Carlos, and F. Polo-Sifontes, Instituto de An-
tropologia y Historia, graciously assisted and per-
mitted my work in 1979. J. R. Martinez R. kindly
provided a copy of his thesis.

The courtesies rendered to me by the personnel
of the following institutions while providing either
information, the loan of specimens, or assistance
during visits are greatly appreciated: American
Museum of Natural History; British Museum
(Natural History); Carnegie Museum of Natural
History; Field Museum of Natural History; Flor-
ida State Museum, University of Florida; Museum
of Natural History, University of Kansas; Mu-
seum of Zoology, Louisiana State University; The
Museum, Michigan State University; Royal On-
tario Museum; Texas Cooperative Wildlife Col-

154

HELDIANA: ZOOLOGY

lection, Texas A «& M University; The Museum,
Texas Tech University; United States National
Museum of Natural History. I am grateful to N.
A. Bitar, F. J. Bonaccorso, D. C. Carter, M. D.
Engstrom, A. M. Hutson, J. Kamstra, R. L. Pe-
terson, and E. L. Tyson for sharing information
concerning their field efforts in either Belize, El
Peten, Izabal, or Quintana Roo.

Besides J. R. Choate, J. K. Jones, Jr., K. F.
Koopman, and R. M. Timm, two anonymous re-
viewers enhanced earlier manuscripts with their
comments and editorial skills. E. Mendez kindly
reviewed my Spanish abstract. Verification of the
state boundaries in the Yucatan Peninsula was
provided by T. Dachtera, National Geographic
Society, and G. de la Torre M., Secretaria de Pro-
gramacion y Presupuesto, Mexico. M. A. Schmidt,
R. A. Rollin, V. Risoli, and S. Cramer typed var-
ious versions of the manuscript. S. M. Velez guid-
ed the manuscript during my absence. This report
is a contribution of the Mammals of Belize Pro-
gram.

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Gazetteer

The numbers for localities are plotted in Figure 1 .

MEXICO

QUINTANA ROO

GUATEMALA

El PETfeN

1. Bacalar 18°43'N; 88°22'W

2. Parque Nacional Tikal 1 7°20'N; 89°39'W

3. Poptun 16°2rN; 89''26'W

158

FIELDIANA: ZOOLOGY

Fig. 1. Distribution of localities in the districts of Belize; Quintana Roo, Mexico; and El Peten and Izabal,
Guatemala. The numbers refer to those listed in the Gazetteer. This map does not display the Caribbean lowlands
of Guatemala and Mexico in their entirety.

MCCARTHY: DISTRIBUTION OF BATS

159

IZABAL

4. Puerto Barrios 15°43'N; 88°36'W

BELIZE
CoROZAL District

5. Chan Chen 18°26'N; 88»27'W

6. Corozal 18°24'N; 88°24'W

7. Patchakan 18°24'N; 88''29'W

8. Santa Clara 18°18'N; 88°30'W

Orange Walk District

9. Honey Camp Lagoon 18°03'N; 88°27'W

10. Orange Walk Town 18°05'N; 88°34'W

11. San Antonio, Rio Hondo 18°1 1'N; 88°39'W

12. Tower Hill, Belize Sugar Industries (B.S.I.)

18°02'N; 88''34'W

Belize District

13. Belize City 17°30'N; 88°12'W

14. Churchyard 17°18'N; 88°33'W

15. Mussel Creek 17°39'N; 88°24'W

16. Rockstone Pond (Altun Ha) 17°46'N;

88°22'W

17. San Pedro, Ambergris Caye 17°55'N;

87°58'W

18. Tropical Park 17°28'N; 88''23'W

16°47'N:

30. Teakettle 17°13'N; 88°5rw

31. Xunantunich 17«t)5'N; 89°08'W

Stann Creek District

32. Kendal 16°49'N; 88°22'W

33. Melinda 17°00'N; 88°18'W

34. Quam Bank, Cockscomb Basin

88°28'W

Toledo District

35. Aguacate 16°10'N; 89°06'W

36. Big Fall 16''15'N; 88°53'W

37. Blue Creek 16°12'N; 89°03'W

38. Crique Jute and Salamanca Camp (Forestry

Camp) 16°16'N; 89°0rw

39. Crique Negro, Columbia Forest 16°17'N;

89°02'W

40. Cuevas Creek and Jacinto Creek Bridges, at

Punta Gorda Road 16°09'N; 88°53'W

41. Forest Home 16'^8'N; 88°50'W

42. Nimli Punit 16°20'N; 88''48'W

43. Orange Point 16°04'N; 88°49'W

44. Pueblo Viejo 16°13'N; 89°09'W

45. Punta Gorda 16°07'N; 88°48'W

46. Rice Station (Agricultural Station) 16°08'N;

88°51'W

47. San Antonio 16°15'N; 88°02'W

48. San Lucas (deserted) 16°05'N; 89°06'W

49. San Pedro Columbia 16°17'N; 88°58'W

50. Santa Elena 16°14'N; 89'X)6'W

51. Union Camp 16°24'N; 89°08'W

Cayo District

19. Augustine 16°58'N; 88°59'W

20. Baldy Beacon, Bald Hills 17°0rN; 88°47'W

21. Banana Bank 17°15'N; 88''48'W

22. Barton Creek at Western Hwy. 17°13'N;

89''57'W

23. Central Farm and Listowel 17°irN;

89°00'W

24. C.I.T.A.,Sibun River at Indian Creek 17''16'N;

88''34'W

25. Macaw Bank 17°05'N; 89'^4'W

26. Roaring Creek 17°15'N; 88''47'W

27. San Antonio 17°05'N; 89°0rW

28. San Luis 16°54'N; 89°00'W

29. SibunCamp 17°05'N; 88°39'W

Appendix

This district checklist of the bat fauna of Belize
is based on published accounts. The citations refer
to the initial taxonomic treatments of specimens.
Districts are arranged from north (left) to south
(right). Abbreviations are as follows: Cz = Coro-
zal; OW = Orange Walk; Bz = Belize; Cy = Cayo;
SC = Stann Creek; Td = Toledo.

In order to give an accurate list of bats, it is
necessary to present certain discrepancies that have
appeared in the literature. Dobson (1878) referred
to certain early sp>ecimens that may have origi-
nated from Belize. One reference (Dobson, 1878,

160

FIELDIANA: ZOOLOGY

p. 520) to "Half-Moon Key, Honduras" for a spec-
imen of Artibeus perspicillatus (= jamaicensis) be-
longs to Belize, since this specimen was collected
by O. Salvin (see Salvin, 1864). Sanderson (1941,
p. 228) recalled "Anoura sp." in his descriptive
narrative of a visit to then British Honduras. This
species was not identified (Hershkovitz, 1951) in
the Sanderson bat collection. Diaemus youngi was
cited from Belize (Villa-R., 1966, p. 340), but R.
L. Peterson (pers. comm.) stated that the specimen
in question was actually from Guyana. Specimens
of 17 species of bats were listed in Disney (1968)

without locality data. These specimens, which are
housed in British Museum (Natural History) and
Royal Ontario Museum, are all from Cayo Dis-
trict. J. L. Eger (pers. comm.) identified the ques-
tionable specimen of Molossus bondae in Disney
( 1 968, p. 7) as M. molossus. Quinones et al. ( 1 978,
p. 559) reported six species, which I collected and
identified, without the exact locality information
other than "the Maya Mountains region." This
locality is 1 km NW Augustine, Cayo District.
Sixty-six bat species are recognized in Belize.

Species

Cz OW

References

Rhynchonycteris naso

Murie, 1935, pp. 17-18; Disney, 1968, p. 7

Saccopteryx bilineata

Sanborn, 1937, p. 331; Hershkovitz, 1951,
p. 553; Disney, 1968, p. 7; Pendergast,
1979, p. 10

Saccopteryx leptura

This paper

Peropteryx kappleri

Cartwright & Kirkpatrick, 1977, p. 466

Peropteryx macrotis

Hershkovitz, 1951, p. 553

Centronycteris maximilliani

Sanborn, 1941, p. 372

Balantiopteryx io

Kirkpatrick et al., 1975, p. 330; Cartwright
& Kirkpatrick, 1977, p. 466

Diclidurus virgo

This paper

Noctilio leporinus

This paper

Pteronotus davyi

Disney, 1968, p. 7; this paper

Pteronotus parnellii

Hershkovitz, 1951, pp. 553-554; Smith,
1972, p. 74; Cartwright & Kirkpatrick,
1977, p. 466; Quinones et al., 1978, p.
559; Pendergast, 1979, p. 10

Pteronotus personatus

This paper

Mormoops megalophylla

This paper

Micronycteris brachyotis

This paper

Micronycteris megalotis

X X

This paper

Micronycteris nicefori

This paper

Micronycteris schmidtorum

X X

This paper

Lonchorhina aurita

This paper

Macrophyllum macrophyllum

This paper

Tonatia bidens

...

This paper

Tonatia evotis

Sanborn, 1941, pp. 372-373; Disney, 1968,

p. 7
Disney, 1968, p. 7; this paper

Tonatia minuta

Mimon cozumelae

This paper

Mimon crenulatum

Ruiz, 1983, p. 374; this paper

Phyllostomus discolor

This paper

Phylloderma stenops

This paper

Trachops cirrhosus

Sanborn, 1941, p. 374; Pendergast, 1979,
p. 10; this paper

Chrotopterus auritns

This paper

Vampyrum spectrum

This paper

Glossophaga commissarisi

Webster & Jones, 1982, p. 5; this paper

Glossophaga soricina

Villa-R., 1966, p. 231; Disney, 1968, p. 7;
Cartwright & Kirkpatrick, 1977, p. 466;
Howell, 1977, p. 510; Quinones el al.,
1978, p. 559; Pendergast, 1979, p. 10

Continued on next page

MCCARTHY: DISTRIBUTION OF BATS

161

Species

Cz OW Bz Cy SC Td

References

CaroUia brevicauda

Carollia perspicillata

Sturnira lilium
Uroderma bilobatum

Vampyrops helleri
Vampyrodes caraccioli
Vampyressa pusilla
Chiroderma villosum
Artibeus intermedius*
Artibeus jamaicensis*

Artibeus lituratus*

Artibeus phaeotis

Artibeus toltecus
Artibeus watsoni

Centurio senex
Desmodus rotundus

X
X

Diphylla ecaudata
Natalus stramineus
Thyroptera tricolor
Myotis elegans
Myotis keaysi

X
X

Eptesicus furinalis
Lasiurus horealis
Lasiurus ega
Lasiurus intermedius
Rhogeessa tumida

X
X
X

X
X

X
X
X

X
X

Bauerus dubiaquercus
Nyctinomops laticaudatus
Eumops auripendulus
Eumops bonariensis
Eumops glaucinus
Eumops underwoodi
Molossus ater

X
X

Molossus molossus
Molossus sinaloae

X
X

Hershkovitz, 1951, pp. 555-556; Villa-R.,
1966, p. 270; Disney, 1968, p. 7; Pine,
1972, p. 42; Quinones et al., 1978, p.
559; Pendergast, 1979, p. 10

Hershkovitz, 1951, p. 555; Disney, 1968,
p. 7; Pine, 1972, p. 42; Quinones et al.,

1978, p. 559; Pendergast, 1979, p. 10
Disney, 1968, p. 7; Pendergast, 1979, p. 10
Disney, 1968, p. 7; Davis, 1968, p. 697;

Pendergast, 1979, p. 10

This paper

Peterson, 1966, p. 676; this paper

This paper

Davis, 1984, p. 14

Dobson, 1878, p. 520; Andersen, 1908, p.
263; Gaumer. 1917, p. 298; Disney,
1968, p. 7; Davis, 1970b, p. 118; Pen-
dergast, 1971, p. 102

Murie, 1935, p. 19; Hershkovitz, 1951, pp.
556-557; Disney, 1968, p. 7; Pendergast,

1979, p. 10

Disney 1968, p. 7; Davis, 1970a, p. 398;

Pendergast, 1979, p. 10
This paper
Davis, 1970a, p. 393; Pendergast, 1979, p.

10
This paper
Hershkovitz, 1951, p. 557; Disney, 1968,

p. 7; Cartwright & Kirkpatrick, 1977, p.

466; Quinones et al., 1978, p. 559; Pen-
dergast, 1979, p. 10
This paper
This paj)er

Sanborn, 1941, p. 382
This paper
Hershkovitz, 1951, pp. 557-558; LaVal,

1973a, p. 25; Quinones et al., 1978, p.

559
Disney, 1968, p. 7; this paper
This paper
This paper
This paper
LaVal, 1973b, p. 29; Kirkpatrick et al.,

1975, p. 331
This paper
Murie, 1935, p. 19
Eger, 1974, p. 5; this paper
This paper
Eger, 1977, p. 42
Eger, 1977, p. 55
Murie, 1935, p. 19; Pendergast, 1979, p.

10
Murie, 1935, p. 19
Hershkovitz, 1951, p. 559; Disney. 1968,

p. 7

* Davis (1984) examined the Artibeus "lituratus" complex in Middle America and restored Artibeus intermedius
J. A. Allen to specific status. Specimens cited in the publications listed for A. jamaicensis and A. lituratus should be
reevaluated.

162

HELDIANA: ZOOLOGY

New Species of Mammals from Northern South America:
Fruit-Eating Bats, Genus Artibeus Leach

Charles O. Handley, Jr.

ABSTRACTS

The larger species of Artibeus of the Amazon Basin are defined, and a new giant species is
named and described from Venezuela and Colombia. Artibeus fallax, A. Hercules, and A. pla-
nirostris are regarded as subspecies of Artibeus jamaicensis, by far the most variable of the
larger Artibeus of the region.

The smaller Artibeus are keyed and arranged in six species groups. A new dwarf species is
described from Brazil, Ecuador, Guyana, Peru, and Venezuela. Distribution and diversity of
the smaller species are discussed. Artibeus cinereus, once thought to range throughout Central
America and much of South America and to include all of the smaller taxa except A. concolor
and A. hartii, is restricted to include only the nominate form and A. quadrivittatus of the lower
Amazon Basin and adjacent coastal areas.

With these additions and changes in status, at least nine species of Artibeus now are known
to occur in northeastern South America.

Las especies de gran tamaiio de Artibeus de la Cuenca del Rio Amazonas son definidas y una
nueva especie gigante de Venezuela y Colombia es nombrada y descrita. Artibeus fallax, A.
hercules, y A. planirostris son consideradas como subespecies de Artibeus jamaicensis, que es
el mas variable de los grandes Artibeus de la region.

Una clave es preparada para las especies de Artibeus menores, y las especies son arregladas
en seis grupos. Una nueva especie enana de Brasil, Ecuador, Guyana, Peru, y Venezuela es
descrita. La distribucion y la diversidad de las especies menores son discutidas. Artibeus cinereus,
que antes se penso estaba distribuida en Centro America y una gran parte de Sudamerica, y
que incluyera todas las taxa mas pequeiias (a excepcion de A. concolor y A. hartii), es ahora
restringuida para incluir solamente la especie nominal y A. quadrivittatus a la Cuenca baja del
Rio Amazonas y a las areas costeras adyacentes.

Con estas adiciones y cambios de "status," por lo menos nueve especies de Artibeus ya son
conocidas y se encuentran en el nordeste de Sudamerica.

Sao definidas as especies maiores de Artibeus que ocorrem na Bacia Amazonica, e uma especie
nova, gigante, e descrita. Artibeus fallax, A. hercules, e A. planirostris sao consideradas subes-
pecies de Artibeus jamaicensis, certamente a especie mais variavel dos Artibeus maiores da
regiao.

Uma chave para os Artibeus menores, os quais foram designados a seis grupos de especies,
e fomecida. Uma especie nova aiia e descrita do Brasil, Equador, Guiana, Peru, e Venezuela.
A diversidade, e as distribui^oes geograficas destas especies, sao discutidas. Artibeus cinereus,
o qual acreditava-se abranger toda America Central e grande parte da America do Sul, alem
de incluir todos taxa menores com excessao de A. concolor e A. hartii, e reduzido a um unico
taxon, restrito ao sul da Bacia Amazonica e as suas areas adjacentes.

Incluindo as adi9oes e mudan^as de status propostas neste trabalho, sao reconhecidas, atual-
mente, ao menos nove especies de Artibeus na regiao nordeste da America do Sul.

From the National Museum of Natural History,
Smithsonian Institution, Washington, D.C. 20560.

HANDLEY: NEW SPECIES OF ARTIBEUS 163

Introduction

Mammals and their ectoparasites were collected
in Venezuela between 1965 and 1968 by the
Smithsonian Venezuelan Project (SVP), supported
in part by a contract (DA-49-MD-2788) of the
Medical Research and Development Command,
Office of the Surgeon General, U.S. Army. Nu-
merous papers have described the ectoparasites
and mammals of the Project. Throughout these
papers undescribed species of mammals have been
referred to by alphabetical designations. Some of
these have been named subsequently by Handley
and Ferris (1972), Handley and Gordon (1980),
and Handley (1984). This paper deals with fruit-
eating bats of the genus Artibeus Leach.

The cranial measurements reported here were
taken as outlined by Handley (1959, p. 98). Hind
foot, tibia, calcar, and forearm were measured on
dry museum si)ecimens or on specimens preserved
in alcohol; all other external dimensions were
measured on fresh specimens in the field. All mea-
surements are in millimeters. Coloration was de-
termined under Examolites (Macbeth Corp., New-
burg, NY 12533) with natural light excluded.

A New Giant Artibeus

It is now generally agreed that in and around
the Amazon Basin there are three large species of
Artibeus. Handley (1976) recognized them as: (1)
/1./m//^;>jo51« Gray— smaller, molars 3/3, rostrum
arched, postorbital process poorly developed, fur
long, coloration blackish, facial stripes faint or ab-
sent, interfemoral membrane (IM) naked; (2) A.
jamaicensis Leach — larger, molars 3/3, rostrum
arched, postorbital process poorly developed, fur
short, coloration gray-brown, facial stripes present
but not sharply defined, IM naked; and (3) A. li-
turatus Olfers— larger, molars 2/3, rostrum flat-
tish, postorbital process well developed, fur short,
coloration chocolate brown, facial stripes promi-
nent and well defined, IM hairy. However, as shown
by Koopman (1978) and Honacki et al. (1982),
there is no consensus on the delimitation of these
species.

The difficulty in defining the species arises pri-
marily from the fact that Artibeus jamaicensis is
unusually variable geographically in morphology.
The other species show very little variation in this
region. Artibeus jamaicensis is large in the Ama-

zon Basin, so large in fact that the subspecies there,
A. j. fallax Peters and A. j. Hercules Rehn, until
recently have been aligned by most authors with
the universally large A. lituratus, although they
differ from it in many characteristics other than
size. To the southeast of the Amazon Basin {A. j.
planirostris Spix) and to the north of it (A. j. trin-
itatis Andersen), A. jamaicensis is dramatically
smaller, in fact similar in size to A. fuliginosus.
Everywhere east of the Andes A. jamaicensis has
3/3 molars; west of the Andes and in Central
America it has 2/3 molars.

Specimens in the SVP collection show that the
large Artibeus jamaicensis fallax and small A. j.
trinitatis apparently intergrade in the Llanos of
Venezuela where the habitat is marginal for A.
jamaicensis and where it is an uncommon bat.
Furthermore, intergradation between the small,
1 2-molar A. j. trinitatis and the slightly larger, 10-
molar A. j. aequatorialis Andersen of the north-
west coast of South America can be seen in spec-
imens from northern Colombia.

These two zones of intergradation are of crucial
importance in the nomenclature oi Artibeus, for
they serve to link "/I. jamaicensis^'' of the West
Indies and Central America and "A. planirostris"
of eastern South America. They are especially im-
portant in the present context because of the dis-
covery of a fourth large Artibeus, superficially sim-
ilar to but larger than A. j. fallax, occurring together
with it in southern Venezuela and with the small
A. j. trinitatis in western Venezuela and northern
Colombia. It can be recognized as follows:

Artibeus amplus new SF>ecies

HoLOTYPE— USNM 440932, adult female with
suckling young, skin and skull, collected 1 5 April
1968 by Norman E. Peterson, F. P. Brown, Jr.,
and J. O. Matson at Kasmera, 2 1 km SW Machi-
ques, Estado Zulia, Venezuela, 270 m, in a damp
cave in a cliff across the Rio Yasa from the Kas-
mera Biological Station, eastern foothills of the
Sierra de Perija. Original number, svp 22086.

Etymology— Latin amplus, large, referring to
the large size of this bat, one of the largest Artibeus.

Distribution— Northern foothills of the Cen-
tral Andes in Colombia; lower eastern slopes of
the Sierra de Perija and the Venezuelan Andes in
western Venezuela; and the vicinity of Cerro Dui-
da and the low mountains of southeastern Bolivar
in southern Venezuela. It probably occurs in ad-

164

HELDIANA: ZOOLOGY

jacent parts of Guyana and Brazil as well. The SVP
collectors found A. arnplus near streams and in
other moist areas (98%); in evergreen forest (90%)
and in forest openings such as yards and orchards
(10%). Most specimens were mist netted (86%),
but some ( 1 4%) were found roosting in the twilight
zone of caves. Elevational range, 24-1200 m.
Holdridge life zones (Ewel & Madriz, 1 968): Trop-
ical humid forest (10%), Tropical very humid for-
est (22%), Premontane humid forest (12%), Pre-
montane very humid forest (2%), Premontane rain
forest (4%), Lower montane very humid forest
(10%), and Lower montane rain forest (40%). Ridge
slopes and valley floor in the area where the ho-
lotype was collected were clothed with second
growth evergreen forest, while lawns, shrubbery,
banana and papaya plants, and scattered grapefruit
trees characterized the grounds of the biological
station.

Description— Size large (forearm 70.0, greatest
length of skull 31.3, maxillary toothrow 11.2 —
averages of Venezuelan specimens). Coloration of
fur as in sympatric Artibeus jamaicensis (dorsum
blackish brown to brown; facial stripes present but
obscure; underparts blackish brown, usually frost-
ed with white; underarms with abundant long,
usually whitish hairs); ears dark fuscous to black,
paler basally; lips and noseleaf blackish; mem-
branes blackish; wing tips undifferentiated or gray-
ish, never white. Horseshoe of noseleaf bound
down mediobasally; legs and interfemoral mem-
brane slightly hairy, the latter particularly medio-
ventrally, where hairs extend as a short fringe be-
yond edge of membrane; forearm long.

Skull superficially like that of Artibeus jamai-
censis, but relatively longer and narrower; rostrum
long and flattish; supraorbital ledges subparallel
and together with postorbital processes often poor-
ly develojDed or even ill-defined; zygomata not very
flared from skull, usually subparallel to one another,
and in side view, thin and fragile; posterolateral
angle of skull not particularly flared; palate rela-
tively narrow and toothrows ovoid in outline;
postpalatal extension usually long, narrow, and
parallel sided; dentition as in A. jamaicensis, ex-
cept I' only weakly bilobed; dental formula 2/2-
1/1-2/2-3/3 X 2 = 32. This bat is the only known
host of Strebla paramirabilis Wenzel and Tri-
chobius assimilis Wenzel (Diptera: Streblidae), so
it can be distinguished from other Artibeus by its
parasites as well as its morphology.

Measurements of the holotype, an adult female:
total length 101, tail vertebrae 0, hind foot (dry)
17, ear from notch 25, forearm 69.2, tibia 24.1,

calcar 6.2, weight 70.4 g. Greatest length of skull
31.9, zygomatic breadth 18.3, postorbital breadth
7.7, breadth of braincase 13.3, depth of braincase
1 1.6, length of maxillary toothrow 1 1.2, postpal-
atal length 9.8, palatal breadth outside of M^ 12.9,
rostral breadth at base of canines 8.3. See Table
1 for additional measurements.

Comparisons— Four large species of Artibeus
occur in Venezuela, all of them together in the
southern part of the country. Among these, Arti-
beus amplus and A. jamaicensis are most alike;
but despite the superficial resemblance, the two
can be distinguished by many characters, both ex-
ternal and cranial. All A. amplus examined have
the lower edge of the noseleaf horseshoe bound
down, while about 95% of A. jamaicensis from the
same localities have it free; all A. amplus have the
interfemoral membrane slightly hairy and fringed
medially, hwX A. jamaicensis never does; and while
A. jamaicensis often has the wings white-tipped,
A. amplus never does. Cranially, A. amplus differs
from A. jamaicensis in having a longer, narrower
skull; longer, somewhat more ffattened rostrum
(most easily seen in dimensions of rostral shield);
less arched nasals; margins of supraorbital nearly
parallel, rather than converging posteriorly, and
usually not as well developed; zygomata thinner
and more fragile and usually subparallel rather
than diverging markedly posteriorly; posterolater-
al angle of skull not so flaring; palate narrower and
toothrows usually less nearly circular in outline;
and postpalatal extension usually longer and nar-
rower, parallel sided (not flaring posteriorward).
The two species are hosts of different species of
parasitic streblid ffies.

Specimens Examined— Total 55. COLOMBIA.
Antioquia: La Tirana, 33 km SW Zaragoza, 520
m (2 usnm). VENEZUELA. Apure: Nulita, Selvas
de San Camilo, 29 km SSW Santo Domingo, 24
m (2 usnm). Bolivar: 21 to 33 km NE Icabani,
775-851 m (6 usnm); Km 125, 85 km SSE El
Dorado, 826-1165 m (5 usnm). T.F. Amazonas:
Belen, Rio Cunucunuma, 56 km NNW Esmeralda,
1 50 m (9 usnm); Cabecera del Caiio Culebra, Cerro
Duida, 40 km NNW Esmeralda, 1 1 40-1 200 m (2 1
usnm); Caiio Culebra, Cerro Duida, 50 km NNW
Esmeralda, 800 m (2 usnm); Tamatama, Rio Ori-
noco, 2 km above Boca del Casiquiare, 135 m (2
usnm). Zulia: Kasmera, 21 km SW Machiques,
270 m (3 USNM, 1 ucv); 15 km W Machiques (1
amnh); Novito, 19 km WSW Machiques, 1 135 m
(1 usnm).

Remarks— In previous publications of SVP, Ar-
tibeus amplus has been known as ""Artibeus sp. D".

HANDLEY: NEW SPECIES OF ARTIBEUS

165

Table 1 . Measurements of adult Artibeus ampins and A. jamaicensis. For each measurement, line 1 includes the
mean plus or minus two standard errors, line 2 the extremes, and line 3, in parentheses, the number of specimens
measured. All specimens are from Venezuela unless otherwise stated.

Total length

100.4 ± 3.88
93-104
(5)

89.9 ± 2.22

80-100

(21)

86.4 ± 2.90

77-95

(19)

83.8 ± 4.14

73-91

(10)

88.1 ± 1.62

80-93

(17)

86.3 ± 1.60

82-90

(8)

Hind foot
(dr>)

Ear

Forearm

Greatest
length

Zygomatic
breadth

15.9 ± 0.40
14-18
(19)

15.4 ± 0.32
15-16
(10)

17.8 ± 0.32
16-19
(17)

18.3 ± 0.32

18-19

(8)

16.9 ± 0.30

16.0-17.7

(12)

17.0 ± 0.28
16.6-17.6

(6)

19.1 ± 0.14
18.7-19.6

(13)

19.3 ± 0.32

18.6-20.2

(8)

Postorbital
breadth

Artibeus amplus, males and females, Zulia and Colombia
18.4 ± 0.60 23.7 ± 1.28 70.8 ±1.78 31.4 ± 0.26 18.6 ± 0.20
17-19 22-26 68.6-75.3 31.0-31.9 18.1-18.8

(7) (7) (7) (7) (7)

A. amplus, males and females, T.F. Amazonas and Bolivar
18.3 ± 0.30 23.0 ± 0.98 69.1 ± 0.90 31.2 ± 0.24 18.4 ± 0.14
17-20 18-26 65.0-73.2 30.3-32.8 17.4-19.1

(22) (21) (22) (29) (30)

A. jamaicensis, females, Zulia
22.6 ± 0.96 61.1 ± 0.64 27.7 ± 0.24
17-25 58.9-64.2 26.7-28.5

(19) (19) (19)

A. jamaicensis, males, Zulia
22.2 ± 0.98 59.3 ± 0.92 27.4 ± 0.14
20-25 56.2-61.4 27.1-27.7

(10) (10) (10)

A. jamaicensis, females, T.F. Amazonas
24.6 ± 0.46 66.8 ±1.12 30.7 ± 0.28
23-26 62.1-70.1 29.4-31.3

(17) (17) (14)

A. jamaicensis. males, T.F. Amazonas
24.6 ± 0.52 65.4 ± 1.42 30.7 ± 0.26
24-26 62.4-68.6 30.2-31.4

(8) (8) (8)

7.9 ± 0.20

7.6-8.3

(7)

7.8 ± O.IO

7.3-8.4

(31)

6.8 ± 0.08

6.5-7.1

(19)

6.8 ± 0.14

6.3-7.0

(10)

7.5 ± 0.10
7.1-7.7

(14)

7.6 ± 0.16

7.2-7.9
(8)

A New Dwarf Artibeus

The taxonomy of the smaller Artibeus is in a
state of flux. As recently as 35 years ago all of the
smaller species except A. concolor Peters and A.
hart a Thomas were believed to be variants of .-i.
cinerens Gervais. Since then, first one and then
another of the supposed subspecies of .4. cinereus
has been shown to be independent species. Today
only A. bogotensis Andersen, A. glaucus Thomas,
A. pumilio Thomas. A. quadrivittatus Peters. A.
rosenbergi Thomas, and A. watsoni Thomas re-
main associated with A. cinereus (Honacki et al.,
1982). However, except for^. quadrivittatus, these
do not properly belong with A. cinereus either.

Artibeus glaucus and A. bogotensis intergrade in
Ecuador and form an Andean-northern South
American species sympatric with A. cinereus in
southern Venezuela. Artibeus glaucus thus has two
subspecies, the nominate form and A. g. bogoten-
sis. Artibeus watsoni Thomas of northwestern South
Ameinca and Central America is closely related,
but intergradation with A. g. glaucus or A. g. bo-
gotensis has not been obsei^ed.

Artibeus pumilio is an enigmatic taxon. Many
museum specimens bear the name A. pumilio, but
perhaps the only specimen properly associated with
the name is the holotype. This specimen may be
only an odd variant of one of the other species,
but not of the species described here. For the pres-
ent, A. pumilio must be regarded as unplaceable.
The same can be said for A. rosenbergi, charac-
teiized by a curiously long, narrow skull such as
can be found occasionally in large samples of most
species o{ Artibeus. Because of their equivocal sta-
tus, neither A. pumilio nor A. rosenbergi is includ-
ed in the appended list of species and key. The
characteii sties and status of these taxa will be the
subject of another paper.

Thus, .A. cinereus now has been shorn of all of
its supposed subspecies except A. c. quadrivittatus.
Its supposed range has been reduced from encom-
passing most of Central Ameiica and tropical South
America to occupying only the Amazon Basin
(possibly only the lower basin) and adjacent coast-
al areas. Sympatric with A. cinereus in much of its
range is a distinctive dwarf species which can be
known as:

166

HELDIANA: ZOOLOGY

Table 1. Continued.

Braincase
breadth

Braincase
depth

Maxillary Postpalatal
toothrow length

Width at
molars

Width at
canines

Tibia

13.5 ± 0.20

13.2-14.0

(7)

Artibeus amplus. males and females,
1 1.0 ± 0.26 1 1.2 ± 0.16 9.7 ± 0.20
10.6-11.6 11.1-11.5 9.3-10.0
(7) (7) (7)

Zulia and Colombia

13.2 ± 0.26 8.4 ±0.16
12.7-13.5 8.2-8.8
(7) (7)

25.9 ± 1.28

24.1-28.2

(6)

13.3 ± 0.12

12.9-14.0

(30)

A. amplus, males and females, T.F. Amazonas and Bolivar
11.2 ±0.06 11.2 ±0.10 9.8 ±0.14 13.3 ± 0.10 8.6 ± 0.08
10.7-11.5 10.7-11.8 9.1-10.6 12.8-13.9 8.3-8.9
(30) (31) (28) (30) (30)

24.8 ± 0.38

23.1-26.2

(22)

12.3 ± 0.10

12.0-12.8

(19)

10.2 ± 0.16

9.5-10.9

(19)

A. jamaicensis. females
10.0 ±0.12 8.6 ±0.14
9.4-10.5 8.2-9.3
(17) (18)

, Zulia

12.1 ± 0.16

11.2-12.7

(19)

7.6 ± 0.10

7.2-8.0

(19)

22.4 ± 0.50

20.1-23.8

(19)

12.1 ± 0.14

11.8-12.4

(10)

10.3 ± 0.18

10.0-10.9

(10)

A. jamaicensis. males,
10.1 ± 0.16 8.4 ± 0.12
9.7-10.4 8.2-8.8
(8) (10)

Zulia

12.2 ± 0.18

11.8-12.6

(10)

7.7 ± 0.12

7.4-7.9

(9)

21.7 ± 0.52

20.1-23.0

(10)

13.2 ± 0.12

12.9-13.5

(14)

10.8 ± 0.18

10.2-11.5

(14)

A. jamaicensis. females, T.F
11.4 ±0.20 9.4 ±0.18
11.0-12.0 8.9-10.0
(13) (14)

. Amazonas
13.7 ± 0.22
13.2-14.4
(13)

8.6 ± 0.12

8.3-8.9

(13)

24.1 ± 0.46

22.3-25.4

(17)

13.4 ± 0.22

12.8-13.7

(8)

11.0 ± 0.18

10.7-11.4

(8)

A. jamaicensis, males, T.F.
11.4 ±0.18 9.3 ± 0.24
11.1-11.8 8.7-9.8
(8) (8)

Amazonas
13.8 ± 0.30
13.0-14.3
(8)

8.8 ± 0.12

8.6-9.1

(8)

23.3 ± 0.52

22.0-24.4

(8)

Artibeus gnomus new species

HoLOTYPE— USNM 387534, adult female, skin
and skull, collected 14 June 1966 by A. L. and M.
D. Tuttle at El Manaco (= Km 74), 59 km SE El
Dorado, Bolivar, Venezuela, 150 m, in a mist net
in an orchard. Original number, svp 9298.

Etymology— Latin gnomus, diminutive fabled
being, dwarf, alluding to the small size of this
species, one of the smallest Artibeus.

Distribution— The Amazon Basin and bor-
dering regions; from northern Amazonas Territory
(14 km SSE Pto. Ayacucho) and northern Bolivar
State (28 km SE El Manteco) in Venezuela and
northern Guyana, to Para (Belem) and Mato Gros-
so (Serra do Roncador), Brazil, and Loreto (Santa
Rosa), Peru. SVP collectors netted A. gnomus
mostly in moist sites (92%) in evergreen forest
(52%) or openings such as savannas (25%) and
yards and orchards (23%). Elevations range 1 19-
161 m in Venezuela, sea level to 530 m in Brazil.
Holdridge life zones: Tropical dry forest (22%),
Tropical humid forest (67%), Tropical very humid
forest (2%), and Premontane humid forest (9%).

Description— Body size small (forearm aver-
ages 36-38, greatest length of skull 18.5-18.7, and
maxillary toothrow 5.7-6.0). Dorsal coloration
gray-brown to brown; underparts paler; facial
stripes very white and sharply defined. Soft parts
coloration in life (usnm 36 1 742, male, Belem, Bra-
zil): ear narrowly edged with yellow, brightest to-
ward base; antitragus entirely yellow; tragus yel-
low, brightest distally and on posterior basal lobe;
noseleaf and horseshoe gray-brown medially,
cream color laterally; lips and chin gray-brown;
iris brown; forearm and fingers brownish flesh col-
or; wings blackish, except membrane between fin-
gers II and III transparent, grayish; interfemoral
membrane sooty brown; legs and feet dark brown;
claws horn color. Face short; shape and propor-
tions of ears, noseleaf, horseshoe, lips, chin, and
interfemoral membrane as in Artibeus cinereus;
noseleaf minutely hirsute; lower edge of horse-
shoe free; basal part of forearm hairy; hind ex-
tremities (except for short hairs on feet) appear
naked.

Skull small, short, and broad; zygomata sub-
parallel; rostrum narrow, very short, moderately

HANDLEY: NEW SPECIES OF ARTIBEUS

167

Table 2. Measurements of adult male and female (combined) Artibeus gnomus and A. glaucus bogotensis. For
each measurement, line 1 includes the mean plus or minus two standard errors, line 2 the extremes, and line 3, in
parentheses, the number of specimens measured. All specimens are from Venezuela.

Total length

47.5 ± 1.40

44-54

(13)

52.2 ± 0.86

49-56

(19)

Hind foot
(dry)

Ear

Forearm

Greatest
length

Zygomatic
breadth

A. glaucus bogotensis. Km 125, 85 km SSE El Dorado
10.6 ± 0.22 17.4 ±0.38 39.6 ± 0.60 20.3 ± 0.10 11.6 ±0.10
10-11 16-19 36.8-41.9 19.4-21.2 10.8-12.1

(19) (19) (19) (50) (44)

Postorbital
breadth

Artibeus gnomus, Rio Supamo, Los Patos, and EI Manaco
9.5 ± 0.28 16.9 ± 0.62 36.7 ± 0.54 18.5 ± 0.18 11.0 ±0.18
9-10 14-19 34.0-38.3 17.9-19.1 10.4-11.2

(13) (13) (13) (14) (8)

4.9 ± 0.10

4.5-5.2

(14)

5.0 ± 0.04

4.6-5.3

(51)

deep and arched, and much swollen posterolater-
ally (part on rostral shield, part within orbit, above
eye); excavation for orbital nerve large and deep;
braincase short and deep, with swelling at pos-
terodorsal apex interrupting junction of sagittal
and lambdoidal crests; postpalatal extension rel-
atively short; internal edge of pterygoid fossa
strongly ridged, narrowing mesopterygoid fossa and
cupping pterygoid fossa which opens straight back;
vomerine ridge visible in mesopterygoid fossa; va-
cuities in roof of posterior nares much anterior to
mesopterygoid fossa and not easily seen; outline
of maxillarv toothrows nearly circular; upper ca-
nine small (especially in basal diameter); M' with
accessory internal ridge on lateral cusps, and with
relatively wide talon; m, present (75 of 79 speci-
mens examined).

Measurements of the holotype, an adult female:
total length 47, tail vertebrae 0, hind foot (dry) 9,
ear from notch 18, forearm 36.5, tibia 12.6, calcar
4.9, weight 10.5 g. Greatest length of skull 18.2,
zygomatic breadth 10.8, postorbital breadth 4.8,
breadth of braincase 8.5, depth of braincase 7.2,
length of maxillary toolhrow 5.5, postpalatal length
6.5, palatal breadth outside of M' 7.1, rostral
breadth at base of canines 4.6. See Table 2 for
additional measurements.

Comparisons— ^r//Z)e«5 gnomus differs from A.
concolor and A. hartii in many ways, but most
significantly in lack of M\ From all other small
Anibeus {A. anderseni, A. cinereus, and A. glaucus
bogotensis) that occur within its range, A. gnomus
can be distinguished by its possession of mj. Among
the specimens examined, m, is consistently absent
in these other taxa while it is consistently present
in A. gnomus (except in southern Venezuela, where
it is absent from both mandibles in four of 53
specimens and from one mandible only in two
others). In addition, A. gnomus differs from all of

the sympatric taxa in its more prominent white
facial stripes; more colorful ears, noseleaf, and lips;
average browner, less grayish coloration of pelage;
shorter face and rostrum (except when compared
with A. concolor); more swollen supraorbital re-
gion; average larger and deeper orbital nerve ex-
cavation (sometimes equally large and deep in A.
g. bogotensis); and more cupped pterygoid fossa,
with internal ridge so enlarged as to significantly
narrow the mesopterygoid fossa.

Artibeus gnomus differs from the sympatric taxa
individually in several other ways. It is much
smaller than A. concolor (forearm averages 36-38
vs. 46-48). In contrast to A. hartii it has notched
inner upper incisors, brownish rather than dark
chocolate coloration, and a wide, unfringed inter-
femoral membrane. Compared with A. aruierseni
(including the holotype, fmnh 21331), ^. gnomus
is similar in size (slightly larger than Rio Madeira
A. anderseni); has rostrum much deeper, more
arched, narrower, and shorter; face not dished;
orbit larger; zygomata more nearly parallel; and
vacuities in roof of p)OSterior nares far forward of
mesopterygoid fossa, rather than opening in it or
close to it.

At Belem, Brazil, both Artibeus gnomus and A.
cinereus were numerous and were often taken in
the same nets. There, fresh specimens of the two
species were compared. Artibeus gnomus is smaller
in size, and has a smaller head and shorter face;
facial stripes much brighter, more sharply defined,
and more prominent; ears, noseleaf, and lips more
brownish, less grayish; ear edgings, antitragus, and
tragus bright yellow, rather than cream; and nose-
leaf edged with cream, rather than plain gray-
brown. Furthermore, it has zygomata more nearly
parallel; rostrum deeper and shorter; supraorbital
area much swollen and its edges nearly parallel;
and smaller teeth.

168

HELDIANA: ZOOLCX}Y

Table 2. Continued.

Braincase
breadth

Braincase
depth

Maxillary
toothrow

Postpalatal
length

Width at
molars

Width at
canines

Tibia

8.5 ± 0.14

8.1-9.0

(14)

9.0 ± 0.06

8.5-9.5

(48)

Artibeus gnomus, Rio Supamo, Los Patos, and El Manaco

7.4 ± 0.14

5.7 ± 0.06

6.3 ± 0.10

7.5 ± 0.12

4.9 ± 0.08 13.2 ± 0.46

7.1-8.0 5.5-5.9 6.0-6.7 7.1-7.9

(14) (14) (14) (14)

A. glaucus bogotensis, Km 125, 85 km SSE El Dorado
7.9 ± 0.06 6.5 ± 0.04 7.0 ± 0.08 8.0 ± 0.06
7.2-8.3 6.0-6.8 6.5-7.5 7.5-8.7

(48) (51) (47) (50)

4.6-5.1
(14)

5.1 ± 0.06

4.9-5.6

(50)

11.2-14.4
(13)

13.6 ± 0.36

12.3-15.8

(19)

In southern Venezuela Artibeus gnomus is sym-
patric with A. glaucus bogotensis. Compared with
Venezuelan specimens and with the holotype (bm
99.11.4.35) of this taxon, A. gnomus is much
smaller and shorter faced; has a deeper, shorter
rostrum; disproportionately wider zygomatic
spread; and smaller teeth.

In addition to comparisons of /I. gnomus with
sympatric species, two other small Artibeus need
to be considered:

1. Artibeus g. glaucus —This species occurs
nearby in the Andes. It (including the holotype,
BM 94.8.6.13) possesses m„ and its skull has the
basic shape of /i. gnomus. However, it is much
larger and darker in color, has the hind extremities
much hairier, the supraorbital region usually less
swollen, and the pterygoid fossa much less cupped
and opening to the mesopterygoid fossa.

2. Artibeus fvatsoni— West of the Andes and ex-
tending into Central America is another small
species, A. watsoni Thomas, which like A. gnomus
possesses m,. It (including its holotype, bm
0.7. 1 1 . 1 9) is larger than A. gnomus; has larger teeth;
longer rostrum, with reduced supraorbital swell-
ing; shallower and less well-defined orbital nerve
excavation; and like A. glaucus has the pterygoid
fossa not cupped and opening into the mesopter-
ygoid fossa (which consequently is not narrowed
by the inner pterygoid ridge).

Discussion— The ten small species of Artibeus
recognized here can be associated in six species
groups:

1. Artibeus concolor Group— Amazon and up-
per Orinoco basins and Guianas. Includes only
Artibeus concolor.

2. Artibeus hartii Group— Mexico and Central
America, across northern South America to Trin-
idad, and south to Peru east of the Andes and to
Ecuador west of the Andes. Includes only Artibeus
hartii.

3. Artibeus glaucus Group— Mexico, Central
America, and South America to Mato Grosso and
Peru. Includes Artibeus glaucus (with two subspe-
cies, A. g. bogotensis and A. g. glaucus), A. gnomus,
and A. watsoni.

4. Artibeus toltecus Group— Mexico and Cen-
tral America. Includes Artibeus aztecus Andersen
and Artibeus toltecus Saussure, each with several
subspecies.

5. Artibeus cinereus Group— Guiana region,
coastal Brazil, and lower Amazon Basin (dubious-
ly also upper Amazon Basin). Includes only Ar-
tibeus cinereus, with A. c. quadrivittatus as a sub-
species.

6. Artibeus phaeotis Group — Mexico, Central
America, and South America to upper Amazon
Basin and western Ecuador. Includes Artibeus an-
derseni Osgood and Artibeus phaeotis Miller, ' with
several subspecies.

Diversity in the small Artibeus is greatest in east-
ern South America, where representatives of five
of the six groups occur and where three of the
groups are endemic. Altogether six species occur
in and around the Amazon Basin, while only one
is known with certainty in the central portion of
the Basin; there are three in the lower Amazon

' Until recently (Koopman, p. 152, in Honacki et al.,
1982) it has not been generally recognized that Artibeus
phaeotis and A. ravus are conspecific. They inlergrade in
eastern Panama and western Colombia. Both names date
from Miller (1902). Although A. ravus was named first,
on an earlier page, A. phaeotis became embedded in the
literature as the name of this species.

HANDLEY: NEW SPECIES OF ARTIBEUS

169

and on the southern fringes in Brazil and Bohvia,
four or five in southern Venezuela, and five in east-
em Peru, Ecuador, and Colombia. In contrast, only
three of the species groups occur in Central Amer-
ica, and only one of them is endemic there.

Several distributional patterns are represented
in the complex of Amazonian species. Artibeus
concolor is found throughout the Basin but scarce-
ly beyond it; A. cinereus occurs in the lower Am-
azon and along the coast for some distance north
and south of the river; A. anderseni is known from
the upper Amazon and an isolated area in northern
Colombia; A. glaucus is higher up, in the Andes,
and eastward around the northern edge of the Ba-
sin in Venezuela; the range of A. hartii resembles
that of A. glaucus, but extends on into Central
America; and the dwarf /I. gnomus has a p)eculiar
circular range, completely ringing the Amazon Ba-
sin but apparently not extending into its interior.

Key to the Smaller Species of Artibeus

1. Molars 3/3 (mj large) 2

r. Molars 2/3 (mj minute) or 2/2 3

2. V notched; facial stripes absent; coloration
pale brown; interfemoral membrane broad
and naked; forearm 43-52 mm

Artibeus concolor

2'. I' not notched; facial strip>es present; color-
ation dark chocolate brown; interfemoral
membrane narrow and fringed; forearm 36-
42 mm Artibeus hartii

3. Supraorbital region much swollen; molars
2/3 (2/2 m A. g. bogotensis and occasionally
in the others) . . . Artibeus glaucus Group, 4

3'. Supraorbital region little, or not at all, swol-
len; molars 2/2 7

4. Rostrum short and moderately arched; pter-
ygoid fossa cupijed and opening back, causing
mesopterygoid fossa to be narrowed; forearm
34-38 mm Artibeus gnomus

4'. Rostrum long and much or only moderately

arched; pterygoid fossa not cupped, opening

into and not narrowing mesopterygoid fossa

5. Rostrum much arched; orbital nerve exca-
vation shallow and often ill-defined; dorsum
pale brownish; ears pale; forearm 35-41 mm

Artibeus watsoni

5'. Rostrum moderately arched; orbital nerve
excavation deep and well defined 6

6. Molars usually 2/3; dorsum dark grayish or
blackish; ears dark; forearm 38-42 mm . . .

Artibeus glaucus glaucus

6'. Molars 2/2; dorsum pale brownish or grayish;

ears pale; forearm 37-41 mm

Artibeus glaucus bogotensis

7. Interfemoral membrane narrow and fringed;
coloration blackish

Artibeus toltecus Group, 8

7'. Interfemoral membrane broad, naked; col-
oration brownish 9

8. Larger, forearm 42-48 mm

Artibeus aztecus

8'. Smaller, forearm 37-41 mm

Artibeus toltecus

9. Rostrum deep and arched; palate long and
moderately wide Artibeus cinereus

9'. Rostrum shallow and flattened; palate short

and very wide

Artibeus phaeotis Group, 10

10. Maxillary toothrow 5.2-6.2 mm; rostrum

often tilted up anteriorly

Artibeus anderseni

10'. Maxillary toothrow 6.7-7.1 mm; rostrum

usually not tilted up anteriorly^

Artibeus phaeotis

Specimens Examined— /irf/Aeiii anderseni —
BRAZIL. Amazonas: Borba, Rio Madeira (1
amnh). Rondonia: Porto Velho (2 amnh, 2 fmnh,
including holotype of ^. anderseni); Sto. Antonio
do Hauayara (4 amnh). COLOMBIA. Bolivar: Ca-
tival. Upper Rio San Jorge, 120 m (16 fmnh).
Antioquia: Aljibos, 26 km S and 22 km W Zara-
goza, 630 m (2 usnm); nr. La Tirana, 24 km S and
22 km W Zaragoza, 520 m (2 usnm). ECUAIX)R.
Napo: Rio Suno (Abajo) (4 amnh). Pastaza: Mon-
talvo, Rio Bobonaza ( 1 fmnh); Rio Pindo Yacu
(1 fmnh); Rio Yana Rumi (1 fmnh). PERU. Hua-
nuco: Monte Alegre (1 amnh). Loreto: Boca Rio
Curaray (1 amnh); Boca Rio Peruate, Rio Ama-
zonas, 90 m (1 FMNH); Lagarto, Alto Ucayali (1
amnh); Mazan (1 amnh); 59 km W Pucallpa (1
usnm); Puerto Indiana, Rio Amazonas (2 amnh);
Rio Morona (Quebr. Pushaga), Alto Amazonas,
220 m (2 FMNH); Rio Yavari Mirim (Quebr. Es-
peranza), 200 m (2 fmnh); Santa Cecilia, Rio Man-
iti, Iquitos, 110 m (3 fmnh); Santa Luisa, Rio
Nanay, Iquitos, 160 m (1 fmnh); Sarayacu, Rio
Ucayali (1 amnh). Pasco: San Juan, Oxapampa,

^ Couplet 10 will separate Artibeus anderseni and A.
phaeotis in South America and in southern Central
America, but it will not distinguish A. anderseni from
Mexican A. phaeotis nanus. In such a comparison, A.
anderseni can be recognized by its relatively broader
skull.

170

FIELDIANA: ZOOLOGY j

274 m (3 usnm). Departamento (?): Yuhucumayo,
1200 ft [= Puno: Yahuaramayo, 366 m?] (1 mcz).

Artibeus cinereus c/iiere«5— BRAZIL. Amazo-
nas: Sta. Clara, Vila Bela Imperatriz [nr. Parintins]
(1 amnh). Para: Fordlandia, Rio Tapajos (2 amnh);
Maracano, Rio Jamunda [= Nhamunda?], Faro (5
amnh); Rio Yumunda, Faro ( 1 bm).

Artibeus cinereus quadrivittatus — BRAZIL,.
Maranhao: Juryassu [= Turia9u?] (1 bm). Para:
Belem (10 usnm); Benevides (1 bm); Para [= Be-
lem] (1 bm); Ilha do Taiuna, Rio Tocantins (3
amnh). Pemambuco: Pemambuco [= Recife] (2
bm). Rio Grande do Norte: Natal (1 usnm). SUR-
IN AME. Surinam ( 1 bm). VENEZUELA. Bolivar:
Hato San Felipe, Serrania de Nuria (1 ucv); Hato
San Jose, 20 km W La Paragua, 300-324 m (2
usnm).

Artibeus glaucus bogotensis — COLOMBIA.
Cundinamarca: Bogota (2 bm): nr. Bogota ( 1 bm);
Curiche, nr. Bogota (2 bm, including holotype of
A. bogotensis); Fomeque (1 amnh); Fusagasuga (2
mcz); Rio Negro, nr. Bogota (2 bm). GUYANA.
Kanuku Mts. (3 bm). VENEZUELA. Bolivar: El
Manaco, 59 km SE El Dorado, 150 m (3 usnm);
Hato San Jose, 20 km W La Paragua, 300-324 m
(3 USNM); 23 to 45 km NE Icabaru, 824-851 m (3
USNM); Km 125, 85 km SSE El Dorado, 826-1 165
m ( 1 20 usnm); Rio Supamo, 50 km SE El Manteco,
1 50 m (2 usnm). T.F. Amazonas: Belen, Rio Cu-
nucunuma, 56 km NNW Esmeralda, 150 m (1
usnm); Caiio Culebra, Cerro Duida, 50 km NNW
Esmeralda, 800 m (3 usnm).

Artibeus glaucus glaucus— BOIAW A. Santa
Cruz: Buenavista, 400 m (1 fmnh). ECUADOR.
Napo: Baeza (1 bm). PERU. Cuzco: Collpa de San
Lorenzo, Quincemil, 700 m (1 1 fmnh); Hda. Ca-
dena, Quincemil, 1000 m (9 fmnh). Junin: Chan-
chamayo, 1000 m (2 bm, including holotype oi A.
glaucus); Huacapistana ( 1 f>4Nh). Puno: Rio Inam-
bari, 670 m (3 amnh); Santo Domingo (1 amnh);
Yahuaramayo, 366 m (1 bm, 1 usnm).

Artibeus gnomus-ToXdA 104. BRAZIL. Mate
Grosso: Serra do Roncador, 264 km N (by road)
Xavantina, 533 m (17 usnm). Para: Belem, Sta.
A, IPEAN (7 usnm); Belem, Utinga (5 usnm); Be-
lem, Benevides (2 usnm). ECUADOR. Pastaza:
Canelos, upper Rio Bobonaza (1 amnh). GUY-
ANA. E. Berbice District: Wikki River (3 usnm).
Mazaruni-Potaro District: Issano Road, 1 2 mi W
of Bartica-Potaro Road (1 usnm). PERU. Loreto:
59 km SW Pucallpa (1 usnm); Santa Rosa, Alto
Ucayali (10°42'S, 73''50'W) (2 amnh). VENE-
ZUELA. Bolivar: El Manaco, 59 km SE El Do-
rado, 150 m (12 usnm); Km 38, SE El Dorado,

100 m (1 ucv); Los Patos, 28 km SE El Manteco,
1 50 m (4 usnm); Rio Supamo, 50 km SE El Man-
teco, 150 m (1 usnm); Salto Chalimaha, Rio Pa-
ramichi, Rio Paragua (1 ucv); Salto Ichun, Rio
Paragua (2 ucv). T.F. Amazonas: Belen, Rio Cu-
nucunuma, 56 km NNW Esmeralda, 150 m (2
usnm); Boca Mavaca, 84 km SSE Esmeralda, 1 38
m (1 usnm); Caiio Leon, Cerro Duida, 325 m (1
amnh); Capibara, Brazo Casiquiare, 106 km SW
Esmeralda ( 1 usnm); Esmeralda, Cerro Duida, 325
m (3 amnh); 14 to 65 km S, SSE, and SSW Pto.
Ayacucho, 119-161 m (16 usnm); Rio Mavaca,
1 08 km SSE Esmeralda, 1 40 m (7 usnm); San Juan,
Rio Manapiare, 163 km ESE Pto. Ayacucho, 155
m (6 usnm); Tamatama, Rio Orinoco, 135 m (7
usnm).

Artibeus phaeot is— HoXoXyTpcs o{A. phaeotis and
A. ravus, plus hundreds of other specimens from
Mexico, Central America, and NW South Amer-
ica.

Artibeus pumilio— PERU. Loreto: Masisea,
Tushemo, Rio Ucayali, 328 m (1 bm, holotype of
A. pumilio).

Artibeus watsoni— PANAMA. Chiriqui: Boga-
va, 250 m (5 bm, including holotype of A. watsoni);
Progreso (34 usnm); Puerto Armuelles (2 usnm).

Remarks— In previous publications of SVP,
Artibeus gnomus has been known as '''' Artibeus sp.
A".

Acknowledgments

Among the persons who helped me put together
this paper I am especially grateful to Sally DeMott,
who measured the SVP skulls; Linda Gordon, who
compiled the tables and worked with me on the
comparisons; and Jane Ailes Small, who read and
criticized the manuscript and did the word pro-
cessing. Curators of several collections kindly per-
mitted me to study specimens under their care in
the preparation of these descriptions: American
Museum of Natural History (AMNH), British
Museum (Natural History) (BM), Field Museum
of Natural History (FMNH), Museum of Com-
parative Zoology, Harvard University (MCZ), and
Universidad Central de Venezuela (UCV). The
SVP collection is in United States National Mu-
seum of Natural History (USNM); a portion of its
specimens have been returned to Venezuela.

HANDLEY: NEW SPECIES OF ARTIBEUS

171

Literature Cited

EwEL, J. J., AND A. Madriz. 1968. Zonas de Vida de
Venezuela. Ministerio de Agricultura y Cria, Caracas,
265 pp., map.

Handley, C. O., Jr. 1959. A revision of American
bats of the genera Euderma and Plecotus. Proceedings
of the United States National Museum, 110: 95-246.

. 1976. Mammals of the Smithsonian Venezue-
lan Project. Brigham Young University Science Bul-
letin, Biological Series, 20(5): 1-91.

. 1984. Newspeciesof mammals from northern

South America: A long-tongued bat, genus Anoura
Gray. Proceedings of the Biological Society of Wash-
ington. 97: 513-521.

Handley, C. O., Jr., and K. C. Ferris. 1972. De-
scriptions of new bats of the genus Vampyrops. Pro-

ceedings of the Biological Society of Washington, 84:
519-523.

Handley, C. O., Jr., and L. K. Gordon. 1980. New
species of mammals from northern South America.
Mouse possums, genus Marmosa Gray, pp. 65-72. In
Eisenberg, J. F., ed.. Vertebrate Ecology in the North-
em Neotropics. Smithsonian Institution Press, Wash-
ington, D.C.

HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds.
1982. Mammal Species of the World. Allen Press,
Inc., and the Association of Systematic Collections,
Lawrence, Kansas, 694 pp.

Koopman, K. F. 1978. Zoogeography of Peruvian bats
with special emphasis on the role of the Andes. Amer-
ican Museum Novitates, 2651: 1-33.

Miller, G. S., Jr. 1902. Twenty new American bats.
Proceedings of the Academy of Natural Sciences of
Philadelphia, 54: 389-412.

172

HELDIANA: ZOOLOGY

Seasonality of Reproduction
in Peruvian Bats

Gary L. Graham

ABSTRACTS

The reproductive conditions of 3,489 specimens were used to determine seasonal patterns
of pregnancy and parturition in Peruvian bats. More species that are trophic generalists yield
birth records for the dry season than do trophic specialists. Relatively more highland than
lowland species have births recorded for both seasons (dry and wet). Presumably, trophic
generalists and highland species experience less seasonal variation in food supplies compared
to the other groups. A larger percentage of nectarivorous species than frugivores have birth
records for the dry season; the reverse is true for the wet season. These patterns are associated
with greater floral resource abundance during the dry season and a greater abundance of fruit
resources during the wet season.

Las condiciones reproductivas de 3489 especimenes fueron usadas para describir patrones
estacionales de embarazo y alumbramiento en murcielagos peruanos. Mas especies de gener-
alistas alimenticios tienen registros de nacimiento durante los dos estaciones (seca y mojada)
que tienen especialistas. Relativamente mas especies desde tierras altas que desde tierras bajas
tienen registros de nacimiento durante la estacion seca. Possiblamente, generalistas alimenticios
y especies de tierras altas sufrir menos variacion estacional de provisiones alimentos que los
otros grupos. Un mayor porcentaje de especies nectivoras que frugivoras tienen registros de
nacimiento durante la estacion seca pero, el opuesto exista para la estacion mojada. Estos
patrones son asociados con un mayor abundancia de recursos de flores durante la estacion seca
y con un mayor abundancia de recursos de frutas durante la estacion mojada.

As condi96es reprodutivas de 3489 especimes de morcegos p)eruanos foram usadas para
descrever seus padroes de parturi^ao. Os morcegos de habitos alimentares generalizados pos-
suem uma propor9ao maior de especies que possuem registros de partos durante a epoca da
seca, do que os morcegos com habitos alimentares especializados. Proporcionalmente mais
especis de morcegos de areas elevadas, do que das planicies, possuem registros de partos durante
as duas epocas (seca e chuvosa). Morcegos generalizados e os de areas elevadas provavelmente
experimentam menos varia^ao epocal em a quantidade da comida, do que os morcegos das
planicies ou morcegos com habitos alimentares especializados. Uma porcentagem maior das
especies nectarivoras, do que das especies frugivoras, possuem registros de partos durante a
epoca da seca, sao a regra ao inves da exce9ao entre os morcegos que consumen fruta. Estes
padroes podem ser interpretados como adpata96es de individuos a recursos alimentares que
sao regularmente, ou ocasionalmente, obteniveis durante a epoca da seca.

From the Department of Biology, University of New
Mexico, Albuquerque, New Mexico 87131.

GRAHAM: PERUVIAN BATS 173

Introduction

Although the timing of reproductive events is
an important Hfe history adaptation, there are few
studies of the reproductive phenology of South
American bats (Racey, 1982). Most of these stud-
ies are of individual species and cover only part
of the year. Gestation, parturition, lactation, and
weaning should be timed to correspond with vari-
ations in the abundance and diversity of food sup-
plies (Fleming et al., 1972; Bradbury & Vehren-
camp, 1977; Bonaccorso, 1979; Wilson, 1979;
August & Baker, 1 982). These variations are known
to be seasonal in most of the Neotropics (Janzen,
1967, 1973; Foster, 1982; Wolda, 1982; Smythe,
1982; Terborgh, 1983). Which reproductive event
is actually synchronized with peak food supplies
is likely to be determined by the relative cost of
each event. Lactation is the most costly period for
most females, but weaning and dispersal pose the
greatest survival problem for young bats (Wilson,
1979; Racey, 1 982). These authors, and Tuttle and
Stevenson (1982), agree that the weaning of young
is the most critical period for individuals of most
bat populations.

Most species of Neotropical bats that have been
studied are polyestrous (Fleming et al., 1972
Thomas, 1972; Bradbury & Vehrencamp, 1977
Myers, 1977; Wilson, 1979; Bonaccorso, 1979
Humphrey & Bonaccorso, 1979). Each year, fe-
male bats confront the problem of timing two
(rarely three in a few vespertilionids) periods of
lactation and weaning with variations in food
availability. Individuals of most species handle
this problem by producing their young so that the
first is weaned at the beginning of the wet season
as fruit supplies are reaching their peak, and the
second later in the wet season when fruit avail-
ability reaches a second peak or remains relatively
high (Wilson, 1979; Tuttle «& Stevenson, 1982).
Exceptions to this general pattern of seasonal poly-
estry have been observed in most studies (Fleming
et al., 1972; Thomas, 1972; Myers, 1977; Myers
& Wetzel, 1983; Bonaccorso, 1979; see Mares &
Wilson, 1 97 1 ; Bradbury & Vehrencamp, 1 977; and
August & Baker, 1982 for good discussions of dry
season birth periods).

The purpose of this study is to examine some
of the details of dry season reproduction in Pe-
ruvian bats. I demonstrate that the different feed-
ing assemblages and the faunas of different zoo-
geographical regions differ in the proportion of
species with birth records during the dry season.

I also offer suggestions as to how dry season births
may be adaptive.

Methods

The bat specimens from Peru housed in the col-
lections of the American Museum of Natural His-
tory, Field Museum of Natural History, Louisiana
State University Museum of Zoology, National
Museum of Natural History, and the Texas Co-
operative Wildlife Collection were examined in
late 1977 and early 1978 for information on re-
production. Data were taken directly from Tuttle
(1970) and Bowles et al. (1979) for specimens not
in the above collections but included in their re-
ports. Information on reproductive condition of
bats was also obtained from the notes of A. L.
Gardner and J. L. Patton. Fieldwork at several
localities in Peru, conducted from June through
August 1977 and from June through November
1978, enabled me to collect and record the repro-
ductive data for many of the bats now in the Lou-
isiana State University collection. Diet informa-
tion was taken from the literature (Heithaus et al.,
1975; Paradiso, 1975; Gardner, 1977) and is given
in Graham ( 1 983). Nomenclature follows Graham
(1983) except for Artibeus glaucus, which I now
consider a distinct species.

Recorded data consisted of species identifica-
tion, locality, elevation, sex, age, and macroscopic
reproductive condition. Individuals with incom-
pletely ossified phalangeal epiphyses were classi-
fied as juveniles. Females were designated as preg-
nant or lactating if this information was included
on specimen tags or indicated by dissection of fluid
preserved females. The crown-rump lengths of
embryos (including extra-embryonic membranes)
were either measured (in mm) or taken from spec-
imen tags. Length of testes was also noted when
it was recorded on sp>ecimen tags, but was not used
to establish reproductive patterns because of un-
certainties in the relationship between testes size
and sexual activity (Taddei, 1976; Thomas, 1972).
Monthly pregnancy frequencies are based only on
those female SF>ecimens for which I am confident
that the presence (or absence) of embryos had been
properly recorded. This limitation excludes most
of the specimens collected prior to 1960.

I classified each species with adequate data (those
with evidence of at least one birth period) into
those that can give birth in the dry (May-Septem-
ber) and/or wet (October-April) seasons. I as-

174

FIELDIANA: ZOOLOGY

sumed that a birth between 1 May and 3 1 August
also indicates weaning of the young during the dry
season. Species with births recorded only for the
month of September (usually a dry month) were
not included in either breeding season, since these
young could be weaned either as the dry season
ended or as the wet season began. The above clas-
sifications were accomplished by determining the
distribution of births across all months, as indi-
cated by the distribution of juveniles, lactating fe-
males, and embryo sizes. If a single birth was in-
dicated for a given season, then that species was
identified as having the potential to produce young
during that season. The presence of juveniles or
lactating females was assumed to indicate partu-
rition during the month prior to capture. If a fe-
male was carrying a large, near-term embryo, par-
turition was assumed to take place in that month.
On the other hand, if the embryo was very small
relative to the adult body length, birth was as-
sumed to take place four months later for most
species, three months later for small insectivorous
species (Findley & Wilson, 1 974), and five months
later for emballonurids (Bradbury & Vehrencamp,
1977) and Desmodus rotundus (Wilson, 1979).

Classification of a species as one that can give
birth during the dry season does not necessarily
mean that it does so each year. I was unable with
this method to determine the actual frequency of
births during either season. A dry season classi-
fication simply means that individuals can at least
occasionally produce young during the dry season.

The lack of birth records for a given species for
either season can be the result of an actual absence
of birthing during that season, or it may be due to
an inadequate sample from that season. If I as-
sume that the lack of a breeding record for a given
season is not due to inadequate sampling, then I
can use the G test (Sokal & Rohlf, 1981) to assess
the significance of the differences between the
groups of species compared. This is a safe as-
sumption, since most of the specimens were col-
lected during the Peruvian dry season (the season
with which this study is principally concerned) and
because the groups of bats predicted to have the
greatest proportion of dry season births are the
groups most poorly sampled (see Appendix and
below).

Regions and Climate

I follow Koopman's ( 1978) division of Peru into
three zoogeographical regions. He lists SF)ecies as

lowland if they were collected below 1000 m east
of the Andes and highland if collected above this
elevation. Pacific coastal species are those col-
lected either along the arid coast or adjacent An-
dean foothills (including the mesic areas of north-
western Peru).

The Pacific coastal plain is characterized by low
(< 50 mm) annual rainfall, most of which falls
between December and April. The mesic areas on
the northwestern Andean slopes experience great-
er rainfall over an extended period. Rainfall east
of the Andes (fig. 1 ) is seasonal and abundant, with
the greatest amounts deposited between the ele-
vations of 1 000 and 3000 m (Bowman, 1916). The
wet season begins in October and continues through
April and the dry season begins in May and ex-
tends through September.

Most investigators agree that food resources are
primarily modulated by rainfall seasonality (Jan-
zen, 1967, 1973; Frankie et al., 1974; Ricklefs,
1975; Buskirk & Buskirk, 1976; Wolda, 1978a,b,
1982; Foster, 1982; Smythe, 1982; Terborgh,
1983). Many of the following generalizations on
seasonal changes in food supplies for bats of mid-
dle and higher elevations are based on the as-
sumption that Peruvian plants and insects respond
to environmental conditions in a manner similar
to closely related organisms in Costa Rica, Pan-
ama, or Puerto Rico (as described by the above
authors). Terborgh's (1983) study of the changes
in fruit, flower, and insect supplies in Manu Na-
tional Park indicates that in the lowlands of south-
eastern Peru, fruit abundance and diversity usually
increase in October with the onset of the wet sea-
son, peak in November and December and again
in March, and decrease to the lowest level in May
and June. Not only is the first peak greater but it
also includes a greater proportion of plants in the
family Moraceae (Terborgh, pers. comm.), which
have fruit favored by bats (Gardner, 1 977). Flower
abundance peaks in the dry season, but some flow-
ers are present throughout the year (Terborgh,
1983). At middle to higher elevations, the phe-
nology of fruit and flower species is probably sim-
ilar to lowland species, but less seasonal (Ter-
borgh, 1977; Nevling, 1971). Changes in insect
resources are more difficult to generalize because
the species in different size classes (Smythe, 1982;
Wolda, 1982) and in different habitats (Bradbury
& Vehrencamp, 1976; Terborgh, 1983; Janzen,
1983) have different periods of peak abundance
and diversity. If the supply across size classes and
habitats is considered, then it is probably rather
high throughout the year. I will not discuss the

GRAHAM: PERUVIAN BATS

175

600.

500.

400.

5 300.

I 200_

100.

Month

Fig. 1. Average rainfall amounts at
Iquitos (lowland) and Yurac (middle el-
evation), Department of Loreto, Peru.
Data are taken from a map published in
1971 by the Servicio Nacional de Me-
teorologia e Hidrologia in Peru and are
averaged from 1 years of records.

seasonal changes in food levels west of the Andes
in Peru because less is known of this region.

Results

More than 3,400 female specimens divided
among 109 species were included in this study.
Twins were recorded for only one female, a Car-
ollia perspicillata, collected in November bearing
two well-developed embryos (24 and 29 mm).
Specimens were collected almost exclusively be-
tween May and December, with June through Au-
gust having the largest samples (see sample sizes
in Appendix). The reproductive records are also
unevenly distributed among the species (Appen-
dix). These sampling problems made it difficult to
identify the reproductive patterns for most species.
If, however, the percentages of all the females that
were pregnant are determined for each species for
each month, many species have data that fit the
pattern of seasonal polyestry. This pattern is well
illustrated by the pregnancy curve ofCarollia per-
spicillata (fig. 2). Pregnancy levels peak in Septem-
ber and December and are followed by periods of

parturition, as suggested by the October drop in
the frequency of pregnant females and by the ju-
veniles collected in January (Appendix). The be-
ginning of the second reproductive cycle of the
season is also indicated by the females of Carollia
castanea (l), Vampyrops dorsalis (7), and Artibeus
planirostris (1) that were simultaneously lactating
and pregnant in November and December (Ap-
pendix). On the other hand, the pregnancy curve
of Myotis (fig. 2) and the records of juveniles and
lactating females of Carollia (fig. 2) and Artibeus
(Appendix) for May-September indicate that births
can occur outside of the wet season. Individuals
of some species (i.e., Desmodus rotundus, Myotis
nigricans, and perhaps Glossophaga soricina and
Artibeus planirostris; see Appendix) may be able
to produce young during any month of the year.
Table 1 lists the number of species in each fam-
ily or subfamily with births recorded during the
seasons. I was able to identify birth periods for 79
(72.5%) of the 109 species listed in the Appendix.
Most species (63 of the 79. 79.7%) have birth rec-
ords for the wet season but a surprisingly large
proportion of all of the species (46 out of 79, 58.2%)
have records of dry season parturition. The ab-
sence of birth records during the wet season

176

HELDIANA: ZOOLOGY

Carollia perspiclllata

60-1 3 3 24 38 100 18 15 23 15

Fig. 2. Changes in pregnancy levels of Carollia perspiclllata and Myotis nigricans. The numbers across the top
refer to the sample sizes for each month. The ordinate represents the percentage of all females for each month that
were recorded as pregnant.

(Mormoopidae) and the dry season (Furipteridae
and Thyropteridae) may represent real periods of
no births but may also be the result of inadequate
sampling.

Are there groups of bats that have a greater ten-
dency than other groups toward parturition during
the dry season? If seasonal fluctuations in food
supplies are less in the highlands than in the low-
lands, then proportionately more highland than
lowland species could be expected to produce and
wean young during the dry season and, if most
species are polyestrous, during both seasons. Thir-
ty-one of the 58 lowland species (53.4%) and 17
of the 27 highland species (63.0%) have records
of dry season births (table 2). Although the 9.6%
difference is in the predicted direction, it is not
significant {P > 0.25). The highland region has a
greater proportion of species with birth records
from both seasons (51.9% vs. 34.5%, P < 0.05).
The highland Sturnira bogotensis that were lac-

tating and pregnant in June and August (Appen-
dix) provide evidence that some species with dry
season birth records are polyestrous, becoming
pregnant again during the dry season. Bats of the
Pacific coastal and slope region also show a strong
tendency toward dry season births.

If trophic generalists (those that consume more
than one type of food, such as fruit and insects)
can switch to another resource when one type be-
comes scarce, they should be less vulnerable to
seasonal fluctuations in their food supply than are
trophic specialists (those species that use only one
major type of food). This reduced seasonality of
food resources should be reflected by a tendency
for births to occur during the dry season and, if
polyestry is common, for both seasons. Nine of
the 1 1 generalist species (81.8%) and 23 of the 68
specialists (33.8%, table 2) have records of indi-
viduals that have given birth during the dry sea-
son, and relatively more generalists (63.6%) than

GRAHAM: PERUVIAN BATS

177

Table 1. Number of species of Peruvian bats with
births recorded for each season.

Family

Dry

Season*
Wet

Both

Total
spe-

ciest

Emballonuridae

Noctilionidae

Mormoopidae
Phyllostomidae
Phyllostominae
Glossophaginae
Carolliinae

6
6

6
4
5

3
3

Stumirinae

Stenodermatinae

Desmodontinae

Furipteridae
Thyropteridae
Vespertilionidae
Molossidae

7
5

1
1
7
5

Totals

* The number of species with records of birth in either
the dry or the wet season or in both. A species is listed
in all three columns if it has birth records for both sea-
sons.

t The total number of species that have data indicating
birth periods. The sum of the dry and wet season records
does not equal the total, because some species have rec-
ords for both seasons.

specialists (29.7%) have records for births during
both seasons (table 2). These large proportional
differences are significant (P < 0.01 and P < 0.05,
respectively).

Since floral resources are greatest in the dry sea-
son when fruit resources are lowest, nectarivorous
species should show a greater tendency for indi-
viduals to produce and wean young during the dry
season compared to frugivores. Six of the seven
(85.8%) bats that are at least partially nectarivo-
rous have birth periods in the dry season com-
pared with less than half (1 7 of 40, 42.5%) of those
that are partially frugivorous (table 2). This large
relative difference is significant (P < 0.05). Dry
season production and weaning of young is prob-
ably the common pattern for most bats that eat
nectar. This is indicated by the large number of
juveniles (13) and lactating females (35) recorded
for this group from May through August. Since
the resource abundances are reversed during the
wet season, the pattern of frugivorous sp>ecies hav-
ing relatively more birth records (39 of the 40
species that are partially frugivorous, 97.5%) than
nectarivores (3 of the 7 species that are partially
nectarivorous, 42.9%) was expected {P < 0.001).
Percentages of the species in these trophic groups
that have birth records for both seasons are similar

(40.0% and 42.9% for frugivores and nectarivores,
resp)ectively, P > 0.50).

The number of insectivorous species that have
birth records for each season is almost equal (17
for the dry, 1 8 for the wet). This is the expected
pattern if insect abundance and diversity remain
high throughout the year.

If a species is seasonally polyestrous, the prob-
ability of recording a birth for the wet season for
that species is greater, since both young are pro-
duced during that season. If most species are sea-
sonally polyestrous, then the majority of species
should have wet season birth records. All of the
ecological groups mentioned above (except the
nectarivores) show considerable evidence of par-
turition during the wet season (table 2).

Discussion

Seasonal polyestry is the dominant pattern in
other studies of Neotropical bat reproduction
(Hemingetal., 1972; Thomas, 1972;Taddei, 1976;
LaVal & Fitch, 1977; Myers, 1977; Bonaccorso,
1979; Wilson, 1979; August & Baker, 1982). The
majority of bats in Peru, especially frugivores,
give birth during the wet season. Although this
pattern is expected if most species are seasonally
polyestrous most of the time, this conclusion is
compromised by the possibility that some species
may be monestrous, producing and weaning their
young during the wet season. I was unable, for
most species, to distinguish between these two re-
productive patterns. However, the females col-
lected from November and December that were
simultaneously lactating and pregnant provide
evidence for seasonal polyestry. Females that fol-
low this pattern wean their first young concur-
rently with the onset of the heavy rains, as fruit
supplies are reaching their peak, and wean their
second young near the middle of the wet season
when food resources remain plentiful.

In Peru, as in many of the other Neotropical
areas discussed in the above studies, young bats
are at least occasionally bom and weaned in the
dry season. These bats are members of zoogeo-
graphical and ecological groups whose tendencies
toward dry season births can be predicted. If food
availability for a particular group of bats does not
fluctuate greatly throughout the year relative to a
second group, then the first group should include
a larger percentage of species with birth records
for the dry season and, if the species are polyes-

178

HELDIANA: ZOOLOGY

Table 2. Number of species of Peruvian bats with births recorded for each season, by geographical region and
ecological group.

Season*

Region

Dry

Wet

Both

Total

and group

% (N)

species*

Geographical Regions

Lowlands

53.4(31)

79.3 (46)

34.5(19)

Highlands

63.0(17)

88.8 (24)

51.9(14)

Pacific coastal

80.0 (8)

60.0 (6)

30.0 (3)

Trophic Groups

Insectivores

58.6(17)

62.1 (18)

22.2 (6)

Frugivores

38.7(12)

100 (33)

38.7(12)

Nectarivores

66.7 (2)

33.3(1)

...

Piscivores

100 (1)

...

Vampires

50.0(1)

100 (2)

50.0(1)

Frugivore/insectivores

60.0 (3)

80.0 (4)

40.0 (2)

Nectarivore/insectivores

100 (3)

66.7 (2)

Piscivores/insectivores

100 (1)

Omnivorest

100 (2)

Trophic Specialists:}:

33.8 (23)

79.4 (54)

29.7(19)

Trophic Generalists§

81.8(9)

90.9(10)

63.6 (7)

* See notes to Table 1 . N = Number of species; % = % of total species.
t Includes species that eat more than two major types of items,
t Includes insectivores through vampires (see text, p. 1 77).
§ Includes frugivores/insectivores through omnivores.

trous, for both seasons. Two groups of Peruvian
bats, highland species and trophic generalists, pos-
sibly experience less seasonal fluctuation in food
supplies relative to lowland species and trophic
specialists. However, the reasons for the damp-
ened fluctuations differ with each group.

At high elevations in Peru, even during the dry
season, clouds cover the forests most of the time
(Terborgh, 1977; pers. obs.), providing moisture
that is largely unavailable to lowland plants. This
yearlong availability of moisture presumably keeps
plant productivity and ultimately food availability
from fluctuating as greatly as in the lowlands. This
reduced fluctuation in food supplies is reflected by
a weak trend toward relatively more highland birth
records for the dry season and by a stronger high-
land trend for birth records for both seasons.

Species with generalized feeding habits are
thought to be able to switch resources when one
becomes reduced. This dampens the effect of sea-
sonal changes in food availability and explains
their larger percentages of birth records for the dry
season and for both seasons. Fleming et al. ( 1 972),
Heithaus et al. ( 1 975), and Bonaccorso ( 1 979) pro-
vide data from Costa Rica and Panama that il-
lustrate seasonal switches made by several of the
same species included in my study as feeding gen-
eralists (i.e., Phyllostomus discolor and Glossoph-
aga soricina).

Relatively more nectarivorous than frugivorous
species have individuals with dry season birth and
weaning records, but the records for both seasons
are not different. These were the expected patterns,
since both groups presumably experience seasonal
variations in their food supplies; floral resources
are greatest and fruit resources lowest during the
dry season (Terborgh, 1983). Hence, dry season
births may actually be the rule for bats that con-
sume nectar but the exception for frugivores. The
reverse is true for the wet season. These seasonality
differences explain the almost equal proportions
of species with birth records for both seasons.

Thus, for trophic generalists, highland, and nec-
tarivorous bat species, many females produce their
first young in the dry season and become pregnant
again, producing their second young during the
wet season. Obviously, many more data are need-
ed to clarify these patterns and to address the pos-
sibility of differential survival of young between
the two seasons.

In Peru, the onset and termination of the rainy
season can vary annually (Terborgh, 1983). Food
supplies track these variations (Terborgh, 1983;
Foster, 1982), and so may bat reproduction. I do
not believe that climatic variability is the principal
factor responsible for the births recorded in Peru
between May and August because the ecological
groups that I compared (frugivores and nectari-

GRAHAM: PERUVIAN BATS

179

vores, trophic specialists and generalists) showed
different seasonal birth patterns. If unusual dry
season rains were the principal factor, then equal
proportions of the different ecological groups
should have responded by producing young during
the normally dry season.

My data support the hypothesis that dry season
births are adaptations for producing and perhaps
weaning young when food resources are available
predictably or occasionally during this time. These
patterns need to be confirmed by long term studies
of actual bat communities, and of the resources
used by individuals at both highland and lowland
localities and during both seasons.

Acknowledgments

I am grateful to the many people who assisted
me in Peru and in the United States. Antonio Brack
E., Eric Cardich Briceno, Richard Bustamante M.,
and Susana Moller-H. of the Direccion General
Forestal y de Fauna, Ministerio de Agricultura,
Lima, provided the necessary collecting permits.
Hernando de Macedo R. and Ramon Ferreyra of
the Museo de Historia Natural "Javier Prado"
were also helpful. I thank Arturo Koenig R., Man-
uel A. Plenge, and Gustavo del Solar for their
hospitality and help in making this study possible.
I appreciate the assistance provided by Faucet and
Aero Peru. Without the reliable assistance of my
Peruvian associates, Manuel Sanchez, Klaus Wehr,
and Reyes Rivera, the fieldwork would have been
much more difficult.

Karl F. Koopman (American Museum of Nat-
ural History), Don E. Wilson (National Museum
of Natural History), David J. Schmidly and Wil-
liam B. Davis (Texas Cooperative Wildlife Col-
lection), and Patricia W. Freeman (formerly of
Field Museum of Natural History) were very help-
ful during trips to their museums. Koopman and
James L. Patton (Museum of Vertebrate Zoology)
loaned fluid-preserved bats. James B. Cope (Jo-
seph Moore Museum), Alfred L. Gardner (Na-
tional Museum of Natural History), and Patton
were kind enough to send me unpublished manu-
scripts and field notes. For valuable assistance in
the field I thank Linda J. Barkley, J. William Eley,
Gary R. Graves, John P. O'Neill, Theodore A.
Parker, III, Thomas S. Schulenberg, and Morris
D. Williams. The excellent suggestions on the
manuscript by Philip Myers were appreciated. I

thank Susan T. Graham for her support during the
several revisions of this work.

1 gratefully acknowledge the financial support
of the LSUMZ Peruvian fieldwork by John S.
Mcllhenny, Irving and Laura R. Schweppe, E. W.
Mudge, and the late Babette M. Odom. Travel to
other museums was made possible in part by an
LSU Foundation-Graduate Student Travel Award.

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GRAHAM: PERUVIAN BATS

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186

FIELDIANA: ZOOLOGY

Tent Construction by Bats

of the Genera Artibeus and Uroderma

Robert M . Timm

ABSTRACTS

Herein, I describe new styles of tents cut and utilized by Artibeus anderseni, A. glaucus, A.
gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, and U. magnirostrum; review
and summarize the literature on tent use by Artibeus and Uroderma; and discuss the effectiveness
of tents as diurnal roosts. Artibeus anderseni alters the shape of Heliconia leaves by cutting the
lateral nerves and interconnected tissue extending out from the midrib. Artibeus glaucus cuts
the basal lateral nerves in Xanthosoma, causing the two sides of the leaf to collapse downward
around the midrib. Artibeus phaeotis cuts the lateral nerves and interconnected tissues in both
banana and Heliconia imbricata; the basal cuts veer out from the midrib such that a distinctive
V-shaped enclosure is formed by the hanging leaf Artibeus toltecus cuts the basal nerves on
Anthurium, causing the sides of the leaf to fold down around the midrib to form a pyramid-
shaped tent. Artibeus watsoni was found to make four distinctive styles of tents, including simple
V-shaped cuts on bifurcated palms, cuts of a few side veins on aroids to produce a rounded
pyramid, elongate J-shaped cuts on banana and Heliconia, and polygonal cuts on Carludovica
pal mat a. Artibeus watsoni has the greatest repertoire in tent styles, and uses the most diverse
array of plant species and leaf shapes. Two styles of tents constructed by Uroderma bilobatum
are reported for the first time, one on the large pinnately leafed palm Scheelea rostrata and the
second on banana. The common denominator between the Uroderma bilobatum tents reported
herein and those previously described is that all are on large, broad leaves and all have a
distinctive V-shaped pattern cut by the bats. Uroderma magnirostrum also creates an inverted
elongate V-shaped tent on pinnately leafed palms.

All New World tent-makers described to date are tropical members of the phyllostomid
subfamily Stenoderminae. Each species of tent-making bat has one or more distinctive style of
tent. Bats select leaves of specific shapes, sizes, and angles for tent construction. Most species
appear to be obligate tent-roosters. Tents provide bats with a cryptic diurnal roost site, in
addition to providing shelter from both the sun and rain and an early warning to the approach
of predators.

Aqui yo describo nuevos estilos de carpas cortadas y utilizadas por Artibeus anderseni, A.
glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, y U. magnirostrum
reviso y hago un sumario de la literatura acerca del uso de carpas por filostomidos; y discuto
la efectividad de las carpas como perchas diumas. Artibeus anderseni altera la forma de las
hojas de Heliconia cortando las nervaduras centrales y tejido interconectado que se extiende

From the Division of Mammals, Field Museum of
Natural History, Chicago, Illinois 60605-2496. The au-
thor's present address is Museum of Natural History and
Department of Systematics and Ecology, University of
Kansas, Lawrence, Kansas 66045.

TIMM: ARTIBEUS AND URODERMA 187

exteriormente desde la nervadura central. Artibeus glaucus corta la base de las nervaduras
laterales en Xanthosoma, causando asi que los dos lados de la hoja colapsen hacia abajo al
rededor de la nervadura central. Artibeus phaeotis corta las nervaduras laterales y tejidos in-
terconectados en banana y Heliconia imbricata; los cortes basales viran hacia afuera desde la
nervadura central de tal modo que una distintiva cavidad en forma de V es formada por la
hoja colgante. Artibeus toltecus corta las nervaduras basales de Anthurium, adi que los lados
de la hoja doblen en redor de la nervadura central de tal modo que una cavidad es formada
en forma de una piramide. Artibeus watsoni fue encontrada haciendo cuatro distintos estilos
de carpas, incluyendo simples cortes en forma de V en palmas bifurcadas, cortes en unas pocas
nervaduras laterales en araceas para producir una piramide redondeada, cortes alargados en
forma de J en banana y Heliconia. y cortes poligonales en Carludovica pal mat a. Artibeus watsoni
tiene el mas grande repertorio en estilos de carpas y usa la mas diversa serie de plantas y formas
de hojas. Dos estilos de carpas construidas por Uroderma bilobatum son reportadas por primera
vez; una en la larga y pinnada hoja de palma Scheelea rostrata y la segunda en banana. El
comun denominador entre carpas de Uroderma bilobatum repnirtadas aqui y aquellas previa-
mente descritas es que todas usan hojas grandes y anchas y todas tienen un distintivo patron
en forma de V cortado por los murcielagos. Uroderma magnirostrum tambien corta una carpa
en forma de V-invertido en hojas pinnadas de palmas.

Todos los filostomidos cortadores y utilizadores de carpas descritos del Nuevo Mundo son
membros tropicales de la subfamilia Stenoderminae. Cada especie tiene uno o mas estilos
distintos de carpas. Los murcielagos escogen hojas de formas especificas, y constroen sus carpas
en angulos peculiares. La mayor parte de las especies parecen utilizar las carpas obligatoriamente.
Las carpas oferecen una percha oculta durante el dia, asi como un abrigo del sol, de la lluvia,
y de predadores.

Neste trabalho, (1) descrevo novos estilos de tendas cortadas e utilizadas por Artibeus an-
derseni, A. glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, e U.
magnirostrum; (2) reviso e resumo a literatura sobre o uso de tendas pelos morcegos da familia
Phyllostomidae, e (3) discuto a eficiencia de tendas como alojamentos diumos. Artibeus an-
derseni altera a forma das folhas de Heliconia, cortando as veias laterais e os tecidos interligados
que estendem da veia central. Artibeus glaucus corta as veias basilares laterais em Xanthosoma,
causando com que os dois lados da folha caiam contra o centro. Artibeus phaeotis corta as veias
laterais e os tecidos interligados nas folhas de bananas e de Heliconia imbricata. Os cortes
basilares partem da veia central, formando um abrigo distinto em forma de "V". Artibeus
toltecus corta as veias basilares de Anthurium, causando com que os lados da folha dobrem em
volta da veia central, criando um abrigo em forma de piramide. Artibeus watsoni constroi
quatro tipos diferentes de tendas, incluindo simples cortes em forma de "V" em folhas de
palmeiras bifurcadas; cortes em algumas das veias laterais em folhas de trepadeiras (resultando
em piramides redondas); cortes alongados, em forma de "J", em folhas de bananas e de
Heliconia; e cortes poligonos em fdlhas de Carludovica palmata. Artibeus watsoni possue o
maior repertorio de estilos de tendas, e usa o conjunto mais di verso de especies de plantas e
de configura96es de folhas. Dois estilos de tendas construidas por Uroderma bilobatum sao
descritos pela primeira vez; um nas fdlhas grandes da palmeira Scheelea rostrata, e outro nas
folhas de bananas. Fatores comuns entre as tendas construidas por Uroderma bilobatum aqui
descritas, e as descritas previamente, sao a forma distinta em "V" cortada pelos morcegos, e
o uso de folhas grandes e largas para a construfao das tendas. Uroderma magnirostrum tambem
constroe tendas em forma de "V" invertido nas folhas de palmeiras.

Todos morcegos construidores de tendas no Novo Mundo p)ertencem a subfamilia Steno-
derminae (familia Phyllostomidae), e cada especie exibe um ou mais estilos caracteristicos de
constru9ao. A maioria destas especies de morcegos parecem alojar-se obrigatoriamente em
tendas, as quais oferecem nao so um abrigo diumo camuflado, mas tambem prote9ao contra
sol, chuva, e predadores.

188 HELDIANA: ZOOLOGY

Introduction

The use of cut leaves for diurnal roosting sites
by bats was first described by Thomas Barbour
(1932), who reported on diurnal roosts of Uro-
derma bilobatum near the Panama Canal. He found
these bats roosting under the leaves of two culti-
vated palms identified as Livistona chinensis and
Prichardia pacifica. Not only were these bats found
roosting under palm fronds, but they had also al-
tered the leaf to produce a diurnal roosting struc-
ture. Barbour (1932, p. 307) stated that "by nip-
ping the ridges of the plications on the under side
the leaf is weakened and as the bitten spots are
skillfully and serially distributed the leaf finally is
sufficiently weakened so that the distal portion
droops sharply downward." Chapman (1932, p.
555) discovered Artibeus watsoni roosting under
the cut veins of a bifurcated palm, Geonoma cu-
neata (reported as G. decurrens), on Barro Colo-
rado Island, Panama, and first called these mod-
ified leaves "tents." Ingles (1953) also reported A.
watsoni constructing tents in the palm Geonoma
oxycarpa (reported as G. binervia) on Barro Col-
orado Island. Goodwin and Greenhall (1961, p.
262) found Artibeus cinereus roosting "under cut
leaves of palm trees and on the under side of ba-
nana leaves" and Uroderma bilobatum roosting
under cut leaves of the carat palm, Sabal glau-
cescens, and coconut palm, Cocos nucifera, on
Trinidad. Ectophylla alba was reported by Timm
and Mortimer (1976) to alter the leaves of five
species of Heliconia in Costa Rica; the bats se-
lected specific leaves for both size and angle to the
ground. Artibeus jamaicensis was found by Foster
and Timm (1976) roosting under the cut leaflets
on a pinnately leafed palm, Scheelea rostrata, in
a tropical dry forest in Costa Rica. Recently, Timm
( 1 984) reported tent construction by another phyl-
lostomid, Vampyressa pusilla, in Costa Rica, and
Koepcke (1984) found Mesophylla macconnelli
utilizing similar tents in Peru. Only one Old World
bat, Cynopterus sphinx (Pteropodidae), has been
reported to alter the shape of palm leaves to pro-
duce a diurnal roosting structure (Goodwin, 1 979).
Reviews of roosting site selection by bats were
provided by Tuttle (1976) and Kunz (1982).

To date, seven species of phyllostomids {Arti-
beus cinereus, A. jamaicensis, A. watsoni, Ecto-
phylla alba, Mesophylla macconnelli, Uroderma
bilobatum, and Vampyressa pusilla) have been re-
ported to modify leaves of plants to produce diur-
nal roosting sites herein referred to as tents. All

are tropical members of the phyllostomid subfam-
ily Stenodermatinae.

The phyllostomid genus Artibeus, which in-
cludes some 1 5 species, is widespread in the Noo-
tropics from northern Mexico southward to Ar-
gentina and Chile. These bats range in size from
10 g (A. anderseni and A. watsoni) to 70 g (A.
lituratus). Uroderma, a closely related genus of
medium-sized stenodermatines that includes only
two species, is found from southern Mexico through
the Amazon Basin of South America. The better
known of the two species, U bilobatum, weighs
from 13 to 21 g.

Herein I describe tent construction and utili-
zation by Artibeus anderseni, A. glaucus, A. gno-
mus, A. phaeotis, A. toltecus, A. watsoni, Uroderma
bilobatum, and U. magnirostrum; describe several
new styles of tents; review and summarize the lit-
erature on tent use by Artibeus and Uroderma;
discuss the effectiveness of tents as diurnal roosts;
and suggest directions for future research.

Methods

Descriptions of Study Areas

Costa Rica— Bosque Brrancia, also known lo-
cally as Bosque Blanco, is located 0.8 km west of
Cuarto Cruces on the south side of the Pan Amer-
ican Highway (Route 1) in Guanacaste Province,
in the Pacific lowlands of western Costa Rica. Bos-
que Brrancia is a restricted area of undisturbed
lowland forest classified as Tropical Moist Forest;
the dominant vegetation includes A nacardium ex-
celsum and Scheelea rostrata. This stand of forest
probably has not been logged previously, and rep-
resents a close approximation to the original (pre-
Columbian) forests of this part of Guanacaste.
Further descriptions of this forest can be found in
Janzen (1971) and Wilson and Janzen (1972).

Parque Nacional de Corcovado is located on
the Osa Peninsula of southwestern Costa Rica,
Puntarenas Province (between 08°27'N and
08°39'N, and 83°25'W and 83°45'W); the eleva-
tion ranges from sea level to 400 m. Corcovado
lies within the Tropical Wet Forest Life 21one
(Holdridge, 1967), with lowland evergreen forest
being the dominant forest type. Mean annual rain-
fall is 3,800 + mm and the wettest months are from
August through November; mean monthly tem-
peratures range from 25.0°C to 26.5°C. Vegetation
and habitat types at Corcovado have been de-

TIMM: ARTIBEUS AND URODERMA

189

scribed by Herwitz (1981) and Hartshorn (1983).
Areas surveyed included both primary forest and
secondary scrub along the coast.

The La Selva Biological Station is the field sta-
tion of the Organization for Tropical Studies lo-
cated 1 km SW of Puerto Viejo de Sarapiqui,
Heredia Province in the Caribbean lowlands of
northeastern Costa Rica (10°27'N,84°00'W); ele-
vation ranges from 29 to 100 m. Mean annual
rainfall is 3,990 mm, with the wettest months being
November, December, and February; mean
monthly temperatures range from 24.5°C (Decem-
ber) to 26.1°C (April). La Selva lies within the
Tropical Wet Forest Life Zone, with lowland ev-
ergreen forest being the dominant forest type.
Vegetation and habitat types of La Selva have been
described by Slud (1960), Holdridge et al. (1971),
Sawyer and Lindsey (1971), and Hartshorn ( 1 983).
One unusual feature of the subcanopy of the La
Selva forest is the diversity and abundance of dwarf
palms (Hartshorn, 1983), especially the broad-
leaved species, Geonoma congesta and Asterogyne
martiana. These species are regularly utilized by
Artibeus watsoni for tent construction. The tran-
sect surveys, which included all Artibeus tents ob-
served, were restricted to primary forest. The Uro-
derma tents described from this site were restricted
to an open banana patch.

Palo Verde (Refugio Nacional de Fauna Sil-
vestre Dr. Rafael Lucas Rodriguez Caballero) is a
wildlife refuge located 2 km S and 12 km E of
Bolson, in the Pacific lowlands of Guanacaste
Province, northwestern Costa Rica (10''30'N,
85°20'W); the elevational range is from 3 to 183
m. Palo Verde lies within the Tropical Dry Forest
Life Zone, with lowland deciduous forest and riv-
erine swamp forest being the dominant forest types.
The vegetation and habitat types of Palo Verde
have been described by Slud ( 1 980) and Hartshorn
(1983). Mean annual rainfall is 1,700+ mm, with
the wettest months being April, May, September,
and October; mean monthly temperatures range
from 26.0°C (November) to 29.7°C (April). The
immediate vicinity of the survey at this site was
in a mosaic of primary forest and secondary scrub
that included considerable bananas scattered
throughout.

The newly expanded Parque Nacional Braulio
Carrillo is located in northeastern Costa Rica (be-
tween 10°05'N and 10°25'N, and 83°54'W and
84'^5'W); the elevation of the park ranges from
100 to 2600 m. Braulio Carrillo is located on the
eastern, Caribbean slope of Volcan Barba in He-

redia Province. The elevational range at which Ar-
tibeus toltecus and associated tents were observed
was from 700 to 1400 m, within the Premontane
Rain Forest Zone with midelevational evergreen
forest and tall palms being the dominant forest
types. Mean annual rainfall at this elevational range
is perhaps as much as 5,000 mm, although no
exact measurements are yet available. The vege-
tation and habitat types at the midelevational
ranges also have yet to be described. On 14-15
April 1986, 3 km of trail ranging from 700 to 1 100
m were surveyed for bat tents. Additionally, an
intensive netting effort with Richard K. LaVal and
Don E. Wilson was conducted in this area over a
7-day period to determine what species of bats
were present and their relative abundance.

At Finca Las Cruces (2 km S of San Vito, Pun-
tarenas Province, 08°45'N, 82°58'W, 1200 m) in
the Premontane Wet Forest-Rain Forest transition
area, approximately 2 km of trail leading down to
the Rio Jaba was surveyed for tents on 1 3 August
1982.

Ecuador— Cascada San Rafael lies 17 km (by
road) west of the village of Reventador, Napo
Province, in northeastern Ecuador (00°5.8'S,
77°34.4'W). Rainfall averages 1,500 to 2,000 mm;
average temperatures range from 18°C to 22°C.
The elevation is 1200 m. Reventador lies within
the Humid Subtropical Life Zone.

Lagarto Cocha and Zancudo Cocha are military
encampments along the Rio Aguarico of Ama-
zonian Ecuador named for prominent lagoons.
Both areas are undisturbed primary lowland rain
forests classified as Moist Forest, with an annual
rainfall averaging from 2,000 to 3,000 mm; the
elevation is approximately 200 m.

Peru— Hacienda Amazonia lies just north of
the confluence of the Alto Rio Madre de Dios and
the Rio Pinipini in the department of Madre de
Dios, southeastern Peru (12°58'S, 71'>09'W), just
north of Atalaya. The Hacienda is located just east
of Parque Nacional del Manu in the Upper Trop-
ical Zone on the eastern foothills of the Andes. On
25 July 1985 Barbara L. Clauson searched the
ridge above the Hacienda for bat tents at an ele-
vation of 825 m in primary rain forest that had
received some selective timber harvest; she re-
turned to this site again on 3 November 1985.

Cerro de PantiacoUa lies above the Rio Palotoa,
10-15 km NNW of Shintuya, in the department
of Madre de Dios, southeastern Peru (12°35'N,
71''18'W). On 15 November 1985 Clauson
searched a steep sloping ridge at 600 m. The sur-

190

HELDIANA: ZOOLOGY

Table 1 . Individual measurements (in cm or degrees) collected from four tents constructed by Artibeus anderseni
on Heliconia in Ecuador.

Blade

Petiole

Basal
height

Length

Width

Angle

Length

Angle

Remarks

176
158
154
140

34
38
36

40
15
50

80
75
88
60

80
55
80
70

200
220
260
140

1 adult male A. anderseni
3 male A. anderseni

rounding forest was primary rain forest in the Up-
per Tropical Zone on the eastern Andean foothills.
Reference specimens of the bats have been de-
posited at Field Museum of Natural History, Chi-
cago; Escuela Politecnica Nacional, Quito, Ecua-
dor; and Universidad Nacional de Costa Rica and
Servicio de Parques Nacionales, San Jose, Costa
Rica. Voucher specimens of plants have been de-
posited in the herbaria at Field Museum, Duke
University, Missouri Botanical Garden, and/or
University of Wisconsin.

Accounts of Species

Artibeus

Artibeus anderseni Osgood, 1916

Artibeus anderseni occupies an extensive range
in western Amazonia; however, little is known of
its biology. This species has long been considered
a junior synonym of A. cinereus. In resurrecting
A. anderseni as a distinct species, Koopman (1978,
p. 14) stated, "Besides its shorter face and more
abrupt forehead, A. anderseni apparently always
lacks the last lower molars, which A. cinereus in
western Amazonia almost always has." I concur
with Koopman in recognizing A. anderseni as a
distinct species.

In late October and early November of 1983,
the trails and forest surrounding the military en-
campments were searched at Lagarto Cocha and
Zancudo Cocha in eastern Ecuador. Artibeus an-
derseni was found to alter the shape of leaves of
several small, forest Heliconia species to produce
diurnal roosting structures. To create a tent from
a Heliconia leaf, the bat severs the lateral nerves
and interconnecting veins that extend along both
sides of the midrib. The cuts ran along the central

90% of the leaf from near (0 to 14 cm) the base
to near (10 to 20 cm) the tip. Nerves and inter-
connected tissues were severed, but not complete-
ly, so that they did afford some support for the
sides of the leaf. Cuts ran parallel to the midrib
for most of its length, but did flare outward slightly
toward the base. The lateral nerves were cut from
3 to 8 mm from the midrib; the midrib was not
cut. Claw marks where the bats roosted started 50
cm from the base in one tent and ran for 1 6 cm
distally; in another they started at 70 cm and ran
for 10 cm distally. Measurements of the blade
length, blade width, blade angle, petiole length,
petiole angle, and basal height of four tents are
provided in Table 1 .

At Zancudo Cocha one Heliconia tent was un-
occupied for two days in succession, then on the
third day was occupied by an adult male with
enlarged testes, an adult lactating female, and a
juvenile male Artibeus anderseni (fig. 1). Another
Heliconia tent was unoccupied.

At Lagarto Cocha 13 tents were found in Hel-
iconia. One was occupied by three subadult males
not in breeding condition; a second tei t contained
a single adult male with enlarged testes.

Artibeus cinereus (Gervais, 1856)

Artibeus cinereus, Gervais's fruit-eating bat, is
found on the islands of Grenada, Trinidad, and
Tobago, and throughout the Amazon Basin and
adjacent coastal areas. Surprisingly little has been
published on roosting behavior or ecology of this
widely distributed species. On Trinidad, Goodwin
and Greenhall (1961, p. 262) stated, "It roosts in
small colonies of a few individuals under the cut
leaves of palm trees and on the under side of ba-
nana leaves." On Tobago, Husson (1954, p. 64)
reported a single male Artibeus cinereus "hanging
in a banana tree in cultivated country near the
shore."

TIMM: ARTIBEUS AND URODERMA

191

Fig. 1. Photograph of an adult male, adult female, and juvenile Artibeus anderseni roosting in a Heliconia leaf
tent. Details of the cut side nerves can be seen along the midrib of the leaf

Artibeus glaucus Thomas, 1 893

Artibeus glaucus is found at midelevalions along
the eastern slopes of the Andes from Venezuela to
Peru. The status of the name glaucus has long been
in a state of flux; it often has been considered a
subspecies of A. cinereus. I concur with Handley
(1987) in regarding .4. glaucus as a distinct species.

A single old adult male Artibeus glaucus (fmnh
124844) was observed roosting under a cut Xan-
thosoma leaf at Cascada San Rafael, Ecuador, on
21 September 1983. Four cut leaves on separate
plants were observed in close proximity to each
other; a fifth cut leaf was observed approximately
10 to 1 5 m to the south. Only the one cut leaf was
occupied by the single bat. An adult female A.
glaucus was netted in the vicinity that evening. All
tent leaves were cut down at the time and four
were measured. On 26 November, two more cut
leaves were found in this Xanthosoma population;
one contained three A. glaucus. The Xanthosoma
in which the bats were roosting were part of a

population of Xanthosoma that occupied approx-
imately 1 ha on a steep east-facing hillside.

The basal lateral nerves from 2 to 5 (usually 3)
of the Xanthosoma leaves were severed near the
midrib, causing the two sides of the leaf to collapse
downward around the midrib. The midrib was not
cut. Four of the five cut leaves were measured in
September; one found in November was measured
(table 2). The leaves that were selected by Artibeus
glaucus for tents all had the midrib running ap-
proximately parallel to the ground, whereas the
majority of unaltered leaves in the p>opulation stood
at more vertical angles.

Artibeus gnomus (Handley, 1987)

Artibeus gnomus, the dwarf fruit-eating bat, is
found in a peculiar circular range ringing the west-
em edge of the Amazon Basin. Although this
species has a wide distribution from Venezuela
and Guyana to Peru, it was only recently recog-

192

HELDIANA: ZOOLOGY

Table 2. Individual measurements (in cm or degrees) collected from five tents constructed by Artibeus glaucus
on Xanthosoma in Ecuador.

Blade

Length Width Angle

Petiole
length

Basal
height

Remarks

57 68 60 140 230 3 basal cut veins on each side, 2-3 cm from midrib

53 • • • 40 • • • 230 3 basal cut veins on each side, 1.5-3.0 cm from midrib

55 62 40 • •■ 230 2 veins cut left, 4 cut right side, 2-3 cm from midrib

64 36 45 137 300 3 basal veins cut on each side, 1.5-3.0 cm from midrib;

adult male A. glaucus hanging 16 cm from base
61 66 20 145 275 5 basal veins cut on each side, 3.5-4.0 cm from midrib; 3 ^.

glaucus

nized as a distinct species, and little biological in-
formation is available (Handley, 1987).

On 15 November 1985 Barbara L. Clauson
found a single adult male Artibeus gnomus roost-
ing under a cut Monstera lechleriana leaf. The
Monstera was growing as an epiphyte approxi-
mately 10 m off the ground on a tree on a sloping
hillside at 600 m elevation at Cerro de Pantiacolla,
southeastern Peru. The single cut leaf was green
and healthy and hung horizontally. No other cut
leaves were observed in the immediate vicinity.

The altered leaf was 70 cm long and 38.5 cm at
its widest point. All lateral nerves along the basal
nearly two-thirds of the leaf were severed imme-
diately adjacent to the thick midrib; this included
the basal 12 nerves on one side and 14 on the
other. The midrib was severed at 44.5 cm from
the base which caused the apical third of the leaf
to droop downward perpendicular to the midrib.
All nerves proximal to the midrib cut were sev-
ered. The lateral nerves along the apical, drooping
25.5 cm were unaltered.

The tent resulting from these cuts was quite en-
closed, being formed by the sides of the leaf col-
lapsing downward around the midrib and the dis-
tal third of the leaf folding down, perpendicularly
to the midrib. The lone Artibeus gnomus was
roosting 9 cm toward the base from the severed
midrib of the leaf.

Artibeus jamaicensis Leach, 1821

Artibeus jamaicensis, the Jamaican fruit-eating
bat, is found throughout much of tropical Central
America, the northern half of South America, and
the Greater and Lesser Antilles. At many localities
this is one of the most common species of bats
encountered and consequently has received more
study than any other phyllostomid.

Artibeus jamaicensis was reported roosting un-
der the cut leaflets of Scheelea rostrata in Costa
Rica by Foster and Timm (1976). Scheelea ros-
trata is a large, pinnately leafed palm with the
leaflets extending out at right angles from the hor-
izontal rachis. Leaflets within the middle 1.3 m
region of the frond were cut at varying distances
that increased going up to the center of the cut
area, then decreased. "As a result . . . the distal
parts of the leaflets folded perpendicularly, hung
vertically below the frond, and formed a broadly
lanceolate tent" (Foster & Timm, 1976, p. 266).

Although several Artibeus jamaicensis occupied
the roost, only two males were captured, one an
adult with enlarged testes, and the second a smaller
male not in breeding condition. Artibeus jamai-
censis apparently has a harem mating system, in
which a single breeding male defends a roost used
by several females and their offspring; nonbreed-
ing males may be found either singly or in small
groups (Morrison, 1979; Kunz et al., 1983). Ar-
tibeus jamaicensis has been found roosting in a
wide variety of situations, including caves, hollow
trees, buildings, and under unaltered leaves (Tut-
tle, 1976), and thus is certainly not an obligate
tent-roosting species, as apparently are the smaller
species oi Artibeus.

Artibeus phaeotis (Miller, 1 902)

Artibeus phaeotis, the pygmy fruit-eating bat, is
found from central Mexico to northern South
America (Timm, 1985). Most accounts of habitat
for pygmy fruit-eating bats mention their being
netted in close proximity to stands of bananas,
Musa X para^/5/aca (Ramirez-Pulidoetal., 1977;
Watkins et al., 1972). Davis (1970) suggested that
they might roost under the leaves of bananas.

During the summer of 1982, Artibeus phaeotis

TIMM: ARTIBEUS AND URODERMA

193

Fig. 2. Top, Dorsal view of a banana leaf {Midsa x paradisiacd) showing the cut nerves running aloQg the midrib
and flaring out toward the base; bottom, tent of Art ibeus phaeotis made from a banana leaf

was observed roosting under the leaves of banana
and Heliconia imbricata at Palo Verde and La
Selva, respectively, in Costa Rica. In all cases the
leaves had been altered to produce tents.

Art ibeus phaeotis constructs roosts in both ba-
nanas and Heliconia imbricata by biting the lateral
nerves and interconnecting tissue that extend at
right angles from the midrib, causing the blade to
fold over in a V-shaped enclosure. The two sides
of the leaf collapse downward, hanging beneath
the midrib (fig. 2). Nerves and interconnected tis-
sues are not completely severed, thus the sides of
the leaf provide some support for the entire length
of the cut. The cuts ran from the base of the leaf
to near the tip (table 3). Near the base, the cuts
flared out from the midrib to the sides to form an
elongate J-shaped pattern. The uncut tip and basal
portion of the leaf provide additional strength. The
undersides of roost leaves are obscured from view
from almost all angles except from directly be-
neath the tent.

To characterize Musa clumps, each of which

presumably represented an individual plant, the
number of stalks (ramets) per clump was counted;
the height of each ramet was estimated to the near-
est half meter. This was done for all clumps of
Musa in the patches that could be located, both
those with tents and without bat tents. When a bat
tent was located, the following measurements were
taken in centimeters: the angle of the petiole (mid-
way from stalk to blade), angle of blade (midway
on blade), petiole length, blade length, blade width,
height of base of blade, height of tip of blade,
height of roost, isolation distance (distance from
nearest solid object on the same vertical level),
length of uncut basal portion of blade, and length
of uncut distal portion. I also noted whether the
roosts were in direct sunlight or in shade (table 3).
Also for all tents the number of stalks on that
column was counted, and the age of the leaf rel-
ative to other expanded leaves was recorded. The
oldest (lowest leaf) on a plant was assigned the
number 1 , then the rest counted up from there.
I scored 232 individual stalks of Musa, which

194

HELDIANA: ZOOLOGY

Table 3. Individual measurements (in cm or degrees) collected from 19 tents constructed by Artibeus phaeotis
on leaves of banana, Musa x paradisiaca, in Costa Rica.

Petiole

Length

Blade
Angle

Width

Baiwl .
height

Height
Tip Roost

Isolation

Uncut

Length

Angle

Base

Tip

110

-27

196

156

182

136

-14

230

146

225

120

146

-40

213

170

228

125

1.5

120

-55

250

180

240

100

-16

205

200

120

-3

200

165

210

142

-36

235

160

230

115

+ 15

190

195

206

0,25

175

-12

320

260

320

>300

1.5

195

+25

360

400

> 300

0,35

130

+23

250

260

290

120

140

155

170

0,40

-22

205

175

200

100

130

180

220

230

147

-37

260

180

240

-12

205

215

220

+28

130

140

129

+ 15

250

220

260

120

+ 15

170

180

220

were in 4 1 clumps scattered in 1 00 m of second
growth forest. The mean number of ramets per
clump was 5.4; the range from 1 to 23. I located
26 leaves that were cut by bats, a ratio of roughly
one tent per 8. 1 ramets. Fourteen clumps had cut
leaves; if a clump had cut leaves it had a mean of
1.9 cut leaves (range 1 to 2). Of the 26 altered
leaves, complete data were taken on 19. The re-
maining tents were decomposing; in a few cases
the bats had not completed the tents. On the par-
tially completed tents, only one side of the midrib
was cut along one-quarter to one-third the distance
of the blade, but did not cause the side to collapse.
Only 2 of the 1 9 complete tents were located di-
rectly in the sun. Bat tents were found in clump
sizes ranging from 1 to 23 (mean = 6.2). The av-
erage height of all plants (N = 232) was 2.3 m.
The average height of plants with tents was 2.0 m
(N = 16).

One adult male with enlarged testes and six
pregnant females were watched over a 3 -day p)e-
riod in one of the roosts. Additionally, two solitary
nonpregnant female Artibeus phaeotis roosted un-
der separate leaves (fig. 3).

Artibeus phaeotis appears to select banana leaves
with specific characteristics. Usually these are the
oldest fully expanded leaves, just over 2 m above
the ground, with the center of the midrib nearly
horizontal to the ground and positioned far enough
from nearby stems and branches to limit access
by predators. Roost sites generally are located in

the shade of surrounding forest trees where ap-
parently they are protected by the forest overstory
from wind, blowing rain, and sunlight.

Although bananas are not native to the New
World, they are now common throughout the range
of Artibeus phaeotis and probably provide roost
sites in other localities. Heliconia, Calathea, and
broad-leafed palms are uncommon at Palo Verde,
hence are not readily available to A. phaeotis for
tent sites there. Artibeus phaeotis used only banana
leaves for tent making at Palo Verde, but con-
structed similar tents in Heliconia imbricata at La
Selva. Heliconia imbricata is the largest species of
Heliconia in Costa Rica, and its leaves are similar
in size and shape to banana. The tents in Heliconia
were similar in all respects to those in banana
leaves. An adult male and adult female were found
roosting together under a single Heliconia leaf in
late June.

Villa-R. ( 1 967) found a single specimen roosting
near the mouth of a small cave in Mexico.

Artibeus toltecus (Saussure, 1 860)

Artibeus toltecus, the lowland fruit-eating bat, is
found along the coasts of eastern and western Mex-
ico from Nuevo Leon and Sinaloa south through
Central America and perhaps to extreme north-
western Colombia. This species appears to be re-
stricted to midelevational slopes, and in Costa Rica

TIMM: ARTIBEUS AND URODERMA

195

Fig. 3. Photograph of an adult female Artibeus phaeotis roosting in a banana leaf tent. Details of the cut side
nerves can be seen along the midrib of the leaf.

I have found it only from 650 to 1 500 m in ele-
vation.

In April 1986 I observed a single Artibeus tol-
tecus roosting under a cut leaf of Anthurium ca-
peratum in Braulio Carrillo National Park, north-
eastern Costa Rica. The Anthurium was growing
as an epiphyte on a small tree at 800 m (IVi km
S, 11 km E of San Miguel, 10°17'N, 84'>05'W).
One leaf on the plant was altered; it was IVi m off
the ground and the midrib hung parallel to the
ground. Four or five lateral nerves were cut basally
on each side, causing the sides of the leaf to fold
down around the midrib. A break of the midrib
at its midpoint caused the distal half of the leaf to
droop down (fig. 4). Seven additional tents of this
style were observed on Anthurium in this area,
ranging in elevation from 700 to 1400 m. It is
assumed that they were made by A. toltecus, the
only small species of Artibeus we netted there, al-
though these tents were not occupied. Six tents
were observed in a 3-km transect ranging from
700 to 1 100 m in elevation.

Davis (1944) reported that Artibeus toltecus

roosts imder banana leaves, although he did not
indicate that the bats were modifying the leaves.
Davis (1944, p. 378) stated:

. . . they had regularly established roosts tin-
der the large, drooping leaves of the banana
trees, each one easily recognized by the man-
ner in which the vane of the leaf hung limply
suspended from the midrib. The closely ap-
pressed vanes of the leaf, plus the natural
darkness within the depths of the grove, af-
forded good concealment. These bats, too,
were wary and that feature coupled with the
nature of their retreat caused considerable
difficulty in procuring specimens.

In light of Davis's description of the roost sites of
A. toltecus in banana leaves and my own obser-
vations on A. toltecus, I suspect that this species
was creating tents similar to those I observed for
A. phaeotis in Costa Rica. The tents formed by A.
phaeotis in Musa (see fig. 2) are similar in ap-

196

HELDIANA: ZOOLOGY

Fig. 4. Top, Ventral view of an Anthurium caperatum leaf showing the cut nerves along the base of the leaf and
the broken midrib; bottom, tent of Art ibeus toltecus.

TIMM: ARTIBEUS AND URO DERMA

197

pearance to those described by Davis (1944) for
A. toll ecus.

Artibeus toltecus has also been reported in caves
(Davis et al., 1964; Jones, 1966; Jones & Alvarez,
1 964), and Goodwin ( 1 934, p. 1 2) reported a single
specimen collected "in one of the buildings
(church?) at San Lucas . . . [the] rest of the con-
gregation seemed to be Glossophaga."

Artibeus watsoni Thomas, 1 90 1

Artibeus watsoni, Thomas's fruit-eating bat, is
one of the smaller members of the genus Artibeus
and found from southern Veracruz south through
Central America to northern South America. It
appears to be restricted to lowland and midele-
vation humid forests.

During the summers of 1974, 1982, 1984, and
1 986, numerous individuals of Artibeus watsoni
were seen roosting under 19 different sj)ecies of
broadleafed plants at several localities in Costa
Rica. At Parque Nacional de Corcovado, Costa
Rica, trails were surveyed on three separate oc-
casions for the presence of tents made by Artibeus
watsoni, in June and August 1982 and again in
August 1984. In mid-June 1982, the following
groupings of Artibeus watsoni were observed: two
(both adults, a male and pregnant female), two
(pregnant female and one not captured), two (not
captured), and six hanging singly (of which two
were captured and found to be adult males). Ad-
ditionally, several tents on banana and Heliconia
were marked for relocation later in the summer.
A tent marked on Heliconia imbricata was relo-
cated 60 days later. The tent was still intact, al-
though it was beginning to break down; a single
A. watsoni v/zs using it. All other marked tents had
decomposed.

On 10 August 1982 I found 90 tents constructed
by Artibeus watsoni along the trail through Cor-
covado's "Monkey Woods." These tents were made
from the following species of plants: Musa x par-
adisiaca (49, 54%), Anthurium ravenii (13, 14%),
unidentified aroid (11, 12%), Heliconia imbricata
(9, 10%), Heliconia latispatha (1, 1%), Heliconia
sp. (3, 3%), and Calathea insignis (4, 4%). Tents
located on Anthurium ravenii were most often
found clumped, with an average of 2.6 tents per
plant, whereas in the other species of plants it was
uncommon to find more than one tent per indi-
vidual plant. Bats were found singly (five) or in

three groupings of four, three, and two individuals.
Four of the single bats were all adult males.

A trail running up to a ridge top was surveyed
from 9 through 1 1 August 1982, with the following
results: 25 tents found of which 16 were on Hel-
iconia imbricata (64%), 8 on Calathea insignis
(32%), and 1 on Carludovica palmata (4%). Three
tents were occupied by two (sexes unknown), one
male, and one female. Near the mouth of the Rio
Llorona on 8 and 9 August I counted the following
groups of bats: eight (three adult females, three
young, and two not caught), two (adult female with
volant young), two (sexes unknown), and three
singles (one a nonreproductive adult female). Ad-
ditional tents were observed in banana, coconut
palm (Cocos nucifera), Calathea insignis, and Car-
ludovica cf. drudei.

In August 1 984, 1 found 63 tents constructed by
Artibeus watsoni. These were distributed on the
following plants: Anthurium ravenii (36, 57%),
Heliconia sp. (14, 22%), Musa x paradisiaca (7,
1 1%), Calathea insignis (3, 5%), Carludovica pal-
mata (\, 1.6%), IVelfia georgii (1 , 1.6%),and(/eo/i-
oma sp. (1, 1.6%). Only 3 of the 63 tents were
occupied; one had two bats and two each had sin-
gle bats. As I noted in 1982, tents on Anthurium
ravenii were often clumped on the same plant with
an average of 2.5 tents per plant.

At La Selva in July 1982, 43 Artibeus watsoni
tents were located over a 5 -day period in the fol-
lowing species of plants: Asterogyne martiana (33,
77%), Geonoma congest a (6, 14%), Geonoma cu-
neata (2, 5%), and an unidentified species of Cy-
clanthaceae (2, 5%). One adult male A. watsoni
was found under an Asterogyne martiana tent on
the first day. On the fifth day an adult female with
young was found under another A. martiana tent
that had been unoccupied for the previous fovu"
days, as was a third adult (not captured) under
another A. martiana tent. All other tents were un-
occupied.

In 1974 1 surveyed approximately 10 km of trails
at La Selva and found 29 tents on the following
species of palms: Asterogyne martiana (19, 66%),
Bactris wendlandiana ( 1 , 3%), Geonoma congesta
(2, 7%) and Geonoma cuneata (7, 24%); all were
unoccupied. Foster and Timm (1976) reported
tents in these palms, although they were not able
to associate bats with the tents. My recent studies
at La Selva have confirmed that these tents were
made by A. watsoni.

At Finca Las Cruces in mid-August 1982, 13
tents constructed by Artibeus watsoni were located;

198

HELDIANA: ZOOLOGY

e
o

•a

eo|
C *

(/J c*
. k»

a T
•2 o

I a

c g
. u

C f
3 ■->
P «

« to

x: CO

♦^

o S

-3 e

o V

TIMM: ARTIBEUS AND URODERMA

199

Fig. 6. Top, Dorsal view of the cyclanth, Carludovica palmata, showing the polygonal cuts; bottom, tent of Artibeus
waisoni on C. palmata.

200

HELDIANA: ZOOLOGY

Fig. 7. Photograph of adult female Artibeus watsoni and her subadult offspring roosting in a Carludovica leaf tent.
Details of the polygonal cuts and folds can be seen in the background.

one was occupied by an adult male. The tents were
distributed on the following species of cyclanths:
Asplundia euryspatha (6, 46%), Carludovica drudei
(4, 31%), and Cyclanthus bipartitus (3, 23%).

Artibeus watsoni uses a variety of species of plants
and a wide array of leaf shapes for diurnal roosts.
I have found four distinct styles of tents at a single
locality (Corcovado). These styles include the sim-
ple V-shaped cuts on bifurcated palms (fig. 5), cut-
ting a few side veins on aroids to produce a round-
ed pyramid, the elongated J-shaped cuts on banana
and Heliconia leaves, and the polygonal cuts on
Carludovica (figs. 6-7). For each distinct leaf shape,
the cuts create a well-concealed diurnal roost. Ar-
tibeus watsoni probably is an obligate tent-rooster,
as it has only been found roosting under cut leaves.

On several instances a bat occupied the same
tent, or tents in close proximity, for two to three
days in succession. Those tents might then remain
unoccupied for several days in succession. Dis-
turbed bats generally flew directly to another tent
from 20 to 50 m away, or attempted to return to
the tent where originally found.

Tents generally are found clumped, both on a
single plant if leaf morphology and age are appro-
priate, and in restricted areas. Up to five tents have
been found on a single Anthurium ravenii, and
when present the mean number of tents was 2.5.

At Parque Nacional de Corcovado, Choe and
Timm (1985) found that Artibeus watsoni showed
strong preference for Anthurium ravenii leaves that
were medium sized, low within the plant, and grew
closer to the ground than average A. ravenii leaves.
Also at this site, Boinski and Timm (1985) doc-
umented that squirrel monkeys {Saimiri oerstedi)
were major predators on A. watsoni, with the adult
male monkeys being the most successful at cap-
turing bats. Additionally, double-toothed kites
{Harpagus bidentatus) followed troops of foraging
squirrel monkeys, using them as "beaters." When
tent-making bats were flushed by the monkeys and
escap)ed, they were routinely captured and con-
sumed by the attending double-toothed kites.

Artibeus watsoni has long been known to cut
palm tents for diurnal roosts, although prior to
this study little had been published on roosting

TIMM: ARTIBEUS AND URODERMA

201

Fig. 8. Dorsal view of Scheelea rostrata showing the leaflets cut by Uroderma bilobatum to form a tent

202

HELDIANA: ZOOLOGY

behavior of this species. The elongate J-shaped
cuts made on bifurcated palms were first described
and illustrated by Chapman (1932, p. 555). He
stated that "both vanes of the leaf whence the bat
flew were cut diagonally to the midrib of the leaf,
so that their terminal portions drooped downward
to form a tentlike shelter." Chapman appropri-
ately termed these three-sided diurnal roosts
"tents," and I have expanded the use of the word
tents to include all modified leaves by bats.

Barbour's (1932) original description of Uro-
derma bilobatum cutting palm leaves for roosts
also provides a secondhand report (p. 308) by H.
C. Clark stating that "Clark has just found for the
first time a youngish coconut palm, a single leaf
of which was being cut by bats of the genus Uro-
derma in a very similar way." The common use
of young coconut palms {Cocos nucifera) by Ar-
tibeus watsoni in Costa Rica, coupled with the total
lack of evidence that Uroderma bilobatum uses
juveniles of this palm, leaves that small, or roosts
that close to the ground, suggests that the tents
seen by Clark were in fact made by A. watsoni.
Artibeus watsoni also is abundant on Barro Col-
orado Island, Panama. Apparently no voucher
specimens of the bats were preserved at the time.
Allen (1939, p. 69) reported "a specimen of A.
watsoni that was hanging by day from the under-
side of a banana leaf" Perhaps the natural-looking
folds caused by the cuts running parallel to the
midrib were not noticed at the time. Ingles (1953)
reported on tents of A. watsoni in two species of
Geonoma on Barro Colorado Island; one tent was
occupied by three individuals. Thomas's fruit-eat-
ing bat has been found roosting in an artificial tent,
an inverted hanging box. Wilson (1970) reported
that several females raised young in the comer of
a suspended box on Barro Colorado Island.

Uroderma

Uroderma bilobatum Peters, 1865

Uroderma bilobatum has been given the dis-
tinctive "common" name of Peters's tent-making
bat. Tents constructed by Uroderma bilobatum
were seen at three separate localities in Costa Rica
during the summers of 1982, 1984, and again in
1986, the first at Bosque Brrancia near Cuarto
Cruces in the Pacific lowlands of northwestern
Costa Rica, the second at Corcovado on the Osa
Peninsula, and the third at La Selva in the Carib-
bean lowlands.

On 25 August 1982 a colony of Uroderma bi-
lobatum was roosting under a modified frond of
the palm Scheelea rostrata at Bosque Brrancia.
The colony included an adult male with enlarged
testes and four adult females.

The Scheelea rostrata frond in which the colony
of Uroderma bilobatum roosted was a mature leaf,
approximately 6.5 m in length. The bats were
hanging approximately 4.5 m off"the ground; most
were clustered together, although a few were spread
out over 50 cm of the frond. The cut leaflets started
at about 3.5 m off" the ground and proceeded up
the frond for the next 2.5 m (fig. 8). The general
pattern of the cut leaflets was a tapering effect, with
the cuts on the lowermost leaflets being farthest
from the midrib. Leaflets along the proximal 2 m
and the distal 50 cm were unmodified. Only the
midrib of the leaflets was cut. Each leaflet had a
distinctive V-shaped fold at its base where it was
attached to the midrib. The bats were hanging
from the leaflets rather than the midrib.

From the dorsal aspect of the leaf, the proximal
portion of the tent (cut leaflets) extended 50 cm
further down on the right side than on the left to
include 10 basal leaflets whose opposites on the
left were unaltered. The basalmost cut leaflet was
cut 34 cm from the midrib. Proceeding distally,
the length of the unmodified basal portion of each
leaflet decreases. The basalmost cut leaflet on the
left was cut 19 cm from the midrib. The overall
appearance of the tent was a sharp, convergent
taper for the next meter. Following this section,
there was a 7 5 -cm section in which the cuts were
close to the midrib (within 3 cm). On the distal-
most 30 cm of the tent, the leaflets were cut closer
to the midrib on the left side than on the right.
Similar tents, each housing a colony of Uroderma
bilobatum, were found in a large stand of Scheelea
rostrata at Corcovado in 1984, and William A.
Haber (pers. comm.) informed me that he has seen
similarly cut leaves in the same species of palm at
Cahuita (09°44'N, 82°49'W) in the Caribbean low-
lands of southeastern Costa Rica in 1 984.

In June of 1982 and again in March of 1986, I
found numerous banana leaves cut by Uroderma
bilobatum just to the north of the field station at
La Selva. The midrib on vertical leaves was cut
to the extent that the distal portion of the leaf
collapsed downward to form a two-sided tent (fig.
9). Severing the midrib on vertical leaves had the
effect of folding the leaf back upon itself creating
a tight, dark crevice at the fold where the bats
roosted (fig. 1 0). In addition to severing the mid-
rib, the bat cut a large V-shaped pattern running

TIMM: ARTIBEUS AND URODERMA

203

Fig. 9. Left, Dorsal view of a banana leaf showing the cut midrib and the large V-shaped cuts through the side
nerves; right, tent of Uroderma bilobatum made from a banana leaf.

from the midrib to the base of the leaf. The side
veins and interconnected tissues were partly sev-
ered. However, because the leaf stood nearly ver-
tical, these V-shaped cuts did not cause further
folding of the leaf The only cut creating the tent
was that of the midrib.

In 1982 five tents in widely separated banana
leaves were located (table 4). One was occupied
by eight Uroderma bilobatum, which included one
adult male with enlarged testes and seven females.
In 1986 eight tents were observed in the same
banana patch. On this occasion eight U. bilobatum

204

FIELDIANA: ZOOLOGY

TIMM: ARTIBEUS AND URODERMA

205

tents were clustered in three clumps of bananas.
Only one tent was occupied; it contained 1 3 bats.

This folded broad-leaved style of tent is un-
doubtedly the tent style illustrated by Walker ( 1 960,
p. 30) in his photograph of roosting Uroderma
bilobatum, although he did not describe it nor
mention where it was observed. Interestingly, I
have searched several dozen banana and larger
Heliconia groves throughout Costa Rica and Ec-
uador specifically looking for this style of tent, and
none were observed. Uroderma bilobatum is an
abundant and widespread species in the lowlands,
but employment of this particular style of tent
appears spotty as I have not observed it elsewhere.

In Panama, Barbour (1932) found that colony
size under a single cut leaf of Prichardia pacifica
varied from a few bats to 56. Prior to Barbour's
discovery that Uroderma altered leaves, Goldman
(1920, p. 199) stated of these bats in Panama.

In the forest near Gatun Uroderma biloba-
tum was located several times, a few in a
place, clinging during the day in clusters to
the midribs on the under sides of large palm
leaves. They usually choose darkened spots
where the leaf was folded over, or over-
hanging pinnae shut out much of the light.

Burt and Stirton (1961) reported U. bilobatum in
El Salvador roosting beneath the leaves of bananas
and coconut palms. I strongly suspect that the bats
had altered these leaves and that these authors had
failed to notice it or failed to associate the hanging
bats with the damage to the leaves. Bloedel (1955,
p. 234) stated of Uroderma bilobatum in Panama
that:

I observed these bats only in their palm-leaf
tents. ... In the latter part of March most
females have nursing young, and are roost-
ing in clusters of 20 to 40, while the males
are separated from them, usually solitary or
in small groups of from 2-5.

In Trinidad, Goodwin and Greenhall (1961, p.
254) found them

on the under side of the fan-shaped leaves
of certain palm trees, especially the carat
palm (Sabal glaucescens). . . . The bat makes
a series of cuts across the pleated surface of
a leaf, causing half of the leaf to bend at an
angle to form a protected retreat.

Table 4. Individual measurements (in cm or de-
grees) collected from four tents constructed by Uroderma
bilobatum on banana, Musa x paradisiaca, in Costa
Rica.

Blade

Petiole

Cut
midrib

Length Width

angle

from base

Remarks

210 95

117

Not occupied

220 90

8 U. bilobatum

1 U. bilobatum

Not occupied

210 95

Not occupied

They were found roosting in colonies of 1 or more
individuals. In Suriname, Husson (1962, p. 161;
1978, p. 143) collected three pregnant female U.
bilobatum "in a plantation where they were found
hanging on the imder side of a leaf of [a] so-called
'paloeloe,' Ravenala guyariensis." In Nicaragua,
Jones ( 1 964, p. 507) collected four female U. bi-
lobatum that "hung together about 10 feet above
the ground in the 'tent' formed by a cut palm frond.
Each was pregnant with a single embryo." In Gua-
temala, Dickerman et al. (1981, p. 409) reported,
"Palm leaf tents were frequently found occupied
by one to seven individuals, but nursing females
were usually found alone or with juveniles."

Leaves selected by Uroderma bilobatum for tents
are all large, and of a variety of shapes. The large
V-shaped pattern cut into the leaves is a charac-
teristic of Uroderma tents. Artibeus watsoni also
uses a variety of leaf shapes for tents; however,
the nature of the cuts and style of tent created
vary with leaf shape. Uroderma bilobatum, on
the other hand, makes patterned cuts that appear
to be an innate response to large leaves, regardless
of the shape. As noted in the banana tents, a single
cut across the midrib creates the tent, and the
V-shaped pattern had no effect upon tent shape;
this perhaps represents wasted effort by the bats.

There are a few records of Uroderma bilobatum
being found roosting in a hollow tree and one "un-
der the eave of house" (Davis, 1968, p. 695). In
all cases these have been of single individuals, and
I suspect represent either recently dispersed young
that have yet to join a breeding colony or bachelor
males. Uroderma bilobatum roosting under a cut
Prichardia leaf was illustrated by Kunz (1982). A
colony of U. bilobatum roosting under a banana
leaf tent was illustrated by Keller ( 1 986). Mac-
donald's ( 1 984, p. 806) photograph of two tent-
making bats under a Heliconia tent is erroneously
labeled Uroderma bilobatum. These bats are ac-

206

HELDIANA: ZOOLOGY

Fig. 11. Tent of Vroderma
magnirostrum on the pinnately
leafed palm Astrocaryum muru-
muru.

tually a small species of Artibeus, probably A.
phaeotis; the tent style and size and coloration of
the bats are typical oi A. phaeotis.

Uroderma magnirostrum Davis, 1968

Although Vroderma magnirostrum is a widely
distributed bat found from Mexico to Bolivia, it
was not recognized as a species distinct from U.
bilobatum until 1968 (Davis, 1968), and few spec-
imens are represented in collections. When W. B.
Davis described this new species he commented

that although it was widely distributed only seven
specimens had been collected prior to the wide-
spread use of mist nets in the 1960s, and that all
specimens available to him had come from local-
ities less than 1000 feet in elevation. He stated
that "These facts strongly suggest basic differences
in the habits of the two species and that those bats
with a deep rostrum are not 'tent-makers' as are
members of the species Uroderma bilobatum"
(Davis, 1968, p. 678). There have been no reports
on the behavior or ecology of U. magnirostrum.
On 25 July 1985 Barbara L. Clauson discovered
a colony of two male and three female Uroderma

TIMM: ARTIBEUS AND URODERMA

207

magnirostrum roosting under the cut leaflets of the
pinnately leafed palm, Astrocaryum murumuru.
The single occupied tent found was on the ridge
above Hacienda Amazonia at 825 m in south-
eastern Peru. The entire colony was collected by
John W. Fitzpatrick. On 3 November Clauson re-
turned to the site to measure ,»ie leaf and noted
an additional cut leaf in the same plant.

When first observed, four bats were hanging to-
gether and one was hanging several centimeters
away. When observed an hour later, all five were
hanging together in a tight cluster approximately
7.5 m off the ground. The bats were hanging from
the leaflets rather than the midrib, approximately
200 cm from the tip of the leaf The colony in-
cluded one adult male with enlarged testes, two
adult females, and two subadults, one female and
one male.

The roosting structure of Uroderma magniros-
trum was in a pinnately leafed palm (fig. 1 1 ). The
bats severed the leaflets along the upper two-thirds
of the leaf; those along the lower third were un-
altered, as were the leaflets at the very tip. As the
leaflets proceeded up the tent they were severed
closer to the midrib forming an elongate, conver-
gently tapering tent (fig. 1 1). The general appear-
ance of the U. magnirostrum tent is similar to that
described herein for U. bilobatum on the pinnately
leafed palm Scheelea rostrata.

The Astrocaryum frond in which the colony of
Uroderma magnirostrum roosted was a mature leaf,
approximately 6.1 m in length and 1.9 m in width
at the widest point, with the petiole 1.1m long.
The leaf left the trunk (d.b.h. .4 m) at 3 m from
the ground and hung at an angle of approximately
50°. The bats were hanging approximately 7.5 m
off" the ground. The cut leaflets started at 1.5 m
from the lowest leaflet and proceeded up the frond
for the next 2.9 m to nearly the tip (fig. 1 1). The
cuts on the lowermost leaflets were furthest from
the midrib. The lowest severed leaflets were cut
up to 34 cm from the midrib, whereas the distal
leaflets were severed only 2 cm from the midrib
of the leaf The midribs of the leaflets were cut
causing the distal portion of the leaflets to fold
downward. Leaflets along the proximal 1.5 m and
the distal .5 m were unmodified. The trunk, pet-
iole, and midrib of this palm were covered with
sharp, penetrating spines several centimeters in
length. After 14'/2 weeks this tent was still alive
and green, most of the leaflets appearing as fresh
in November as they did in July.

This Astrocaryum contained a second cut leaf
that was unoccupied. This roost was also in a ma-

ture leaf which was an older leaf than the occupied
tent, with many broken, yellowed, and brown leaf-
lets. The leaf was approximately 6 m in length and
2 m in width at the widest point, with a petiole .8
m long. The leaf left the trunk at 2.9 m from the
ground and hung at an angle of approximately 60°.
Cuts were distributed asymmetrically along the
length of the leaf The cut leaflets on the left side
of the leaf started 3.05 m from the lowest leaflet
and proceeded up the frond for the next 1 .3 m, to
.88 m from the tip. The cut leaflets on the right
side of the leaf started 3.25 m from the lowest
leaflet and proceeded up the frond for the next
1.49 m, to .49 m from the tip. The cuts on the
lower leaflets were farthest from the midrib. The
lowest severed leaflets were cut up to 35 cm from
the midrib, whereas the distal leaflets were severed
as close as 1.5 cm from the midrib. Leaflets along
the proximal 3.05 m and 3.25 m and the distal
.88 m and .49 m were unmodified.

I propose the common name of Davis's tent-
making bat for this species.

Conclusions

A review of the literature on tent-making bats
contains some 32 primary references covering the
55-year period from 1932 through 1986. Surpris-
ingly, we actually know very little about the bi-
ology of these bats.

As late as 1975 Eisentraut was yet doubting th^t
bats were cutting leaves to make tents, stating:

. . . observers maintain that the bats form
these tentlike structures themselves, by
making a series of holes running across the
middle of a large palm leaf The bats then
supposedly bend the outer half of the leaf
around, so they can then rest inside this
'tent'. . . . On the basis of personal obser-
vations in tropical regions in Africa, I tend
to believe instead that these holes were made
by insect larvae while the leaves were still
rolled up. A storm can then easily break the
leaf along the line of holes and form the tent
roof which is so convenient for the bats (Ei-
sentraut, 1975, p. 142).

Eisentraut, by his own admission, had never seen
a bat tent. I believe that if he had, he would have
come to the same conclusion Thomas Barbour did

208

HELDIANA: ZOOLOGY

nearly a half century earlier, that the bats and not
insects were making the cuts.

Although we have yet to actually observe bats
cutting leaves to form the roosting structures de-
scribed herein, I hope the volume of data pre-
sented here and in my other works establishes for
a fact that many species of small and medium sized
stenodermines are indeed tent-makers. The ob-
servations presented represent data collected from
several hundred tents located over a 1 5 -year pe-
riod. Several facts consistently emerge between my
observations and those independently corrobo-
rated by others. Bats of the genera Artibeus and
Uroderma (as well as Ectophylla, Mesophylla, and
Vampyressa) roost under cut leaves. These leaves
may be on a wide variety of species of plants, but
generally the shape of the leaves is similar. The
shape of the cuts is very characteristic for each
species of bat and the patterns and styles of tents
created by the bat species are consistent.

The behavioral repertoire associated with tent-
making in bats certainly evolved more than once,
as evidenced by the patchy distribution of tent-
making species with the chiropteran suborders
Megachiroptera and Microchiroptera. Within the
Megachiroptera, a single species of tent-maker is
known, Cynopterus sphinx. Within the Microchi-
roptera, tent-makers are known only from one
subfamily of the Phyllostomidae, the Stenoder-
minae. The Stenoderminae constitute an ex-
tremely speciose and diverse group of bats, with
more than 30 species currently recognized. Tent
construction within stenodermines may be a trait
that evolved once, twice, or as many as three times.
The Artibeus- Uroderma group are sister genera and
form one clade of the tent- making repertoire. Sec-
ondly, the Mesophylla- Vampyressa group are sis-
ter genera (and perhaps should be considered con-
generic) and would constitute the second clade.
Finally, Ectophylla would constitute a third lin-
eage. The relationship between these three lineages
is uncertain and warrants further investigation.

Knowing that bats modify the leaves of several
species of plants to produce diurnal roosting struc-
tures led to the following questions: (1) Are bats
selecting specific species of plants for tents? (2)
What styles of tents are cut by bats and do these
differ between species? (3) Do bats select for a
particular angle, size, or shape of leaf for diurnal
tents? (4) Are leaves selected preferably in larger
clumps or smaller clumps? (5) Are older or youn-
ger leaves selected? (6) Are leaves of a particular
height class selected? (7) Are leaves that are not
adjacent to solid objects selected? (8) What do

typical tents look like? (9) How and why did tent
construction evolve?

On occasion I have found "cheaters," species of
bats roosting in a tent made by another species.
Is cheating an evolved strategy of roost site selec-
tion of some bats?

Bats of the genera Artibeus and Uroderma ac-
tively modify leaves to produce diurnal roosting
structures, but by biting the tissue between veins
along the midrib and leaving the midrib and most
veins intact, do not kill the leaves. The resulting
tent is available for use as a roost for an extended
period of time; one was observed in use for more
than 60 days. Bats select for specific sizes and
shapes of leaves. Tents provide concealment from
predators and protection from the rain, wind, and
sun. This type of roost offers the additional ad-
vantage that the bats are warned about the ap-
proach of a potential predator, because even slight
movements of the leaf stem or the leaf itself are
transmitted as magnified vibrations to the roosting
bat. Tents may provide bats with suitable roosting
sites that would not otherwise be available in close
proximity to prime food resources.

One of the most productive areas for future re-
search will be exploring aspects of the biology of
these bats from an evolutionary perspective. Fu-
ture subjects I will be addressing include the role
of tent roosting in controlling ectoparasites and
the correlation between complexity of tents and
social systems in these bats. I believe that tent-
making originated as an antipredation strategy and
has since, secondarily, evolved to play a major
role in controlling ectoparasites and in social be-
havior.

Many factors influence the choice of roost site
selection by bats. Included among these are vul-
nerability to predation, physical stability of the
site, proximity to food sources, and general ap-
propriateness of the nest microenvironment for
the rearing of young. It seems likely that tent con-
struction requires considerable time and energy
expenditure by bats, attesting to intense selection
pressures involved.

Acknowledgments

I thank Eduardo Lopez Pizarro and El Depar-
tamento de Vida Silvestre and Fernando Cortes
and Servicio de Parques Nacionales of Costa Rica
for making this study possible. The Organization
for Tropical Studies (OTS), Rebecca Butterfield,

TIMM: ARTIBEUS AND URODERMA

209

William A. Haber, Gary Hartshorn, Charles E.
Schnell, and Joe M. Wunderle are gratefully ac-
knowledged for assistance with logistics. Robert J.
Izor assisted with field logistics in Costa Rica and
Peru, and John W. Fitzpatrick assisted in Peru. In
Ecuador I thank the Comandancia General del
Ejercito Ecuatoriano, the Corporacion Estatal Pe-
trolera Ecuatoriana, and the Ministerio de Agri-
cultura y Ganaderia for making our studies there
possible. Luis Albuja, Ramiro Barriga, Angelitos
Garrett, Myriam Ibarra, Gustavo Orces, and Don-
ald J. Stewart provided logistic assistance in Ec-
uador. Kerry A. Barringer, William C. Burger,
Thomas B. Croat, Robin B. Foster, Barry Ham-
mel, and Timothy Plowman provided identifica-
tions or confirmed identifications of the plants.
Barbara L. Clauson, Alfred L. Gardner, Lawrence
R. Heaney, Karl F. Koopman, Thomas H. Kunz,
Bruce D. Patterson, and Timothy Plowman pro-
vided valuable suggestions on earlier drafts of the
manuscript. My wife, Barbara, provided superb
assistance with all aspects of this project, including
providing several of the photographs used and all
data on Uroderma magnirostrum. Rosanne Mie-
zio prepared the illustrations. Nina Cummings,
Ron Testa, and Diane White expeditiously and
cheerfully executed my photography requests. This
project was funded in part by grants from the Rice
Foundation of Chicago, the National Science
Foundation [INT-8303 194], National Geographic
Society, and Field Museum of Natural History. I
especially thank Mr. and Mrs. Arthur A. Nolan,
Jr. for their continuing support of my research.

This paper is dedicated to Philip Hershkovitz
in recognition of his contributions to Neotropical
mammalogy, and most especially for the friend-
ship he has shown me.

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Appendix

List of scientific names of plants mentioned in the text and used by Neotropical bats for tent con-
struction.

Anacardiaceae

Anacardium excelsum (Bertero & Balbis) Skeels

Araceae (cont'd.)
Monstera lechleriana Schott
Xanthosoma sp.

Araceae

Cyclanthaceae

Anthurium caperatum Croat &. Baker
Anthurium ravenii Croat & Baker

Asplundia euryspatha Hari.
Carludovica dmdei Masters

TIMM: ARTIBEUS AND URODERMA

211

Cyclanthaceae (cont'd.)
Carludovica palmata R. & P.
Cyclanthus bipartitus Poit.

Marantaceae

Calathea insignis Petersen

Musaceae

Heliconia imbricata (Kuntze) Baker
Heliconia latispatha Benth.
Musa X paradisiaca L.

Plenakospermum guyanense Endl. (syn. Ravenala
guyanensis Petersen)

Palmae

Asterogyne martiana (H. Wendl.) H. Wendl. ex

Hemsley
Astrocaryum murumuru Mart.
Bactris wendlandiana Burret
Cocos nucifera L.

Geonoma congesta H. Wendl. ex Spruce
Geonoma cuneata H. Wendl. ex Spruce (syn. G.

decurrens H. Wendl.)
Geonoma oxycarpa Martius (syn. G. binervia

Oerst.)
Livistona chinensis (Jacq.) R. Br. ex. Mart.
Prichardia pacifica Seem. & H. Wendl.
Sabal mauritiiformis (Karsten) Griseb. &. H.

Wendl. ex Griseb. (syn. S. glaucescens Lodd. ex

H. E. Moore)
Scheelea rostrata (Oersted) Burret
Welfia georgii H. Wendl. ex Burret

212

FIELDIANA: ZOOLCX5Y

Comparative Ultrastructure and Evolutionary Patterns

of Acinar Secretory Product

of Parotid Salivary Glands in Neotropical Bats

Carleton J. Phillips, Toshikazu Nagato, and Bernard Tandler

ABSTRACTS

Secretory products produced by acinar cells of the parotid salivary glands of 1 5 species of
Neotropical bats {Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, T.
sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium,
Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis,
and Molossus molossus) were compared by transmission electron microscopy. Extensive inter-
and intrageneric differences were found in the ultrastructure of the mature acinar secretory
granules. This variation in secretory cell product exceeded any previously reported intraordinal
phenotypic variation at the cellular level, but was in keeping with previously reported bio-
chemical data on salivary protein polymorphism in primates and rodents. Data from molecular
biology and systematics lend support to the hypothesis that the microscopic variations are
directly representative of genie differences among species. It also is postulated that intrageneric
microscopic differences at least partly are due to neutral (nonfunctional) differences in molecular
structure or charge (or both) rather than evolutionary selection. Among the phyllostomids, a
general trend in parotid acinar cell product was found in Artibeus and Ariteus, in which a
decrease in enzymatic content of the product could be correlated with ultrastructural differences.
The secretory product in Artibeus and Ariteus also differed significantly from that of the genus
Sturnira. and it is proposed that the phenotypic differences between Sturnira and the other two
stenodermatines represent a major genetic difference of systematic importance. The ultrastruc-
tural appearance and substructure of the parotid acinar secretory granules could not be con-
sistently correlated with diet alone, although insectivorous-animalivorous SF)ecies have enzyme-
rich, mostly electron-dense granules, whereas two fruit bats, Artibeus and Ariteus, have pale,
enzyme-poor parotid granules.

Productos de secrecion producidos por celulas acinares de las glandulas salivales de 1 5 esF>ecies
de murcielagos neotTopicales (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia
bidens, T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira
lilium, Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida bra-
siliensis y Molossus molossus) fueron comparados mediante microscopio electronico de trans-
mision. Extensivas diferencias inter- e intragenericas fueron encontradas en la estructura de
granulos glandulares acinares maduros. Esta variacion en productos de celulas secretoras excedio

From the Department of Biology, Hofstra University,
Hempstead, NY 11550 (Phillips); and Department of
Oral Biology, School of Dentistry, Case Western Reserve
University, Cleveland, OH 44106 (Nagato and Tandler).
Dr. Nagato's present address is Department of Oral and
Maxillofacial Surgery, Ehime University School of Med-
icine, Shizukawa, Shigenobu, Onsen-gun, Eshime 79 1 -02,
Japan.

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 213

cualquier variacion fenotipica intraordinal previamente reportada a nivel celular, pero estuvo
en armonia con datos bioquimicos previamente reporlados de polimorfismo en proteinas sa-
livales de primates y roedores. Datos de biologia molecular y sistematica proveen apoyo a la
hipotesis de que las variaciones microscopicas son direclamente representativas de diferencias
geneticas entre especies. Es lambien postulado que diferencias microscopicas intragenericas, al
menos parcialmente, son debidas a diferencias neulrales (no funcionales) en estructura molecular
o carga (o ambas) mas bien que a seleccion evolutiva. Entre los filostomidos, una tendencia
general en el producto parotido de las celulas acinares fue encontrado en Artibeus y Ariteus, en
los cuales una disminucion en contenido enzimatico del producto p>odria estar relacionado con
diferencias ultraestructurales. El producto secretado en Artibeus y Ariteus tambien se diferencio
significativamente de aquel del genero Sturnira y es propuesto que las diferencias fenotipicas
entre Sturnira y los otros dos stenodermatinos representan una mayor diferencia genetica de
importancia sistematica. La apariencia ultraestructural y subestructura de los granulos secre-
torios parotidos acinares podria no estar consistentemente correlacionada solo con la dieta,
aunque especies insectivoras-animalivoras tienen granulos ricos en enzimas mayormente densos
en electrones, mientras que los murcielagos frugivoros, Artibeus y Ariteus, tienen granulos
parotidos palidos y pobres en enzimas.

Produtos secretorios produzidos por acinos das glandulas parotidas de 15 especies de mor-'
cegos neotropicos (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens,
T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium.
Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis,
e Molossus molossus) foram comparados atraves da microscopia de transmissao eletronica.
Vastas diferengas intra- e intergenericas foram encontradas na ultra-estrutura dos granulos
maduros dos acinos secretorios. Esta variafao no produto das celulas secretorias supera qualquer
varia^ao fenotipica intraordinal previamente relatada para o nivel celular, porem concorda
com as rela96es publicadas sobre dados bioquimicos do polimorfismo de proteinas salivares
em primatas e em roedores. Dados sistematicos, e de biologia molecular, apoiam a hipotese
que variances microscopicas sao diretamente representativas das diferen^as geneticas entre
especies. Propoese tambem, que estas diferen^as microscopicas intragenericas sao ao menos
parcialmente causadas por diferen9as neutras (i.e., naofuncionais) nas estruturas moleculares
ou nas suas cargas eletricas (ou ambas), ao inves de serem consequencias da sele^ao evolutiva.
Entre os morcegos da familia Phillostomidae, foi encontrado em Artibeus e em Ariteus um
padrao geral nos produtos dos acinos parotideos, onde uma redu9ao do conteudo enzimatico'
e correlacionado a diferencas nas ultraestruturas dos granulos produzidos. Os produtos secre-
torios em Artibeus e em Ariteus sao significantemente diferentes dos produtos do genero Sturnira.
e prop6e-se que as diferen9as fenotipicas entre Sturnira e os outros dois stenodermatinos
representam uma grande diferen9a genetica, de importancia sistematica. A aparencia ultraes-
trutural, e a subestrutura dos granulos secretorios dos acinos parotideos, nao se correlacionam
consistentemente com a dieta por si, apesar de que as especies insetivoras-animalivoras possuem
granulos ricos em enzimas e densos em eletrons, equanto que dois morcegos frugivoros, Artibeus
e Ariteus, (wssuem granulos parotideos palidos e com poucas enzimas.

Introduction

Neotropical bats are extremely diversified; ex-
isting species represent perhaps the most outstand-
ing mammalian example of ecomorphological ad-
aptation. Dentitions, jaw morphology, brains,
kidneys, tongues, and digestive tracts are only a
few examples among the anatomical features that
have been investigated in recent years (e.g., Phil-
lips, 1971; Forman, 1972; Phillips et al., 1977,

1984; Freeman, 1979, 1981; Eisenberg «t Wilson,
1978; Studier et al., 1983). The major salivary
glands are yet another system that has attracted
attention, primarily because histological, ultra-
structural, and histochemical investigations have
consistently revealed striking interspecific differ-
ences and unusual histological and secretory fea-
tures (Wimsatt, 1956; DiSanto, 1960; Junqueira
& Fava de Moraes, 1965; Junqueira et al., 1967,
1973; Phillips et al., 1977; Mineda, 1977, 1978;

214

HELDIANA: ZOOLOGY

Pinkstaff et al., 1982; Tandler & Cohan, 1984;
Nagatoetal., 1 984; Tandler & Phillips, 1985; Phil-
lips & Tandler, 1985, 1987; Tandler et al., in press).

Mammalian salivary glands are highly complex
organs that not only secrete digestive enzymes but
also can secrete hormones, antibodies, and toxins,
to name but a few known products (e.g., Tandler,
1972; Hand, 1980b). Data are available on the
biochemistry of saliva in humans and common
laboratory species; but very little is known about
the specific biochemistry of the parotid acinar se-
cretory granules except for laboratory rats in which
some of the proteins have been characterized (Ro-
binovitch & Sreebny, 1969; Ball, 1974; Wallach
et al., 1975). The complex structure and function
of salivary glands is underscored by data from
studies of Neotropical bats, which recently have
been analyzed by both transmission electron mi-
croscopy and histochemistry. For example, the ac-
cessory submandibular gland of Trachops cirrho-
sus has been shown to differ histologically from
any known mammalian salivary gland, with the
exception of the same gland in Megaderma lyra
and M. spasma. Megaderma lyra is an Old World
ecological equivalent of Trachops (Phillips &.
Tandler, 1985, 1987; Phillips et al., 1987). Both
of these unrelated species feed on frogs (Lekagul
& McNeely, 1977; Tuttle & Ryan, 1981), which
possibly has been a factor in the convergent evo-
lution of their submandibular glands. A previously
unknown cellular organelle has been described in
another Neotropical bat, Tonatia sylvicola (Na-
gato et al., 1984). In this species, the presence of
xmique crystalloid smooth endoplasmic reticulum
in seromucous acinar cells is sex-linked (being
found only in submandibular acinar cells in males);
a steroid product produced by this organelle pos-
sibly serves as a species-isolating mechanism or
as part of a chemo-behavioral system, or both.
Lastly, a comparative investigation of the secre-
tory product in seromucous acinar cells in sub-
mandibular glands of five species ofArtibeus has
revealed that the ultrastructural characteristics of
secretory products can have systematic implica-
tions (Tandler et al., 1983, 1986). This study was
of particular interest because the salivary gland
data matched genie data independently derived by
Koop and Baker (1983).

For the present investigation we surveyed par-
otid acinar cell secretory products in a selected
group of 1 5 species of Neotropical bats. Our group
comprised four families (Mormoopidae, Phyllo-
stomidae, Vespertillionidae, and Molossidae) and
included a group of species in which dietary habits

ranged from insectivory and animalivory to om-
nivory and frugivory. This investigation is the first
comprehensive interspecific survey of secretory
product ultrastructure, and addresses the follow-
ing questions: ( 1 ) what is the range of variation in
secretory products; (2) what are the evolutionary
patterns in parotid secretory product; and (3) what
systematic conclusions can be reached by com-
parative ultrastructural analysis?

Materials and Methods

Numbers and sex of specimens used in the pres-
ent study are given in the Appendix. All of these
bats were collected during fieldwork in Mexico,
Jamaica, and Suriname. Voucher specimens for
all species and collecting localities are deposited
in the mammal collections of either the Carnegie
Museum of Natural History or The Museum, Tex-
as Tech University. Bats typically were collected
at night with mist nets, and were kept overnight
without food until they were killed between 0900
and 1 200 the following morning. The animals were
anesthetized with 0.25 ml of sodium pentobarbital
(50 mg/ml, intraperitoneally) and the salivary
glands removed, placed on dental wax, flooded
with freshly mixed fixative, and diced into pieces
measuring approximately 1 mm'.

Two fixation protocols were used at different
times during the project. Specimens collected in
Mexico and Jamaica were fixed in 2% glutaral-
dehyde in 0. 1 M phosphate buffer and then stored
unrefrigerated in fresh fixative. The specimens col-
lected in Suriname were fixed in a modified tri-
aldehyde-dimethylsulfoxide (dmso) mixture, first
described by Kalt and Tandler (1971) and later
modified slightly for fieldwork (Phillips, 1 985). The
trialdehyde fixative, consisting of 3% glutaralde-
hyde, 1% paraformaldehyde, 0.5% acrolein, 2.5%
DMSO, and 1 mM CaClj in a 0.05 M cacodylate
buffer and sucrose at pH 7.2, proved superior to
the simple glutaraldehyde fixative in that (1) a
higher percentage of tissues proved to be ade-
quately fixed for study, and (2) the mitochondria
tended to remain intact instead of being disrupted.
Although the composition of the fixatives was dif-
ferent, we have not found any evidence that these
differences introduced undesirable artifacts that
would influence our analysis. Additional details
about the techniques can be found in both Phillips
(1985) and Nagato et al. (1984).

To remove unbound aldehydes, the tissue blocks
were subjected to prolonged washing in phos-

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

215

acinar lumen.

acinar cell

secretory
graauLej

striated Lntercalated acinus

duct duct

Fig. I . Schematic diagram of a secretory unit in a typical parotid gland. A cluster of secretory cells, arranged
around a lumen to form an acinus, drains into an intercalated duct that in turn empties into a striated duct. In reality,
more than one acinus may be associated with an intercalated duct, which itself may be branched. The acinar cells
elaborate the primary saliva, which has plasma-like concentrations of electrolytes, and add their organic secrelory
products to it. The precise role of the intercalated duct cells has not been established, but they probably play a role
in secretion and transport. The principal function of the striated ducts is resorption of electrolytes, mainly sodium,
from the primary saliva, rendering it hypotonic.

phate-buffered sucrose. The blocks were postfixed
for two hours in phosphate-buffered 2% OSO4,
rinsed in distilled water, soaked overnight in cold
aqueous 0.25 uranyl acetate, rinsed again in dis-
tilled water, dehydrated in ascending concentra-
tions of ethanol, passed through propylene oxide,
and embedded in Epon-Maraglas (Tandler & Wal-
ter, 1977). Thin sections were doubly stained with
methanolic uranyl acetate (Stempak & Ward, 1 964)
and lead citrate (Venable & Coggeshall, 1965). All
sections were examined in a Siemens Elmiskop la
transmission electron microscope (tem). Semithin
sections ( 1 nm) used for orientation were stained
with toluidine blue (Bjorkman, 1962) and exam-
ined in a Zeiss Ultraphot.

Tissue samples used for comparisons were se-
lected carefully from among available tissue blocks.
We generally avoided edges of tissues where me-
chanical trauma often affects not only the cellular
ultrastructure but also the microscopic appearance
of the secretory granules. In selecting representa-
tive "mature" granules we took into account the
full array of inter- and intracellular variability as
well as the often complex substructural geometry
of the granules. Selection of the most representa-
tive granules admittedly was subjective, but based
on our experience, four principal criteria were
applied: (1) cells selected for analysis showed no
signs of shrinkage or swelling, and sensitive or-
ganelles, such as mitochondria, were not distorted;

(2) the appearance of the granule was not altered
in any meaningful way by variations in fixation;

(3) the appearance of the secretory granules had
to be unrelated to their location within the spec-
imen block; and (4) the development of the gran-
ules could be traced (from Golgi complex to apical
cytoplasm) without any major breaks in devel-
opmental sequence.

Results '

The basic histology of the parotid salivary gland
was similar in all 15 sp)ecies examined; in each
species the acinus was formed by a cluster of cells
connected to the striated duct by intercalated ducts
of varying lengths (fig. 1). At the tem level, acinar
cell secretory products were found to differ in all
species examined (figs. 2-5). These differences
could not be related to granule ontogeny by com-
paring "immature" Golgi-GERL (Golgi-endoplas-
mic reticulum-lysosomes)-associated granules with
"mature" granules in the apical cytoplasm. In our
samples that consisted of both males and females,
we found no evidence of sexual dimorphism in
secretory granule substructure, and in our largest
samples (5 to 1 specimens) we found no examples
of individual or geographic variation that could
not be attributed to typical inter- or intracellular
variation or to granule geometry. Although vari-

216

FIELDIANA: ZOOLOGY

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

217

53 F

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c
u

i/i

■<t

•S

S-

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CJ,

o
>

•i

■^

r<^

r-'

**^

i/i

k
Q

■S.

S.,

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218

FIELDIANA: ZOOLOGY

ation may occur at the individual level, we found
no unequivocal evidence that it is detectable by
transmission electron microscopy. Last, the ex-
amples selected to illustrate each species (figs. 2-
5) should be regarded as "average" or represen-
tative of the most common, well-fixed granule
morphology.

Pteronotus parnellii— The secretory granules
consisted of a moderately dense, homogeneous,
variably thick outer zone and a slightly lighter in-
terior in which were embedded a collection of lin-
ear densities, usually randomly disposed but
sometimes showing a degree of orderliness. Each
of these densities was outlined by a thin, barely
discernible, lucent layer (fig. 2).

Phvllostomus elongatus— The matrix of the
secretory granules was uniformly dense and con-
tained a series of irregular shell-like densities, which
in thin section appeared as ribbons consisting of
a dense line flanked by thin lucent plies. In certain
granules, the ribbons were aligned in concentric
fashion, imparting a layered appearance to the pe-
riphery of these cell structures.

Phvllostomus latifolius— These granules
bore some resemblance to those in P. elongatus in
that some had one or two ribbon-like layers im-
mediately subjacent to their limiting membrane,
but many granules lacked these layers. Instead, the
prevailing inclusion was a lucent, coreless strand
disposed in a complex skeinlike or twisted pretzel
conformation (fig. 2).

Ton ATI A BiDENS— These secretory granules con-
tained several groups of hexagonally-packed tu-
bules just beneath the limiting membrane. In fa-
vorable transverse sections, such packets appeared
as honeycomb structures, whereas in longitudinal
section they appeared as parallel linear densities.
The dense matrix shows a vermiculate pattern (fig.
2).

ToNATiA SYLVicoLA— The secretory granules in
this species differed from those in T. bidens. The
granules consisted of light and dark material that
displayed two principal patterns. Either the dark
material was centrally placed, where it often con-
tained one or several lucent spaces, or the light
and dark material each occupied one hemisphere
of the granule. In either case, crystalloid tubules
of the sort found in T. bidens never were present
(fig. 2).

Trachops cirrhosus— In this species, the gran-
ules consisted of a large, central, very dense spher-
ule surrounded by a lucent cortex. The cortical
material contained linearly arranged punctate den-
sities that, at low to moderate magnifications, gave

the appearance of several continuous dense layers
in the outer portion of the granules (fig. 2).

Glossophaga soricina— The bulk of each
granule was occupied by homogeneously dense
material, which formed a smooth interface with
the remainder of the granule. The other portion
of each granule was pale and contained minute,
dense specks (fig. 3). The lighter material occa-
sionally intruded into the dense zone in the form
of a tortuous ribbon.

Leptonycteris sanborni— The secretory gran-
ules contained an amorphous, dense, central in-
clusion in a paler matrix. A small bundle of short
filaments was occasionally present in the matrix
(fig. 3).

Sturnira liuum— The secretory granules in this
species resembled those in Trachops cirrhosus.
They consisted of a large, central, dense core sur-
rounded by a rim of lucent material in which were
embedded distinct punctate densities often dis-
posed in layers (fig. 3).

Artibeus jamaicensis— The granules were large
and pale and contained some faintly discernible,
twisted fibrils. A few granules possessed a small,
dense, usually peripheral inclusion (fig. 3).

Ariteus FLA vescens— These secretory granules
had a pale matrix in which were numerous, flat,
narrow, moderately dense lamellae in a random
arrangement. Viewed on edge, the lamellae ap-
peared as dense lines; face on, as gray, irregular
structures (as seen in fig. 3). In a few granules,
there was a dense polygonal plate that lacked pe-
riodicity.

Eptesicus lynni— These granules had a com-
plex substructure. The limiting membrane was un-
derlaid by a layer of moderately dense material
that made incursions, arcuate and anfractuous, into
a dense matrix, producing a variety of patterns.
These extensions of the F>eripheral light material
had a layered structure (fig. 4, top).

Eptesicus brasiliensis— As in E. lynni. the lim-
iting membrane was subtended by layered mate-
rial, usually in several plies, that extended into the
dense matrix. These extensions were less tortuous
than those in E. lynni and appeared to subdivide
the granule interior (fig. 4, bottom).

Tadarida brasiliensis— The secretory granules
in this species exhibited a spectrum of patterns.
The most common was relatively simple, with
either short, flat, dense prisms or dense dots being
suspended in a slightly less dense matrix. A few
granules contained dense hollow spheres, whereas
others had a mazelike configuration based on light
and dark laminations (fig. 5, top).

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

219

Fig. 4. Higher magnification of mature secretory granules in parotid acinar cells in Eptesicus lynni (top) and E.
hrasiliensis (bottom). Note the subtle but consistent differences in the granule substructure. Eptesicus lynni, x 63,000;
E. brasiliensis, x 76,800.

220

HELDIANA: ZOOLOGY

Fig. 5. Parotid acinar cell granules in Tadarida brasiliensis (top) are compared to those found in Molossus molossus
(bottom). Note the range of variation in granule substructure in T. brasiliensis. Tadarida brasiliensis, x 26,000;
Molossus molossus, x 24,000.

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

221

MoLOSSUs MOLOSSUS— The variable morphol-
ogy of the granules in this species appeared to
depend on their stage of maturation. Early gran-
ules were small, with a moderately dense matrix
in which were some prominent dense particles. As
the granules matured (based on size, density, and
spatial relationship to the Golgi complex), they
enlarged and the particles decreased somewhat in
density. The mature granules had a farinaceous
matrix probably resulting from comminution of
the dense particles (fig. 5, bottom).

Discussion

The idea of using comparative ultrastructural
analysis of homologous, morphologically differ-
entiated cells to study evolutionary pathways and
to explore systematic relationships is new to the
study of mammalian orders, but has been used in
broader studies of vertebrates and invertebrates
(Eakin, 1968; Rieger & Tyler, 1979; PhiUips &
Tandler, 1987). Nevertheless, the potential value
of ultrastructural comparisons has been demon-
strated recently by studies of gastric mucosa, ret-
ina, and submandibular salivary glands (Phillips
et al., 1984; Feldman & Phillips, 1984; Tandler et
al., 1983, 1986; Phillips & Tandler, 1987). The
discovery of patterns of intersF>ecific differences in
cellular architecture and cellular secretory prod-
ucts is in keeping with previously successful use
of histology in evolutionary and systematic mam-
malogy (e.g.. Quay, 1954; Forman, 1972; Phillips
& Oxberry, 1972; Sands et al., 1977; Naumova,
1981; Hood & Smith, 1982, 1983).

Secretory cells, such as the parotid acinar cells
used in the present study, seem to hold special
promise for comparative investigation. Firstly, the
entire secretory process— from nuclear dna to syn-
thesis of proteins and complex carbohydrates and
packaging of materials into secretory granules and
their subsequent discharge— has been studied in-
tensively over the past several decades (e.g.,
Jamieson & Palade, 1971; Castle et al., 1975; Pa-
lade, 1975). Secondly, the secretory process seems
to be relatively conservative— in the sense that
basic pathways are the same in virtually all secre-
tory cells— and, therefore, interspecific compari-
sons are facilitated and interpretation is somewhat
simplified (Phillips & Tandler, 1987).

In the present study we limited our descriptive
comparisons to "mature" secretory granules. These
are the secretory granules that accumulate within
the apical cytoplasm of the acinar cell; this product

either is discharged from the cell into the acinar
lumen (thus becoming part of the formative saliva)
or, after an unknown storage interval, is broken
down through autophagy and recycled within the
cell. The glycoprotein components of the acinar
secretory product are elaborated by the rough en-
doplasmic reticulum acting in concert with the
Golgi complex and are perhaps the most interest-
ing feature from a comparative point of view. Syn-
thesis of these glycoproteins begins with transcrip-
tion of a very small segment of the genome into
mRNA. The exportable proteins of the secretory
granules could be regarded as providing a "win-
dow" on the genome because, although all somatic
cells have the same genome (Briggs & King, 1 952,
1957), only a small portion actually is operational
in any given fully differentiated cell. A protein
synthesized for export thus directly reflects a por-
tion of the operational genome. Any polysaccha-
ride components of the secretory granules prob-
ably are at least one more step removed from the
genome because synthesis of complex carbohy-
drates is enzymatically determined and usually
takes place within saccules in the Golgi complex
(Tandler, 1978). Even so, the polysaccharides also
have considerable potential for comparative anal-
ysis.

Production of secretory granules can be consid-
ered in terms of ontogeny. The first "granules"
typically are seen in direct proximity to the Golgi
complex, where carbohydrates are linked to pro-
teins and the membrane that will encase the gran-
ule is synthesized in a process involving both the
GERL and the Golgi complex itself (Tandler, 1978;
Hand, 1 980a; Hand & Oliver, 1 984). Newly formed
("immature") granules differ greatly from "ma-
ture" granules (see Castle et al., 1975) and were
not used by us in describing the product for each
of our species. Nevertheless, in view of species
differences found by us, it is interesting to ask
whether or not immature granules of one species
might resemble mature granules in another species.
Such similarities might be expected if hetero-
chronic differences accounted for differences in
mature product. However, because no such cross-
species similarities were found by us between
immature and mature secretory granules, hetero-
chrony does not seem to account for sp>ecies dif-
ferences, at least among the 1 5 bats examined here.

No two Neotropical sfiecies examined here ex-
hibited the same mature secretory granules in their
parotid gland acinar cells, although in a few cases
there was a degree of resemblance. This extreme
variability is easily the greatest ever reported for

222

HELDIANA: ZOOLOGY

homologous cells within an order or, as in the case
of the Phyllostomidae, within a family of mam-
mals (or any other vertebrates). To what can we
attribute this striking finding?

One consideration is fixation, which certainly
affects the appearance of any cellular feature as
viewed with transmission electron microscopy. In-
deed, it can be said that the appearance of cells
and their products essentially is the consequence
of their intrinsic chemistry combined with the
chemistry of the fixative at the moment in time
when fixation occurred. Different fixatives and tis-
sue processing can have profound effects on the
microscopic appearance of secretory granules in
salivary glands (Simson et al., 1978). However, we
used consistent processing techniques and two
similar fixatives. Our specimens of Ariteus and
Leptonycteris both were fixed with 2% glutaral-
dehyde, whereas all of our other specimens were
fixed with a trialdehyde-DMSO fixative (Kalt &
Tandler, 1971; Phillips, 1985). We were able to
eliminate fixation as a source of variation because
we also have examined specimens of Artibeus that
had been fixed in both solutions, coincidentally
with Leptonycteris, Ariteus, and all of the other
species examined (see A. phaeotis parotid in Phil-
lips et al., 1977).

Generally speaking, given consistent prepara-
tion techniques, microscopic differences in secre-
tory granule substructure can be ascribed to bio-
chemical differences among the granules.
Microscopically detectable sequestration of indi-
vidual types of macromolecules within secretory
granules has been demonstrated only rarely (Ra-
vazzola & Orci, 1980; Kousvelari et al., 1982) but
clearly is the best available explanation of intra-
granule substructure. Based on fundamental prin-
ciples of biochemistry, it thus can be concluded
that macromolecules packaged in the secretory
granules most likely sort themselves out according
to charge and steric effects, as well as chemical
interactions, to yield a characteristic pattern for
each species. However, it also should be noted that
a homogeneous appearance of intragranular sub-
stance does not in itself preclude sequestration of
different enzymes within the granule. Separate lo-
calization of different enzymes (a-amylase and
chymotrypsinogen A or a-amylase and trypsino-
gen) within pancreatic cell zymogen granules has
been demonstrated with a combination of hrp
(horseradish-p)eroxidase)-labeled and ferritin-la-
beled antibodies (Ono et al., 1980).

At present it is impossible to correlate exactly
secretory granule substructure with particular

chemical components such as certain enzymes or
mucosubstances, so from microscopic images alone
we cannot say precisely how the granules in our
species differ chemically from one another. How-
ever, some conclusions can be inferred from the
literature. For example, an electron-dense image
(see, for example, fig. 2), labeled classically as "se-
rous," can be associated with granules rich in en-
zymes. Such an image is typical in species such as
laboratory rodents and primates, for which some
data are available on the biochemistry of parotid
saliva (e.g., Jacobsen «Sc Hensten-Pettersen, 1974).
On the other hand, the presence of electron-dense
"serous" granules does not preclude the presence
of mucosubstances within the secretory granules.
Pinkstaff'et al. (1982) reported that, although the
parotid product in the little brown bat, Myotis
lucifugus, was "serous" with standard histological
techniques, both neutral and acidic mucosub-
stances could be demonstrated histochemically.
The parotid granules in Artibeus and Ariteus are
interesting in this regard because electron-dense
material is scarce (especially in Artibeus, fig. 3) and
enzyme production is extremely low, at least in
Artibeus (Junqueira et al., 1973). By way of con-
trast, Sturnira lilium has largely electron-dense
granules (fig. 3) and thus differs considerably from
the other two stenodermatines; Sturnira produces
saliva rich in enzymes (Junqueira et al., 1973).

Although the parotid granules in Artibeus (and,
by extension, Ariteus) are low in enzyme content,
to what can we attribute their tem image? This is
an intriguing question because Wimsatt (1956) re-
ported that the gland was negative for mucosub-
stances, whereas Radtke (1972) reported the pres-
ence of sialomucins in parotid acinar cells. In part,
this apparent disagreement is the result of differ-
ences in techniques that cannot be resolved by
transmission electron microscopy.

What can be determined about the parotid sal-
ivary glands in Neotropical bats that relates to
their diets, evolutionary history, or systematic re-
lationships? Clearly the ultrastructure of parotid
secretory granules has systematic significance, be-
cause no two genera (or species either) are exactly
alike. This finding is in keeping with our studies
of the submandibular gland in five species of Ar-
tibeus (Phillips et al., 1977; Tandler et al., 1983,
1986). The seromucous cells in this gland were
found to contain granules that allowed for three
groupings of Artibeus species {A. cinereus-A.
phaeotis, A. jamaicensis-A. lituratus, and A. con-
color) that matched the independently derived
genie data (from isozyme analysis) reported by

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

223

Koop and Baker ( 1 983). The differences in parotid
acinar product in Eptesicus lynni and E. brasi-
liensis (fig. 4) also are interesting from this per-
spective because independent genie data from 1 9
presumptive loci show that E. lynni probably orig-
inated from the E. fuscus species complex, sepa-
rate from the origin of E. brasiliensis (Arnold et
al., 1980). Eptesicus lynni was found to share only
62% of analyzed alleles with E. brasiliensis.

It might seem surprising that the microscopic
images of salivary gland secretory products are
generically (and often specifically) distinctive, but
variation of such a fine resolution actually is in
keeping with numerous genetic studies of mam-
malian saliva. Several genetic markers have been
found in human saliva (Azen & Opp)enheim, 1 973;
Ashton & Balakrishinan, 1974; Tan & Ashton,
1976), and salivary proteins in particular tend to
be polymorphic (Azen, 1972, 1973). Additionally,
both sex and strain differences in salivary proteins
have been reported in laboratory mice (Ikemoto
& Matsushima, 1984). Although none of these dif-
ferences has been demonstrated microscopically
(all are based on biochemical analysis alone), their
occurrence nevertheless is significant to compar-
isons among mammalian species.

Although the sometimes subtle but consistent
microscopic differences between species within the
genera Phyllostomus, Tonatia, and Eptesicus (figs.
2, 4) are in keeping with our hypothesis of the
sensitivity of secretory product to genie differ-
ences, they cannot readily be related to any known
ecological differences between species within each
genus. It is altogether possible that such species
differences do not represent direct evolutionary
selection. Instead, the differing images might rep-
resent species-specific protein polymorphisms of
a type that would not significantly affect "perfor-
mance" of the saliva even though differences in
primary molecular structure or surface charges (or
both) could be indirectly detected by transmission
electron microscopy (Phillips & Tandler, 1987).
The existence of such "nonfunctional" (and pre-
sumably nonselected, i.e., "neutral") interspecific
variation in a protein molecule has been demon-
strated previously in the otherwise conservative
hemoglobin molecule (Perutz, 1983). In this ex-
ample the tertiary and quaternary structures ap-
parently are conserved regardless of large numbers
of functionally neutral amino acid substitutions in
the primary structure. Although these substitu-
tions in themselves most often have no effect on
the functional capacity of the molecule, they
nevertheless are known sometimes to change mo-

lecular surface charges (Perutz, 1983). For the
present we postulate that similar nonselected, non-
functional variations could account for the micro-
scopically detectable intrageneric differences in
salivary proteins in Tonatia bidens and T. sylvi-
cola, Phyllostomus latifolius and P. elongatus, and
Eptesicus lynni and E. brasiliensis. This explana-
tion seems more parsimonious than the alterna-
tive, which would be to assume that the acinar cell
component of the parotid saliva is "functionally"
different in closely related, ecologically similar
species. The extent or degree of microscopic dif-
ferences in the mature acinar cell product ulti-
mately might tell us more about relative times of
divergence than about ecological differences among
congeneric species of bats.

Microscopic comparisons of salivary gland se-
cretory products could be valuable to cladistic
studies of chiropteran families, at least to judge
from our data. The phyllostomines generally are
regarded as the most primitive (least derived) of
the Phyllostomidae (Smith, 1976), and their par-
otid acinar products thus would qualify as plesio-
morphic (following Henning, 1 966). The products
in all five species examined (Phyllostomus elon-
gatus, P. latifolius, Tonatia bidens, T. sylvicola,
and Trachops cirrhosus) contained large amounts
of electron-dense material (fig. 2), as did the se-
cretory product in Pteronotus parnellii (a closely
related mormoopid), Eptesicus lynni and E. bra-
siliensis (Vespertilionidae), and Tadarida brasi-
liensis and Molossus molossus (Molossidae), all of
which serve as "outgroups."

If the electron-dense, enzyme-rich secretory
granules are regarded as plesiomorphic in micro-
chiropteran bats, then it would be reasonable to
regard a granule with less electron-dense material
as apomorphic (derived). The glossophagine gen-
era examined (Glossophaga and Leptonycteris) are
representative of a phyllostomid evolutionary trend
in which dentition, tongues, and associated mus-
culature were modified for feeding on fruit, pollen,
and nectar (Park & Hall, 1951; Phillips, 1971;
Greenbaum & Phillips, 1974; Griffiths, 1982, 1983;
Smith 8l Hood, 1 984). In these genera the electron-
dense component has been reduced in comparison
to the phyllostomines. Carollia perspicillata, which
is omnivorous, fits into this category in that the
parotid secretory granules contain relatively little
electron-dense material. In this bat the secretory
granules are very distinctive because the electron-
dense inclusions are often in the form of cagelike
geodesic structures (Phillips & Tandler, 1987;
Tandler et al., in press).

224

FIELDIANA: ZOOLOGY

Perhaps the most interesting systematic finding
in our data lies within the nominal subfamily
Stenodermatinae. These bats represent an evolu-
tionary trend toward frugivory that includes ex-
treme gastric adaptation at the gross, histological,
histochemical, and cellular levels (Forman, 1972;
Forman et al., 1979; Phillips & Studholme, 1982;
Phillips et al., 1984). The pale parotid secretory
granules in Artibeus and Ariteus are synapomor-
phous, whereas the parotid granules in Sturnira
are electron-dense, enzyme-rich, and more nearly
like the plesiomorphic granules of the phyllosto-
mines, in particular Trachops cirrhosus (figs. 2-3).
This example is interesting because Sturnira al-
ways has been something of an enigma. Although
de la Torre (1961) allied this genus with Vampy-
rops-\\\it stenodermatines and Smith (1976) in-
cluded it with the "long-faced" stenodermatines,
others (e.g., Walton & Walton, 1968) previously
had placed the genus in a separate subfamily (Stur-
nirinae) based on a variety of morphological fea-
tures that seemed inconsistent with the other, more
traditional, stenodermatines such as Artibeus. In-
deed, Slaughter (1970) pointed out that Sturnira
has some distinctive dental features that possibly
link the genus to the glossophagines. The differ-
ences in the parotid secretory granules thus are in
keeping with a variety of other phenotypic differ-
ences. While it is reasonable to suggest that secre-
tory granule differences of this magnitude repre-
sent a major interspecific difference in the
operational segment of the genomes of homolo-
gous parotid acinar cells, the eventual systematic
value of such data will await availability of data
about still other phyllostomid genera.

Although our comparative data clearly docu-
ment great microscopically detectable plasticity in
the secretory product of parotid acinar cells, many
questions about their evolution remain unan-
swered. For example, what has been the role of
diet? At first glance, our data suggest that insec-
tivorous and animalivorous species have enzyme-
rich, electron-dense granules, whereas frugivores
have enzyme-poor, pale secretory granules. Yet,
what about Sturnira lilium, which certainly in-
cludes large amounts of fruit in its diet (Gardner,
1977)?

At least three factors interfere with any effort to
correlate parotid acinar cell product with diet. First,
acinar cells are but one cell type among several
that influence the biochemistry of parotid saliva.
Second, and possibly more important, the par-
otid is but one salivary gland among a host of
glands (the submandibular and sublingual and mi-

nor glands) that are located throughout the oral
region and that contribute substances to the saliva.
Some data suggest that different glands in different
species might have been more responsive, in an
evolutionary sense, to changes in diet. For ex-
ample, it is the accessory submandibular glands
of Trachops and Megaderma that are unique in
histology and might correlate with feeding on frogs
(Phillips &. Tandler, 1985; Phillips et al., 1987);
the parotid in Trachops is similar to that of other
phyllostomines.

A third factor, for which we presently have no
data, is the possibility that parotid acinar cells can
respond in some way to diet at the individual level.
Our sample sizes are large enough to convince us
of the near uniformity of secretory granule mor-
phology within a population. However, Schick et
al. (1984) have recently published the first report
of a secretory cell responding to dietary intake by
a shift in enzyme production. This is the first such
detailed molecular data known to us and, although
pancreatic acinar cells in laboratory rats were the
source of the data, the implications for compar-
ative studies of salivary glands are worth noting.

A final factor for consideration is the complex
role that salivary glands play in the lives of mam-
mals. Salivary glands in Artibeus, Chiroderma, and
Ametrida might contribute to gastric cytoprotec-
tion (Studier et al., 1983; Phillips et al., 1984). In
Tonatia sylvicola an unusual organelle found in
submandibular seromucous cells of males might
be related to species isolation or sex recognition,
or both (Nagato et al., 1984). If these examples
are typical for bats, then digestion is but one of
several major functions of salivary glands.

In conclusion, this first systematic microscopic
survey of a secretory cell product has demonstrat-
ed a previously unknown, extraordinary degree of
variation within a group of related sjjecies. Based
on our findings, one might conclude that Neo-
tropical microchiropteran bats will serve as a sig-
nificant model for study of how secretory cells
have evolved in mammals; meanwhile, data from
comparative investigations will contribute to our
knowledge of genie relationships among these an-
imals.

Acknowledgments

Financial support for field and laboratory re-
search that led to the data reported here came from
a variety of sources, which we are pleased to ac-
knowledge: Research Corporation Grant C-1251

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

225

(to Phillips); NIH Grant DE 03455-02 (to Phil-
lips); NSF Grant CDP-80 1-8653 (to Phillips);
Hofstra University HCLAS Executive Committee
grants (to Phillips); NIH Grant AM-08305 (to
Tandler); NIH Grant RO DE 07648-01 Al (to
Tandler & Phillips); the Alcoa Foundation (through
Dr. Hugh H. Genoways, Carnegie Museum of Nat-
ural History); and the Graham Netting Research
Fund via a grant from the Cordelia S. May Char-
itable Trust (through Genoways). In addition to
recognizing our financial support, we also wish to
thank several colleagues whose field companion-
ship in such places as Suriname and Jamaica, hard
work, and willingness to share time and ideas in
their laboratories played a major part in our study.
These are: Hugh H. Genoways, Carnegie Museum
of Natural History; Gary W. Grimes and Dorothy
E. Pumo, Hofstra University; Robert J. Baker,
Texas Tech University; and Henry A. Reichart,
formerly of STINASU, in Suriname. Field assis-
tance was provided by K. M. Studholme, S. L.
Williams, N. M. Sposito, J. Groen, R. L. Honey-
cutt, B. Koop, M. Arnold, B. A. Oxberry, P. Bil-
leter, J. Bickham, and J. Patton. We also acknowl-
edge technical assistance of Carol Ayala, Case
Western Reserve University, and typing by Linda
Cossen, Special Secretarial Services, Hofstra Uni-
versity. Artwork and lettering were done by Helen
Tandler. All specimens were collected and im-
ported under proper permits issued by the relevant
authorities in Mexico, Jamaica, Suriname, and the
United States.

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228

FIELDIANA: ZOOLOGY

Appendix

Specimens Examined

Voucher specimens of each species and each
locality are deposited in the collections of the Car-
negie Museum of Natural History and The Mu-
seum, Texas Tech University. All specimens ex-
cept those of Leptonycteris sanborni (Jalisco,
Mexico) and Ariteus flavescens and Eptesicus lynni

(Jamaica) were collected in Brokopondo Province,
Suriname.

Pteronotus parnellii, 10 (5 S6, 5 29); Tonatia bi-
dens, 3 {2 SS, 1 2); T. sylvicola, 2 {66); Trachops
cirrhosus, 2 (1 3, 1 2); Phyllostomus elongatus, 3
(\6,2 22); P. latifolius, 1 (2); Glossophaga soricina,
3 (2 $6, 1 2); Leptonycteris sanborni, 2 (3<5); Sturnira
lilium, 5 (2 35, 3 22); Artibens jamaicensis, 3 (22);
Ariteus flavescens, 1 (2); Eptesicus lynni, 2 {$$); E.
brasiliensis, 3 (22); Tadarida brasiliensis, 1 (S); Mo-
lossus molossus, 8 (4 66, 4 22).

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS

229

Distribution of the Species and Subspecies of
Cebids in Venezuela

Roberta Bodini and Roger Perez-Hernandez

ABSTRACTS

Thirteen species of Primates representing nine genera in the family Cebidae are found in
Venezuela. The geographic distribution of these species exhibits four main patterns. Alouatta
and Cebus are widely distributed in all parts of the country. Four genera, Saimiri, Callicebus,
Cacajao, and Chiropotes are restricted to south-central Venezuela; their distribution is centered
in the Amazonian lowlands. Aotus and Ateles are each represented by one species or subspecies
in south-central Venezuela and another in northwestern Venezuela. Pithecia is mainly restricted
to the Guianan highlands in eastern Venezuela, with a single known outlying locality in south-
central Venezuela. Of the nine cebid genera in Venezuela, all occur in south-central Venezuela,
four occur in northwestern and north-central Venezuela, and only three occur in eastern Ven-
ezuela. Distribution maps for the 1 3 species in Venezuela are presented with exact localities of
specimens.

Nueve generos de cebidos representados por trece especies se hallan en Venezuela. La re-
particion geografica de las especies se resuelve en cuatro patrones geograficos principales. De
este modo, Alouatta y Cebus son extensamente repartido por todo el pais. La distribucion de
los generos, Saimiri, Callicebus, Cacajao y Chiropotes esta restringida al sur-central de Vene-
zuela con concentracion en las tierras bajas Amazonicas. Aotus y Ateles son representatados,
cada cual, por una especie (o subespecie), en el sur-central, y otra especie in el noroeste del
pais. Pithecia se reparte en las alturas guayanas venezolanas del este, y es conocido, a la vez,
por un solo dato de captura del sur-central venezolano. En resumen, la totalidad de los nueve
generos esta presente en el sur-central de Venezuela, cuatro de ellos en el noroeste y norte-
central, y tres en la Venezuela oriental. La reparticion de las trece especies esta documentado
por mapas.

Treze especies de Primatas, representando nove generos da familia Cebidae, sao encontradas
na Venezuela. A distribuifao geografica destas especies exibem quatro padroes principais. Al-
ouatta e Cebus espalham-se por todas partes do pais. Quatro generos, Saimiri, Callicebus,
Cacajao, e Chiropotes, limitam-se ao centro-sul de Venezuela; a Bacia Amazonica sendo seus
focos de distribuifoes. Aotus e Ateles sao representados por uma especie ou subespecie cada
no centro-sul da Venezuela, e por outra especie no noroeste do pais. Pithecia limita-se ao
planalto Guianense, no leste da Venezuela, com apenas uma localidade conhecida no centro-
sul do Pais. Em suma, todos nove generos de Cebidae ocorrem no centro-sul da Venezuela,
quatro ocorrem no centro-norte e noroeste do Pais, e apenas tres ocorrem no regiao leste da
Venezuela. Apresentam-se mapas de distribui96es das 13 especies na Venezuela, com locali-
dades exatas dos especimes colecionados.

From the Institute de Zoologia Tropical, Universidad
Central de Venezuela, Apartado 47058, Los Chaguara-
mos, Caracas 1041 -A, Venezuela.

BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 231

Introduction

Primates of the family Cebidae are some of the
most conspicuous mammals in the Neolropics, as
they are diurnal, often forage in large troops, and
may be quite vocal. However, we actually know
very little about the exact distributions of most
species of cebids. In Venezuela, several authors
have discussed cebids as part of generic revisions
(see Elliot, 1912; Hershkovitz, 1949, and later
works; HiU, 1960, 1 962; Kellogg & Goldman, 1944)
and in works on the status or collections of specific
species (i.e., Bodini, 1983; Handley, 1976; Mon-
dolfi & Eisenberg, 1979; Rudran & Eisenberg,
1982).

Herein we summarize the distribution of cebids
in Venezuela based on recent collections, speci-
men records from several museums, and the lit-
erature. Specimens reported on are housed in the
following collections: Estacion Biologica Rancho
Grande (EBRG); Museo de Biologia, Universidad
Central de Venezuela (MBUCV); Museo de Cien-
cias Naturales (MCN); and Museo de Historia
Natural La Salle (MHNLS). Our objectives are to
provide accurate locality records and distribution
maps that may be utilized in future investigations.

Generic Distributions

The nine genera of cebids that inhabit Venezuela
exhibit four main patterns of geographic distri-
bution (figs. 1-7). (1) Two geneT^—Alouatta and
Cebus— arc widely distributed in all paris of the
coimtry. (2) Four genera— Sa/m/r/, Callicebus,
Cacajao, and Chiropotes—UTe restricted to south-
central Venezuela, centering on the Amazonian
lowlands (see Eisenberg & Redford, 1 979). (3) Two
genera— .4orw5 and Ateles—are each represented
by one species or subspecies in south-central Ven-
ezuela and another in northwestern Venezuela or
northwestern and north-central Venezuela, which
is the region of the northeastern spurs of the An-
dean chain and the enclosed Maracaibo basin. (4)
One genus— Pithecia— is mainly restricted to the
Guianan highlands in eastern Venezuela, with a
single known outlying locality in south-central
Venezuela. In summary, of the nine cebid genera
in Venezuela, all occur in south-central Venezuela
(including the outlier record of Pithecia), four oc-
cur in northwestern and north-central Venezuela,
and only three occur in eastern Venezuela.

Specific and Subspecific Distributions
Saimiri

In a recent revision of the squirrel monkeys,
Hershkovitz (1984) recognized four species: Sai-
miri boliviensis, S. oerstedi. S. sciureus, and S.
ustus. All those squirrel monkeys found in north-
em South America he referred to the single species
S. sciureus (fig. 1 ). Squirrel monkeys from Ama-
zonian Venezuela and adjacent Brazil and Colom-
bia he regarded as 5. sciureus cassiquiarensis.
Hershkovitz (1984) plotted several localities in
Territorio Federal Amazonas, but listed only a
single specific locality: "Casiquiare, Rio (mouth),
2°01'N, 67"X)7'W." Hill (1960) previously had
mapped the distribution of 5. sciureus as occurring
throughout all of Venezuela, although he listed
only a single locality in the state of Bolivar (Camp
Canaracuni, 4''36'N, 64''10'W). The name cassi-
quiarensis Lesson is based on Humboldt's descrip-
tion of a captive female from the banks of the Rio
Casiquiare, Amazonas, Venezuela.

Specimens Examined— Total 29. Bolivar: Ca-
naracuni (4°36'N, 64«'10'W). Territorio Federal
Amazonas: Campo Cacuri (4°49'N, 65°26'W); Cano
Yureba, Rio Ventuari (3''35'N, 66°46'W); Rio Pu-
runame, 40 km from union with Rio Orinoco
(3°19'N, 65°15'W); Rio Ventuari (3°59'N,
67'^2'W); San Fernando de Atabapo (4°02'N,
67»37'W); San Juan de Manapiare (5°14'N,
66'^2'W).

Aotus

Night monkeys are found from Panama
throughout much of Amazonian South America
to Paraguay. Historically, it has generally been as-
sumed that all night monkeys represented the sin-
gle species Aotus trivirgatus. However, in a recent
revision of the genus, Hershkovitz (1983) recog-
nized nine allopatric species. A tenth species, Aotus
hershkovitzi from Colombia, has recently been
proposed by Ramirez-Cerquera (1983). Two
species of Aotus, A. lemurinus and A. trivirgatus.
have been reported from Venezuela (fig. 2).

Aotus lemurinus griseimembra Elliot is known
in Venezuela only fi-om the extreme northwestern
region, the states of Cojedes and Merida (Hersh-
kovitz, 1983), and the states of Tachira, Trujillo,
and Zulia (Handley, 1976).

Specimens Examined— Total 10. Zulia: Campo
a Rosario, EHstrito Catatumbo (l''44'N, 67°03'W);

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Reserva San Manuel, Los Canaguatos (9*t)4'N,
71''56'W); Kunana, Rio Negro, Sierra de Perija
(9*75'N, 72*25'W); Rio Tocuco (72''25'W).

Aotus trivirgatus trivirgatus Humboldt is known
from the region south of the Rio Orinoco in Ter-
ritorio Federal Amazonas and the state of Bolivar.
The Venezuelan national collections contain
specimens from Cano Manapiare and Rio Anta-
vari, and two literature records report specimens
from the right bank of the Rio Caroni (INP-
ARQUES, 1982;MARNR-DGS-POA-SFS, 1982).

Our recent discovery of Aotus trivirgatus in
Guyana to the east of the Rio Caura, well outside
of the Rio Orinoco basin, led to an investigation
of geographic variation in the species. Preliminary
results suggest that these A. trivirgatus represent a
new geographic race (Bodini & Ferreira, in prep.).

Specimens Examined— Total 1 1. Bolivar: Cano
Manapiare (7»1 1'N, 66°40'W); Rio Antavari
(S'^O'N, 63°10'W). Territorio Federal Amazonas:
Alto Manapiare (S'^S'N, 66°02'W); San Juan de
Manapiare (S^M'N, 66°02'W); near Isla Cigarron,
Rio Negro (r44'N, 67°03'W); recently collected
in La Neblina, east of Rio Varia (0°59'N, 66°10'W).

Callicebus

The genus Callicebus is the sole representative
of the subfamily Callicebinae and is found from
northern South America south to northern Para-
guay. Hershkovitz (1963) and Kinzey (1982) rec-
ognized three species, Callicebus moloch, C per-
sonatus, and C. torquatus, of which only C.
torquatus is found in Venezuela.

Callicebus torquatus lugens Humboldt is known
in Venezuela south of the Rio Orinoco (fig. 3). Hill
(1960) originally suggested that C. torquatus is
found as far east as Guyana, although this was
questioned by both Handley (1976) and Hersh-
kovitz (1963), who reported Venezuelan speci-
mens only from Territorio Federal Amazonas.
Handley (1976) reported 31 specimens from
southern Territorio Federal Amazonas. Bodini
(1981) reported specimens from Maripa, Camp
Canaracuni, and the Rio Antavari in Venezuela.
Recent reports by Kinzey ( 1 982) and Bodini (1981)
confirm the presence of C torquatus in Guiana
Region.

Specimens Examined— Total 26. Bolivar: Cana-
racuni (4'36'N, 64°10'W); Maripa, 150 km from
Ciudad Bolivar (7*22'N, 65'^9'W); Rio Antavari
(5»20'N, 63M0'W); Alto Paragua (4°30'N,
63*WW). Territorio Federal Amazonas: Alto Cano
Caname, Departamento Atabapo (3*'33'N,

67"06'W); Alto Ventuari (3°50'N, 67°04'W); Boca
Cano Maica, Rio Ventuari (66°30'W); Cano Ya-
gua, Cerro Cucurito, Departamento Atabapo
(3''38'N, 66»25'W); Cacuri, Alto Ventuari (4°49'N,
65"26'W); La Esmeralda (3"X)8'N, 65°32'W); Rio
Cunucunuma (3°10'N, 66°0rW); Rio Puruname
(3°19'N, 65''15'W); La Neblina, east of Rio Varia
(0°59'N, 66°10'W).

Alonatta

Alouatta is the only genus within the subfamily
Alouattinae and is represented by some six species
widely distributed in Central and South America.

A single species, Alouatta seniculus, is abundant
and widely distributed in Venezuela. Hill (1962)
recognized three subspecies of A. seniculus as oc-
curring in Venezuela. Alouatta seniculus seniculus
Linnaeus is foimd in extreme northwestern Ven-
ezuela, primarily in the states of Apure, Tachira,
and Zulia, with a single record from Barinas (Ti-
coporo Forest, 8'X)6'N, 70*'40'W). Alouatta seni-
culus arctoidea Cabrera, called the "Caracas howler
monkey" by Humboldt, inhabits all the coastal
region from Falcon to the state of Miranda. Ca-
brera (1 958) proposed Caracas as the type locality.
A third subspecies, A. seniculus stramineus Hum-
boldt, inhabits all of Venezuela south of the Rio
Orinoco in Territorio Federal Amazonas and the
state of Bolivar (fig. 4).

In addition to these three subspecies, we believe
a fourth, previously unrecognized, form exists and
is widely distributed throughout the Venezuelan
llanos. This undescribed subspecies is character-
ized by coloration and size. The southern limit of
its distribution is clearly defined by the Rio Ori-
noco, but its northern and western limits in the
Andean piedmont are as yet undetermined.

Howler monkeys are extremely adaptable to a
wide array of environments, and we believe they
are found throughout Venezuela. Their apparent
absence in certain regions probably reflects lack of
collecting rather than true distributional gaps, as
our records demonstrate for the state of Anzoa-
tegui. The problem of current and historical dis-
tribution and systematic relationships of the var-
ious populations in Venezuela is in need of study.

Specimens Examined— Total 80. Anzoategui:
Los Cocos, Rio Caris (8''30'N, 64'^5'W). Apure:
Caiio San Agustin, Selva de San Camilo (7"'19'N,
7r57'W); Hato El Frio (7°44'N, 68°54'W); Las
Raicitas, El Saman (7°55'N, 68°40'W). Aragua:
Asentamiento Los Castillos, Turagua (10°09'N,
67»3rW); Cumbre de Guacamaya (10°2rN,
67'40'W); Los Picachos, Rancho Grande ( 1 0*2 1'N,

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6T4l'W). Barinas: Reserva Forestal Ticoporo
(8°09'N, 70°40'W). Bolivar: Guayoba, Rio Caura,
4 km from Maripa (7°20'N, 65°10'W); Hato Bella
Vista, El Palmar (8°00'N, 62°00'W); Represa del
Guri, Operacion Rescate (7°40'N, 63°00'W); San
Martin de Turumban, Anacoco-Cuyuni (6°42'N,
61°02'W). Carabobo: Las Quiguas, San Esteban
(10''25'N, eS^OrW). Cojedes: Hato Barbasco
(9°04'N, 68°08'W); Hato Itabana, 38 km from Las
Vegas (9°17'N, 68*'13'W); Las Queseras, El Baul
(8'*25'N, 68°17'W); Montaria Las Lomas, San Car-
los (9°38'N, 68°34'W). Distrito Federal: Hacienda
El Limon (10°28'N, 67''1 7'W). Falcon: Sanare, De-
partamento Silva (10°23'N, 68°25'W); Sierra San
Luis (1 1°15'N, 69°10'W). Guarico: Corozo Pando
(8°14'N, 67°17'W); Hato Mapurite, 40 km N of
Calabozo (9°17'N, 67°24'W); Manapire, near San
Antonio (9°1 7'N, 66''1 1 'W); Rio Tiznado (8°1 8'N,
67°48'W). Lara: Cumbre de Las Trojas, 45 km S
of Cabudare (9°45'N, 69°07'W). Miranda: Rio Ne-
gro (10°20'N, ee'lS'W); La Guzmanera, Guatopo
(10'X)0'N, 66°15'W). Portuguesa: Agua Blanca
(9°40'N, 69°07'W). Tachira: Cerro El Teteo, Bura-
gua (7°30'N, 7 r57'W); La Fria (8°1 3'N, 72°14'W);
Paramo Tama (7°27'N, 72°26'W). Territorio Fed-
eral Amazonas: Cacuri, Alto Ventuari (4°49'N,
65°26'W); Caiio Yureba, Ventuari (3°35'N,
66°46'W); Rio Hacha, Alto Ventuari (3°47'N,
65''38'W); Rio Puriname, 40 km from confluence
of Rio Orinoco (3°19'N, 65°15'W); San Juan de
Manapiare (9°05'N, 66°02'W); Cerro La Neblina,
E of Rio Varia (0°59'N, 66°10'W). Territorio Fed-
eral Delta Amacuro: Caiio Caneima (9°05'N,
60°55'W); Guiniquina (9°10'N, 61°06'W). Zulia:
Laguna de Manaties, Departamento Catatumbo
(9°27'N, 72°02'W); La Victoria, Rio Negro (9°36'N,
72°1 5'W); Rio Guasare ( 1 1°02'N, 72°05'W); Sierra
de Perija (9°00'N, 72°00'W).

Chiropotes

The bearded sakis comprise two species limited
to northern South America (Cabrera, 1958; Hersh-
kovitz, 1977;Mittermeier«Sc.Coimbra-Filho, 1981).
One species is found in southern Venezuela rep-
resented by a single subspecies, Chiropotes satanas
chiropotes Humboldt (fig. 5). All of our records
and those reported by Handley (1976) are from
Territorio Federal Amazonas. Cruz Lima (1945)
and Rudran and Eisenberg (1982) proposed the
occurrence of this species in the state of Bolivar
on the basis of Humboldt's description; Mondolfi
(1976) reported observations of Chiropotes at

Maripa and Caiio Maniapure (Bolivar), but we are
aware of no specimens from this region.

Specimens Examined— Total 29. Territorio
Federal Amazonas: Cacuri, Alto Ventuari (4°49'N,
65''26'W); Caiio Yureba, Departamento Atabapo
(3°3rN, 66''44'W); Caiio Yagua, Cerro Cucurito
(3''3 1 'N, 66°44' W); Laguna de Chiripo, Caiio Blan-
co (3°27'N, 66°40'W); Rio Ocamo, Alto Orinoco
(2°44'N, 65°11'W); Puerto Ayacucho (5°36'N,
67''35'W); Rio Orinoco, S of San Fernando de Ata-
bapo (4'^0'N, 67°38'W); San Fernando de Ata-
bapo (4°02'N, 67°37'W); San Juan de Manapiare
(5°14'N, 66'^2'W).

Cacajao

The genus Cacajao, or uakaris, contains two
species which are found in northern South Amer-
ica (Cabrera, 1958; Hershkovitz, 1972; Mit'ter-
meier & Coimbra-Filho, 1981). One species, Ca-
cajao melanocephalus, is found in Venezuela
restricted to the upper Orinoco region of southern
Territorio Federal Amazonas (fig. 5). Although few
specimens exist in collections, they appear to be
abundant.

Specimens Examined— Total 2. Territorio Fed-
eral Amazonas: Alto Cano Atacavi, Departamento
Casiquiare (3°05'N, 67'>02'W); La Nebhna, E of
Rio Varia (0°59'N, 66°10'W).

Cebus

The capuchin monkeys are found from Hon-
duras south through Central America and the
northern two-thirds of South America. Four species
currently are recognized (Cabrera, 1958; Hersh-
kovitz, 1972). Three species of Cebus are repre-
sented in Venezuela: Cebus albifrons, C apella,
and C. «/^rm//a/z^s (Hershkovitz, 1949, 1958).

Three subspecies of Cebus albifrons are found
in Venezuela (fig. 6). Cebus albifrons adustus
Hershkovitz was described on the basis of three
specimens from "near head of Rio Cogollo (Apon)
at eastern base of Sierra de Perija, about 5 kilo-
meters northwest of Machiques, Lake Maracaibo
region, Zulia" (Hershkovitz, 1949, p. 369). We
report additional specimens from Rio Guasare and
Kasmera. This subspecies is restricted to the Sierra
de Perija of extreme northwestern Venezuela and
adjacent Colombia. Cebus albifrons leucocephalus
Gray is found in extreme western Venezuela, in
the region south of the Lago de Maracaibo basin.

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and the states of Apure, Merida, Tachira, and Zu-
lia. Cebus albifrons unicolor Spix is found in ex-
treme southern Venezuela in Territorio Federal
Amazonas. Hershkovitz (1949) reported speci-
mens from Marimonda, Rio Orinoco, and from
Solano, Rio Casiquiare. Handley (1976) reported
specimens from Rio Mavaca and Tamatama. All
locality records for this subspecies in Venezuela
are south of the Rio Ventuari.

Specimens Examined— Total 15. Apure: Caiio
San Augustin, Selva de San Camilo (7°19'N,
7r57'W). Merida: Palmichoso, S of Las Virtudes
(9'^9'N, 70°57'W). Tachira: La Fria (8°13'N,
72°14'W). Territorio Federal Amazonas: Cano
Yagua, Cerro Cucurito, Departamento Atabapo
(B^SS'N, 66''25'W); near Boca Padamo, left side of
Rio Orinoco (3°02'N, 65°13'W). Zulia: Kasmera,
Perija (10°05'N, 72°45'W); Kunana (9°36'N,
72°15'W); Rio Bravo, Distrito Catatumbo (9°05'N,
72°22'W); Rio Guasare (1 1°02'N, 72°05'W).

Cebus apella is represented in Venezuela by two
subspecies (fig. 6). Cebus apella apella Linnaeus is
restricted to Amazonian Venezuela, the Territorio
Federal Amazonas, and is found along both banks
of the upper Orinoco. Cebus apella margaritae
HoUister is endemic to and restricted to Margarita
Island. The 800-km gap between the ranges of
these two subsp>ecies is striking and unexplained.

Specimens Examined— Total 10. Nueva Espar-
ta: Sierra de Copey (1 1°03'N, 63°56'W). Territorio
Federal Amazonas: Alto Caiio Caname (3''22'N,
67°08'W); Caiio Yapacana (3°30'N, 66°45'W); San
Fernando de Atabapo (4°02'N, 67°37'W).

Cebus nigrivittatus is widely distributed in Ven-
ezuela and represented by perhaps five subspecies
(fig. 6) (Hershkovitz, 1949; Cabrera, 1958). Cebus
nigrivittatus apiculatus was described by Elliot
(1912) on the basis of specimens from La Union,
Rio Caura, near its confluence with the Rio Ori-
noco. It is distributed throughout central Vene-
zuela south of the Orinoco between the Rio Caroni
and the Rio Ventuari.

Cebus nigrivittatus brunneus was described by
J. A. Allen (1914) from specimens from Aroa, a
station on the Bolivar Railway, Yaracuy, north-
western Venezuela. Hershkovitz (1949) reported
an additional specimen from the Paria Peninsula.
These records plus our specimens suggest that C.
n. brunneus is continuously distributed through-
out the Cordillera de la Costa of extreme northern
Venezuela. Cebus nigrivittatus nigrivittatus Wag-
ner is restricted in Venezuela to the Amazonian
region of Territorio Federal Amazonas. Cebus ni-
grivittatus olivaceus Schomburg is found in south-
eastern Venezuela. Hershkovitz (1949, p. 348) re-

ports the type locality as "Vicinity of 'Our Village,'
said to be situated at latitude 4°57'N., 61°rW.,
altitude 3,100 feet above sea level, southern foot
of Mount Roraima."

We believe that an undescribed subspecies, Ce-
bus nigrivittatus subsp., is widely distributed
throughout central and northern Venezuela north
of the Orinoco. The status of this population is
currently under study.

The Orinoco Delta region, which Eisenberg and
Redford ( 1 979) excluded from their consideration
of biogeographic regions due to insufficient data,
is now shown by the distributions of Alouatta,
Cebus, and Pithecia to be clearly aligned with the
Guyana highlands. The Llanos region now extends
up to the western edge of the delta, but does not
include it, as demonstrated by the distributions of
Alouatta and Cebus.

Specimens Examined— Total 75. Anzoategui:
Los Cocos, Rio Caris (8°30'N, 64°05'W); 10 km
W of Laguna de Unare (10°02'N, 65°12'W); Mor-
ichal Largo [between Anzoategi and Monagas]
(8°18'N, 63°15'W). Aragua: Rancho Grande
(10°10'N, 67°19'W). Barinas: Reserva ForestalTi-
coporo, on Barinas-Pedregal road (8°03'N,
70°18'W). Bolivar: Caiio La Urbina (7°15'N,
66°25'W); Carretera Caicara-S. Juan de Mana-
piare, km 1 75 (6°02'N, 66°29'W); Carretera El Do-
rado-Santa Elena, km 33 (6°12'N, 6ri4'W); Cur-
aima. El Palmar (8°01'N, 61°26'W); El Dorado-
Santa Elena, km 121 (5°18'N,6ril'W);Guayopo,
Rio Caura, 14 km from Maripa (7°09'N, 65°10'W);
Canaracuni (4°17'N, 64°05'W); Guri, Operacion
Rescate (7°18'N, 63°00'W); Rio Antavari (5'i09'N,
63°05'W); Rio Marajano, Meseta de Jaua (4°08'N,
64°1 1'W); Rio Villacoa, 4 km N of mouth (6°16'N,
67'^5'W); San Martin de Turumban, Rio Cuyuni
(6"'19'N, 61'^9'W). Carabobo: Bahia de Patanemo
(10"'12'N, 67''26'W); Urama (10°12'N, 68°08'W).
Cojedes: Cerro Azul, La Blanquera (8°26'N,
68°07'W); Montana Las Loma, San Carlos (9°1 7'N,
68°16'W); Pica, Las Vegas (9°1 5'N, 68°1 7'W). Dis-
trito Federal: El Avila, Caracas (10° 14'N, 66°13'W);
Hacienda El Limon (lO^B'N, 67°08'W). Falcon:
Sanare, Distrito Silva (8''23'N, 68°12'W). Guarico:
Hato Flores Moradas, Calabozo (8°23'N, 67"! 3'W);
Hato Mapurite, 40 km N of Calabozo (9°08'N,
67°irW); Parmana (7°28'N, 65°18'W); San Jose
de Tiznados (9°16'N, 67''16'W). Lara: La Pastora,
1 1 km SSW of Sanare (9°2rN, 70°07'W). Miran-
da: Rio Negro (10''20'N, 66°17'W); La Guzma-
nera, Guatopo (10°00'N, 66°15'W). Territorio
Federal Amazonas: Alto Manapiare (5''13'N,
66°0rW); Alto Ventuari (4°45'N, 65°20'W); Caiio
Yureba, Rio Ventuari (3°16'N, 66''21'W); Cano

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Mayaba, Rio Ventuari (4°07'N, 66°16'W); Cano
Morrocoy, Alto Ventuari (5°08'N, 66°02'W); La
Esmeralda (S-^S'N, 65°32'W); Ocamo, Rio Oca-
mo (2°20'N, 65°15'W); Puruname, 40 km from
Rio Orinoco (3°19'N, 65°15'W); La Neblina, E of
Rio Varia (0°59'N, 66°10'W). Territorio Federal
Delta Amacuro: Cano Araguabisi (9°12'N,
60*'27'W); Guiniquina (9''10'N, 61'X)3'W); Tobe-
juba, Guayo (9°09'N, 6r25'W). Yaracuy: Agua
Negra (10°14'N, 68''14'W); Carretera Boca de
Aroa, 20 km from Palmasola (lO^Ol'N, 69°27'W).

Ateles

Spider monkeys are widely distributed from
northeastern Mexico throughout tropical South
America. Four species are recognized, of which
only Ateles belzebuth occurs in Venezuela (fig. 7).

Ateles belzebuth belzebuth GeofFroy is found in
southern Venezuela, south of the Rio Orinoco;
most records are from Territorio Federal Ama-
zonas. Kellogg and Goldman ( 1 944) indicate a wide
distribution in Guyana and report the Venezuelan
localities of La Union, Rio Mato, and El Llagual
(on both banks of the Rio Caura).

Ateles belzebuth hybridus Geoffroy is known
from northern and western Venezuela (Hershko-
vitz, 1949; Cabrera, 1958; Kellogg & Goldman,
1944). Handley (1976) reported specimens from
Apure and Trujillo in western Venezuela. Our
specimens are from the states of Barinas, Tachira,
and Zulia. Mondolfi and Eisenberg (1979) report-
ed it from Cupira and Guatopo, state of Miranda,
suggesting a discontinuous distribution on the
coast.

Specimens Examined— Total 16. Barinas: Re-
serva Forestal de Ticoporo, Sabana de Anare
(8°06'N, 70°40'W). Bolivar: Canaracuni (4°06'N,
64°10'W). Miranda: Cupira (10°10'N, 65°44'W).
Tachira: La Fria (8°13'N, 72°14'W). Territorio
Federal Amazonas: Cacuri, Rio Ventuari (4°49'N,
65°26'W); Rio Ocamo, Alto Orinoco (2°44'N,
65°11'W); Salto del Oso, Alto Ventuari (4''55'N,
65°25'W); San Juan de Manapiare (S^M'N,
66°02'W). Zulia: Rio Guasare ( 1 1°02'N, 72°05'W).

Lagothrix

Woolly monkeys have not been collected in
Venezuela; however two subspecies of Lagothrix
lagothricha are to be expected: L. lagothricha la-
got hricha should be found in Territorio Federal

Amazonas south of the Rio Ventuari, and L. I.
lugens, in the Selva de San Camilo, state of Apure
(Fooden, 1963; Hemandez-Camacho & Cooper,
1976).

Pithecia

Sakis are found only in northern South America;
Hershkovitz (1979) recognized four monotypic
species, of which only Pithecia pithecia Linnaeus
is found in Venezuela. P. pithecia is found in Ven-
ezuela south of the Rio Orinoco and throughout
the Guianas and northeastern Brazil. In Venezuela
specimens have been reported primarily from the
extreme northeastern region, the state of Bolivar
and Territorio Federal Delta Amacuro (fig. 5), with
a single outlier locality record for Belen, Rio Cunu-
cunuma, Territorio Federal Amazonas (3°39'N,
65°46'W) (Handley, 1976). If the distribution pro-
posed by Hershkovitz (1979) and Mittermeier and
Coimbra-Filho (198 1) is correct, P. pithecia should
inhabit the entire region between the upper Rio
Orinoco and the Rio Caroni, an enormous area
for which no specimens have been recorded.

Specimens Examined— Total 24. Bolivar: Rio
Curumo (7°15'N, 61°20'W); Rio Grande (8°16'N,
6 ri 7'W); Gurisoco, El Palmar (8°62'N, 6 1°26'W);
La Trinidad, El Palmar (7°12'N, 6r23'W); Guri,
Operacion Rescate (7°18'N, 63°00'W); Rio Boto-
namo, near Rio Cuyuni (6°59'N, 6rirW); San
Martin de Turumban, Rio Cuyuni (6''59'N,
61°02'W). Territorio Federal Delta Amacuro: Al-
tiplanicie de Nuria (7°50'N, 61°18'W); Yotacuay,
SW of Cupiare (8°30'N, 61°00'W).

Literature Cited

Allen, J. A. 1914. New South American Monkeys.
BviUetin of the American Museum of Natural History,
33: 653.

BoDiNi, R. 1981. Musculatura locomotora de la viudita
(Callicebus torquatus lugens). Sus implicaciones fun-
cionales y iilogeneticas. Memoria de la Sociedad de
Ciencias Naturales La Salle, 16(XL): 1-165.

. 1983. Distribuci6n, status de la investigacion

y conservaci6n de los cebidos en Venezuela, p. 1 39.
In Symjwsio sobre Primatologia en Latinoamerica. IX
Congreso Latinoamericano de Zoologia, Arequipa,
Peru.

Cabrera, A. 1957(1958]. Calalogo de los mamiferos
de America del sur. Revista Museo Argentine de Cien-
cias Naturales, "Bernardino Rivadavia," 4(1): xviii +
138 pp.

Cruz Lima, E. 1945. Mammals of Amazonia. I. Gen-
eral introduction and Primates. Contribution from the

BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA

243

Museu Paraense Emilio Goeldi de Historia Natural e
Etnografia, Belem do Para, Rio de Janeiro, Brasil, 274
pp., 42 pis.

EiSENBERG, J. F., AND K. Redford. 1979. A biogeo-
graphic analysis of the mammalian fauna of Vene-
zuela, pp. 31-36. In Eisenberg, J. F., ed., Vertebrate
Ecology in the Northern Neotropics. Smithsonian In-
stitution Press, Washington, D.C., 271 pp.

Elliot, D. G. 1912. A Review of the Primates. Vols.
I and II. American Museum of Natural History, New
York.

FooDEN, J. 1963. A revision of the woolly monkey
(genus Lagothrix). Journal of Mammalogy, 44: 213-
247.

Handley, C. O., Jr. 1976. Mammals of the Smith-
sonian Venezuelan project. Brigham Young Univer-
sity Sciences Bulletin, Biological Series, 20(5): 1-91.

Hernandez-Camacho, J., and R. W. Cooper. 1976.
The nonhuman Primates of Colombia, pp. 35-69. In
Thorington, R. W., Jr., and P. G. Heltne, eds.. Neo-
tropical Primates: Field Studies and Conservation.
National Academy of Sciences, Washington, D.C.

Hershkovitz, P. 1949. Mammals of northern Colom-
bia. Preliminary report No. 4: Monkeys (Primates),
with taxonomic revisions of some forms. Proceedings
of the United States National Museum, 98(3232): 323-
427.

. 1958. Type localities and nomenclature of some

American primates, with remarks on secondary hom-
onyms. Proceedings of the Biological Society of Wash-
ington, 71: 53-56.

1963. A systematic and zoogeographic account

of the monkeys of the genus Callicebus (Cebidae) of
the Amazonas and Orinoco river basins. Mammalia,
27: 1-79.
. 1972. The Recent mammals of the Neotropical

region: A zoogeographic and ecological review, pp.
31 1-431. In Keast, A., F. C. Erk, and B. Glass, eds..
Evolution, Mammals, and Southern Continents. State
University of New York Press, Albany, N.Y. 543 pp.
1977. Living New World Monkeys (Platyrrhi-

ni) with an Introduction to Primates. University of
Chicago Press, Chicago, 1117 pp.

1 979. The species of sakis, genus Pithecia (Ce-

bidae, Primates), with notes on sexual dichromatism.
Folia Primatologica, 31: 1-22.

1983. Two new species of night monkeys, ge-

nus Aotus (Cebidae, Platyrrhini): A preliminary report
on Aotus taxonomy. American Journal of Primatol-
ogy, 4: 209-243.

. 1984. Taxonomy of squirrel monkeys genus

Saimiri (Cebidae, Platyrrhini): A preliminary report
with description of a hitherto unnamed form. Amer-
ican Journal of Primatology, 7: 155-210.

Hill, W. C. O. 1 960. Primates. Comparative Anatomy
and Taxonomy. IV. Cebidae, Part A. Edinburgh Uni-
versity Press, London, 523 pp.

. 1962. Primates. Comparative Anatomy and

Taxonomy. V. Cebidae, Part B. Edinburgh University
Press, London, 537 pp.

INPARQUES. 1982. Guia de los Parques Nacionales
y Monumentos Naturales de Venezuela. Ediciones
Fundacion de Educacion Ambiental, Caracas.

Kellogg, R., and E. A. Goldman. 1944. Review of
the spider monkeys. Proceedings of the United States
National Museum, 96(3186): 1-45.

Kjnzey, W. G. 1982. Distribution of primates and for-
est refuges, pp. 455-482. In Prance, G. T., ed.. Bio-
logical Diversification in the Tropics. Columbia Uni-
versity Press, New York, 7 1 4 pp.

MARNR-DGS-POA-SFS. 1982. Informe Nacional de
Fauna Silvestre. Ministerio del Ambiente y de los Re-
cursos Naturales Renovable, Caracas.

Mittermeier, R. a., and A. Coimbra-Filho. 1981.
Systematic: Species and subspecies, pp. 29-109. In
Coimbra-Filho, A. F., and R. A. Mittermeier, eds..
Ecology and Behavior of Neotropical Primates. Aca-
demia Brasileira de Ciencias, Rio de Janeiro.

MoNDOLFi, E. 1976. Fauna silvestre de los bosques
humedos tropicales de Venezuela, pp. 113-181. In
Hamilton, L. S., J. Steyermark, J. P. Veillon, and E.
Mondolfi, eds., Conservacion de los bosques humedos
de Venezuela. Sierra Club— Consejo de Bienestar Ru-
ral, Caracas.

MoNDOLR, E., AND J. F. EiSENBERG. 1 979. Ncw records
for A teles belzebuth hybridus in northern Venezuela,
pp. 93-96. In Eisenberg, J. F., Vertebrate Ecology in
Northern Neotropics. Smithsonian Institution Press,
Washington, D.C, 271 pp.

Ramirez-Cerquera, J. 1983. Reporte de una nueva
especit de Primates del genero Aotus de Colombia, p.
146. Symposio sobre Primatologia en Latinoamerica.
IX Congreso Latinoamericano de Zoologia. Arequipa,
Peru.

RuDRAN, R., AND J. F. Eisenberg. 1982. Conservation
and status of wild primates in Venezuela, pp. 52-59.
In Olney, P. J. S., ed.. International Zoo Yearbook,
Vol. 22. The Zoological Society of London, London,
488 pp.

244

FIELDIANA: ZOOLOGY

Host Associations and Coevolutionary Relationships
of Astigmatid Mite Parasites of New World Primates
I. Families Psoroptidae and Audycoptidae

Barry M. OConnor

ABSTRACTS

Coevolutionary patterns among mites of the families Psoroptidae and Audycoptidae and
New World Primates are reviewed. Host records for primate parasites originally described from
artificial situations are compared with field collections from Peruvian primates, with most host
associations verified. A new sp)ecies of Audycoptidae, Saimirioptes hershkovitzi, is described
from Cebus apella. The psoroptid subfamily Cebalginae is hypothesized to be a monophyletic
group based upon 10 synapomorphies. Phylogenetic relationships within the Cebalginae are
reviewed, with cospeciational histories supported for the genera Alouattalges and Schizopodalges
and their hosts, and a more diffuse cospeciational pattern observed for the genera Cebalgoides,
Cebalges, and Fonsecalges and their hosts. Historical relationships of the genus Procebalges
remain problematical.

Patrones coevolutivos entre acaros de las familias Psoroptidae y Audycoptidae y primates
del Nuevo Mundo son revisadas. Registros de huespedes para parasitos de primates originar-
iamente descritos de situaciones artificiales son comparados con colecciones de campo de
primates peruanos, con la mayoria de asociaciones de huesped verificadas. Una nueva especie
de Audycoptidae, Saimirioptes hershkovitzi, es descrita de Cebus apella. Los psoroptidos de
la subfamilia Cebalginae son hipotetizados ser un grupo monofiletico basado en 10 sinapo-
morfias. Relaciones filogeneticas entre los Cebalginae son revisadas, con historias coesp>ecia-
cionales soportadas F>or el genero Alouattalges y Schizopodalges y sus huesp>edes, y un patron
coespeciacional mas difuso observado en los generos Cebalgoides, Cebalges y Fonsecalges y sus
huesjjedes. Relaciones historicas del genero Procebalges p>ermanecen problematicas.

Padroes coevolucionarios entre os acarinos das familias Psoroptidae e Audycoptidae, e os
primatas do Novo Mundo, sao revisados. Os registros de hospedes primatas de parasitas, que
foram descritos em situa^oes artificials (cativeiros), sao comparados com cole96es de campo
de primatas peruanos, e a maioria das associagoes hospedeiras atualmente registradas foram
averiguadas. Uma nova esjjecie de Audycoptidae, Saimirioptes hershkovitzi, encontrada em
Cebus apella e descrita. Baseando-se num estudo de 1 sinapomorfias, prop6e-se ser a subfamilia
psoroptidea, Cebalginae, um grupo monofiletico. As rela96es filogeneticas entre os Cebalginae
sao revisadas, e as historias de coesF>ecializa9ao entre os generos Alouattalges e Schizopodalges,
e seus respectivos hospedes, sao confirmadas. Os padroes de coespecializa9ao entre os generos
Cebalgoides, Cebalges e Fonsecalges, e seus hospedes, sao mais difusos. Os relacionamentos
historicos do genero Procebalges permanecem incertos.

From the Museum of Zoology and Department of Bi-
ology, The University of Michigan, Ann Arbor, MI 48 109.

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 245

Introduction

Coevolutionary patterns among hosts and par-
asites have been the focus of much recent study
and discussion (Brooks, 1979, 1981, 1985; Brooks
& Glen, 1982; Futuyma & Slatkin, 1983; Nitecki,
1983). Phylogenetic analysis of evolutionary re-
lationships among parasite groups or host groups
can provide additional data sets (e.g., treating par-
asite distributions as character states of their hosts),
or such analyses can be used as tests for hypotheses
regarding evolutionary relationships of the other
lineages of associated organisms. These phyloge-
netic analyses are especially useful when the par-
asite groups are host specific and indicate little
history of secondary colonization of new hosts.
The associations between mites and primates are
particularly amenable to such analysis because a
number of acarine groups are specifically associ-
ated with primates, and these lineages exhibit
enough within-group diversity to allow the con-
struction and comparison of phylogenetic hypoth-
eses. I have previously detailed such hypotheses
for several groups of astigmatid mites parasitic on
primates (OConnor, 1984).

Formulation of hypotheses regarding the his-
tory of associations between primates and their
associated mites requires three steps. First, the
parasite taxa must be described and their natural
host and geographic ranges discovered. Second,
taxa above the species level in classically derived
classifications must be tested for naturalness (i.e.,
monophyly). Finally, phylogenetic relationships
among all taxa must be elucidated. In the present
paper, each of these questions will be addressed
for the associations among certain groups of as-
tigmatid mites and New World Primates. For the
reader interested in summary information regard-
ing known host-parasite relationships for New
World primates, an exhaustive list of literature
records may be found in Hershkovitz (1977).

One major difficulty in the application of this
methodology to the study of the history of mite-
primale associations is the scanty knowledge of
the distribution and identity of the parasite sp)ecies
and their natural host ranges. Although many aca-
rine parasites of primates have been described,
much of the material has originally come from
zoos and primate research centers where the f>os-
sibility of unnatural interspecific contact between
host species makes the transfer of parasites a real
problem. Many other sjjecies have been described
from preserved host specimens in museum col-

lections where contamination in the field or in the
museum may also have been a problem. Finally,
the actual field locality from which either parasites
or hosts were collected is known for extremely few
of the known primate parasites. The uncertainties
involved in the host range and geographic distri-
bution of so many of the known species of primate
parasites render the phylogenetic analyses pro-
posed earlier (OConnor, 1984) subject to some
doubt.

In 1 98 1 , 1 was invited by Philip Hershkovitz to
collect parasites from specimens of a number of
primate species which had been collected or ob-
tained during field studies in Peru in 1 980. A very
large number of mites was collected, providing a
unique survey of the primate parasites in a small
area in Peru and a test for previously reported
host-parasite associations. These collections yield-
ed specimens belonging to three families of astig-
matid mites: Psoroptidae, Audycoptidae, and
Atopomelidae. In the present paper, the collec-
tions of Psoroptidae and Audycoptidae will be
discussed. The collections of Atopomelidae, con-
sisting of a number of described and undescribed
species of the genus Listrocarpus, will be studied
separately.

Materials and Methods

Primate specimens were collected in the field by
Hershkovitz or obtained from local individuals.
These specimens were prepared by removing and
simply drying the skins at the time of collection.
Upon their arrival in the United States, one or
more skins of each species collected were made
available to me for parasite removal before the
skins were sent for tanning. The following species,
all identified by Hershkovitz, were examined
(number examined in parentheses): Cebus apella
(3); C. albifrons (1); Lagothrix lagothricha (1); L.
flavicauda ( 1 ); Alouatta seniculus ( 1 ); Pithecia hir-
suta (3); Callicebus moloch (1); Aotus nancymai
(4); and Saimiri sciureus (5).

Parasites were removed in two ways. First, all
skins examined were vigorously brushed over white
paper, with the dislodged parasites collected under
a dissecting microscop>e. Finally, one skin of each
species was soaked in water and mild soap until
soft (the single specimen oi L. flavicauda was not
soaked). These skins were then gently washed, the
wash water filtered through a 200-mesh sieve (mesh

246

HELDIANA: ZOOLOGY

openings 75 micrometers), and the residue ex-
amined under a dissecting microscope. Mites col-
lected were preserved in 70% ethanol for subse-
quent study.

In the laboratory, mites were cleared in lacto-
phenol and mounted in Hoyer's medium, with
some specimens retained in alcohol in the cases
of large series. Voucher specimens of nominal
species are deposited in Field Museum of Natural
History, Chicago, and, when available, will be
placed in the following institutions: Museum of
Zoology, The University of Michigan, Ann Arbor;
The United States National Museum of Natural
History, Washington, D.C.; LTnstitut Royale des
Sciences Naturelles, Brussels, Belgium; and the
collection of F. S. Lukoschus, Katholieke Uni-
versiteit, Nijmegen, Netherlands.

Species Accounts

tritonymphs. Among 1 1 protonymphs in the pres-
ent collection, eight exhibit characteristics similar
to the male tritonymphs (i.e., posterior opistho-
somal lobes sclerotized; coxal fields III well scler-
otized) while three exhibit characteristics of female
tritonymphs (posterior lobes unsclerotized; scler-
otization of coxal fields III much weaker). I inter-
pret these differences as evidence for sexual di-
morphism at the protonymphal instar in this
species.

Material Examined— Total 133. Twenty-nine
females, 43 males (of which 3 1 were in tandem
with female tritonymphs), 38 female tritonymphs,
8 male tritonymphs, 3 female protonymphs, 8 male
protonymphs, 4 larvae from Cebus albifrons.
PERU, Loreto: Nauta, Rio Tigre, 6 km above Rio
Tigrillo; 18 December 1980; P. Hershkovitz (9264).
Host now a tanned skin (fmnh 122795). Mites
labeled bmcx: 81-081 1-3. No specimens were re-
covered from Cebus apella.

Family PSOROPTIDAE

Seven species of mites in the family Psoropti-
dae, subfamily Cebalginae, are known to parasitize
New World primates. These species, with their
known hosts and distributions, are listed below,
with new records from the Peruvian collections of
Hershkovitz indicated under "Material Exam-
ined." Keys to most of these species may be found
in Fain (1963c).

Cebalgoides cebi Fain, 1963

Cebalgoides cebi Fain, 1 963, Bull. Ann. Soc. Roy. Ent.
Belg., 99: 331. Fain, 1963, Bull. Inst. Roy. Sci. Nat.
Belg., 39(32): 91.

This species was briefly diagnosed by Fain
(1963a) from specimens collected from Cebus al-
bifrons which originated in "Amerique du Sud"
and died in the Antwerp (Belgium) Zoo. It was
more thoroughly described and illustrated by Fain
(1963c), who listed specimens from Cebus albi-
frons from Venezuela (type collection), C apella
from "Amerique du Sud," and Leontocebus (Oed-
ipomidas) oedipus (= Saguinus oedipus) from Co-
lombia. All these hosts had died in the Antwerp
Zoo.

Fain (1963c) noted sexual dimorphism in tri-
tonymphs of C cebi. The two protonymphs he
examined exhibited the characteristics of the male

Alouattalges corbeti Fain, 1963

Alouattalges corbeti Fain, 1963, Bull. Inst. Roy. Sci.

Nat. Belg., 39(32): 122. Fain, 1966, Acarologia 8:

103.
Rosalialges cruciformis Lavoipierre, 1 964, Acarologia,

6: 348.

This species was briefly described from the ho-
lotype female collected from a preserved specimen
of Alouatta seniculus macconnelli which had been
collected at Paramaribo, Surinam and preserved
in the British Museum (Natural History) (Fain,
1 963c). Although only the holotype was described,
several specimens were apparently recovered from
this host. Almost simultaneously, Lavoipierre
(1964a) described and figured the female based
upon two specimens collected from an "Aotes''''
(sic) sp. which had died in San Francisco, Cali-
fornia, after its importation from Peru. Fain (1966)
provided illustrations of the female and illustrated
but did not describe the male beyond length and
width measurements.

Material Examined— Total 15. Five females,
8 males, 2 female tritonymphs from Alouatta se-
niculus. PERU, Loreto: Nauta, Rio Samiria; 18
November 1980; P. Hershkovitz (9050). Host now
a tanned skin (fmnh 122789). Mites labeled bmoc
81-0809-5. No specimens were recovered from
Aotus nancymai, suggesting the possibility that the
two specimens collected by Lavoipierre (1964a)
represent contamination.

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES

247

Schizopodalges lagothricola Fain, 1 963

Schizopodalges lagothricola Fain, 1963, Bull. Ann. Soc.
Roy. Ent. Belg., 99: 469. Fain, 1963, Bull. Inst. Roy.
Sci. Nat. Belg., 39(32): 100.

This species was described from numerous spec-
imens from two juvenile Lagothrix lagothricha
which died in the Antwerp Zoo. The origin of the
hosts was stated as "Amerique du Sud" (Fain,
1 963b). Fain ( 1 963c) provided illustrations of male
and female and indicated that the hosts originated
in Colombia.

Material Examined— Total 30. Eleven fe-
males, 1 1 males, 3 tritonymphs, 2 protonymphs,
3 larvae from Lagothrix lagothricha. PERU, Lo-
reto: Nauta, Rio Samiria; 15 November 1980; P.
Hershkovitz (9032). Host now a tanned skin (fmnh
122790). Mites labeled bmcx: 81-0809-10.

Cebalges gaudi Fain, 1 962

Cebalges gaudi Fain, 1962, Rev. Zool. Bot. Afr., 66:
160. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg.,
39(32): 81.

This species was briefly described from numer-
ous sp)ecimens collected from a preserved speci-
men of Cebus capucinus with no locality infor-
mation (Fain, 1962). Full descriptions and figures
were provided later (Fain, 1963c).

Material Examined— Four females from Ce-
bus apella. PERU, Loreto: Nauta, Rio Samiria; 18
November 1980; P. Hershkovitz (9049). Host now
a tanned skin (fmnh 122792). Mites labeled bmoc
81-0811-2.

Fonsecalges johnjadini Fain, 1962

Fonsecalges johnjadini Fain, 1 962, Rev. Zool. Bot.
Afr., 66: 161. Fain, 1963, Bull. Ann. Soc. Roy. Ent.
Belg., 99: 468. Fain, 1963, Bull. Inst. Roy. Sci. Nat.
Belg., 39(32): 86.

This species was briefly described from several
Caliithrixjacchus which had died in captivity (Fain,
1 962). Fain ( 1 963b) provided an illustration of the
male, and Fain (1963c) gave a complete descrip-
tion with figures of both male and female. The
hosts, which were listed as eight Hapale jacchus
(= Callithrix jacchus) imported from the "bassin
de I'Amazone," died in Antwerp. Additional sjiec-

imens were recorded from a specimen of Hapale
jacchus leucocephalus (= Callithrix jacchus geof-
froyi) collected in Bahia, Brazil, and preserved in
Brussels.
Material Examined— None.

Fonsecalges saimirii Fain, 1963

Fonsecalges saimirii Fain, 1963, Bull. Ann. Soc. Roy.

Ent. Belg., 99: 330. Fain, 1963, Bull. Inst. Roy. Sci.

Nat. Belg., 39(32): 90. Fain, 1966, Acarologia, 8:

107.
Dunnalges lambrechti Lavoipierre, 1 964, Acarologia,

6: 343.

This species was briefly described from numer-
ous specimens collected from two Saimiri sciureus
from "Amerique du Sud" and which died in the
Antwerp Zoo in 1959 and 1963 (Fain, 1963a).
Fain (1963c) provided further descriptive infor-
mation and indicated the original hosts had come
from "Amazonie" and had died shortly after their
arrival at the Antwerp Zoo. He also mentioned
additional specimens from "Tamarins spp. ori-
ginaires d' Amazonie." The inexact locality infor-
mation leaves the actual identity of the type host
in doubt following the revision of the genus Sai-
miri by Hershkovitz (1984). Lavoipierre (1964a)
described and illustrated this species as Dunnalges
lambrechti, from "a long series of sjiecimens com-
prising all stages" from several Tamarinus nigri-
collis (= Saguinus nigricollis) from eastern Peru
which died in San Francisco, California. Fain
( 1 966) illustrated parts of this species from the
type specimens.

Fain (1963c) did not observe sexual dimor-
phism in the nymphs of this species, as only two
tritonymphs and some shed tritonymphal cuticles
were examined. Sexual dimorphism was noted in
the tritonymphs during the present study. All lar-
vae and protonymphs examined bear three pairs
of sclerotized apophyses in the ventrolateral re-
gion: a rounded apophysis immediately posterior
to trochanter II, a pointed apophysis lateral to the
base of leg III, and a pointed apophysis poster-
iolaterally near the posterior lobes. Tritonymphs
in which these apophyses are retained are here
interpreted as males, while those in which all
apophyses are lost are interpreted as females.

Material Examined— Total 59. Three females
from Saimiri sciureus macrodon. PERU, Loreto:
Nauta, Rio Tigre, 5 km above Rio Tigrillo; 17
December 1980; P. Hershkovitz (9257). Host now

248

FIELDIANA: ZOOLOGY

a tanned skin (fmnh 122810). Mites labeled bmoc
81-0809-15. Two females, 1 male from same host
species and locality; P. Hershkovitz (9258, fmnh
122811, BMOC 8 1 -0809- 1 6). Twenty-four females,
8 males, 5 female tritonymphs, 8 male trito-
nymphs, 5 protonymphs, 1 larva from same host
species. PERU, Loreto: Nauta, Rio Tigre, 6 km
above Rio Tigrillo; same date; P. Hershkovitz
(9268, FMNH 1 228 1 6, bmoc 8 1 -0809- 1 8). Two fe-
males from same host sjjecies and locality; P.
Hershkovitz (9267, fmnh 1 228 1 5, bmoc 8 1 -0809-
19).

Procebalges pitheciae Fain, 1963

Procebalges pitheciae Fain, 1963, Bull. Ann. Soc. Roy.
Ent. Belg., 99: 332. Fain, 1963, Bull. Inst. Roy. Sci.
Nat. Belg., 39(32): 96.

This species was briefly described from speci-
mens collected from a Pithecia monachus im-
ported from "Amerique du Sud" and which died
in the Antwerp Zoo (Fain, 1963a). The descrip-
tions were completed and illustrations provided
later (Fain, 1963c).

Material Examined— Total 242. Nineteen fe-
males, 28 males (7 in tandem with female trito-
nymphs), 15 female tritonymphs, 1 protonymph
from Pithecia hirsuta. PERU, Loreto: Nauta, Rio
Samiria; 30 November 1980; P. Hershkovitz
(9 1 1 5). Host now preserved as a tanned skin (fmnh
122797). Mites labeled bmoc 81-0809-7. Forty-
seven females, 8 1 males ( 1 7 in tandem with female
tritonymphs), 22 female tritonymphs, 8 male tri-
tonymphs, 13 protonymphs, 8 larvae from same
host and locality; P. Hershkovitz (9088, fmnh
122796, BMOC 81-0809-9).

Family AUDYCOPTIDAE

Three species of hair follicle inhabiting mites of
the family Audycoptidae have been previously de-
scribed from New World primates, all from squir-
rel monkeys identified as Saimiri sciureus. With
the recent recognition of several valid species in
the genus Saimiri (Hershkovitz, 1984), the exact
identification of the reported hosts is problemat-
ical. Specimens representing a fourth sr>ecies were
recovered from Cebus apella from the present col-
lections.

Audycoptes greeri Lavoipierre, 1 964

Attdycoptes greeri Lavoipierre, 1 964, Ann. Natal Mus.,
16: 194.

This species was described from females col-
lected from the sinus-hair follicles of Saimiri sci-
ureus collected in eastern Peru and kept in captiv-
ity in California (Lavoipierre, 1964b). In the
absence of more detailed collection information,
the true host may have been either S. sciureus or
S. boliviensis, as both occur in eastern Peru (Hersh-
kovitz, 1984).

Material Examined— Five females from "5fl/-
muri sciurea" (sic), without further collection data,
from the Lavoipierre collection. University of Cal-
ifornia, Davis.

Audycoptes lawrencei Lavoipierre, 1964

Audycoptes lawrencei Lavoipierre, 1964, Ann. Natal
Mus., 16: 199.

This species was described from the same hosts
and habitat as Audycoptes greeri (Lavoipierre,
1964b), so the actual specific identity of the host
remains uncertain, as indicated for greeri.

Material Examined— Two females from "^a/-
muri sciurea"" (sic), without further collection data,
from the Lavoipierre collection, University of Cal-
ifornia, Davis.

Saimirioptes paradoxus Fain, 1968

Saimirioptes paradoxus Fain, 1968, Acarologia, 10:
286.

This species was described from a single female,
containing a larva, collected from a Saimiri sci-
ureus which had died in the Antwerp Zoo (Fain,
1968). No information concerning the geographic
origin of the host was given, making the specific
identity of the host uncertain.

Material Examined— None.

Saimirioptes hershkovitzi, new species

In the following description, all measurements
are given in micrometers and are presented as ho-
lotype (range of three measured specimens).

Female (figs. 1-2)— Body elongate, cylindrical

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES

249

Fig. 1 . Saimirioptes hershkovitzi, female. A, dorsum; b, venter.

to somewhat flattened dorsoventrally; length in-
cluding gnathosoma 4 1 5 (4 1 5—42 1 ), width at level
of transverse coxal apodemes III 146 (135-146);
entire body with transverse striations which are
very thick anteriorly, thin posteriorly.

Dorsum (fig. la)— Prodorsal sclerite narrow an-
teriorly, much widened medially, narrowed pos-
teriorly and fused internally with apodemes of cox-
al fields II in most posterior region. Sclerite
traversed by anterior transverse striations later-
ally. Length of sclerite 70 (70-77). Paired dorsal
protuberances present mesal to setae h and sh. with

striations encircling protuberances along most of
their length; lengths 29 (28-29). Idiosomal chae-
totaxy as follows: scapular setae filiform, in a
transverse line posterior to prodorsal sclerite,
lengths sci 4 (4-5), see 39 (29-39); setae J, and /,
very short and rounded, subdivided into two dis-
tinct lobes; setae h and sh filiform, lengths 9 (6-
9); setae di subdivided into three parts, a ventral
filiform part, length 26 (23-26), and two dorsal
rounded lobes; setae d^ similar to ^2 but filiform
part shorter, length 19 (14-19), and middle lobe
more elongate; remaining setae filiform; /j 29 (28-

250

FIELDIANA: ZOOLOGY

< >

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES

251

29); /, ventrally positioned, 18 (18-19); ^^4 24 (19-
24); ^5 21 (no variation); l^ ventral, 5 (4-5); l^
ventral, 30 (27-32); a, ventral, 14 (no variation);
a, 30 (27-30). Four pairs of idiosomal cupules
present: ia dorsal between setae h and sh; im dorsal
between and lateral to setae d2 and d^; ip ventral
slightly anterior to seta /j; ih ventral, between and
mesal to setae U and /j. Opisthosomal glands pres-
ent, small, opening between and lateral to setae d^
and /,. Anus terminal. Opening to bursa copulatrix
terminal, above anal opening.

Venter (fig. lb)— Coxal apodemes well devel-
oped. Anterior apodemes of coxal fields I straight,
length 66 (57-66), fused medially to form a ster-
num of length 23 (23-24). Anterior apodemes of
coxal fields II curving mesally, length 75 (75-79);
posterior apodemes of coxal fields II transverse,
fused ventromedially with anterior apodemes of
coxal fields III, extending dorsally, length of ven-
tral portion 50 (46-50), length of dorsal portion
31 (28-31). Anterior apodemes of coxal fields III
extending almost longitudinally from trochanters
III to F>osterior apodemes of coxal fields II, then
bending at right angles to fuse with the latter, length
of longitudinal portion 49 (49-51), transverse
portion 39 (39-44). Anterior apodemes of coxal
fields IV very wide, anteriomedially directed, length
50 (50-53). Ovipore located between coxal fields
III and IV; a crescentic epigynal apodeme posi-
tioned anterior to ovipore; genital valve elongate,
disappearing posteriorly under transverse flap; a
pair of small thickenings along posterior, internal
portion of valves; two pairs of vestigial genital
papillae lateral to genital ojiening. Three pairs of
filiform coxal setae present: cxI at posterior end
of coxal fields I between coxal apodemes II, length
12 (9-12); cxIII on median apex of coxal apo-
demes IV, length 9 (6-9); cx/P'mesad of the base
of leg IV, length 7 (6-7). Genital setae absent.

Gnathosoma (fig. 2g)— Gnathosoma elongate,
somewhat widened posteriorly, then tapering to a
point ventrally, length (excluding chelicerae) 62
(59-62), maximum width 26 (24-26). Chelicerae
elongate, with small, toothed digits, length 46 (44-
46). Palps apparently two-segmented; basal seg-
ment bearing two very short, filiform setae, 1 prox-
imodorsal, 1 medioventral; distal segment with
short solenidion and 5 sclerotized, pxjinted pro-
jections. Subcapitulum with pair of short, filiform
setae medioventrally; with pair of sclerotized,
pointed projections positioned slightly anterior to
subcapitular setae, and unpaired projection ante-
riomedially; median rutellar lobes with sclerotized
points medially.

Legs (figs. 2a-f) — Legs I-II similar in structure,
with all segments free, lengths 84 (80-90). Femora
I-II bearing two ventral apophyses, one filiform
seta vF. lengths: I, 9 (8-9); II, 33 (30-33). Genua
I-II each with large ventral apophysis; setae cG
expanded basally, then tapering, lengths 22 (20-
25), setae mG filiform, lengths 43 (40-45). Tibiae
I-II each with large ventral apophysis; setae gT
short, filiform, lengths 6 (5-6); solenidia api-
codorsal, lengths: I, 20 (18-20); II, 22 (21-23).
Tarsi I-II each with pointed dorsal apophysis, two
distal hooked apophyses; chaetotaxy of tarsi I-II
similar; setae d as strongly hooked, apical claws,
lengths 5 (no variation); e filiform, lengths 40 (43-
50); /filiform, lengths 7 (6-7); la filiform, lengths
6 (5-6); wa very short, not measurable; ra filiform,
lengths 6 (5-6); solenidia «, blunt apically, lengths:
I, 14 (no variation); II, 26 (24-26). Pretarsi I-II
similar, total lengths 40 (37-40); pretarsi divided
into long ambulacral stalk and rounded ambula-
cral disc; ambulacral disc with condylophore guide
and ventral "rays" (fig. 2b); condylophores atten-
uate, appearing to divide at base of ambulacral
disc.

Legs III-IV similar, each with fused trochanter-
femur bearing a large, ventral apwaphysis. Genua
III-IV glabrous. Tibiae III-IV each with setae kT
enlarged, bifurcate, and deeply rooted; tibia III
with solenidion 0, length 6 (6-7), IV absent.
Tarsi III-IV each with apical, pointed apophysis,
three setae: fi^ apical, lengths: III, 100(91-100); IV,
97 (86-97); r filiform, lengths 7 (6-7); w filiform,
lengths 3 (no measurable variation).

Types— Holotype and two paratype females from
Cebus apella. PERU, Loreto: Nauta, Rio Samiria;
18 November 1980; P. Hershkovitz (9049). Host
now a tanned skin (fmnh 122792). Mites labeled
BMOC 81-0811-2. Holotype and 1 paratype de-
posited in Field Museum of Natural History, Chi-
cago; 1 paratype in Museum of Zoology, Univer-
sity of Michigan, Ann Arbor.

Systematic Position— Saimirioptes hershko-
vitzi shares with 5". paradoxus, the tyF>e-sp)ecies and
only other member of the genus, the presence of
dorsal lobes between setae ^1 and t/j^ the similar
form of setae t/, and /, (rounded, bifurcate, and
without filiform part), and the expansion of the
bases of setae cG of genua I-II. The new sF>ecies
differs from the type-species in the relative lengths
of the scapular setae (sce:sci 5-6:1 in S. hershko-
vitzi, 1.5:1 in S. paradoxus); the shorter filiform
fKJrtion of setae f/, (not reaching dy in S. hersh-
kovitzi, extending past d^ in S. paradoxus); the
shorter lengths of setae 4 (29 in S. hershkovitzi.

252

HELDIANA: ZOOLOGY

67 in S. paradoxus); the presence of setae U (absent
in S. paradoxus); the longer pretarsi I-II (37-40
in S. hershkovitzi, 17 in S. paradoxus); and the
greater lengths of setae d on tarsi III-IV (at least
twice as long as the entire leg in S. hershkovitzi,
less than half the leg length in S. paradoxus).

Discussion

The Peruvian collections reported upon here
support the hypothesis that most of the records
of primate-Cebalginae associations previously re-
ported reflect natural host-parasite associations.
The association between Alouattalges corbeti and
Aotus species was not verified and remains ques-
tionable. As there were no field collected repre-
sentatives of the Callitrichidae examined during
this study, the occurrence of cebalgine mites on
these hosts remains generally untested in natural
situations, with only Fonsecalges johnjadini re-
corded from noncapti ve callitrichids (Fain, 1 963c).

Having established that most records of pri-
mate-cebalgine associations reflect natural asso-
ciations, two additional hypotheses must be pro-
posed and tested before any coevolutionary
hypotheses may be tested. These preliminary hy-
potheses concern the monophyly of the Cebalginae
and the phylogenetic relationships of the taxa
within the group. I have previously discussed these
questions (OConnor, 1 984) but, due to space con-
straints, was unable to detail the reasoning behind
my conclusions.

Monophyly of the Cebalginae

Before any hypothesis of historical associations
may be tested, at least one of the lineages must be
hypothesized to be monophyletic (Brooks, 1981).
In this study, the psoroptid subfamily Cebalginae,
which comprises all psoroptid mites parasitizing
New World Primates, must be tested for mono-
phyly. In last defining this subfamily. Fain ( 1 963c)
listed 21 character states for the group. Because
the comprehensive morphological studies of Fain
(1963c) were carried out before the methods of
phylogenetic systematics became widely discussed
and utilized, no distinction between ancestral and
derived states was made in the diagnosis of the
Cebalginae. In order to test whether the Cebalgi-
nae represents a monophyletic group, I have ex-
amined the characters listed by Fain (1963c) using

outgroup comparison to polarize the states. I have
previously hypothesized that the taxa comprising
the four subfamilies of Psoroptidae which para-
sitize the Primates form a monophyletic group
(OConnor, 1984). These taxa include the Makial-
ginae (sensu OConnor, 1984; i.e., including the
Cheirogalalginae and Galagalgidae of Fain), par-
asites of the Strepsirrhini; the Paracoroptinae, par-
asites of African Cercopithecidae and Hominidae;
the Nasalialginae, parasites of Asian Cercopithe-
cidae; and the Cebalginae, from New World pri-
mates. This grouping of taxa is regarded as the
ingroup in the following analysis. Outgroups used
in defining the character state polarities were the
other subfamilies of Psoroptidae and earlier de-
rivative groups in the Astigmata.

Character Analysis

1 . Presence of retrograde apophyses on coxa!
fields III. This state is unique to the cebalgine
genera, with such apophyses not occurring in other
taxa in the ingroup or outgroup.

2. Male body size much smaller than female.
In the outgroups and other ingroups, males are
similar in size to females or somewhat larger or
smaller. The substantial reduction in body size of
male Cebalginae is unique.

3. Female opisthosoma more or less squared
posteriorly and laterally, reinforced with sclero-
tized areas. This condition is exhibited by all Ce-
balginae, but not in any outgroup. Among the in-
groups, the squared body is unique to the
Cebalginae, but sclerotized reinforcement is also
found in the Nasalialginae and some Paracorop-
tinae (Pangorillalges). Among outgroup taxa,
sclerotized reinforcement is present in some Pso-
ralginae (Edentalges), but again without the squared
body form.

4. Male with legs III very modified, with 3-4
terminal segments fused and bearing medially di-
rected projections. The modification of the third
pair of legs in male Cebalginae is unique and pres-
ent in all taxa. No similar modification occurs
anywhere in ingroup or outgroup.

5. Reduction or loss of paranal suckers in males.
Paranal suckers are present in males in the out-
groups and ingroups. In the Cebalginae, the suck-
ers are very reduced or absent.

6. Tarsus I with 2 apical solenidia. I regard the
apical displacement of solenidion oj, in the Ce-
balginae as derived. In the other ingroups and most
outgroups, this solenidion is median or basal on

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES

253

the tarsus. Apical displacement of this solenidion
also occurs in the subfamily Psoroptinae and the
monobasic Marsupialginae, conditions I regard as
convei^ent.

7. Loss of dorsal seta ^,. Seta d^ is absent in all
Cebalginae and retained in the other ingroups and
most outgroups. This seta is also lost in the Lis-
tropsoralginae and the psoroptid parasites of ro-
dents (Echimyalges. Myoproctalges, and Coen-
dalges), conditions I regard as convergent.

8. Reduction or loss of apophyses from the pos-
terior tarsi of the female. Apophyses are present
on the posterior tarsi of some of the outgroups
such as the Listropsoralginae and are retained in
some ingroups, the Makialginae and Paracorop-
tinae. I consider the presence of these apophyses
to be plesiomorphic for the primate-associated
psoroptid lineage. In the Cebalginae, taxa in which
females have well-developed posterior legs retain
vestiges of these apophyses, while taxa in which
the legs are reduced retain no traces. I consider
this reduction to be a derived state for the Cebal-
ginae and regard the loss of these structures in the
Cebalginae and Nasalialginae as convergent.

9. Posterior edge of female opisthosoma with
2 pairs of long, strong setae. In most ingroups and
psoroptid outgroups, seta /, is long and strong while
seta di is shorter and thinner. Of the ingroups,
setae l^ and d^ are equally well developed in the
Cebalginae and Nasalialginae. However, in the Ce-
balginae, setae /, and d^ are closely associated, usu-
ally on a single projection, while in the Nasalial-
ginae these setae are separate, on distinctly different
projections. I regard these two conditions as con-
vergent. I also regard as convergence the elonga-
tion of seta di in some Psoralginae.

10. Dorsal seta l^ sometimes absent. Seta I4 is
absent in all Cebalginae except Cebalgoides cebi.
The seta is retained in that species, the other in-
groups, and most outgroups. The loss of seta I4
could be regarded as a synapomorphy defining a
group containing all Cebalginae except Cebal-
goides. However, this hypothesis conflicts with
groupings suggested by all other characters (see
page 256). At this point, it is more parsimonious
to regard the loss of seta U as a synapomorphy for
the Cebalginae, with a reversal in Cebalgoides.

1 1 . Loss of retrograde apophyses on the ante-
rior legs. Retrograde apophyses are present on the
anterior legs in some of the outgroups, notably the
Listropsoralginae and Marsupialginae in the Pso-
roptidae and in the related families Audycoptidae,
Rhyncoptidae, and Myocoptidae. Among the in-
groups, these apophyses are retained in the Ma-

kialginae but lost in the Cebalginae, Paracorop-
tinae, and Nasalialginae. I regard this loss as
characterizing these latter three groups as a natural
unit and thus plesiomorphic for the Cebalginae.

12. Female bursa copulatrix subterminal or
ventral. In the outgroups, the female bursa cop-
ulatrix is terminal, as it is in the Cebalginae, Ma-
kialginae, and Nasalialginae among ingroups. I re-
gard the dorsal position of the bursa in the
Paracoroptinae as derived. Thus, this character
retains the plesiomorphic condition in the Cebal-
ginae.

13. Base of the gnathosoma with retrograde
apophyses. In some outgroups as well as in the
Cebalginae, Makialginae and some Paracordpti-
nae (i.e., Pangorillalges), the base of the gnatho-
soma bears retrograde apophyses. These are ab-
sent in certain outgroups (e.g., the Psoroptinae)
and in Paracoroptes and Nasalialges among in-
group taxa. I regard the retention of these apoph-
yses in the Cebalginae as plesiomorphic.

14. Absence of retrograde apophyses on coxae
I-II. Retrograde apophyses occur on coxae I-II
only in the Makialginae. They do not occur on
both coxal fields in any other ingroup or outgroup
taxa, although the Listropsoralginae and Myocop-
tidae have apophyses on coxae II. I regard, these
structures as synapomorphies characterizing the
Makialginae, and their absence in the Cebalginae
as plesiomorphic.

15. Well-developed "claws" on tarsi I-II.
Clawlike apophyses on the apices of tarsi I-II are
present in many outgroup taxa, including both
mammal and bird parasites. I regard the presence
of this character state as ancestrally characterizing
all Psoroptidia, and thus plesiomorphic for the
Cebalginae.

16. Tarsus II with solenidion apically dis-
placed. Solenidion oj is apically displaced in the
Cebalginae, Paracoroptinae, and Nasalialginae. In
most outgroups and in the Makialginae, the so-
lenidion retains its ancestral, basal position. I re-
gard this displacement as diagnosing a natural
group consisting of the above three primate-as-
sociated subfamilies and thus plesiomorphic for
the Cebalginae.

17. Female tarsal chaetotaxy 7-7-6—4 or -5.
Possession of 7 setae on tarsi I-II involves the loss
of seta ba in the Paracoroptinae and Cebalginae
and probably the Nasalialginae. Retention of 8
setae on these segments (including ba) is charac-
teristic for most outgroup taxa and the Makial-
ginae. I regard this state as diagnosing a natural
group comprising the Cebalginae, Paracoroptinae,

254

HELDIANA: ZOOLOGY

and Nasalialginae and thus plesiomorphic for the
Cebalginae.

1 8. Angles of female opisthosoma generally with
strong hooks. Such hooks are found in the cebal-
gine genera Cebalges, Cebalgoides. and Fonse-
calges, but not in other taxa in the outgroup or
ingroup. I regard the presence of these structures
as characterizing a monophyletic unit within the
Cebalginae but not the group as a whole.

19. Posterior legs of female normal or atro-
phied. Legs III-IV of the female are reduced in
size and may exhibit fusion of segments in the
genera Cebalges and Fonsecalges but not in other
Cebalginae nor other members of the ingroup.
Well-developed legs are characteristic of most out-
group taxa, although similar reduction of the pos-
terior legs occurs in some Psoroptinae and Pso-
ralginae. I regard the atrophied legs of some
Cebalginae as characterizing a smaller monophy-
letic unit within the group and convergent with
the outgroup taxa noted above.

20. Posterior legs of nymphal stages normal or
short and atrophied and bearing a long seta. The
reduction of the legs in the nymphs is found in all
cebalgine genera except Procebalges. It is also found
in a few outgroup taxa (e.g., the Psoroptinae and
Psoralginae), but not in most outgroups or in other
ingroup taxa. I regard this state as diagnosing a
group within the Cebalginae and not characteriz-
ing the group as a whole.

2 1 . Dorsal seta d^ sometimes absent. Seta d^ is
absent in Schizopodalges lagothricola and Fon-
secalges johnjadini (but not F. saimirii), and pres-
ent in other Cebalginae, other ingroups, and most
outgroups. The distribution of this derived state
within the Cebalginae strongly suggests the inde-
pendent loss of this seta in the two species sharing
the state. This character at best diagnoses a group
within the Cebalginae, but is more likely a case of
convergence. In no way does this character diag-
nose the group as a whole.

Conclusions

On the basis of the above analysis, I conclude
that, of the 21 character-states listed by Fain
(1963c) as diagnosing the Cebalginae, five repre-
sent unique synapomorphies which diagnose the
Cebalginae and occur in no other group (character-
states 1-5), five are synapomorphies which diag-
nose the Cebalginae but which also occur as con-
vergent states in certain other taxa (character-states
6-10), seven are symplesiomorphies diagnosing

larger groups which include the Cebalginae (char-
acter-states 11-17), and four represent within-
group apomorphies diagnosing smaller groups
within the Cebalginae (character-states 1 8-2 1 ). The
ten synapomorphies diagnosing the Cebalginae
leave no doubt that the group is a natural one.

Phylogenetic Relationships
Within the Cebalginae

I have previously presented a hypothesis of phy-
logenetic relationships among genera in the Ce-
balginae based upon 17 characters (OConnor,
1 984). This cladogram is reproduced here (fig. 3),
with numbers on the cladogram referring to the
derived states of the characters listed below. In the
earlier study, space limitations prevented discus-
sion of the different states of these characters and
the reasons for interpreting their polarity. Addi-
tionally, an error in character 1 2 appeared in the
list of character-states. As I have hypothesized that
the Cebalginae form a monophyletic group within
a larger lineage comprising the Makialginae, Par-
acoroptinae, and Nasalialginae, taxa in these three
lineages were used as outgroups to polarize the
character-states within the Cebalginae.

1. Female with coxal apodemes III-IV fused.
In all taxa in the outgroups, coxal apodemes III-
IV end freely. Within the Cebalginae, these apo-
demes are fused together on either side in the gen-
era Procebalges, Schizopodalges, and Fonsecalges.
Although I regard the fused condition to be de-
rived, conflicts with many other characters suggest
that these apodemes have fused independently in
the three genera.

2. Female epigynum fused with coxal apo-
demes I. In most outgroup taxa, the female ovi-
pore is located between coxal fields II and III.
There has been a trend toward anterior displace-
ment of the ovip)ore in a number of psoroptid
lineages (e.g., Psoralginae, Listropsoralginae, Pso-
roptinae), and the more anterior p>osition occurs
in certain taxa among the primate associated lin-
eage as well (e.g., Lemuralges in the Makialginae).
Within the Cebalginae, this derived state occurs
in the genera Procebalges and Schizopodalges. Be-
cause this character-state distribution conflicts with
many other characters, I regard the presence of
this state in the two genera as convergence.

3. Male with paranal suckers lost. In all out-
groups, the male retains paranal suckers. Within
the Cebalginae, the suckers are retained only in

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES

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■ 16

■ 1

Fig. 3. Phylogenetic relationships among genera in the subfamily Cebalginae. Numbers refer to derived states of
characters discussed in the text. Hosts of each mite taxon given in parentheses.

Procebalges. I regard the loss of these structures
as derived and characterizing the sister group of
Procebalges.

4. Immature stages with pretarsi III-IV lost.
Described immature stages of outgroup taxa all
retain the pretarsus on legs III-IV. Within the Ce-
balginae, only Procebalges retains this ancestral
state. In all other Cebalginae, pretarsi III-IV are
lost in the immature stages, a condition I regard
as derived.

5. Gnathosomal apophyses lost in both sexes.
As discussed in the previous section, I regard the
presence of gnathosomal apophyses as ancestral
in the Cebalginae. These structures are lost in both
sexes in the genera Schizopodalges and Alouat-
talges within the Cebalginae, a condition I regard

as derived. I regard this state as a synapomorphy
for these two genera and convergent with the sim-
ilar state occurring in the genera Lemuralges
(Makialginae), Paracoroptes (Paracoroptinae), and
Nasalialges (Nasalialginae). I have treated loss of
these apophyses in the female only as a separate
character (16, below).

6. Female with seta 5 of tarsi I-II in the form
of a strongly hooked claw. In the outgroup taxa,
seta s is simple and filiform. Within the Cebalgi-
nae, this state is retained in all taxa except Schi-
zopodalges and Alouattalges where the seta is en-
larged and hooked. I regard the latter state as
derived and a synapomorphy for the two genera.

7. Male with a large spur on tarsus III. This
condition is unique to the cebalgine genera Schi-

256

FIELDIANA: ZOOLOGY

zopodalges and Alouattalges. In the outgroups and
other ingroup taxa, such a spur does not exist. I
regard the presence of this character-state as in-
dicative of common ancestry of these two genera.

8. Male with ridges on leg III restricted to tibial
element. Ridges on the distal portion of leg III of
the male characterize the Cebalginae (see above).
Such ridges do not occur in any outgroup. I regard
the restriction of the ridges to a spur extending
from the tibial portion of the leg as a derived state
defining the group Schizopodalges + Alouattalges
because this condition forms a functional complex
with the spur on tarsus III (character 7) and a
dorsal spur on femur IV. This latter character was
not considered in the previous analysis (OConnor,
1984). The conjunction of the three spurs serves
to lock legs III and IV together into a unit.

9. Female with recurved hooks on posterior
border of opisthosoma. Strong hooks are present
in this position in females of Cebalgoides, Ce-
balges. and Fonsecalges. Weaker development in
this area is characteristic of other cebalgine genera,
while in the outgroups, no projections exist. I re-
gard the possession of strong hooks as a derived
condition defining a lineage comprising the three
genera noted above.

10. Male with pretarsus III lost. Males retain a
pretarsus on leg III in most outgroup taxa. This
pretarsus is lost in the genera Galagalges and Chei-
rogalalges (Makialginae), and also in the cebalgine
genera Cebalgoides, Cebalges, and Fonsecalges. I
regard the loss of pretarsus III as a synapomorphy
for the latter three genera within the Cebalginae,
with independent loss in the makialgine lineage
comprising the former two genera.

1 1 . Female without opisthosomal sclerite. A
median sclerite is present in females of most out-
group taxa and is retained in all Cebalginae except
Schizopodalges. I regard this loss as derived in the
latter genus. Convergence in this character occurs
with some outgroup taxa. This sclerotization is
also lost in Gaudalges caparti, and Lemuralges
(Makialginae), and Nasalialges (Nasalialginae).

12. Female with seta s of tarsi III-IV enlarged
and clawlike. In the previous study (OConnor,
1984), a lapsus occurred in that the state "seta 5
of tarsi III-IV reduced" was listed as a derived
condition for the genus Alouattalges. In fact, pos-
session of a small, filiform seta s on these tarsi
must be regarded as the ancestral condition, as it
occurs in all outgroup taxa as well as in most Ce-
balginae. In the genus Alouattalges, seta 5 is en-
larged and recurved on tarsi III-IV, a condition I
regard as the true derived state.

13. Male with opisthosomal lobes widely
spaced. In most outgroup taxa, the opisthosomal
lobes of the male are close together, a condition
also found in most Cebalginae. These lobes are
widely spaced in the genus Cebalgoides within the
Cebalginae, a condition I regard as derived. Con-
vergence occurs, with this state also occurring in
Nasalialges (Nasalialginae).

1 4. Male with apodeme between genital and anal
region strongly reduced. In the outgroup taxa, males
do not possess a transverse apodeme between the
genital and anal regions. A large apodeme is pres-
ent in this position in most male Cebalginae. In
the prior analysis (OConnor, 1 984), I stated that
this apodeme was absent in Cebalgoides as indi-
cated by Fain (1963c) and regarded this as a re-
versal. Closer examination of a number of speci-
mens indicates that a very small apodeme is present
in some males of this taxon. I regard the presence
of the apodeme as a derived condition for the
Cebalginae, and its reduction or loss in Cebal-
goides as a further derived state.

1 5. Female with tibiae-tarsi III-IV fused. In the
outgroups, the tibia and tarsus of legs III-IV are
freely articulated. This condition is present in most
Cebalginae as well. In the genera Cebalges and
Fonsecalges, these segments are fused in the fe-
male, a condition I regard as derived within the
Cebalginae. Similar fusions occur in some other
psoroptid subfamilies but not within the primate-
associated lineage.

16. Female with gnathosomal apophyses lost.
Gnathosomal apophyses are retained in females
of most outgroup taxa and are retained in some
Cebalginae. These apophyses are lost in the female
but retained in the male in the genus Fonsecalges.
I regard this loss as independent of the loss of
apophyses in the Schizopodalges-Alouattalges lin-
eage, where the apophyses are lost in both sexes
(character 5).

17. Female with pretarsi III-IV lost. Pretarsi
III-IV are retained in the female in all outgroup
taxa and in all Cebalginae except Fonsecalges. I
regard this loss as derived and convergent with the
similar loss in females in other psoroptid groups
(e.g., some Psoroptinae, Psoralginae).

History of Primate-Cebalgine Evolution

The present knowledge of the diversity and host
associations of the family Audycoptidae is not suf-
ficient for formulating hypotheses concerning the

OCO^fNOR: MITE PARASITES OF NEW WORLD PRIMATES

257

history of their host associations. However, given
the phylogenetic relationships among cebalgine
taxa presented in the cladogram (fig. 3), two meth-
ods exist for using these relationships to make hy-
potheses regarding the history of the associations
between the New World primates and the Cebal-
ginae. The first method would involve comparing
the phylogenetic hypothesis for the parasite group
with a similar hypothesis for the hosts. The hy-
pothesis tested by this comparison is whether the
current pattern of host-parasite associations di-
rectly results from strict cospeciation between the
hosts and their parasites. Given a phylogenetic
hypothesis for each group, such a hypothesis is
easily tested. The second type of conclusion which
might be drawn from the parasite cladogram is a
phylogenetic hypothesis concerning host relation-
ships. This method is dependent upon the as-
sumption of cospeciation, or at least noncoloni-
zation, between the two lineages.

The current state of knowledge concerning the
phylogenetic relationships among New World pri-
mates is somewhat confused. Many early hypoth-
eses were based upon classical methodology in
which ancestral and derived character-states were
not differentiated in proposing hypotheses of re-
lationships. Schwartz et al. (1978) summarized the
state of knowledge only a few years ago by saying
that "Platyrrhini apF>ears in and of itself to be a
natural group, although both its wider relation-
ships and the relationships among its members
remain unclear" (p. 128). Much new information
relating to this question was presented in symp>osia
in 1978-1979 and published in a volume edited
by Ciochon and Chiarelli (1980). In the following
discussion, I refer to phylogenetic hypotheses for
New World primates suggested by Rosenberger
(1977) and contributors to the Ciochon and Chi-
arelli volume as representing the most modem
phylogenetic thinking regarding Platyrrhine rela-
tionships.

Returning to the question of cospeciation be-
tween primates and cebalgine mites, the first dif-
ficulty encountered is the lack of a consensus con-
cerning phylogenetic relationships among all New
World primates. Phylogenetic hypotheses based
on the following types of data sets are in strong
disagreement: dentition (Rosenberger, 1977; 3
separate hypotheses); integumentary characters
(Perkins & Meyer, 1980); karyology (Chiarelli,
1980); immunological methods (Sarich & Cronin,
1980); and other immunological data and protein
sequence data (Baba et al., 1 980). Despite the strong
disagreement among these data sets, some patterns

are common to several or all of these. These may
be compared with the cladogram of mite relation-
ships to test the cospeciational hypothesis.

One host relationship which is supported by al-
most all data sets is the relationship between Al-
ouatta and the Atelinae (including Lagothrix). Of
the hypotheses cited above, only the karyological
evidence did not support this relationship, and
then only because the karyotype of Lagothrix is
so derived that Chiarelli ( 1 980) made no hypoth-
esis as to its relationships. This relationship be-
tween Alouatta and Lagothrix is mirrored by the
sister group relationship between their parasites,
Alouattalges and Schizopodalges, which is among
the most strongly supported relationships among
the Cebalginae.

A second host relationship which is partially
mirrored in the mile phylogeny is the relationship
between the genera Cebus and Saimiri, which was
supported by all data sets except the integumen-
tary data (Perkins & Meyer, 1980). In this case,
the mite phylogeny suggests a cospeciational pro-
cess, but some additional hypotheses are required
to explain the distributions of the genera Cebal-
goides, Cebalges, and Fonsecalges. The relation-
ships among these three parasite genera are com-
patible with a cospeciational scenario if an early
sjjeciation event between the Cebalgoides and the
Cebalges-Fonsecalges lineages occurred in con-
junction with a common ancestor of Cebus and
Saimiri. Extinction of the Cebalgoides lineage on
the Saimiri line, with cospeciation of the Cebalges-
Fonsecalges lineage in both host lines, leads to the
present distribution on the Cebidae. In all cases,
colonization events are required to explain the
presence of both of these lineages on the Callitrich-
idae, given the probable monophyly of that taxon.

Certain other relationships among the parasite
taxa are not mirrored by host phylogenies. The
sister group relationship between Procebalges and
all other cebalgines is not reflected in any hyp)oth-
esis of host relationships. The relationship of Pi-
thecia to the Alouatta-Lagothrix lineage was sug-
gested by both dental data (Rosenberger, 1977)
and karyology (Chiarelli, 1 980). Interestingly, the
integumentary data (Perkins &. Meyer, 1980) sug-
gest that Pithecia retains the most plesiomorphic
skin characteristics of any mite-bearing New World
primate (Aotus was regarded as even more plesio-
morphic). As cebalgine mites are skin inhabitants,
it might be suggested that plesiomorphic skin re-
tains plesiomorphic mites, a hypothesis which
would require the ancestral possession of a Pro-
cebalges lineage on other cebids with subsequent

258

HELDIANA: ZOOLOGY

extinction on all but Pithecia in order to retain a
basically cospeciational history. Discovery of ce-
balgine mites on hosts related to Pithecia (Chi-
ropotes, Cacajao) will provide a test of these hy-
potheses.

The utility of using the proposed mite phylogeny
to infer a host phylogeny at higher levels than
mentioned above depends upon the degree to which
cospeciational patterns outweigh colonizations or
extinctions in the hypothesized historical relation-
ships. Among the entire psoroptid lineage para-
sitizing all Primates, cospeciational patterns ap-
pear to be supported in a large majority of cases
(OConnor, 1984). However, noncosj)eciational
patterns such as the distribution of the genus Le-
muralges in the Makialginae, or the cebalgine par-
asites of the Callitrichidae, make the use of these
mites as consistent indicators of host phylogeny
at least somewhat suspect. Given the large dis-
parity in phylogenetic hypotheses generated from
subsets of the overall character matrix for the New
World Primates, this parasite data deserves to be
at least considered by future workers in this area.

Acknowledgments

I would like to thank Philip Hershkovitz, with-
out whose cooperation and interest this study could
not have been attempted. I also thank Robert
Timm and Bruce Patterson, Field Museum of Nat-
ural History, for their hospitality and cooperation
during the processing of host specimens and for
their critical review of the manuscript. I thank J.
H. S. Klompen, University of Michigan, for his
comments on the manuscript. I thank Margaret
van Bolt, Museum of Zoology, University of
Michigan, for assistance with the illustrations.

Literature Cited

Baba, M., L. Darga, and M. Goodman. 1980. Bio-
chemical evidence on the phylogeny of Anthropoidea,
pp. 423-443. In Ciochon, R. L., and A. B. Chiarelli,
eds.. Evolutionary Biology of the New World Monkeys
and Continental Drift. Plenum Press, New York.

Brooks, D. R. 1979. Testing the context and extent of
host-parasite coevolution. Systematic Zoology, 28: 299-
307.

studying the evolution of ecological associations. An-
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Brooks, D. R., and D. R. Glen. 1982. Pinworms and
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85.

Chiarelli, A. B. 1980. The karyology of South Amer-
ican Primates and their relationship to African and
Asian species, pp. 387-398. In Ciochon, R. L., and
A. B. Chiarelli, eds.. Evolutionary Biology of the New
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Ciochon, R. L., and A. B. Chiarelli, eds. 1980. Evo-
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Fain, A. 1 962. Diagnoses d'acariens nouveaux. Revue
de Zoologie et de Botanique Africaines, 66: 154-162.

. 1 963a. Nouveaux acariens psoriques parasites

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Royale d'Entomologie de Belgique, 99: 322-332.

. 1 963b. Un nouvel acarien producteur de gale

chez un singe sud-americain. Bulletin et Annales de
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1 963c. Les acariens producteurs de gale chez

les lemuriens et les singes avec une etude des Psorop-
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Sciences Naturelles de Belgique, 39(32): 1-125.

1 966. Les acariens producteurs de gale chez les

lemuriens et les singes II. Nouvelles observations avec
description d'une espece nouvelle. Acarologia, 8: 95-
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. 1968. Notes sur trois acariens remarquables

. 1981. Hennig's parasitological method: A pro-
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. 1985. Historical ecology: A new approach to

(Sarcoptiformes). Acarologia, 10: 276-291.

FuTUYMA, D. J., AND M. Slatkin, EDS. 1983. Cocvo-
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pp.

Hershkovitz, P. 1977. Living New World Monkeys
(Platyrrhini), vol. 1 . University of Chicago Press, Chi-
cago, 1117 pp.

. 1984. Taxonomy of squirrel monkeys genus

Saimiri (Cebidae, Platyrrhini): A preliminary report
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Lavoipierre, M. M. J. 1964a. A note on the family
Psoralgidae (Acari: Sarcoptiformes) together with a
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parasitic on primates. Acarologia, 6: 342-352.

. 1964b. A new family of acarines belonging to

the Suborder Sarcoptiformes parasitic in the hair fol-
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NiTECKi, M. H., ED. 1983. Coevolution. University of
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OConnor, B. M. 1984. Co-evolutionary patterns be-
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Perkins, E. M., and W. C. Meyer. 1980. The phy-
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259

331-346. In Ciochon, R. L., and A. B. Chiarelli, eds., R. L., and A. B. Chiarelli, eds., Evolutionary Biology

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260 HELDIANA: ZOOLOGY

Notes on Bolivian Mammals

2. Taxonomy and Distribution of Rice Rats

of the Subgenus Oligoryzomys

Nancy Olds and Sydney Anderson

ABSTRACTS

There are at least three kinds of small, long-tailed rice rats of the subgenus Oligoryzomys
(genus Oryzomys) in Bolivia. We use the names Oryzomys microtis (including O. fornesi), O.
chacoensis, and O. longicaudatus for these taxa. The correctness of these names is less certain
and resolution of the nomenclatorial questions awaits study of specimens from outside Bolivia.
The ranges of O. chacoensis and O. microtis are at low elevations and probably overlap to some
degree geographically. The range of O. longicaudatus is in the highlands. No sample from one
locality includes specimens of more than one species; therefore, ecological and microgeographic
differences at places where any two of the three species meet are unknown. Further study may
reveal other and more cryptic species within Bolivia. No one measurement or other characteristic
that we have studied will unequivocally distinguish all adult specimens of any one of the three
kinds from Bolivia. Geographic variation probably occurs within as well as beyond Bolivia in
at least two of the three species (the most uniform seems to be O. chacoensis), but more material
is needed to describe such patterns. We refrain from using subspecies names in consideration
of an ignorance of both geographic patterns of variation and the status of available names. The
only name in the subgenus with a Bolivian type locality, O. chaparensis, is tentatively considered
a synonym of O. microtis.

Existen en Bolivia por lo menos tres clases de pequefias ratas arroceras de cola larga del
subgenero Oligoryzomys (genero Oryzomys). Nosotros usamos los nombres de Oryzomys mi-
crotis (incluyendo O. fornesi), O. chacoensis, y O. longicaudatus para estos taxa. La exactitud
de estos nombres es menos cierta y la resolucion de problemas de nomenclature necesita estudio
de esp>ecimenes de fuera de Bolivia. Los rangos de distribucion de O. chacoensis y O. microtis
estan en bajas elevaciones y probablemente se superponen geograficamente en algun grado.
Ninguna muestra de una localidad contiene especimenes de mas de una especie. Diferencias
ecologicas y microgeograficas son desconocidas en lugares donde cualquiera de las dos o tres
especies se encuentran. Ulterior estudio puede revelar otras y mas cripticas especies en el interior
de Bolivia. Ninguna medida u otra caracteristica que nosotros hemos estudiado separara cla-
ramente todos los especimenes adultos de Bolivia. Variacion geografica ocurre probablemente
dentro asi como fuera de Bolivia en al menos dos de las tres especies (la mas uniforme parece
ser O. chacoensis), pero mas material es necesario para describir tales patrones de variacion.
Nos abstenemos de usar nombres subespecificos en consideracion de la ignorancia de los
patrones de variacion y el estado de los nombres disponibles. El unico nombre en el subgenero
con una localidad tipica en Bolivia, O. chaparensis, es tentativamente considerado como O.
microtis.

From the Department of Mammalogy, American Mu-
seum of Natural History, New York, NY 10024.

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 261

Existem no minimo tres tipos de pequenos ratos-de-arroz, de cauda longa, pertencentes ao
subgenero Oligoryzomys (genero Oryzomys) na Bolivia. A estes taxa damos os nomes de Ory-
zomys microtis (incluindo O. fornesi), O. chacoensis, e O. longicaudatus. A precisao destes
nomes e incerta, e resolu^oes da nomenclatura aguardam estudos de especimes nao bolivianos.
O. chacoensis e O. microtis ocorrem em eleva?6es baixas, e as duas esjiecies provavelmenle
coocorrem em algumas areas. Oryzomys longicaudatus ocorre em areas montanhosas. Amostras
de um so local nao incluem mais do que uma esF)ecie, e nao se conhecem diferen9as ecologicas
ou microgeograficas em areas onde duas ou mais especies possam coocorrer. Com futuros
estudos, novas especies, mais ocultas, poderao ser encontradas na Bolivia. Nenhum unico
carater que estudamos pode inequivocamente separar especimes adultos dos tres tipos de
Oligoryzomys na Bolivia. Varia96es geograficas provavelmente ocorrem na Bolivia, bem como
em outras areas, mas mais material e necessario para poder documentar quaisquer padroes
geograficos (O. chacoensis parece ser a esp>ecie mais uniforme em aparencia). Dada a falta de
conhecimento, ambos dos padroes geograficos, e da disponibilidade dos nomes, nao usamos
nomes de subespecies. O unico subgenero com localidade de tipo na Bolivia, O. chaparensis,
e aqui lentativamente considerado como sinonimo de O. microtis.

Introduction

Bolivian landscapes range from less than 300 m
to more than 4000 m, and habitats range from the
humid lowland Amazonian tropical forests and
subtropical savannahs to the high barren plains
and snow-capped peaks of the Andean altiplano
(fig. 1 ). Habitats may change abruptly, often within
only a few kilometers. The mammals of Bolivia
are also diverse, and provide excellent opportu-
nities for ecological and taxonomic studies of broad
scope. However, the animals are poorly known
(Mares & Genoways, 1982); before satisfactory
general conclusions can be reached, the Bolivian
species need to be clearly delimited, both mor-
phologically and geographically.

Mice of the genus Oryzomys occur throughout
South America and are important members of
small mammal communities (Myers & Carleton,
1981; Mares et al., 1981; Alho, 1982; O'Connell,
1982;Streilein, 1982a-c; Viega-Borgeaud, 1982).
We examined critically one subgenus of Oryzo-
mys, Oligoryzomys, in Bolivia to determine how
many species are present and where they occur.
For a general description of the subgenus, see Myers
and Carleton (1981, pp. 9-12).

The subgenus Oligoryzomys needs revision.
There are few published studies of the more than
45 named forms (Tate, 1932; Ellerman, 1941; Ca-
brera, 1961; Myers & Carleton, 1981; Honacki et
al., 1982). Myers and Carleton (1981) studied
Oligoryzomys from Paraguay, where they recog-
nized three species: Oryzomys nigripes. O. cha-

coensis, and O. fornesi. They also clarified no-
menclatorial questions relating to the 'name
Oryzomys nigripes. Since Paraguay borders Boliv-
ia, this recent study was used as our starting point.
We used the same measurements and comparable
analyses. We assumed that O. chacoensis and O.
fornesi occurred also in southeastern Bolivia, near
the Paraguayan border. If O. nigripes occurs in
Bolivia, the most probable place for it is in eastern
Santa Cruz, from which no specimens are now
available.

At least six names have been used in the liter-
ature or in museum collections for Bolivian sjdcc-
imens of the subgenus Oligoryzomys: Oryzomys
longicaudatus and O. stolzmanni (of the high-
lands); O. chaparensis (from the lowlands of Co-
chabamba); O. nigripes (used for all forms); O.
delicatus (used for a few specimens at middle el-
evations); and 0./7ave5ce/ts (lowland). These names
have been used with considerable uncertainty in
the past (see summaries in Tate, 1932, and espe-
cially Myers & Carleton, 1981). One cause of this
problem is that the original descriptions are vague
or apply equally well to more than one species of
Oligoryzomys.

The taxonomic confusion can be resolved by
detailed study of adequate numbers of museum
specimens, and the nomenclatorial confusion re-
solved by comparisons with type specimens. Re-
newed interest in South American mammals has
resulted in more specimens, which will help in
these tasks.

262

FIELDIANA: ZCX)LOGY

Brasi

Chile

Argentina

V _

- II

Fig. 1 . Map of Bolivia showing the departments and the 500- and 3000-m contour lines. Be, Beni; Ch, Chuquisaca;
Co, Cochabamba; LP, La Paz; Or, Onaro; Pa, Pando; Po, Potosi; SC, Santa Cniz; Ta, Tarija.

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

263

Methods

We examined specimens in the collections of
the American Museum of Natural History
(AMNH); Academy of Natural Sciences in Phila-
delphia (ANSP); California Academy of Sciences
(CAL): Field Museum of Natural History (FMNH);
Museum of Vertebrate Zoology, University of Cal-
ifornia (MVZ); Museum of Zoology, University of
Michigan (UMMZ); and United States National
Museum of Natural History (USNM).

Measurements were obtained as follows: exter-
nal measurements are from the original labels or
field notes, when available, or remeasured (fluid-
preserved specimens only). The following cranial
measurements were taken to the nearest 0.0 1 mm
on a craniometer (see Anderson, 1 968) following
Myers and Carleton (1981) and Musser (1979):

ZN = depth of zygomatic notch
LR = length of rostrum from tip of nasals to
posterior edge of zygomatic notch
GLS = greatest length of skull
ZB = zygomatic breadth
BB = breadth of braincase
BIC = breadth of interorbital constriction
LIF = length of incisive foramen
LPB = length of palatal bridge
LMl, LM2, LM3 = crown lengths of upF>er mo-
lars
WMl, WM2, WM3 = crown widths of upjjer
molars
MM = greatest breadth across molars (labial
edges)
LMX = crown length of upp>er toothrow
LNP = length of nasal projection
LD = length of diastema
LB = length of bulla
HB = height of bulla
TL = total length, including tail

T = tail length
HP = length of hind foot, including claw
E = length of ear, from notch

Specimens were sorted by age, using dental cri-
teria as outlined here (following Myers & Carleton,
1981):

Age Class I: M' not erupted or newly erupted,

M- unworn.
Age Class II: M^ slightly to moderately worn,

but not flat; M- slightly worn; enamel island

formed by the isolation of the internal part of

the mesoflexus of M-.

Age Class III: M ' flat or slightly concave; enam-
el island of above well isolated; M' and M-
substantially worn.

Age Class IV: M^ concave; enamel island oblit-
erated; teeth well worn, but main cusps still
discernible.

Age Class V: M ' and M- flat or concave; folding
pattern obliterated.

Individuals in age class I were examined, mea-
sured, and included in mapping geographic ranges,
but were excluded from statistical treatment of
character variation.

Sp)ecimens examined are listed by locality and
museum catalogue number in the Appendix. Lo-
calities are plotted in Figure 2. Statistical analyses
were done using the computer facilities of the City
University of New York (CUNY) and programs
from SAS (Statistical Analysis System) Institute,
Inc. (1982).

Taxonomy

Our knowledge of geographic variation and the
status of some of the names is sketchy at best, so
we do not use subspecific names. The name Ory-
zomys (Oligoryzomys) longicaudatus stolzmanni
was first used by Hershkovitz (1940, p. 81), by
inference for Bolivian populations, but we are not
certain that O. stolzmanni (Thomas, 1894; type
locality Huambo, 3700 ft, department of Ama-
zonas, Peru) and O. longicaudatus (type locality
restricted to Valparaiso, Chile) are conspecific or
that the Bolivian specimens are conspecific with
either. The resolution of these problems awaits
further study beyond Bolivia.

We noticed no diflerence between Bolivian spec-
imens from Beni and Brazilian SF)ecimens from
the vicinity of the typ>e locality of O. microtis (Low-
er Rio Solimoes, 50 mi above mouth), and thus
the subspecific name Oryzomys microtis microtis
might be applied to Bolivian specimens. However,
we need to know more about geographic variation
in the species (see Remarks under O. microtis).
Some other names that may refer to consjsecific
populations and thus be relevant as possible sub-
specific epithets are as follows:

1. Oryzomys destructor from lowland eastern
Peru has been assigned to longicaudatus but may
prove to be consp)ecific with microtis; if so, de-
structor is the senior synonym. If destructor and
microtis are conspecific, the northern Bohvian mice

264

HELDIANA: ZOOLOGY

Fig. 2. Distribution of Bolivian Oligoryzomys: • = Oryzomys microtis; O = O. longicaudatus; X = O. chacoensis.

might be known as Oryzomys destructor destruc- Bolivia on geographic and ecological grounds, al-

tor, or perhaps O. destructor microtis. though no specimens are presently available. Myers

2. Oryzomys fornesi from northern Argentina and Carleton (1981) referred specimens from San

might also be expected to occur in southeastern Joaquin, Beni, to O. fornesi.

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

265

3. Oryzomys mattogrossae from Brazil just to
the east of Bolivia is conspecific with O. microtis
and has its type locality nearest the department of
Beni, where most of the Bolivian specimens have
been taken; thus, for geographic reasons, it must
be considered in any future subspecific determi-
nations. The uncertain specific status of utiariten-
sis from the same type locality as mattogrossae is
noted below. If it is conspecific with mattogrossae,
utiaritensis should be regarded as a synonym
thereof at the subspecies level also.

We concur with Myers and Carleton (1981) in
the recognition of two species groups of Oligory-
zomys. These are the smaller-bodied, small-toothed
Oryzomys microtis. O.Jlavescens, and O. delicatus,
and the larger-bodied, larger-toothed O. chacoen-
sis, O. longicaudatus, and O. nigripes.

In the Species Accounts section, synonymies
cover only Bolivian records. Other relevant names
are discussed elsewhere in the text.

Results of Statistical Analyses
Sex and Age Variation

We examined sex and age variation in Oryzomys
microtis. O. longicaudatus. O. chacoensis. and O.
flavescens (from Uruguay). Males are slightly larg-
er in general; the average size differences, consid-
ering all characters for each species, are 2.7%, 1 .6%,
2.5%, and 2. 1%, respectively. We examined sexual
dimorphism in all measurements for each species
by one-way and two-way analyses of variance of
sex and of sex and age for each species. Results of
two-way analyses of variance of sex and age (using
all specimens of tooth wear class II or greater) on
specimens of O. microtis from the department of
Beni, Bolivia, are presented in Table 1 . Our results
are roughly comparable to those of Myers and
Carleton (1981). We did not separate sexes in fur-
ther statistical analyses, although we watched for
unbalanced sex ratios in samples when interpret-
ing results.

Age variation is more difficult to assess, as no
detailed study has been published. Mice of this
subgenus probably continue to grow for most of
their lives (see Myers &. Carleton, 1981), although
the rate slows with age. We performed least-squares
regression analyses (General Linear Models pro-
cedures of SAS) on five variables: greatest length
of skull (GLS), total length (TL), zygomatic breadth

(ZB), length of the first upper molar (LMl), and
length of the hind foot (HF), by species, to plot i
graphically the relationship between relative age I
and size. Based on study of these graphs, we de-
cided to include age classes II-V in further anal-
yses, as did Myers and Carleton (1981). Most spec- I
imens were juveniles (age class I) or young adults
(age classes II-III). Few specimens of age classes
IV-V were present among the species we exam-
ined. See Tables 2 and 3 for the mean adult ages
of the specimens studied.

Principal Components

We analyzed principal components (with the
Princomp procedure of SAS on a correlation ma-
trix) using measurements taken on individuals of
all sp)ecies (the approximate numbers of specimens
are in tables 2 and 3, some specimens were ex-
cluded because of missing measurements). When
plotted, the first principal component tends to sep-
arate the smaller species {O. microtis and O. fla-
vescens) from the larger species {O. chacoensis and
O. longicaudatus). The percentage of the totJil vari-
ance accounted for by the first three components
is 68.7%.

Discriminant Analysis

We performed several different discriminant
analyses (using the SAS programs Discrim, Step-
disc, and Candisc). A stepwise discriminant anal-
ysis chose the following characters (in order of
selection): tail length, breadth of interorbital con-
striction, diastema length, length of incisive fo-
ramina, rostral length, bullar length, length of
hind foot, molar breadth, length of nasal projec-
tion, length of zygomatic notch, total length, zy-
gomatic breadth, length of palatal bridge, and length
of maxillary toothrow.

Using the SAS program Discrim, we were able
to test the posterior probability of group mem-
bership. In all analyses, we used as "known" groups
samples of the species Oryzomys microtis from
Beni, Bolivia; O. flavescens from Uruguay; O. lon-
gicaudatus from the department of La Paz. Boliv-
ia; and O. chacoensis from Paraguay. Plots of the
individual mice on the first two canonical corre-
lates showed little overlap among the four species.
We assumed that the following "unknowns" be-
longed to one of the four "known" groups. We
submitted as "unknowns" the holotypes of O. mi-

266

HELDIANA: ZOOLOGY

crotis, O. mattogrossae, and O. chaparensis; the
paratype of O. chaparensis; and two specimens
from Beni assigned by Myers and Carleton (1981)
to O. chacoensis (see Remarks under O. microtis).
The holotypes of O. microtis and O. mattogrossae
were assigned to the O. microtis species sample,
with a posterior probability of more than 95%.
The holotype of O. chaparensis was also assigned
to O. microtis (P = 0.987). The paratype of O.
chaparensis was assigned to O. microtis (P = 0.985).
Several other specimens that we wanted to test,
including the type of O. delicatus, had missing
values and could not be used in the analysis. We
recognize that the validity of the taxonomic con-
clusions based on these analyses depends on the
correctness of the initial assumption, and that it
needs further testing. Not all measurements were
available for all skulls, but in each comparison
the largest possible subset was used.

Species Accounts

Subgenus Oligoryzomys Bangs, 1 900

Diagnosis— Within the genus Oryzomys, the
subgenus Oligoryzomys is distinguished by small
size and delicate structure throughout, tail rela-
tively long, hind foot long and slender; skull small,
delicate, interorbital region narrow, outer edges of
frontals squarish but unbeaded, braincase smooth
and unridged, zygomatic plate narrow and with
slight forward projection, molar teeth and incisors
small and delicate but with cusp pattern like other
Oryzomys (description adapted from Bangs, 1 900).
Oryzomys of the subgenus Microryzomys are also
small and delicate, but they differ from Oligory-
zomys in having a more slender rostrum, shallow-
er zygomatic notch, shorter and more rounded
braincase, sphenofrontal foramen and squamo-
soalisphenoid groove present, and karyotype with
a low FN/2n ratio (Myers & Carleton, 1981, p.
12).

Oryzomys chacoensis Myers and Carleton, 1981

Oryzomys chacoensis Myers and Carleton, 1 98 1 , p. 19
(typ)e locality "419 km by road NW Villa Hayes
[alongside the Trans Chaco Highway], Dept. Bo-
queron, Paraguay").

Diagnosis— C>ryzc>my5 chacoensis Myers and
Carleton (1981, p. 20) was diagnosed as "A me-

Table 1 . Nongeographic variation in Oryzomys mi-
crotis. Results of analysis of variance of Bolivian O. mi-
crotis.

Char-

Error

F (inter-

acter

d.f.

F(sex)

F (age)

action)

3.75

0.44

0.52

26.31***

1.87

1.35

GLS

22.69***

1.77

0.75

13.65***

1.19

1.33

6.57*

0.41

2.55*

BIC

0.74

1.79

1.22

LIF

4.64*

1.71

0.51

LPB

4.32*

0.81

1.01

LMl

0.08

2.38*

0.76

LM2

1.38

5.37***

5.57***

LM3

4.23*

1.93*

1.82

WMl

0.51

1.38

1.68

WM2

0.82

1.43

1.06

WM3

0.17

1.90*

1.24

3.07

3.08**

1.81

LMX

1.53

2.38*

1.36

LNP

15.05***

1.26

1.73

10.87**

1.23

1.58

5.45*

0.48

0.71

1.77

0.41

0.83

5.36*

1.38

1.21

0.02

1.65

2.34*

6.41*

0.95

0.67

0.02

1.53

0.84

* P < 0.05; ** P < 0.01; *** P < 0.001.

dium-sized species of the subgenus Oligoryzomys
unique in its whitish underside with hair white to
the base on the chin and throat, relatively long
ears having hairs on inner surface with unusually
short or absent dark basal bands, small but dis-
tinctive tufts of orangish hairs anterior to the ears,
and karyotype with 2n = 58, FN = 74."

Distribution in Bolivia— We examined spec-
imens (see Appendix) from the departments of
Cochabamba, Santa Cruz, and Tarija (fig. 2). Myers
and Carleton (1981) reported two specimens from
Beni. These specimens have been restudied and
are here reassigned to Oryzomys microtis (see Re-
marks under that species). Habitat of O. chacoen-
sis is grassland and thomscrub. The highest known
elevation is 640 m (Rio Lipeo).

General Description and Comparisons—
Cranial and external measurements are listed in
Table 2. The dorsal pelage is rufous and heavily
lined with black hairs. The venter is white and
sharply distinguished from sides, which are clearer
than dorsum. Frequently a thin orange line sep-
arates the sides from the belly. The cheeks are
slightly paler than the rest of the face. The tail is
dark gray, weakly bicolored, and long relative to
the body. The hind feet are whitish above. Juve-

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

161

Table 2. Mean, standard deviation, and range (in mm) for Oryzomys chacoensis and O. longicaudatus.

O. chacoensis

O. longicaudatus

Bolivia

Paraguay

Argentina

Bolivia

Character

(N = 15)

(N = 16)

(N = 5)

(N = 34)

1.15 ± 0.28

1.30 ± 0.20

1.15 ± 0.12

1.02 ± 0.20

(0.88-1.59)

(0.90-1.61)

(0.94-1.24)

(0.57-1.36)

7.27 ± 0.45

7.39 ± 0.57

7.67 ± 0.34

7.14 ± 0.72

(6.62-7.94)

(6.57-8.86)

(7.25-8.07)

(4.49-8.15)

GLS

24.90 ± 0.82

24.52 ± 1.09

26.12 ± 0.32

25.01 ± 1.39

(23.80-26.69)

(22.52-27.12)

(25.79-26.43)

(21.06-27.29)

13.10 ± 0.59

12.76 ± 0.62

13.64 ± 0.09

13.17 ± 0.65

(11.94-13.98)

(11.91-14.27)

(13.54-13.71)

(11.55-13.98)

11.33 ± 0.41

11.16-0.34

11.39 ± 0.06

11.25 ± 0.37 .

(10.42-11.87)

(10.59-11.59)

(11.35-11.45)

(10.58-11.96)

BIC

3.89 ± 0.20

3.76 ± 0.10

3.86 ±0.13

3.63 ± 0.22

(3.52-4.27)

(3.64-3.95)

(3.76-4.04)

(3.35-4.17)

LIF

4.62 ± 0.26

4.71 ± 0.39

5.07 ± 0.29

4.68 ± 0.27

(4.25-5.20)

(4.17-5.56)

(4.64-5.31)

(4.23-5.26)

LPS

4.14 ± 0.22

3.98 ± 0.26

4.22 ± 0.31

3.97 ± 0.21

(3.65-4.54)

(3.50-4.65)

(3.75-4.52)

(3.49-4.41)

LMl

1.68 ± 0.10

1.63 ± 0.07

1.74 ± 0.5

1.59 ± 0.09

(1.47-1.84)

(1.54-1.79)

(1.70-1.83)

(1.42-1.81)

LM2

1.09 ± 0.06

1.10 ± 0.07

1.14 ± 0.07

1.05 ± 0.04

(0.96-1.18)

(1.00-1.28)

(1.08-1.26)

(0.97-1.15)

LM3

0.84 ± 0.05

0.80 ± 0.04

0.84 ± 0.04

0.81 ± 0.05

(0.74-0.92)

(0.73-0.88)

(0.78-0.88)

(0.67-0.92)

WMl

1.07 ± 0.07

1.03 ± 0.03

1.11 ± 0.05

1.04 ± 0.06

(0.93-1.24)

(0.96-1.08)

(1.05-1.17)

(0.91-1.15)

WM2

1.02 ± 0.07

1.00 ± 0.04

1.04 ± 0.06

0.99 ± 0.06

(0.86-1.14)

(0.93-1.06)

(0.97-1.11)

(0.87-1.09)

WM3

0.87 ± 0.05

0.85 ± 0.03

0.90 ± 0.05

0.83 ± 0.06

(0.78-0.97)

(0.78-0.92)

(0.84-0.94)

(0.67-0.95)

4.66 ± 0.20

4.57 ± 0.17

4.70 ± 0.13

4.61 ± 0.27

(4.32-4.99)

(4.34-5.09)

(4.58-4.87)

(4.03-5.08)

LMX

3.60 ±0.12

3.50 ± 0.13

3.71 ± 0.15

3.46 ± 0.14

(3.29-3.78)

(3.23-3.68)

(3.61-3.98)

(3.20-3.77)

LNP

1.43 ± 0.24

1.59 ± 0.17

1.49 ± 0.17

1.40 ± 0.25

(1.04-1.77)

(1.35-2.07)

~ (1.29-1.64)

(0.99-2.04)

5.66 ± 0.34

5.63 ± 0.44

5.89 ± 0.24

6.04 ± 0.47

(5.17-6.33)

(5.02-6.44)

(5.63-6.18)

(4.96-7.07)

3.54 ± 0.14

3.60 ± 0.14

3.75 ± 0.03

3.47 ± 0.20

(3.28-3.81)

(3.28-3.82)

(3.72-3.78)

(3.07-3.84)

2.75 ± 0.18

2.73 ± 0.15

2.72 ± 0.06

2.76 ± 0.32

(2.43-3.04)

(2.40-2.96)

(2.66-2.76)

(2.23-3.32)

227.93 ± 12.45

219.50 ± 12.29

229.00 ± 8.34

217.68 ± 16.88

(204-252)

(194-241)

(215-237)

(172-252)

134.27 ± 10.26

126.06 ± 8.46

138.00 ± 5.70

124.15 ± 12.18

(112-150)

(112-139)

(130-145)

(87-145)

25.33 ± 1.33

24.38 ± 1.54

23.40 ± 2.30

25.13 ± 1.20

(23-28)

(23-29)

(21-26)

(22-28)

16.70 ± 1.33

15.07 ± 1.69

18.40 ± 1.14

15.03 ± 1.72

(14-18)

(12-17)

(17-20)

(12-18)

AGE

2.65 ± 0.82

2.31 ± 0.68

2.55 ± 0.87

3.04 ± 0.88

(2-4)

(2-5)

268

HELDIANA: ZOOLOGY

niles are grayer, as are all young Oligoryzomys in
Bolivia, but this is especially noticeable on the
venter, which has mixed gray and white hairs. In-
cisive foramina extend posteriorly to the anterior
edge of M' or slightly beyond. Alisphenoid strut
(a strut of the alisphenoid bone that covers the
lateral part of the alisphenoid canal, see Musser,
1 982, p. 29) is generally absent (table 4). The sides
of the interorbital constriction are divergent pos-
teriorly (less parallel-sided than in O. longicau-
datus) (fig. 3). The following quotes are from Myers
and Carleton (1981, first and second quotes, p. 2 1 ,
third quote, p. 24):

Oryzomys chacoensis differs from O. cha-
parensis Osgood (1916) primarily in color
pattern: the type of chaparensis is much
darker and less hispid dorsally, yellowish
ventrally, lacks the orange tufts anterior to
the ears, and has a grayish throat. The distal
portions of the nasals of the holotype flare
laterally to an extent not seen in chacoensis.

amined the two darker ones. When skulls are com-
pared, the two darker mice are among the older
and larger individuals in the series. They seem to
have relatively broader braincases than most, but
there is no character shared by these two that is
not also seen in one or more of the others. In a
discriminant analysis (SAS program Discrim) the
posterior probability of membership (in the four
reference species) allies these two mice with O.
chacoensis (ansp 18187, P = 0.997, and ansp
18188, P = 0.954 with chacoensis). We have as-
signed these mice to O. chacoensis. Since Rio Li-
pto is in the area where these two species meet,
further study there should reveal whether sym-
patry exists and if species differences remain dis-
tinct.

Specimens from Argentina (listed in the Ap-
pendix) are similar in coloration to both Para-
guayan and Bolivian specimens, but are woolier.
This sample extends the known range of O. cha-
coensis into the department of Jujuy, northwestern
Argentina.

Oryzomys chacoensis can be distinguished
from fornesi, with which it occurs sympat-
rically, by its larger size (maxillary toothrow
usually > 3.3mm, ears usually > 15mm),
characteristic karyotype, lack of preputial
glands, and in most specimens by its lack of
buff" on the belly.

. . . the hind feet of chacoensis are relatively
short compared to those of the more terres-
trial fornesi.

In reference to the last point, however, our cal-
culations of length of hind foot relative to length
of head and body (using data from tables 2 and 3)
are about 26% for both O. chacoensis and O. mi-
crotis (including fornesi). Comparison of speci-
mens also reveals no noticeable difference.

Remarks— This species is clearly distinct from
O. microtis (here including O. fornesi). In most
morphometric characters, O. chacoensis grossly
resembles O. longicaudatus from Bolivia and O.
nigripes from Brazil.

Our specimens from Santa Cruz show some
variation in coloration: two specimens (amnh
247772-247773) are pale. The specimens from
Tarija resemble the Paraguayan samples more
closely than do the specimens from Santa Cruz.

A series of 13 sp)ecimens from Rio Lipeo in-
cludes two with darker pelage that resemble O.
longicaudatus. We measured these mice and ex-

Oryzomys longicaudatus (Bennett, 1832)

Mus longicaudatus Bennett, 1832, p. 2 (type locality
"In trees in Chile," restricted to Valparaiso by Ca-
brera, 1961, p. 391).

Oryzomys longicaudatus: Thomas, 1898, p. 3 (Aguai-
renda Mission, San Francisco, perhaps not O. lon-
gicaudatus); Thomas, 1926, p. 194 (Tupiza).

Oryzomys Stolzmanni: Thomas, 1902, p. 130 (Cha-
ruplaya, Choro); Neveu-Lemaire and Grandidier,
1 9 1 1 , p. 9 (Charuplaya, Choro).

Oryzomys ^X>;flavescens groMTp: Thomas, 1925, p. 578
(Carapari, perhaps not O. longicaudatus).

Oryzomys stolzmanni stolzmanni: Sanborn, 1950, p.
2 (Rio Aceramarca, Cocapunco, Nequejahuira,
Okara, Pitiguaya, Pongo).

Diagnosis— Bolivian specimens referred to this
species differ from other Bolivian Oligoryzomys
in that they are larger (especially in the size of the
teeth) than O. microtis and lack the buffy wash on
the venter; are grayer-bellied, darker, and less griz-
zled dorsally than O. chacoensis; and occur at gen-
erally higher elevations (at least in northern Bo-
livia).

Distribution in Bolivia— O. longicaudatus is
found in the valleys and mountains of the Andes
from at least middle elevations (1200 m, Entre
Rios) up to 3720 m (Poopo). Specimens from near
Camiri and Cuyambuyo in southern Bolivia are
from elevations of 780 to 1000 m. Specimens (see
Appendix) have been reported or examined from

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

269

Fig. 3. Dorsal and ventral views of skull of (left), Oryzomys longicaudatus stolzmanni; (middle), O. chacoensis,
and (right), O. microtis microtis. Specimens in the American Museum of Natural History. Scale at lower right represents
10 mm.

the departments of Chuquisaca, Cochabamba, La
Paz, Oruro, Polosi, Santa Cruz, and Tarija (see
map, fig. 2).

General Description and Comparisons—
Cranial and external measurements are presented
in Table 2. The belly is gray, rarely washed with
buff; upper parts are a dark brown lined with black
hairs, and often the sides are more rufous. A thin
orange lateral line may be present. Nose and face

are dark with paler cheeks. Often there are pale
spots just behind the ears. Hind feet are pale above.
The tail is brown above, weakly to strongly bi-
colored, and relatively long compared to that of
any other Bolivian Oligoryzomys. Incisive foram-
ina generally extend to the anterior edge of M' or
slightly behind. Alisphenoid strut is generally ab-
sent (table 4). The interorbital area is slightly more
constricted on the average than that of O. cha-

270

HELDIANA: ZOOLOGY

coensis, and has relatively parallel sides (see fig.
3).

Remarks— Oryzomys longicaudatus may be a
composite of more than one species. Oryzomys
destructor and O. stolzmanni were considered sub-
species of 6>. longicaudatus by Cabrera ( 1 96 1 ) and
full species by Soukup (1961). Gardner and Patton
(1976) found four karyotypic variants in speci-
mens they assigned to O. longicaudatus, and com-
mented that these may represent four separate
species. One of their karyotypic variants, no. 2, is
represented by 1 7 specimens from Balta, depart-
ment of Loreto (now Acayali), Peru, at an eleva-
tion of about 300 m. Myers and Carleton (1981,
p. 26) mentioned that the karyotype of these spec-
imens as reported by Gardner and Patton (1976)
agreed better with results they obtained for O.for-
nesi (= O. microtis) than for either O. chacoensis
or O. longicaudatus. We have examined these
specimens and judge that they are clearly different
from O. longicaudatus and belong with either O.
microtis or O. destructor (Balta is near the type
locality of O. destructor). The inclusion of karyo-
type no. 3 under O. longicaudatus, represented by
one female from Venezuela, also needs verifica-
tion.

We have five specimens from Caracato, de-
partment of La Paz (amnh 248977-248981), that
are quite different from other Bolivian Oryzomys
longicaudatus. The pelage closely resembles that
of O. nigripes from Paraguay in color and texture;
it is paler and more obviously lined with black
hairs dorsally than the pelage of typical O. lon-
gicaudatus. They are also slightly larger bodied,
but four of the five are old individuals (age classes
IV-V) and this may account for the larger size.
Because of the small sample size and the single
locality, we chose not to distinguish this popula-
tion taxonomically.

It is possible that Andean populations may oc-
cupy areas that are rather widely scattered and
physically isolated in different valleys or side can-
yons. If so, there may be considerable geographic
variation. More specimens are needed to detect
and describe such patterns.

The sjjecimens from near Cuyambuyo are at
comparatively low elevations (980 to 1000 m) for
O. longicaudatus, although the species occurs at
progressively lower elevations farther south (Os-
good, 1916). The two localities near Cuyambuyo
are only about 60 km from Rio Lipeo, where O.
chacoensis occurs. The SAS Discrim program
analysis allied the largest adult specimen from near
Cuyambuyo (ummz 155891) with O. longicauda-

tus (F = 0.956). There are problems in distinguish-
ing O. longicaudatus and O. chacoensis solely by
cranial morphology in all areas.

Oryzomys microtis Allen, 1916

Oryzomys (Oligoryzomys) microtis Allen, 1 9 1 6, p. 525
(type locality Lower Rio Solimoes, 50 mi above
mouth [80 km from its confluence with Rio Negro,
Amazonas, Brazil]).

Oryzomys chaparensis Osgood, 1 9 1 6, p. 205 (holotype
and paratype, type locality Todos Santos, on Rio
Chapare, Department of Cochabamba, Bolivia);
Gyldenstolpe, 1932, p. 25 (Todos Santos); Myers
and Carleton, 1981, p. 38 (Todos Santos); all in
reference to the same specimens.

Oryzomys fornesi Massoia, 1973, p. 22 (type locality
Naineck, Dept. Rio Pilcomayo, province of For-
mosa, Argentina); as used by Myers and Carleton,
1981, p. 25 for five specimens fVom San Joaquin,
Beni, Bolivia.

Diagnosis— Allen described Oryzomys microtis
as being readily distinguished from other Olig-
oryzomys by its pale coloration, relatively small
ears, and tail less than half total length. However,
the tail of the holotype is more than half of the
total length, a discrepancy noted by Goodwin
(1953). Massoia (1973) diagnosed O. fornesi, here
considered a junior synonym of O. microtis, as the
smallest species of the subgenus in Argentina, hav-
ing the ears covered with short ochraceous hairs,
the pterygoids short (shorter than the molar se-
ries), the interorbital constriction narrow, and the
incisive foramina not extending to the molar se-
ries. In Bolivia, it can be distinguished by its small
body size, toothrow generally less than 3.3, gray-
buffy color of the belly, and relatively short tail in
proportion to body.

Distribution in Bolivia— Specimens (see Ap-
pendix) from the lowlands of Bolivia, in the de-
partments of Beni, Cochabamba, Pando, La Paz,
and Santa Cruz have been examined. Habitat in-
cludes marshes and wet forests up to an elevation
of 1 800 m (Guanay, see fig. 2).

General Description and Comparisons —
Smallest of the Bolivian Oligoryzomys, with small
teeth, toothrow generally less than 3.3 (averaging
3.10). Massoia ( 1 973) gave general measurements
useful in distinguishing this species from other Ar-
gentine species of Oryzomys (Oligoryzomys) as fol-
lows: length of hind foot generally less than 24,
length of ear generally less than 13, breadth of
braincase less than 10.8, and length of incisive
foramina generally less than 4.5. All of his mea-

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

271

Table 3. Mean, standard deviation, and range (in mm) for Oryzomys microtis and O. flavescens.

O. microtis

O. flavescens

Bolivia

Brazil

Peru

Uruguay

Character

(N = 67)

(N = 20)

(N = 26)

(N = 20)

1.01 ± 0.17

0.98 ±0.18

0.84 ±0.13

1.07 ± 0.15

(0.53-1.46)

(0.78-1.48)

(0.61-1.18)

(0.79-1.32)

6.82 ± 0.51

6.82 ± 0.42

6.60 ± 0.41

6.75 ± 0.43

(5.17-8.16)

(6.16-7.46)

(5.96-7.36)

(5.89-7.39)

GLS

23.54 ± 1.06

23.79 ± 0.88

23.27 ± 0.82

23.41 ± 0.94

(21.24-25.78)

(22.47-25.04)

(21.75-24.53)

(21.66-24.81)

12.28 ± 0.51

12.40 ± 0.44

11.86 ± 0.53

12.05 ± 0.49

(11.42-13.73)

(11.59-13.32)

(11.01-12.93)

(11.25-12.76)

10.78 ± 0.36

10.71 ± 0.23

10.47 ± 0.32

10.70 ± 0.26 •

(9.38-11.78)

(10.35-11.06)

(9.49-10.87)

(10.29-11.22)

BIC

3.76 ± 0.17

3.72 ± 0.14

3.63 ±0.11

3.35 ±0.17

(3.38-4.19)

(3.44-3.93)

(3.45-3.94)

(2.95-3.77)

LIF

3.96 ± 0.27

4.01 ± 0.22

4.00 ±0.18

4.59 ± 0.35

(3.38-4.53)

(3.65-4.37)

(3.73-4.33)

(3.54-5.20)

LPB

4.04 ± 0.32

4.06 ± 0.22

3.98 ± 0.26

3.77 ± 0.19

(3.14-4.76)

(3.64-4.38)

(3.42-4.51)

(3.51-4.06)

LMl

1.46 ± 0.10

1.43 ± 0.07

1.51 ± 0.06

1.47 ± 0.06

(1.22-1.64)

(1.32-1.56)

(1.39-1.66)

(1.37-1.60)

LM2

0.95 ± 0.06

0.95 ± 0.05

0.93 ± 0.08

1.01 ± 0.06

(0.81-1.09)

(0.84-1.06)

(0.67-1.04)

(0.83-1.09)

LM3

0.70 ± 0.07

0.68 ± 0.05

0.66 ± 0.05

0.73 ± 0.04

(0.54-0.85)

(0.59-0.78)

(0.55-0.77)

(0.66-0.80)

WMl

0.94 ± 0.06

0.93 ± 0.06

0.99 ± 0.04

0.97 ± 0.04

(0.78-1.08)

(0.80-1.04)

(0.91-1.10)

(0.93-1.07)

WM2

0.88 ± 0.06

0.89 ± 0.06

0.93 ± 0.04

0.93 ± 0.05

(0.70-1.01)

(0.78-0.99)

(0.82-1.00)

(0.86-1.03)

WM3

0.73 ± 0.06

0.74 ± 0.06

0.80 ± 0.05

0.78 ± 0.04

(0.55-0.85)

(0.58-0.85)

(0.71-0.93)

(0.70-0.85)

4.33 ± 0.21

4.39 ± 0.23

4.33 ± 0.18

4.26 ±0.15

(3.83-4.93)

(3.99-4.81)

(4.07-4.70)

(3.98-4.50)

LMX

3.10 ± 0.17

3.07 ± 0.16

3.14 ± 0.13

3.22 ±0.12

(2.64-3.43)

(2.75-3.34)

(2.93-3.39)

(2.96-3.47)

LNP

1.32 ± 0.22

1.25 ± 0.32

1.20 ± 0.20

1.33 ± 0.14

(0.91-1.74)

(0.75-1.70)

■ (0.81-1.62)

(1.16-1.68)

5.75 ± 0.41

5.78 ± 0.36

5.32 ± 0.40

5.35 ± 0.35

(4.82-6.62)

(5.13-6.51)

(4.49-6.01)

(4.67-5.90)

3.29 ± 0.14

3.18 ± 0.19

3.01 ± 0.12

3.30 ±0.15

(2.94-3.57)

(2.90-3.63)

(2.75-3.27)

(2.91-3.53)

2.46 ± 0.20

2.10 ± 0.23

2.22 ±0.12

2.64 ± 0.23

(1.96-2.87)

(1.72-2.45)

(1.96-2.38)

(2.02-2.94)

185.88 ± 11.12

185.00 ± 10.60

179.77 ± 9.16

198.56 ± 17.66

(165-214)

(164-202)

(160-196)

(160-223)

101.00 ± 6.67

95.44 ± 7.88

101.69 ± 8.13

110.37 ± 11.05

(87-116)

(75-105)

(77-115)

(85-127)

22.81 ± 1.21

21.68 ± 1.34

21.37 ± 1.60

25.21 ± 1.85

(19-25)

(20-25)

(18-25)

(20-28)

14.49 ± 1.04
(12-16)

13.97 ± 1.05
(12-16)

AGE

2.57 ± 0.73

2.35 ± 0.59

2.97 ± 0.88

2.18 ± 0.35

(2-5)

(2-4)

(2-5)

(2-3)

272

FIELDIANA: ZOOLOGY

surements fall within the extremes of our values
for O. microtis, except for measurements of the
toothrow (LMX), in which his are on the upper
end of our range. Values of cranial and external
measurements are listed in Table 3. The venter is
white mixed with gray to gray-bufFy and clear huf-
fy. The dorsum is rufous brown, lined with black
hairs, and has paler sides. Often there is no clear
demarcation between sides and belly. There are
no white spots behind ears. Juveniles are similar
to adults, but show a grayer belly. The tail is dark
above and weakly bicolored. Occasionally orange-
tipped hairs lie anterior to the ear. Hind feet are
white above. Incisive foramina generally extend
posteriorly to the anterior edge of M' and not
beyond. In comparison with adults of O. cha-
coensis and O. longicaudatus, O. microtis has a
shorter skull, longer braincase, shorter incisive fo-
ramina, a relatively broader interorbital constric-
tion with divergent sides, and smaller, narrower
teeth; also, an alisphenoid strut is more often pres-
ent (table 4).

Remarks— This species is similar to Oryzomys
flavescens of Uruguay and Argentina (see Myers
& Carleton, 1981). Specimens of both species were
reported from the same locality, Capitan Solari,
in the province of Chaco, northern Argentina, by
Contreras and Berry (1983). Measurements of 6>.
flavescens from Uruguay are listed in Table 3 (see
also Langguth, 1 963). In comparison with O. mi-
crotis, specimens of O. flavescens have slightly
larger teeth, larger bullae, longer incisive forami-
na, and are generally larger. Oryzomys flavescens
needs to be more clearly defined. Massoia and
Fomes ( 1 967) once synonymized O. flavescens with
O. nigripes, but according to the later analyses of
Massoia (1973) and Myers and Carleton (1981)
these taxa are not consjiecific. Uruguayan speci-
mens oi O. flavescens are clearly different from O.
nigripes from eastern Paraguay. Future studies may
reveal additional relationships, including the pos-
sibility that O. microtis and O.fiilvescens of Cen-
tral America and northern South America are con-
specific (Handley, 1976, referred Venezuelan
specimens to O. fiilvescens).

The name Oryzomys microtis flyrnesi might be
used to reflect the conspecific status oiflyrnesi and
microtis. This subspecific name would apply to at
least the five specimens identified by Massoia
(1973), pending more detailed studies that would
test whether geographic differences warrant con-
tinued subspecific recognition.

Until the relationships of the Andean ""longi-
caudatus -group" are clarified, it is also possible

Table 4. Presence or absence of alisphenoid strut in
five species of Oryzomys (Oligoryzomys) expressed as a
percentage; number of specimens in parentheses.

Alisphenoid strut

Species

Present

Absent

O. chacoensis

2.4(1)

97.6(41)

O. longicaudatus

5.6 (2)

94.4 (34)

O. microtis

35.8 (54)

64.2 (97)

O. flavescens

3.2(1)

96.8 (30) •

O. nigripes

8.3 (3)

91.7(33)

that one or more of the earlier names currently
assigned to this group actually belong with Ory-
zomys microtis, and have priority. For example:
O. stolzmanni (named in 1894, type locality listed
under synonymy) and O. destructor (named in
1 844, type locality: "the house mouse of the 'Plan-
tagen at the border of the forest," eastern Peru,
above 6000 ft, according to Tate, 1932, p. 9; re-
stricted by Cabrera, 1961, to the haciendas along
the Rio Chinchao in the department of Huanuco,
between 900 and 1000 m) have been associated
with O. longicaudatus. These type localities are at
low elevations, which suggests the possible affinity
of these forms with microtis rather than with lon-
gicaudatus. We have examined a series from low
elevation in the department of Pasco, Peru, that
is clearly O. microtis. This locality is near the
type locality of destructor (Cole, 1984). If future
study of the holotype of destructor proves it to be
conspecific with O. microtis, the name of the species
should be O. destructor instead of O. microtis.

We have examined the type specimens of Ory-
zomys microtis (amnh 37091, type locality Lower
Rio Solimoes, 50 mi above mouth, Brazil), O.
mattogrossae (amnh 37542, type locality Utiarity,
Mato Grosso, Brazil), O. utiaritensis (amnh 37541,
type locality Utiarity, Rio Papagaia, Mato Grosso,
Brazil), and O. delicatus (amnh 7317/5925, type
locality Trinidad, West Indies). We judge that the
small lowland Bolivian specimens are O. microtis
and also represent the same species as specimens
referred by Myers and Carleton to O. fornesi (O.
microtis has priority). The Bolivian specimens dif-
fer slightly from Massoia's description of O. for-
nesi in having a shorter average toothrow, other-
wise in being slightly larger. If O. fornesi and O.
microtis are conspecific, it is possible that this dif-
ference in measurements reflects his small sample
size (N = 5), or that there may be significant geo-
graphic variation between Argentine and Bolivian
populations. We further treat O. mattogrossae as
a synonym of O. microtis (O. mattogrossae is not

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

111

included in the synonymy above because the name
has never been used in print for a Bolivian spec-
imen). As mentioned by Myers and Carleton
( 1 98 1 ), O. utiaritensis may belong with O. nigripes.
More specimens from Mato Grosso are needed to
establish the characteristics of the species there.
Oryzomys microtis was considered a synonym of
O. delicatus by Alho ( 1 982) and Pine ( 1 973); how-
ever, because of the slightly damaged condition of
the skin and skull of the holotype of O. delicatus
and the remoteness of its type locality (on the is-
land of Trinidad, Allen & Chapman, 1 897), we
defer any decision on this question.

We have reexamined the two specimens from
Beni that Myers and Carleton (1981) referred to
Oryzomys chacoensis (usnm 39 1 297, 46074 1 ) and
have assigned them to O. microtis. These speci-
mens are young adults (age class II), have external
measurements that fit well with O. microtis and
that are rather small for O. chacoensis (TL 200,
200; T 111, 110; HF 19, 24, respectively), and
have toothrows measuring 3.34 and 3.40, respec-
tively. The toothrows fall on the upper end of the
range for O. microtis and the lower end of the range
for O. chacoensis (tables 2-3). We then compared
their measurements through a discriminant anal-
ysis with the samples ofO.flavescens, O. microtis,
O. chacoensis. and O. longicaudatus. The posterior
probability of membership for both was greatest
for O. microtis (usnm 391297, P = 0.937; usnm
460741,/' = 0.953).

We refer the holotype (fmnh 21330) and the
one paratype (amnh 40787) o^ Oryzomys chapa-
rensis to Oryzomys microtis. The holotype is the
largest and one of the oldest individuals we have
identified as O. microtis. Its greatest skull length
is 26.8, zygomatic breadth 13.4, breadth of brain-
case 1 1.2, crown length of maxillary toothrow 3.5,
tail length 111, and length of hind foot 26. The
skull length, the toothrow, and hind foot length
are larger than other Bolivian O. microtis (see table
3). The pelage coloration of this specimen falls
within the range of variation in O. microtis. For
a comparison of pelage coloration between O. cha-
parensis and O. chacoensis, see the account of O.
chacoensis under General Description and Com-
parisons.

Geographic Variation— We were unable to
detect significant differences in p>elage coloration
between our samples from Brazil, Bolivia, and
Peru. The Brazilian specimens are much like the
Bolivian specimens. A / test showed only three
variables that differ significantly: bullar height (/ =
5.45, P < 0.001), tail length (/ = 2.61, P < 0.05),

274

and length of hind foot (/ = 3.24, P < 0.01); all
have greater average values in Bolivian specimens, |
but the ranges of values overlap. One possible con-
founding factor is that the Brazilian sample is com-
posed of mostly males (ca. 95%) and has a lower
average adult age than the Bolivian or Peruvian '
samples (see tables 2-3). If the degree of sexual
dimorphism in this species is considered (2.72%,
see p. 266), the difference may actually be less than i
noted in our tests. For these reasons the subspecies !
name Oryzomys microtis microtis might be ap- I
propriate for specimens from northern Bolivia.

More differences exist between Peruvian and
Bolivian samples. The Bolivian specimens are
generally larger: the zygomatic notch is deeper,
diastema longer, braincase broader, zygomatic
breadth greater, and interorbital constriction
broader. However, Bolivian specimens have sig-
nificantly narrower teeth. Brazilian specimens, as
noted above, are similar to Bolivian specimens
and differ from the Peruvian material about as
much as Bolivian specimens do. The Peruvian mice
used in this comparison come from a lowland site
in Pasco, the department just south of Huanuco,
which is near the type locality of Oryzomys de-
structor.

One skin (amnh 247776) from near Villa Tunari
in Cochabamba is dorsally paler and ventrally
whiter (not bufly) than any skin from Pando or
Beni. A skin (amnh 246809) from near Buena Vis-
ta in Santa Cruz is comparable to the darker spec-
imens from Beni, but is ventrally less bufl[> .

Concluding Remarks

Given the considerable pelage differences noted
in reference to the few available specimens from
Cochabamba and Santa Cruz in the accounts of
Oryzomys chacoensis and O. microtis, the prob-
lems noted in reference to sF>ecimens from south-
em Tarija in the accounts of O. chacoensis and O.
longicaudatus, the difficulty of identification that
results from overlapping ranges of measurements,
the sketchy knowledge of geographic variation, and
the small samples from most critical areas, the
taxonomy of Bolivian Oligoryzomys will probably
prove to be more complicated than that presented
here. We hojie our summary will provide infor-
mation useful in further taxonomic work on the
subgenus. All of our data and detailed analyses are
deposited in the Department of Mammalogy,

FIELDIANA: ZOOLOGY

American Museum of Natural History, and are
available for the use of other interested workers.
More than 400 additional specimens, many in-
cluding karyological and biochemical prepara-
tions, have been collected in 1984, 1985, and 1986
since this paper was written. Study of these will
certainly help to clarify remaining taxonomic
problems.

Acknowledgments

This study was partially funded by a grant from
the Undergraduate-Graduate Research Fund Pro-
gram, which is supported by the Susan Greenwall
Foundation and administered by the American
Museum of Natural History. We thank Dr. Leslie
F. Marcus for his statistical advice. Masaaki Yone-
da kindly provided us with a list of SF>ecimens in
the Museo Nacional de Historia Natural, La Paz,
Bolivia (MNHN). We are grateful to the curators
of the collections referred to here, especially Dr.
Philip Myers, Dr. Guy G. Musser, and Dr. Michael
D. Carleton, for their generous advice and assis-
tance in various ways. We also thank Dr. Robert
S. Voss, Dr. James L. Patton, and Dr. Robert M.
Timm for reviewing the manuscript.

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. 1982. Results of the Archbold Expeditions.

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Myers, P., and M. D. Carixton. 1981. The species
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Miscellaneous Publications of the Museum of Zool-
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Neveu-Lemaire, M., and G. Grandidier. 1911. Notes
sur les mammiferes des hauts plateaux de I'Amerique
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O'CoNNELL, M. A. 1982. Population biology of North
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Genoways, eds.. Mammalian Biology in South Amer-
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Osgood, W. H. 1916. Mammals of the Collins-Day
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Pine, R. H. 1973. Mammals (exclusive of bats) of Be-
lem. Para, Brazil. Acta Amazonica, 3: 47-79.

Sanborn, C. C. 1959. Small rodents from Peru and
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SouKUP, J. 1961. Materiales para el catalogo de los
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Streilein, K. E. 1982a. Ecology of small mammals in

the semiarid Brazilian caatinga. I. Climate and faunal
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. 1 982b. Ecology of small mammals in the semi-

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1 982c. Ecology of small mammals in the semi-

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population ecology. Annals of Carnegie Museum, 51:
251-269.

Tate, G. H. H. 1932. The taxonomic history of the
South and Central American cricetid rodents of the
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Thallomyscus, and Melanomys. American Museum
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. 1 898. On the small mammals collected by Dr.

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. 1902. On mammals from Cochabamba, Bo-

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. 1926. The Spedan Lewis South American Ex-

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Cricetides) dans le foyer natural de peste de Barracao
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malia, 46: 335-359.

276

HELDIANA: ZOOLOGY

Appendix ^

Specimens Reported or Examined

The latitude and longitude are abbreviated; 1657/6523 means 16*'57'S, 65''23'W, for example. Where
references are given (to Thomas), the specimens have not been examined by us. As noted, there are a
few other specimens that we have not seen.

O. chacoensis

Specimens: 88.

Boquer6n

4 1 9 km by road NW Villa Hayes, 1 amnh (248397).

BOLIVIA
Santa Cruz

1623/6059, San Ignacio, province of Velasco, 10
USNM (390120-390121, 390664-390666,
391522-391526).

1808/6312, 7 km E and 3 km N Ingeniero Mora,
13 AMNH (247758-247761, 247765-247773).

Tarua

2056/6321, 2 km S and 10 km E of Tiquipa, 4

amnh (246799-246801, 246822).
21 19/6325, 8 km S and 10 km E of Villa Monies,

8 AMNH (246802-246806, 246810, 146912,

246914).
2128/6317, 35 km by road SE Villa Montes, Ta-

ringuiti, 5 ummz (155937-155938, 156332-

156334).
2241/6426, Rio Lipeo, 13 ansp (18176-18188).

PARAGUAY

Presidente Hayes

Rio Pilcomayo, 15 mi W Rio Paraguay, 3 amnh
(143892-143894). - -

NuEVA AsuNa6N

19 km by road WSW, km 588 Trans Chaco, 8

amnh (249255-249262).

Chaco

50 km WNW Fortin Madrejon, 9 amnh (248398-
248406).

ARGENTINA

JUJUY

Yuto, 10 AMNH (167855, 179976, 179980, 182570-
182571, 182738-182739, 183312, 185226-
185227, 185269).

Sta. Barbara, 4 amnh (185224-185225, 185228,
186954).

O. longicaudatus

Specimens: 134.

BOLIVIA

Chuquisaca

1927/6407, Tola Orko, Tomina Province, 6 usnm
(271588-271590, 545226-545227, 545229).

1929/6433, Horcus, 80 km SE Sucal, 3 mvz
(134654-134656).

1931/6409, Monte Canto, 1 usnm (271591).

?, Chuyayacu, 1 fmnh (72889).

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

zm.

CCXTHABAMBA

1620/6645, Yungas, 1 cm (5276).

1654/6642, El Choro, 1 amnh (1 19405), 12 fmnh

(74902-74913), 1 bm (Thomas, 1902).
1714/6541, Incachaca, 19 amnh (38525-38542,

38550, not seen), 2 cm (5081-5082).
1734/6621, Parotani, 2 amnh (38543-38544), 1

fmnh (21668).
1742/6509, 20 mi E Totora, 1 mvz (1 19916).
1807/6509, Aiquile, 2 fmnh (50970-50971).

22 1 2/6436, 8 km by road N of Cuyambuyo, 2
UMMZ (156326-156327).

2213/6436, 4 km by road N of Cuyambuyo, 10
UMMZ (155889-155891, 156312-156313,
156315-156317, 156319-156320).

Department Uncertain
?, Zapial, 1 usNM (270911).

La Paz

1530/6824, Cocapunco, 1 amnh (72644).
1535/6843, Tacacoma, 1 amnh (91540).
1535/6843, Tacacoma-Sorata, 2 amnh (91541-

91542).
1539/6824, Okara, 2 amnh (72704-72705).
1543/6840, 10 km by road N of Sorata, 3 ummz

(156301-156303).
1547/6840, Sorala, at base of Ml. Sorata, 4 amnh

(91536-91539).
1600/6516, Charuplaya, 16 bm (Thomas, 1902).
1618/6753, Rio Aceramarca, 2 amnh (72693-

72694).
1619/6752, Nequejahuira, 3 amnh (72722, 72724-

72725).
1620/6808, Mt. Chacaltaya, 1 ummz (1 1571 1).
1620/6756, Pongo, 15 amnh (72702-72703,

72706-72715, 72726, 81283, 241612).
1621/6747, Pitiguaya, 4 amnh (72716, 72729-

72731).
1659/6749, Caracato, 5 amnh (248977-248981).
1823/6659, Poopo, 1 mnlp (not seen).

POTOSi

1 844/6609, 3 km SE of Pocoata, 1 amnh (255946).
2127/6543, Tupiza, 1 bm (Thomas, 1926).

Santa Cruz

1754/6429, 5 mi W Comarapa, 1 mvz (1 19917).
2005/6334, nr. Camiri, 1 cal (13803).

Tarua

2131/6445, 10 mi NW Tarija, 1 mvz (1 19918).
2 1 32/64 1 2, Entre Rios, 3 USNM (27 1 4 1 1-27 1 4 1 2,
271432).

O. microtis

Specimens: 230.

BOLIVIA
Beni

1049/6525, Guayaramerin, 2 amnh (210050-

210051).
1110/6522, 4 km below Santa Cruz, 1 "amnh

(211727).
1 142/65 1 6, 4 km S Santa Rosa, 4 amnh (2 1 1 729-

211730, 211791-211792).
1 200/6502, Rio Itenez, 20 km from mouth, 2 amnh

(211756-211757).
1200/6506, Puerto More, Rio Itenez, 3 amnh

(211758-211760).
1 2 1 3/65 1 3, Mamore River on bank opposite Cas-

cajal, 16 amnh (211724-211725, 211754, not

seen, 211762-211774).
1226/6511, Mamore River, 2 amnh (211721,

211761).
1229/6417, Rio Itenez, opposite Costa Marques,

Brazil, 11 amnh (210122, 210038-210045,

210365-210366).
1229/6418, Rio Itenez, 1.5 km below Costa Mar-
ques, Brazil, 1 amnh (210364).
1230/6415, Pampa de Meio, 2 amnh (210046-

210047).
1 230/64 1 8, Baures River mouth, 6 amnh (2 1 0028-

210031, 210382, 210383, not seen).
1240/6330, Curiche River mouth, 6 amnh

(210032-210037).
1241/6432, 15 km above Horquilla on Rio Ma-

chupo, 1 amnh (210053).
1244/6318, Versalles, 1 amnh (210052).
1244/6428, Las Perias, 3 fmnh (1 17079-1 17081),

1 USNM (460741).

278

FIELDL\NA: ZOOLOGY

1300/6515, Mamore River, 1 amnh (21 1722).
1304/6449, San Joaquin, 1 fmnh (1 17075), 6 usnm

(364735, 364738, 391299, 460273, 460742-

460743).
1312/6410, Cachuelita, 1 usnm (460739).
1 3 1 2/65 1 5, 8 km N Exaltacion, 6 amnh (2 1 1 775-

211780).
1313/6221, 20 km W Larangeira, Bahia de los

Casara, 2 amnh (210048-210049).
1313/6409, Boroica, 1 usnm (460740).
1322/6520, Palacios, province of Yacuma, 1 usnm

(461082).
1324/6442, Chaco Lejo, 20 km SE San Ramon, 2

usnm (391295-391296).
1325/6435, Tacuaral, 1 usnm (391297).
1343/6521, Puerto Caballo, 2 amnh (211785,

214597).
1420/6455, 10 km W San Pedro, on Mamore Riv-
er, 5 AMNH (21 1786-21 1790).
1428/6734, Rurrenabaque, 2 amnh (247774-

247775).
1434/6455, 23 km W San Javier, 1 amnh (214760).
1437/6457, Ibare River mouth, 2 amnh (21 1783-

211784).
1446/6451, Ibare River, 24 km from mouth, 1

amnh (21 1755, not seen).
1454/6422, 6 km W of Casarabe, 7 amnh (255947-

255953).
1503/6658, 1 km E of La Embocada, 2 ummz

(155940, 156292).
1515/6415, El Triunfo, 1 usnm (391298).
1550/6441, 5 km NW Grande River mouth, 2

AMNH (21 1781-21 1782).
?, Centenela, 1 fmnh (1 17074).
?, "Beni" only, 1 amnh (232699).

Pando

1117/6855, Rio Nareuda, 11 amnh (248982-
248990. 248993-248994).

Santa Cruz

1 703/6335, 7 km N Santa Rosa, 1 amnh (25460 1 ).
1724/6344, 7 km N and 17 km W of Buena Vista,
3 AMNH (246809, 246820, 246935).

BRAZIL

Amazonas

Solimoes, 10 amnh (37088-37097, 37091 = ho-

lotype of O. microtis).
Guatsue, 1 amnh (37100).
Lower Solimoes, 1 amnh (37157).

ParA

Capim, 150 mi SE Belem, 1 amnh (188964).
Capim, Est. B.R. 14, km 97, 1 amnh (203400).
Rio Xingu; Porto de Moz, 1 amnh (95983).
Rio Xingu; Villarinho do Monte, 4 amnh (95984-
95986, 95997).

GoiAs: Rio Madeira

COCHABAMBA

1648/6508, Todos Santos, 1 amnh (paratype of
O. chaparensis, 40787), 1 fmnh (holotype,
21330).

1657/6523, 2 km E of Villa Tunari, 4 amnh
(247662-247664, 247776).

Auara Igarape, 5 amnh (91874, 91876-91878,

94245).
Sta. Antonio du Uayara, 4 amnh (92258-92261).
Rosarinho, Lago Miguel, 1 1 amnh (92705-92715).
Rosarinho, Lago Sampaio, 2 amnh (9271 6-927 1 7).

Mato Grosso

La Paz

1 5 1 5/68 1 0, Mapiri, 5 amnh (7272 1 , 730 1 1 , 72732,

72697-72698).
1528/6752, Guanay, 1 amnh (72701).
1528/6818, Ticunhuaya, 1 amnh (72700).
1540/6742, 4 km NW Alcoche, 2 ummz (126777,

127167).

Utiarity, Papagaio River, 3 amnh (37540-37542;
37541 = type of O. utiaritensis, 37542 = type
of 6). mattogrossae).

? Amazonas

S bank R. Amazon: Villa Bella Imperatriz, 2 amnh
(91899-91900).

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS

279

PARAGUAY

Caaguazu

24 km NNW Carayao, 1 ummz (133821).

Presidente Hayes

24 km NW Villa Hayes, 3 UMMZ (133830-1 33831,
133833).

PERU

Ayacucho

Hda. Luisiana on Rio Apurimac, 1 amnh (242484).

Pasco

Ca. 1 km N Puerto Bermudez, 38 amnh (24555 1-
245588).

Maldonado

Barra del Arrojo Maldonado, 9 km ENE Punta
del Este, 1 amnh (205994).

Montevideo

Rio Santa Lucia, 1 km SE Santiago Vasquez, 2
AMNH (205995-205996).

ROCHA

22 km SE Lascano, 18 amnh (205997-206009,
206011-206015).

San Jose

Dept. only, 3 amnh (232216-232218).

Soriano

3 km E Cardona, 1 amnh (206016).

O. flavescens

Specimens: 33.

Treinta y Tres

16 km SSW Boca del Rio Tacuari, 1 amnh
(206017).

URUGUAY

Artigas

6 km NNW Belen, 2 amnh (205986-205987).

Cerro Largo

Sierra de Vaz, Rio Tacuari, 20 km SE Melo, 1
amnh (205988).

Lavalleja

12 km WSW Zapican, 4 amnh (205989-205905).

O. nigripes

Specimens: 31.

BRAZIL

GOIAS

Anapolis, 7 amnh (134528-134530, 134532,
134534, 134538, 134540).

280

FIELDIANA: ZOOLOGY

MiNAS Gerais PARAGUAY

(Serra do Caparao), Rio Caparao, 8 amnh (80369- Ainambay

80370, 80372, 80375-80379).

(Serra do Caparao), Boa Espera, 1 amnh (80391). 4 km by road SW Cerro Cora, 1 ummz (125523).

Mato Grosso CaaguazO

Maracaju, 9 amnh (134541-134542, 134544- N of Coronel Oviedo, 1 ummz (124212).

134547, 134551, 134838, 134902). 24 km NNWCarayao, 3 ummz (133835-133837).

SAo Paulo

Piquete, 1 amnh (36496).

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 281

New Records and Current Status of Euneomys (Cricetidae)
in Southern South America

Jose L. Yaftez, Juan C. Torres-M iira, Jaime R. Rau, and Luis C. Contreras

ABSTRACTS

New records of Euneomys from central and southern Chile are given. Specific assignment of
this material is difficult, owing to the lack of good diagnostic characters, small number of
specimens, and spotty distribution. Absence of clear-cut differences between Euneomys localities
suggests that there is only a single species, E. chinchilloides. Subspecific assignments are deferred,
because at present they can only be made geographically, and the gaps between localities seem
to be more apparent than real.

Se citan nuevos registros de Euneomys para Chile Central y Sur. La determinacion es-
pecifica de este material es dificil debido a la carencia de buenos caracteres diagnosticos, al
bajo niimero de especimenes y a la distribucion localizada con grandes hiatos intermedios. La
ausencia de claras diferencias entre Euneomys de distintas localidades sugiere la existencia de
tan solo una especie, E. chinchilloides. Las nominaciones subespecificas se posponen, ya que
al moment© estas solo se pueden realizar basadas en las localidades geograficas y los hiatos
parecen ser mas aparentes que reales.

Sao citados novos registros de Euneomys para o Sul e Centro do Chile. Dada a falta de bons
carateres diagnosticos, o baixo numero de especimes, e a distribui^ao localizada com hiatos
intermediarios, e dificil a determina9ao esfiecifica deste material. A ausencia de caracteristicas
distintas entre Euneomys de localidades diferentes sugere a existencia de uma especie apenas:
E. chinchilloides. Nao foram designadas nomina96es subespecificas ja que, no momento, estas
so poderiam ser realizadas na base das localidades geograficas, e os hiatos podem ser mais
aparentes do que reais.

Introduction

The genus Euneomys is one of the least studied
cricetid rodents in southern South America. These
herbivorous, volelike rodents have a relatively
large, heavy body and a short tail and ears (Os-
good, 1943). Euneomys has close affinities with
the phyllotine and sigmodontine groups (Hersh-

From Museo Nacional de Historia Natural, Casilla
787, Santiago, Chile (Yanez); Dei>artamento Biologia y
Quimica, Facultad de Ciencias, Universidad de Talca,
Casilla 747, Talca, Chile (Torres-Mura and Contreras);
and Estacion Biologica de 'Doiiana' (Sevilla), Apartado
1056, Sevilla 41013, Espana (Rau).

kovitz, 1 962; Mann, 1978). The genus has a known
distribution from 33°00'S to Cape Horn (fig. 1),
mainly along and to the east of the Andes (Hersh-
kovitz, 1962; and subsequent reports).

In Chile Euneomys is known from Lago Fag-
nano and Hermite Island (Tierra del Fuego), Punta
Arenas (Magallanes), Torres del Paine and Laguna
Lazo (Ultima Esperanza), Puerto Ibaiiez (General
Carrera), Campo Bandera (Aysen), Pino Hachado
(Malleco), and Lo Valdes and Farellones (Santia-
go) (Osgood, 1943; Mann, 1944, 1978; Markham,
1971; Greer, 1965; Reise & Venegas, 1974; Pine
et al., 1979). In Argentina Euneomys is certainly
known from Bahia del Buen Suceso (Tierra del
Fuego), upper Rio Chico (Santa Cruz), Paso de

YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS

283

Fig. 1. Collecting localities of Euneomys. 1, Farellones; 2, Lo Valdes; 3, Bartos del Flaco; 4, Pino Hachado; 5,
Lastarria; 6, Puerto Ibariez; 7, Torres del Paine; 8, Rio Baguales and Cueva del Milodon; 9, Estrecho de Magallanes
and Palli Aike; 10, Punta Arenas; 11, Lago Fagnano; 12, Hermite Island; 13, Grevy Island; 14, San Rafael; 15, Cueva
Traful; 16, Cerro Leones; 17, Paso de los MoUes, Pilcaniyeu; 18, Upper Rio Chico, Santa Cruz; 19, Bahia del Buen
Suceso. Open circles represent new localities.

284

FIELDIANA: ZOOLOGY

Table 1. Mandibular measurements of Euneomys found in bam owl pellets from Lastarria and Baiios del Flaco
and from animals trapped in Farellones near Santiago.

Character

Farellones*

BaAos del Flaco

Lastarria

MH-1

4.07 ± 0.12(3.8-4.1)

4.1, 3.9

2.6,t4.0, 3.1t

MHC

8.8 ± 0.44(8.1-9.1)

9.2, 9.1

7.9,6.1

MAL

6.07 ± 0.08 (5.9-6.2)

6.1, 6.1

5.9,6.0,6.1

MDL

4.03 ± 0.24 (3.6-4.4)

3.6, 4.1

3.6,3.9,3.1

6.24 ± 0.16(6.1-6.5)

6.2, 6.3

5.8, 6.0, 5.5

MH- 1 = Mandibular height at the first molar; MHC = mandibular height from condyle to angular process; MAL =
mandibular alveolar length; MDL = mandibular diastema length; SL = symphysis length.

Values from Farellones near Santiago are X ± 2 SE (and range; N = 7). Individual measurements are given for
Baiios del Flaco and Lastarria.

* LCM collection, f Juvenile, the last cheektooth is not fully erupted.

Los MoUes, Pilcaniyeu, and Cerro Leones (Rio
Negro), Cueva Traful (Neuquen), and San Rafael
(Mendoza) (Hershkovitz, 1962; Pine et al., 1978;
Pearson & Pearson, 1982; Massoia, 1982). In this
paper we report new records of Euneomys in Chile
and review the scant literature on the taxonomy
and geographic distribution of this poorly known
genus.

New Records and Current Status

Some new records are from bam owl (Tyto alba)
pellets from Lastarria (near Temuco), 39°14'S,
70°40'W, and from Bafios del Flaco (near San Fer-
nando), 34°57'S, 70°26'W. In addition, six other
specimens of Euneomys were recently captured on
Grevy Island (Tierra del Fuego; 55°32'S, 67°37'W)
and at Palli Aike (Magallanes; 52°25' S, 69''42'W)
and Rio Baguales (Ultima Esperanza; 51°02'S,
72°30'W); all are deposited in the Museo del Ins-
tituto de la Patagonia (MIP), Punta Arenas. Data
obtained from these new specimens have been in-
corporated by locality into Table 2.

Determination of this material to genus was not
difficult, using Reise's (1973) key and studies by
Hershkovitz (1962) and Mann (1978). However,
determination to species is difficult, owing to a
lack of good diagnostic characters, the small num-
ber of specimens, and their spotty distribution.

Osgood recognized two species of Euneomys in
Chile: E. chinchilloides (with two subspecies) in
Tierra del Fuego and Magallanes and E. petersoni
in Ultima Esperanza and Aysen. Subsequently,
Mann ( 1 944) described E. noei from Lo Valdes in
the Andes near Santiago. Hershkovitz (1962) con-
sidered E. noei as doubtfully separable from E.
mordax, with type locality at San Rafael (Men-

doza Province, Argentina), and thought that mor-
daxwas probably a subspecies of £". chinchilloides.
Greer (1965) referred his four Malleco specimens
to E. chinchilloides petersoni, despite their greater
similarity to noei or mordax in measured char-
acters. Miller and Rottmann (1976) identified all
their Euneomys from near Santiago as E. mordax.
Mann (1978) thought all Chilean specimens rep-
resented a single polytypic species: E. chinchil-
loides chinchilloides from Grande de Tierra del
Fuego Island and the adjacent mainland, E. c.
petersoni from the eastern Andes of Ultima Es-
peranza and Aysen, and E. c. noei from the Andes
outside Santiago. Subsequently, Pine et al. (1979)
suggested that two sympatric species of Euneomys
occur in Farellones near Santiago, but declined to
identify the other species until comparisons could
be made with specimens representing the entire
range of the genus. Karyotypes of nine specimens
from Farellones show insignificant differences
(Spotomo, pers. comm.).

Thus, Hershkovitz ( 1 962) and Mann (1978) rec-
ognized only one polytypic species, whereas Pine
et al. (1979) and Tamayo and Frassinetti (1980)
regarded specimens from different areas as differ-
ent species. Tamayo and Frassinetti ( 1 980) thought
Greer's Malleco specimens also represented Eu-
neomys sp. Unfortunately, this taxonomic uncer-
tainty is difficult to dispel, as only seven intact
skulls of Euneomys from the Andes outside of
Santiago are deposited in Chilean collections (La-
boratorio Citogenetica de Mamiferos [LCM], Fac.
Medicina, Universidad de Chile), and none is de-
posited in collections in Mendoza or San Rafael,
Argentina (R. A. Ojeda, pers. comm.; H. A. La-
giglia, pers. comm.).

Euneomys material recently obtained from owl
pellets and deposited in the Museo Nacional de
Historia Natural de Chile is incomplete, consisting

YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS

285

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« -5
— -c

<N 00 O w O^

f»% —

^^ "^ ^^ o ^^ q

^4^ic

— r~ r~
<N ^-^ <N ^^ «N ^.^

ri^ 4S AE

tN ^..^ m ^«, m ^«,

r^SiS 42
-: fi <N ^

^ — ^ op O

— 00 00

O >r% \0

*Ys 7^ 7°

>0 ^^ CT> ^^ O 3

vO o o

— — fS

Si's 5 ^

o ^

o
3 S

(N ,_ O
^ ^' r<^

1 r<^ 00
>0 ^^ CM

CO ^

o J:;
c 5

286

HELDIANA: ZOOLOGY

mainly of mandibles and maxillae. The limited
samples and their disintegration make a statistical
analysis impossible. Comparisons of the owl pellet
material with the seven specimens collected near
Santiago (table 1) show no differences. The same
results are obtained when comparisons are ex-
tended to include all known Chilean specimens
(table 2). Using these criteria it is impossible for
us to assign specimens from Lastarria and Banos
del Flaco to either E. c. mordax or E. c. petersoni.

Conclusion

The absence of clear-cut differences between lo-
calities in Table 2 suggests that there is only a
single species represented, E. chinchilloides. Be-
cause subspecific assignments at present can only
be made geographically, treatment of subspecies
is deferred until an adequate sample of specimens
is available. The fact that Euneomys are not taken
in traps at the same localities where they are found
in owl pellets (e.g., Reise & Venegas, 1976; Pear-
son &. Pearson, 1982; present study) suggests that
current gaps in the distribution o^ Euneomys might
be attributable to sampling error rather than to
truly disjunct distributions. The new records re-
ported here fill both morphological and geograph-
ical gaps.

Acknowledgments

We thank Angel Spotomo and Claudio Venegas
for making specimens of Euneomys available to
us. Bruce D. Patterson, Ronald H. Pine, and an
anonymous reviewer made valuable comments.
Mrs. Veronica Aguirre typed the manuscript, and
Luz Uribe assisted with the figure. This report is
dedicated to P. Hershkovitz in recognition of his
pioneering contributions to our understanding of
Neotropical mammals.

Literature Cited

Greer, J. K. 1965. Mammals of Malleco Province,
Chile. Publications of The Museum, Michigan State
University, Biological Series, 3: 51-151.

Hershkovitz, P. 1962. Evolution of Neotropical cric-
etine rodents (Muridae). Fieldiana: 2kx)logy, 46: 1-
524.

Mann, G. 1 944. Dos nuevas especies de roedores. Bio-
logica (Santiago), 1: 95-126.

. 1978. Los pequenos mamiferos de Chile. Gay-
ana: Zoologia, 40: 1-342.

Markham, B. 1971. Catalogo de los anfibios, reptiles,
aves y mamiferos de la Provincia de Magallanes, Chile.
Publicaciones Instituto de la Patagonia, Serie Mono-
grafias N2. 3, 64 pp.

Massoia, E. 1982. Restos de mamiferos recolectados
en el paraje Paso de Los Molles, Pilcaniyeu, Rio Negro.
Revista Investigaciones Agropecuarias INTA, Buenos
Aires, 17: 39-53.

Miller, S. D., and J. Rottmann. 1976. Guia para el
reconocimiento de mamiferos chilenos. Ed. Gabriela
Mistral, Santiago, 200 pp.

Osgood, W. H. 1943. The mammals of Chile. Field
Museum of Natural History, Zoological Series, 30: 1-
268.

Pearson, O. P., and A. K. Pearson. 1982. Ecology
and biogeography of the southern rainforests of Ar-
gentina, pp. 129-142. In Mares, M. A., and H. H.
Genoways, eds.. Mammalian Biology in South Amer-
ica. A Symposium Held at the Pymatuning Laboratory
of Ecology, May 10-14, 1981. Sf>ecial Publication Se-
ries, Pymatuning Laboratory of Ecology, University
of Pittsburgh, 6: 1-539.

Pine, R. H., J. P. Angle, AND D. Bridge. 1978. Mam-
mals from the sea, mainland, and islands at the south-
em tip of South America. Mammalia, 42: 105-1 14.

Pine, R. H., S. D. Miller, and M. L. S. Schamberger.
1979. Contributions to the mammalogy of Chile.
Mammalia, 43: 339-376.

Reise, D. 1973. Clave para la determinacion de los
craneos de marsupiales y roedores chilenos. Gayana:
Zoologia, 27: 1-20.

Reise, D. L., and W. Venegas. 1974. Observaciones
sobre el comportamiento de la fauna de microma-
miferos en la Region de Puerto Ibaiiez (Lago General
Carrera), Aysen, Chile. Boletin Sociedad de Biologia,
Concepcion, Chile, 47: 71-85.

Tamavo, M., and D. Frassinetti. 1980. Catalogo de
los mamiferos fosiles y vivientes de Chile. Boletin
Museo Nacional de Historia Natural, Chile, 37: 323-
399.

YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS

287

Morphological Variation, Karyology,

and Systematic Relationships

of Heteromys gaumeri (Rodentia: Heteromyidae)

Mark D. Engstrom, Hugh H. Genoways, and Priscilla K. Tucker

ABSTRACTS

Morphological variation was assessed within and among populations of Heteromys gaumeri
using univariate and multivariate statistical analyses of external and cranial measurements.
Although patterns and amount of nongeographic variation in H. gaumeri were similar to other
heteromyines, geographic variation was relatively conservative. Mean values of most characters
were statistically homogeneous among localities and spatially unpattemed. Consequently, no
association was found between levels of within- and among-sample variation for individual
characters (the "Kluge-Kerfoot phenomenon"). Populations of//, gaumeri were chromosomally
monomorphic. The lack of morphological and chromosomal variation in //. gaumeri contrasts
sharply with patterns in other heteromyines. Heteromys gaumeri is morphologically and chro-
mosomally distinct from the //. desmarestianus species group (to which it is currently assigned)
and appears to share some primitive characters with Liomys (the sister group of Heteromys).
We recommend that H. gaumeri be removed from the H. desmarestianus group.

La variacion morfologica intra e interpoblacional de Heteromys gaumeri fue evaluada usando
analisis estadisticos univariados y multivariados de medidas extemas y craneales. A pesar de
que los patrones y cantidad de variacion intrapoblacional en //. gaumeri fue similar a la de
otros heterominos, la variacion geografica fue relativamente conservadora. Los valores pro-
medio de la mayoria de los caracteres fueron estadisticamente homogeneos entre las localidades,
sin mostrar ningun patron de variacion espacial. En conservencia, no se encontro asociacion
alguna entre los niveles de variacion intra e interpoblacional para caracteres individuals ("fen-
omeno Kluge-Kerfoot"). Las poblaciones de //. gaumeri fueron monomorficas cromosomica-
mente. La falta de variacion tanto morfologica como cromosomica en H. gaumeri contrasta
marcadamente con los patrones encontrados anteriormente para otros heterominos. Heteromys
gaumeri es morfologica y cromosomicamente distinguible del grupo //. desmarestianus (al cual
se asigna actualmente) y aparentemente comparte algunos caracteres primitivos con Liomys (el
grupo hermano de Heteromys). Nosotros recomendamos que se remueva a H. gaumeri del
grupo //. desmarestianus.

Avalia-se a variafao morfologica intra- e interpopulacional de Heteromys gaumeri, atraves
de analises estatisticas uni- e multivariadas de medidas extemas e craniais. Apesar dos padroes,
e da quantidade de varia^ao intrapopulacional em H. gaumeri serem similares aos de outros
heteromideos, a varia9ao geografica e relativamente conservadora. Os valores medios da maior
parte dos carateres examinados sao estatisticamente homogeneos entre as localidades, e nao

From Department of Biology, Angelo State Univer-
sity, San Angelo, TX 76909 (Engstrom); Section of
Mammals, Carnegie Museum of Natural History, 4400
Forbes Avenue, Pittsburgh, PA 1 5206 (Genoways; pres-
ent address. University of Nebraska State Museum, 212
Morrill Hall, Lincoln, NE 68588); and Department of
Wildlife and Fisheries Sciences, Texas A&M University,
College Station, TX 77843 (Tucker, present address, The
Jackson Laboratory, Bar Harbor, ME 04609).

ENGSTROM ET AL.: HETEROMYS GA UMERI 289

surgiu nenhum padrao de variances locals. Consequentemente, nao foram encontradas asso-
cia96es entre os niveis de varaiafoes intra- e interpopulacionais para carateres individuals (o
"fenomeno KJuge-Kerfoot"). Popula^oes de H. gaumeh mostraram-se cromossomicamente
monomorficas. A falta de varia9ao morfologica ou cromossomica em H. gaumeri e altamente
contrastante aos padroes encontrados em outros heteromideos. Heteromys gaumeri dlstingue-
se tanto morfologica quanto cromossomicamente do grupo H. desmarestianus. ao qual esta
atualmente deslgnado, e aparentemente possue carateres primitivos em comum com Liomys—
grupo irmao de Heteromys. Recomendamos que H. gaumeri seja removido do grupo H. des-
marestianus.

Introduction

Spiny mice of the genus Heteromys (Hetero-
myidae: Heteromyinae) are Neotropical rodents
that typically occur in rainforest and cloudforest
habitats from east-central Mexico south to north-
em South America. Currently, the genus is par-
titioned into two subgenera {Xylomys and Hetero-
mys), and two species groups {desmarestianus and
anomalus groups) are recognized in the nominate
subgenus (Goldman, 1911; Hall, 1981; Rogers &
Schmldly, 1982). Hall and Kelson (1959) and Hall
(1981) noted that taxonomlc relationships within
Heteromys and in particular the H. desmaresti-
anus species group were problematical and in need
of revision. Recently, Rogers and Schmldly ( 1 982)
reviewed morphological variation among repre-
sentatives of the H. desmarestianus species group
from northern Middle America, exclusive of H.
gaumeri, and concluded that only two species (//.
desmarestianus and H. goldmani) were represent-
ed in the material they examined.

Heteromys gaumeri, the third species currently
recognized in the desmarestianus group, is endem-
ic to the Yucatan Peninsula. Ecologically this
species characteristically occurs in relatively dry
deciduous and subdeciduous-subperennlal forest.
Systematic relationships of H. gaumeri are enig-
matic. In his review of the subfamily Heteromyi-
nae, Goldman (1911) placed H. gaumeri in his
desmarestianus group for convenience, but noted
(p. 29) that it was "not closely related to any known
species." In this paper, we review morphological
and chromosomal variation within H. gaumeri and
comment on the systematic relationships of this
species to the H. desmarestianus species group and
the subgenus Heteromys.

Materials and Methods

A total of 322 specimens o{ Heteromys gaumeri
was examined in the morphological analyses.

Specimens examined are listed in the Appendix,
and institutions housing those specimens are as
follows (abbreviations for North American col-
lections follow Choate &. Genoways, 1 975): Amer-
ican Museum of Natural History (amnh); British
Museum (Natural History), London (bmnh); Field
Museum of Natural History (fmnh); Museum of
Natural History, University of Kansas (ku); James
Ford Bell Museum of Natural History, University
of Minnesota (mmnh); Royal Ontario Museum
(ROM); Texas Cooperative Wildlife Collections,
Texas A«&M University (tcwc); The Museum,
Texas Tech University (ttu); Museum of Zoology,
University of Michigan (ummz); Unlversidad Na-
cional Autonoma de Mexico (unam); and National
Museum of Natural History (usnm).

Each specimen was assigned to one of five pre-
sumptive age classes (I-V, from youngest to oldest)
based on pelage characteristics and state of erup-
tion and relative wear of the maxillary toothrow,
following Genoways (1973). For each adult and
selected subadult specimens, the following four
external (recorded from specimen labels) and 10
cranial measurements (measured to the nearest 0. 1
mm using dial calipers) were taken, as defined by
Genoways (1973): total length (TL); length of tail
(TV); length of hind foot (HF); length of ear (LE);
greatest length of skull (GLS); zygomatic breadth
(ZB); interorbital constriction (IOC); mastoid
breadth (MB); length of nasals (LN); length of ros-
trum (LR); length of maxillary toothrow (MTR);
depth of braincase (DBC); Interparietal width
(IW); and interparietal length (IL). Each measure-
ment was chosen because it was examined in pre-
vious studies of variation in other heteromyine
rodents and was geographically variable in some
taxa (Goldman, 1911; Genoways, 1973; Rogers &
Schmldly, 1982).

Initially, four qualitative cranial characters and
dorsal coloration also were examined, as described
by Genoways (1973) for Liomys (the sister group
of Heteromys). Unlike patterns found in species
of Liomys, each of these characters was equally or

290

FIELDIANA: ZOOLOGY

more variable within populations of H. guumeri
as among populations, and the characters varied
in no apparent geographic pattern. Consequently,
qualitative cranial characters and dorsal color-
ation were not analyzed further.

Nongeographic variation in the 14 mensural
characters was examined in one sample of 94 spec-
imens collected near Campo Experimental Fores-
tal "El Tormento," 7.5 km W Escarcega, Cam-
peche. These specimens were collected within a
two square kilometer area of transitional tropical
evergreen-tropical deciduous forest and were here
considered to represent a single population. All
calculations were performed using subprograms of
the Statistical Analysis System (SAS; SAS Insti-
tute, Inc., 1982). Standard statistics (mean, range,
standard deviation, standard error, coefficient of
variation, skewness, and kurtosis) were calculated
for each variable within each subgroup (MEANS
and UNIVARIATE procedures). Student's / test
(or an approximation of variances were unequal)
was used to test for significant differences between
sexes within each age class (TTEST procedure). A
model I, one-way analysis of variance (ANOVA)
was used to test for significant differences among
age classes with sexes pooled (GLM procedure).
Subsequently, a Duncan's multiple range test
(DUNCAN) was used to determine maximally
nonsignificant subsets of age classes.

Straney (1978) criticized the use of F tests in
unbalanced ANOVAs to determine patterns of
nongeographic variation. To augment hypothesis
testing, variance partitioning of a model II, two-
way ANOVA (VARCOMP procedure) was used
to estimate the relative contributions of sex, age,
sex by age interaction, and error to within sample
variation. Age and sex were considered random
factors (see discussion in Leamy, 1983) and the
percent contribution of each factor was estimated
from variance components. The main effects (sex
and age) in the two-way ANOVAs were not in-
dependent because the data were unbalanced
(Searle, 1971). Consequently, ANOVAs were run
with sex entered into the model first, then again
with age entered first.

For analysis of geographic variation of mensural
data, adult specimens were assigned to one of 1 1
grouped localities to increase sample size (fig. 1).
In no instance did a grouped locality cross a major
physiographic or previously recognized taxonomic
boundary. The specific geographic composition of
samples is as follows: Group 1— Chuntuqui and
Laguna de Sotz (= Zotz), Peten, Guatemala; 103
km SE Escarcega (= Francisco Escarcega), Cam-

peche, Mexico. Group 2 — 7.5 km W Escarcega,
Campeche, Mexico. Group 3— Apazote, 7 km N,
5 1 km E Escarcega, and La Tuxpeiia, Campeche,
Mexico. Group 4— Dzibalchen and San Jose Car-
pizo, Campeche, Mexico. Group 5— Esmeralda and
Santa Rosa, Yucatan, Mexico. Group 6— Chichen
Itza and Piste, Yucatan, Mexico. Group 7— Ti-
zimin and Tunkas, Yucatan, Mexico. Group 8 —
La Vega, Pueblo Nuevo Xcan, and Puerto Mo-
relos, Quintana Roo, Mexico. Group 9— Felipe
Carrillo Puerto, Quintana Roo, Mexico. Group
10— Bacalar, Quintana Roo, Mexico. Group 1 1 —
Kate's Lagoon and Rockstone Pond, Belize, Be-
lize.

For each mensural character, we analyzed two
aspects of geographic variation: (1) statistical het-
erogeneity of mean values among geographic sam-
ples, and (2) significant departure of means from
spatial randomness (see Sokal & Oden, 1978). The
null hypothesis of homogeneity of means among
grouped localities for each character was tested
using a model I, one-way ANOVA (SAS:GLM).
By ANOVA, the variance of each character was
partitioned into among and within (error) locality
effects, and the percentage of variation attributable
to each effect was estimated from variance com-
ponents (SAS: VARCOMP). Homogeneity among
grouped localities across all characters was ex-
amined using a multivariate analysis of variance
(SAS:GLM, MANOVA).

The null hypothesis of no geographic pattern
among grouped locality means was examined by
testing for significant association between geo-
graphic and phenetic distance matrices. For each
character, a phenetic distance matrix was con-
structed in which the elements were calculated as
the absolute differences between means for all pairs
of localities (Sokal, 1979). Multivariate phenetic
matrices of taxonomic distance (Sneath & Sokal,
1973) were calculated for all 14 mensural char-
acters and a set restricted to those showing sig-
nificant heterogeneity, using the NT-SYS library
of computer programs (Rohlf et al., 1982). Ele-
ments of the geographic distance matrix were the
actual map distances (in km) between all pairs of
grouped localities (taken from the center of each
grouped locality); all connections between pairs of
localities were maintained because there are no
obvious physiographic or ecological barriers to gene
flow among populations of//, gaumeri on the Yu-
catan Peninsula. Three test statistics (Mantel's Z;
Spearman's rho, R; and a component of Kendall's
tau, K^) were used to test for significant association
between each phenetic distance matrix and the

ENGSTROM ET AL.: HETEROMYS GAUMERI

291

"T"

KILOMETERS
50 100

I H r-"

25 50
MILES

Fig. 1 . Geographic distribution of Heteromys gaumeri and approximate geographic areas included in the 1 1
grouped locahties used in analyses of morphological variation. Closed circles denote specimens used in analyses of
geographic variation and open circles denote specimens examined but not included in statistical analyses.

geographic distance matrix (Dietz, 1983). Values
of P associated with each statistic were estimated
from 2,000 random permutations using a FOR-
TRAN program supplied by E. J. Dietz.

In the chromosomal analysis, standard karyo-
types were examined for 26 specimens oi Hetero-
mys gaumeri. Additionally, karyotypes of one
specimen of H. desmarestianus and 20 specimens

292

HELDIANA: ZCX)LOGY

of H. anomalus were examined for comparison.
Standard karyotypes were prepared from speci-
mens sampled from natural populations using the
in vivo bone marrow technique of Patton (1967),
as modified by Lee (1969). Terminology regarding
relative chromosome arm ratios is that of Patton
(1967). In the calculation of Fundamental Num-
bers (FN), relative chromosome arm ratios were
scored conservatively. Several chromosomes re-
corded as acrocentric here had telomeric knobs of
chromatin in elongated preparations. Only chro-
mosomes which consistently displayed second
arms of chromatin, regardless of state of contrac-
tion of the preparation, were scored as biarmed.
The following voucher specimens are deposited in
the Texas Cooperative Wildlife Collection, Texas
A&M University, and the Carnegie Museum of
Natural History (sample size in parentheses): Het-
eromys gaumeri (total 26)— MEXICO. Campeche:
7.5 km W Escarcega (8). Quintana Roo: 2 km N,
8 km W Bacalar (8); 8 km NNE Felipe Carrillo
Puerto (3); 2.5 km NNE Felipe Carrillo Puerto (1).
Yucatan: Chichen Itza (1); Cenote Seco, 2 km E
Chichen Itza (5). Heteromys desmarestianus (total
l)-MEXICO. Chiapas: 9.4 km S Palenque (1).
Heteromys anomalus (total 20)— VENEZUELA.
Miranda: 25 km N Altagracia de Oricuto (6); 24
km N Altagracia de Oricuto (1). Monagas: Cari-
pito (4). Sucre: 40 km SW Caripito (9).

Results

Morphological Variation

NoNGEOGRAPHic VARIATION— The Sample of
Heteromys gaumeri (N = 94) from 7.5 km W Es-
carcega, Campeche, was used to estimate within-
sample variation of the 1 4 mensural characters.
Two approaches were taken, hypothesis testing and
estimation of variance components.

Initially, / tests were used to test for significant
differences between sexes in each of age classes I-
IV. Males average larger than females for most
characters in most age classes; however, significant
differences between sexes were found for only one
measurement in age class I (length of rostrum) and
seven measurements in age class III (table 1).

Results of ANOVA among age classes with sexes
pooled are given in Table 1 . Separate analyses of
the age effect within each sex gave results similar
to those in Table 1 and are not presented. Signif-
icant variation with age was found in each of the

14 measurements. Each measurement tended to
increase with age, although this pattern was not
pronounced for length of maxillary toothrow. In
the a posteriori DUNCAN analysis, age class V
averaged significantly larger than the other age
classes for two characters (interorbital constriction
and interparietal length), whereas age classes IV-
V formed a homogeneous subset for the remaining
12 characters. Patterns of variation among age
classes I-III were less consistent, although age
classes I-III differed significantly from each other
and age classes IV-V in six of 10 cranial mea-
surements.

To complement hypothesis testing, the relative
contributions of sex, age, sex by age interaction,
and error (residual variation) to a two-way AN-
OVA were estimated from variance components
(table 2). Separate analyses with either sex or age
entered into the model first generally yielded re-
sults that differed by 1% to 4%. Because of the
similarity of these estimates (and because sex most
often is analyzed before age in studies of nongeo-
graphic variation), only the results in which sex
was entered into the model first are presented.

Most of the variation (average 97%) in the AN-
OVA was attributable to the effects of age and
error. Age contributed the largest proportion of
variance for most characters (average 53%); error
was nearly as important, contributing an average
of 44%. For length of hind foot, length of maxillary
toothrow, interparietal width, and interparietal
length, age contributed a relatively small propor-
tion of the variance, and variation was mostly
attributable to error. Homogeneity of estimates
across characters was examined using z-transfor-
mations and a subsequent c/z/-square test (Sokal
&, Rohlf, 1981). Estimates of age and error were
significantly heterogeneous across characters.

Sex and interaction were relatively unimportant
factors in the ANOVA for most characters, con-
tributing an average of 1% and 2% of the variance,
respectively. The only noteworthy exception to this
pattern was interparietal length, for which inter-
action accounted for 23% of the variance. Despite
this exception, estimates of the effects of sex and
interaction were statistically homogeneous across
characters. Given that only a small proportion of
the variance of each character was attributable to
the effect of sex, the significant sexual dimorphism
found for seven of 1 4 characters in age class III in
preliminary / tests (table 1) probably was due to
trivial differences accentuated by large sample size
(N = 47). For most characters, the largest pro-
portion of variance was attributable to age and

ENGSTROM ET AL.: HETEROMYS GAUMERI

293

Table 1. Age variation in 14 external and cranial measurements of //er^romy^^aM/wer/ from 7.5 km WEscarcega,
Campeche, Mexico.

Age

class N

Mean (range) ±

2SE

Total Length

265.0 (237.0-272.0)

10.00

2.7

261.2(235.0-288.0)

9.82

7.0

III*

242.6(186.0-275.0)

5.87

7.8

II 1 17

220.6 (200.0-252.0)

6.70

6.3

I 1 4

190.3(179.0-210.0)

13.60

7.1

F= 19.45***

Length of Tail

145.5(137.0-159.0)

4.58

5.9

139.5(139.0-150.0)

1.00

0.5

HI*

132.5

(87.0-152.0)

3.65

8.9

120.1(101.0-136.0)

4.62

7.9

I 1 4

99.8

(96.0-117.0)

6.85

6.9

F = 20.27***

Length of Hind Foot

35.0

34.2

(32.0-36.0)

0.66

3.9

in*

33.2

(30.0-35.0)

0.40

4.1

33.0

(30.0-35.0)

0.70

4.6 "

I 1 6

31.0

(29.0-34.0)

1.71

6.8

F = 6.76***

Length of Ear

17.7

(17.0-18.0)

0.67

3.3

17.4

(17.0-19.0)

0.31

3.6

16.1

(14.0-18.0)

0.26

5.6

15.6

(14.0-18.0)

0.44

6.1

I 1 6

14.8

(14.0-16.0)

0.61

5.1

F= 15.77***

Greatest Length of Skull

V 2

35.8

(35.5-36.0)

0.50

1.0

IV 15

35.2

(33.9-37.5)

0.49

2.7

HI* 1 47

32.9

(30.2-34.5)

0.26

2.7

n 1 19 ,

31.2

(29.6-34.2)

0.50

3.5

I 1 6

28.8

(27.8-29.8)

0.65

2.7

F= 71.20***

Zygomatic Breadth

16.3

(16.1-16.5)

0.23

1.2

16.1

(15.3-16.7)

0.19

2.4

ni* 1 44

15.1

(14.0-16.3)

0.14

3.1

n 1 .19

14.4

(13.7-15.1)

0.20

3.0

I 1 6

13.5

(12.9-13.8)

0.33

3.0

F = 59.28***

Interorbital Constriction

V 1 3

9.3

(9.2-9.5)

0.20

1.9

IV 1 16

8.7

(8.3-9.0)

0.10

2.4

ni* 1 47

8.2

(7.5-9.3)

0.12

4.8

n 1 19

7.8

(7.2-8.6)

0.16

4.3

I 1 6

7.5

(7.0-7.7)

0.20

3.3

F= 29.71***

Mastoid Breadth

15.5

(15.3-15.8)

0.30

1.7

15.3

(14.8-15.9)

0.18

2.3

III 1 47

14.7

(14.0-15.4)

0.10

2.3

294

FIELDIANA: ZOOLOGY

Table 1. Continued.

Age
class

Mean (range) ± 2 SE

19
6

F = 27.94***

14.4 (13.4-15.2) ± 0.21 3.2

13.7 (13.1-14.1) ± 0.29 2.6

III

Length of Nasals

2 14.5 (14.0-15.0) ± 1.00 4.9

16 14.3 (13.2-15.6) ± 0.37 5.1

47 13.0 (11.8-14.2) ± 0.18 4.7

19 11.9 (10.8-13.4) ± 0.29 5.3

6 10.5 (10.0-10.9) ± 0.26 3.0

F = 59.06***

III*

Length of Rostrum

2 15.6 (15.1-16.0) ± 0.90 4.1

16 15.2 (14.2-16.3) ± 0.27 3.6

47 14.0 (12.7-15.3) ± 0.18 4.3

19 12.9 (11.9-13.9) ± 0.23 3.9

6 11.6 (11.0-12.1) ± 0.37 3.9

F = 65.77***

III

Length of Maxillary Toothrow

16 4.9 (4.7-5.2) ± 0.07 3.0

17 4.8 (4.5-5.1) ± 0.08 3.3
3 4.8 (4.7-4.8) ± 0.07 1.2

46 4.7 (4.3-5.2) ± 0.05 3.4

5 4.7 (4.5-4.8) ± 0.17 2.8
F = 3.72**

III

Depth of Braincase

3 9.1 (8.9-9.4) ± 0.29 2.8

16 9.0 (8.5-9.5) ± 0.13 3.0

46 8.8 (8.3-9.7) ± 0.08 3.3

19 8.5 (7.9-8.8) ± 0.13 3.3

6 8.5 (8.3-8.8) ± 0.16 2.3

F= 12.68***

III

Interparietal Width

3 9.3 (8.9-9.8) ± 0.54 5.1

15 8.8 (8.0-10.0) ± 0.31 6.8

19 8.6 (7.6-9.4) ± 0.21 5.3

45 8.5 (7.3-9.7) ± 0.17 6.7

6 8.0 (7.3-8.6) ± 0.36 5.6

F=4.11**

III

Interparietal Length

3 5.5 (5.1-6.1) ± 0.61 9.6

15 5.1 (4.2-5.8) ± 0.21 8.0

45 5.0 (4.2-5.7) ± 0.10 6.8

19 4.9 (4.2-5.3) ± 0.14 6.3

6 4.7 (3.8-5.3) ± 0.45 11.9

F= 3.23*

Vertical lines alongside age classes denote nonsignificant subsets. Asterisks after F statistics indicate levels of
significance (* P < 0.05; ** P < 0.01; *** P < 0.001). Males and females in age classes I-IV were used to test for
significant mean differences due to sex (/ test). Asterisks following age classes indicate significant sexual dimorphism
{P < 0.05).

ENGSTROM ET AL.: HETEROMYS GAUMERI

295

Table 2. Percentage of total variation attributable to
sex (S), age (A), sex by age interaction (S x A), and error
(E) for 14 external and cranial measurements of Hetero-
mys gaumeri from 7.5 km W Escarcega, Campeche,
Mexico.

Char-

Variance components

acters*

S X A

61.3

1.5

37.2

63.6

36.4

4.8

29.9

65.3

50.3

49.7

GLS

84.1

15.9

77.5

4.6

17.9

IOC

66.7

0.5

32.8

64.1

2.0

33.9

79.7

1.4

18.9

82.1

0.8

17.1

MTR

2.2

21.1

76.7

DBC

1.2

42.4

0.6

55.8

3.9

19.1

77.0

1.1

22.7

76.2

Mean

0.9

53.1**

2.4

43.6**

* Abbreviations of characters are defined in text. Es-
timates were calculated from variance components (con-
sidering sex as a random factor).

** Significant heterogeneity {P < 0.01) of estimates
among characters.

(TL) of the total variance. For those characters
that were significantly heterogeneous, an average
of 1 7% of the variance was attributable to locality.
In a MANOVA across all mensural characters,
locaHties were marginally heterogeneous {P < 0.05)
by two test statistics (Hotelling-Lawley Trace and
Wilk's Criterion), but homogeneous by a third test
statistic (Pillai's Trace), suggesting an overall lack
of mensural differentiation among localities.

Because significant spatial patterning is theo-
retically possible (although unlikely) even with sta-
tistically homogeneous means (Sokal & Oden,
1 978), each character was tested for significant de-
partures from spatial randomness. We found no
significant association between matrices of phe-
netic and geographic distance for any character by
any of three test statistics. Multivariate taxonomic
distance matrices calculated by using all characters
and using only those characters that were signifi-
cantly heterogeneous among localities were also
incongruent with geographic distance. According-
ly, mean values of most mensural characters were
homogeneous among grouped localities and not
spatially patterned as a simple function of geo*
graphic distance.

error, and the main effects in the model (sex and
age) were independent.

Variance components also were estimated from
a two-way ANOVA restricted to age classes IV
and V (results available on request). In this anal-
ysis, the pattern of variance partitioning changed
considerably. For each character, error contrib-
uted the largest proportion of variance (average
64%), whereas sex, age, and interaction had small-
er average contributions ( 1 0%, 1 2%, and 1 4%, re-
spectively). These estimates, however, should only
be regarded as approximations because of the small
sample size of age class V (N = 3). Estimates de-
rived from larger subsets of age classes included a
large average contribution (> 40%) attributable to
the effect of age. Based on these results, geographic
analyses were restricted to age classes IV-V with
sexes pooled.

Geographic Variation— Standard statistics
were calculated for each mensural character in each
grouped locality. By ANOVA, the variance of each
character was partitioned into among and within
(error) locality effects. Only six of 14 characters
were significantly heterogeneous among grouped
localities (table 3). For seven characters (HF, ZB,
IOC, MB, DBC, IW, IL), the among-locality vari-
ance component was zero and for the remaining
characters, locality accounted for 6% (LE) to 30%

Karyology

Heteromys gaumeri (2n = 56, FN = 76; fig.
2a)— The autosomal complement comprises grad-
ed series of 1 1 pairs of large- to small-sized meta-
centric and submetacentric chromosomes and 1 6
pairs of large- to small-sized acrocentric elements.
The X chromosome is large and submetacentric
and the Y is medium-sized and subtelocentric. No
variation was noted among individuals.

Heteromys desmarestianus (2n = 60, FN =
66; fig. 2b)— The autosomal complement com-
prises four pairs of submetacentric and metacen-
tric chromosomes, two large, one medium-sized,
and one small, and 25 acrocentric pairs graded
from large to small. The X chromosomes are pre-
sumed to be large and submetacentric (no males
were examined).

Heteromys anomalus (2n = 60, FN = 68; fig.
2c)— The autosomal complement comprises two
large, one medium-sized, and two small pairs of
submetacentric and metacentric chromosomes and
24 acrocentric pairs graded from large to small.
The X chromosome is large and submetacentric
and the Y is medium-sized and subtelocentric. No
variation was noted among individuals.

296

FIELDL\NA: ZOOLOGY

Discussion

r Patterns of nongeographic variation in mensural
characters of Heteromys gaumeri are generally
concordant with those observed in other hetero-
myine rodents (Genoways, 1973; Rogers &
Schmidly, 1982; see also Straney, 1978). In all
species examined, age contributes a large propor-
tion of within-sample variance for most charac-
ters; only a minor component of total variance is
attributable to sex. Male heteromyines generally
average larger than females, however, and with
large sample size secondary sexual dimorphism
often appears significant for some characters (Gen-
oways, 1973; age class III, this study).

I Relative levels of variability of individual char-
acters among species of heteromyines were com-
pared using coefficients of variation of adults for
the 12 external and cranial measurements com-
mon to each study of intralocality variation (total
length, length of tail, and the 10 cranial measure-
ments included in this study; see Genoways, 1 973;
Rogers & Schmidly, 1982). Coefficients of varia-

f tion (CVs) appeared congruent among species for
individual characters, although CVs appeared het-
erogeneous among characters within species (ex-
ternal measurements and interparietal width and
length consistently were more variable than other
characters). There was no indication of a reduction
of within-sample variation for any character in H.
gaumeri relative to other heteromyines.

Although patterns and level of intralocality vari-
ation in Heteromys gaumeri appear similar to oth-
er heteromyines, geographic variation in H. gau-
meri is relatively conservative. In H. gaumeri,
mean values for most characters were homoge-
neous among localities and geographically unpat-
temed; only a small proportion of variance (av-
erage 8%) was attributable to interlocality variation.
We found no positive statistical correlation be-
tween levels of within- and among-sample vari-
ance (the "Kluge-Kerfoot phenomenon," KJuge &
Kerfoot, 1 973) for mensural characters in H. gau-
meri (tested using Kendall's rank correlation be-
tween the W, and A, statistics suggested by Sokal,
1976; but see Rohlf et al., 1983). These data con-
trast with studies of geographic variation within
species of Liomys (Genoways, 1973) and other
members of the H. desmarestianus species group
(Rogers & Schmidly, 1982), in which population
samples were statistically heterogeneous and ap-
peared spatially patterned.
The relative lack of interlocality variance in

Heteromys gaumeri might be attributable to a re-
stricted geographic distribution, to relative envi-
ronmental homogeneity on the Yucatan Penin-
sula, and/or to a lack of genetic divergence among
populations. Compared to wide-ranging species of
Liomys and Heteromys, the geographic area oc-
cupied by H. gaumeri is relatively small, with little
topographic or climatic relief Plant communities
grade gradually from lowland tropical evergreen
forest in the south and east, to deciduous forest in
the northwestern and north-central portions of the
peninsula, to a scrub zone bordering the northern
coast (Paynter, 1955). For statistically heteroge-
neous but spatially unpattemed characters, char-
acter distributions might be determined mainly by
stochastic factors (e.g., genetic drift) in the absence
of strong migration or selective gradients (Sokal
&. Oden, 1978). Peromyscus yucatanicus, which
has a similar geographic range and occupies sim-
ilar habitats to H. gaumeri, is geographically vari-
able in color and cranial size (Lawlor, 1 965; Huck-
aby, 1980), and at least color closely tracks
vegetational changes on the peninsula. Conse-
quently, lack of interlocality differentiation in col-
or, qualitative, and mensural characters in H. gau-
meri might not result solely from selective
responses or stochastic processes in a homoge-
neous environment.

The reduced level of geographic variation in
Heteromys gaumeri also is consistent with a hy-
pothesis of little genetic divergence among pop-
ulations. If populations of H. gaumeri are genet-
ically similar, similarity probably is not a product
of panmixia, but more likely of a reduction of
genetic variation through a genetically depauper-
ate founding population (see Johnston, 1976;
Johnston &. KJitz, 1977; Baker, 1980) or genetic
bottleneck. Although we have little direct evidence
of genetic variation among populations of//, gau-
meri, individuals sampled appear chromosomally
monomorphic. In //. desmarestianus, interlocality
polymorphism in fundamental number is pro-
nounced (D. S. Rogers, pers. comm.). More sen-
sitive estimation of the level of interpopulational
genetic divergence of //. gaumeri awaits study of
genie variation.

Systematic Relationships of H. gaumeri

MorpholcxjV— Allen and Chapman (1897) de-
scribed Heteromys gaumeri from seven individ-
uals from Chichen Itza, Yucatan. In his review of

ENGSTROM ET AL.: HETEROMYS GAUMERI

297

Table 3. Geographic variation in six external and cranial measurements of Heteromys gaumeri.

Grouped
locality

Percentage of variation

Mean (range) ± 2 SE

Locality

Error

1
2
3

4
S

7
8
9

10
11

3
16
10

9
17

4
3
3

2
4

288.7
261.7
276.9
272.0
280.0
286.8
294.5
281.7
263.3
256.5
270.5

Total Length

276.0-298.0) ± 13.13

235.0-288.0) ± 8.63

245.0-302.0) ± 10.98

262.0-286.0) ± 8.83

263.0-295.0) ± 7.51

265.0-324.0) ± 8.29

288.0-300.0) ± 4.93

275.0-290.0) ± 8.82

250.0-280.0) ± 17.64

253.0-260.0) ± 7.00

250.0-290.0) ± 18.65

23.7

76.3

3.75*

Length of Tail

154.7 (

148.0-166.0)

11.39 13.6

86.4

2.02*

144.8 (

129.0-159.0)

4.12

150.4(

124.0-173.0)

9.73

154.4(

141.0-166.0)

9.48

•

161.0(

144.0-193.0)

9.22

154.4(

135.0-183.0)

5.53

162.8 (

160.0-166.0)

2.75

152.7 (

151.0-155.0)

2.40

V-

145.7 (

140.0-150.0)

5.93

147.5 (

147.0-148.0)

1.00

146.5 (

129.0-160.0)

12.92

Greatest Length of Skull

35.8

(35.2-36.2)

0.61 15.6

84.4

2.30*

35.3

(33.9-37.5)

0.44

35.0

(33.5-36.3)

0.57

34.9

(33.6-36.2)

0.88

35.7

(34.3-36.7)

0.52

35.9

(34.1-38.2)

0.41

36.4

(35.8-37.0)

0.41

35.2

(33.8-36.6)

1.62

35.5

(34.2-36.6)

1.10

34.4

(34.0-34.7)

0.40

34.6

(34.3-34.8)

0.50

Length of Nasals

15.2

(14.7-15.7)

0.58 13.8

86.2

2.19*

14.3

(13.2-15.6)

0.34

13.9

(12.8-14.5)

0.40

14.2

(13.6-15.0)

0.61

14.6

(13.7-15.5)

0.39

14.8

(13.6-15.8)

0.27

14.8

(14.0-15.5)

0.47

14.6

(13.7-15.8)

0.87

14.6

(13.9-15.2)

0.54

13.8

(13.2-14.5)

0.75

13.9

(13.3-14.5)

1.20

Length

of Rostrum

15.4

(14.8-16.3)

0.92 10.5

89.5

2.16*

15.4

(14.2-16.3)

0.25

15.1

(14.1-15.8)

0.38

15.2

(14.2-16.0)

0.61

15.5

(14.8-16.0)

0.33

15.6

(14.6-16.6)

0.23

16.0

(15.1-16.7)

0.62

15.4

(14.4-16.4)

0.84

298

FIELDIANA: ZOOLOGY

Table 3. Continued.

Grouped

Mean (range) + 2 SE

Percentage of variation

locality

Locality Error

9
10
11

4
3
2

15.6 (15.2-16.0) ± 0.37
14.8 (14.2-15.5) ± 0.75
14.6 (14.4-14.8) ± 0.40

Length of Maxillary Toothrow

18.9

4.9

(4.7-5.1)

± 0.23

4.9

(4.7-5.2)

± 0.07

4.9

(4.6-5.2)

± 0.13

5.0

(4.9-5.1)

± 0.06

4.8

(4.6-5.0)

± 0.08

5.0

(4.4-5.3)

± 0.08

5.2

(4.9-5.6)

± 0.21

4.9

(4.7-5.1)

± 0.17

4.8

(4.6-5.0)

± 0.17

4.9

(4.8-5.1)

± 0.13

5.2

(5.0-5.4)

± 0.40

81.1

2.76**

Asterisks following F statistics indicate levels of significance (see footnote to Table 1). Percentages of variation are
based on variance components estimated from the entire data set. Grouped localities are defined in text and outlined
in Figure 1.

the Heteromyinae, Goldman (1911) placed H.
gaumeri in his H. desmarestianus group, but noted
that it was aberrant and not closely related to any
known species. Although H. gaumeri possesses
characters diagnostic for Heteromys, including up-
per and lower molars with three lophs and lower
permanent premolars with three lophs, it also
shares some characters with the sister group Lio-
mys not shared with other Heteromys, including
posterior sole of hind foot haired as in Liomys
rather than naked as in other Heteromys and early
disappearance of the enamel island between cin-
gulum and metaloph(id) of upper and lower mo-
lars, a condition somewhat intermediate between
the genera (Goldman, 191 l;Genoways, 1973). As-
suming Heteromys is monophyletic, the characters
shared between H. gaumeri and Liomys probably
are primitive (based on the "ojierational rule" out-
group procedure outlined by Watrous & Wheeler,
1981). Dorsal coloration of H. gaumeri also is
unusual in that the dorsum has a definite ochra-
ceous cast with a bright ochraceous lateral line
resembling Liomys pictus and L. spectabilis more
than other, darker Heteromys. However, dorsal
coloration in H. gaumeri might be convergent on
L/omys through their common occupation of xeric
forest and thorn scrub habitats rather than more
mesic forest characteristic of most other species
of Heteromys.

Karyology — Compared to the nominate
species in the Heteromys desmarestianus and H.

anomalus species groups, the karyotype of//, gau-
meri also is divergent (see fig. 2). Superficially, the
karyotypes of //. desmarestianus (2n = 60, FN =
66) and //. anomalus (2n = 60, FN = 68) appear
similar and might differ by a single rearrangement
(heterochromatic addition/deletion or pericentric
inversion), whereas the karyotype of //. gaumeri
(2n = 56, FN = 76) differs by a minimum of seven
rearrangements from //. desmarestianus and six
from //. anomalus. The karyotype reported herein
for //. desmarestianus differs from that reported
by Genoways (1973; 2n = 60, FN = 82) in having
a larger number of acrocentric elements. Popula-
tions of //. desmarestianus are variable in fun-
damental number and the karyotype reported
herein is among the lowest FN karyotypes known
for the species (D. S. Rogers, pers. comm.).

Conclusions— We agree with Goldman (1911,
p. 30) that Heteromys gaumeri "is a somewhat
aberrant species, presenting characters which set
it off from all the others [species of the H. des-
marestianus group]." The distinctive morpholog-
ical, ecological, and karyotypic features of//, gau-
meri distinguish it from other members of the //.
desmarestianus group (including //. goldmani; D.
S. Rogers, pers. comm.). Shared (probably prim-
itive) characters with Liomys suggest that H. gau-
meri might represent an early branch of the lineage
leading to other Heteromys (which share probable
derived states for these characters). At present, we
believe the distinctness and unresolved phyloge-

ENGSTROM ET AL.: HETEROMYS GAUMERI

299

XI &{ii SI M >(» 99 MM

AiS Mr A«

A^it/s.

OA Af> A* OH Awn. /in Ml Aft
r%^ AA AA 4

X Y

iri B Kn .. 11 a it

9A tl DC «» ai t* It im

DA lid t\ »t tt) U M AA

OA nn 60 Oft Aft »»

X X

10 n QO AA A9 00 to A«

i^a i<( (in ft A #0 #11 ak aq

HA (fS llW 411 HA #14 jf^

X X

Fig. 2. Representative karyotypes of Heteromys: A, karyotype of a male H. gaumeri (2n = 56, FN = 76) from
7.5 km W Escarcega, Campeche, Mexico; B, karyotype of a female H. desmarestianus (2n = 60, FN = 66) from 9.4
km S Palenque, Chiapas, Mexico; C, karyotype of a female H. anomalus {2n = 60, FN = 68) from Caripito, Monagas,
Venezuela. Insert shows sex chromosomes of a male.

300

FIELDIANA: ZOOLOGY

netic position of H. gaumeri would best be em-
phasized by removing it from the H. desmaresti-
anus group, and recognizing it as a divergent species
at the same cladogenic level as the species groups
in the subgenus Heteromys. Further investigation
of heteromyines might indicate that H. gaumeri
warrants subgeneric recognition.

Species Account

I Heteromys gaumeri Allen and Chapman, 1897

Heteromys gaumeri Allen and Chapman, 1897, Febru-
ary. Bull. Amer. Mus. Nat. Hist., 9: 9.

HoLOTYPE— Adult male, skin and skull, amnh
12028/10461; from Chichen Itza, Yucatan, Mex-
ico. Type examined.

Distribution— Northern Belize; Peten, Gua-
temala; and Campeche, Quintana Roo, and Yu-
catan, Mexico (see fig. 1); subdeciduous-subpe-
rennial tropical rain forest, tropical deciduous
forest, and thorn scrub forest from sea level to
100 m.

Description— Dorsal coloration of adults ranges
from dark to medium gray, with heavy admixture
of orange buff hairs lending an overall ochraceous
cast to the otherwise gray dorsum (adults in worn
pelage appear more ochraceous and molting in-
dividuals often have a 'salt and pepper' appear-
ance); rich orange buff lateral line, usually broad
and conspicuous, extending from cheeks to base
of tail, ochraceous hairs often extending onto dor-
sal and ventral surfaces of ankle; margins of fore-
arm orange buff, interrupted on dorsal surface by
white line; venter and feet white; tail well haired,
grayish brown above, dull white below, with con-
spicuous terminal tuft of hairs; ears dusky, lightly
edged with dull white; sole of hind foot haired,
posteriorly. Subadults medium to dark gray above,
dorsum without ochraceous hairs; ochraceous lat-
eral line faint and narrow. Juvenile pelage similar
to that of subadults, but spiny hairs on dorsum
absent. Tail longer than head and body; soles of
hind feet with six tubercules; body size medium
for the genus. Skull size medium, with relatively
large auditory bullae; lower permanent premolar
with three lophs; upp>er and lower molars with
three lophs, enamel island formed between meta-
loph(id) and cingulum disappearing quickly with
wear. 2n = 56, FN = 76.

Couv\v.\so^s— Heteromys gaumeri is geo-
graphically isolated from all other heteromyines

except H. desmarestianus, which it might contact
at the southern edge of the Yucatan Peninsula (see
Jones et al., 1 974). Heteromys gaumeri differs from
H. desmarestianus in averaging smaller in most
external and cranial measurements; having a broad,
bright ochraceous lateral line, extending onto
cheeks and ankles (a narrow, pale ochraceous lat-
eral line often is present in H. desmarestianus, but
seldom extends onto cheeks or ankles); having soles
of hind feet haired posteriorly (this area is naked
in H. desmarestianus and all other Heteromys);
having a relatively well-haired tail with terminal
tuft (the tail is sparsely haired in H. desmaresti-
anus, without a conspicuous terminal tuft); having
relatively large auditory bullae; and in having a
diploid number of 56 chromosomes (compared to
60 in H. desmarestianus). For additional com-
parisons with other heteromyines, see Goldman
(191 1) and Genoways (1973).

Remarks— Laurie (1957, p. 387) assigned eight
specimens from three localities in the state of Yu-
catan, Mexico (Tekom, 2; X-Cala-Koop, 1; Chi-
chen Itza, 5) to Heteromys desmarestianus. One
of us (HHG) reexamined her material, which is
stored in alcohol; based on size, coloration, and a
haired posterior sole of the hind foot, all are as-
signable to H. gaumeri. Consequently, the north-
ernmost locality for H. desmarestianus on the Yu-
catan Peninsula is 85 km W Chetumal, Quintana
Roo, Mexico (Jones et al., 1974). Two other spec-
imens, from Kate's Lagoon, Belize, identified by
Laurie (1957, p. 387) as H. desmarestianus, are
referable to H. gaumeri. These specimens, along
with additional material from Honey Camp La-
goon (reported by Izor &. McCarthy, 1984) and
Rockstone Pond reported here, suggest that H.
gaumeri is distributed throughout northern Belize.

Acknowledgments

Permits to collect specimens in Mexico were
kindly provided by the Director General of the
Departmento de Conservacion de la Fauna Sil-
vestre. Especial thanks are extended to Biol. Fran-
cisco Rodriguez Gallegos, Ing. Antonio Sanchez,
and personnel of Campo Experimental Forestal
"El Tormento" for their hospitality during field-
work near Escarcega, Campeche. We also thank
the many curators who permitted examination of
specimens in their care. E. Jaquelin Dietz provided
the computer program for testing the association
between distance matrices. Kenneth Schoenly pre-
pared the final copy of Figure 1. D. S. Rogers

ENGSTROM ET AL.: HETEROMYS GAUMERI

301

provided information and helpful discussion on
character-states within the Heteromyinae. Por-
tions of this study were funded by the Texas Ag-
ricultural Experiment Station (project H-1977 to
D. J. Schmidly) and a Faculty Organized Research
Grant, Angelo State University. Fieldwork in Ven-
ezuela was supported by NSF Grant DEB 79-2 1519
to J. W. Bickham and the Carnegie Museum of
Natural History through the M. Graham Netting
Research Fund, established by a grant from the
Cordelia S. May Charitable Trust.

Literature Cited

Allen, J. A., AND F. M. Chapman. 1897. Onmammals
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Appendix
Specimens Examined

MEXICO: Campeche: Apazote, near Yohaltun,

6 (usnm); Campeche, 1 (usnm); 5 km S Cham-
poton, 10 m, 3 (ku); Dzibalchen, 1 (ku); 7 km N,
5 1 km E Escarcega (= Francisco Escarcega), 4 (ku);
7.5 km W Escarcega, 65 m, 94 (15 ku, 79 tcwc);

7 km E Escarcega, 4 (ku); 103 km SE Escarcega,
2 (ku); La Tuxpena, Champoton, 1 2 (usnm); San
Jose Carpizo, 3 (fmnh); San Jose Carpizo, 45 km
S Campeche, 19 (ummz); San Juan, 4 (fmnh).
Quintana Roo: 2 km N, 8 km W Bacalar, 1 1 (tcwc);

8 mi NNE Felipe Carrillo Puerto, 4 (tcwc); 4 km
NNE Felipe Carrillo Puerto, 30 m, 9 (ku); 2.5 mi
NNE Felipe Carrillo Puerto, 1 (tcwc); La Vega, 5
(usnm); Pueblo Nuevo Xcan, 10 m, 1 (ku); 1.5
km S, 1 km E Pueblo Nuevo Xcan, 1 (ku); Puerto
Morelos, 4 (usnm). Yucatan: Calcehtok, 55 km SW
Merida, 2 (1 ku, 1 ummz); Chichen Itza, 10 m, 36
(15 amnh, 11 bmnh, 2 fmnh, 1 ku, 7 usnm);
Chichen Itza, Cenote Xtoloc, 1 (tcwc); 2 km E

Chichen Itza, Cenote Seco, 5 (tcwc); 4 km E Dzi-
tya, 1 (mmnh); Esmeralda, Laguna de Chichan-
canab, 2 (1 ku, 1 ummz); Esmeralda, 45 km S Peto,
4 (ummz); 1 km SW Casa de la Esmeralda, Laguna
de Chichancanab, 2 (1 ku, 1 unam); 2 km SSW
Casa de la Esmeralda, Laguna de Chichancanab,
3 (1 AMNH, 2 unam); La Ceiba, 2 km SE Laguna
de Chichancanab, 1 (unam); 66 km NE Merida,

1 (ku); 14 km SW Muna, 1 (rru); Oxkutzcab,
Ebiztic (cave deposit), 6 (usnm); Peto, 3 (ku); 3
km N Piste, 16 (7 ku, 6 mmnh, 3 ttu); 2 km N
Piste, 2 (ku); Piste, 10 m, 2 (ku); Progreso, 1 (usnm);
Santa Rosa, 1 (ummz); Santa Rosa, 20 km S Peto,

2 (ummz); Santa Rosa, 25 km S Peto, 1 (ummz);
1 km SSW Santa Rosa, 3 (1 ku, 1 ummz, 1 unam);
Tekom, 2 (bmnh); 6 km N Tizimin, 1 (ku); Tun-
kas, 7 (usnm). Buena Vista, 1 (fmnh) and X-Cala-
Koop, 1 (bmnh) were not located exactly. GUA-
TEMALA: Peten: Chuntuqui, 2 (usnm); Laguna
de Sotz [= Zotz], 2 (usnm). BELIZE: Belize: Kate's
Lagoon, 2 (bmnh); Rockstone Pond, 1 1 (rom). Or-
ange Walk: Honey Camp Lagoon, 2 (fmnh).

Additional Records (Hatt et al., 1953, p. 64,
unless noted otherwise)— MEXICO: Yucatan: Ac-
tun Chacaljas, 3 km SSW Calcehtok (cave de-
posit); Actun Coyok [= Coyoc], 3.5 km SSE
Oxkutzcab (cave deposit); Actun Has, 3.5 km WSW
Yokat (cave deposit); Actun Lara, 3 km SW Yokat
(cave deposit); Actun Oxkintok, 3 km SW Santa
Cruz (cave deposit); Actun Spukil, 4.5 km SSW
Calcehtok (cave deposit); Loltun, 5 km SW
Oxkutzcab (cave deposit); Xbac (Gaumer, 1917,
p. 13); Yaxcach (Gaumer, 1917, p. 13). GUA-
TEMALA: Peten: 1 1 km NE JHores, 3.2 km inland
SE shore Laguna Peten Itza, ca. 100 m (Ryan,
1960, p. 11).

ENGSTROM ET AL.: HETEROMYS GAUMERI

303

Species Groups of Spiny Rats,

Genus Proechimys (Rodentia: Echimyidae)

James L. Patton

ABSTRACTS

Nine species groups of the spiny rat subgenus Proechimys, family Echimyidae, are defined
on the basis of bacular characters and qualitative features of the cranium. The latter include
the structure of the incisive and mesopterygoid foramina, temporal ridge and infraorbital canal
development, and counterfold pattern of the cheekteeth.

Three groups are apparently monotypic, including the decumanus -group of southwestern
Ecuador and adjacent Peru, the canicollis-group of northeastern Colombia and adjacent Ven-
ezuela, and the s i mo nsi -group of the western Amazon Basin from Colombia to northern Bolivia.
The remaining six groups are polytypic, but the number of species in each remains unclear.
The semispinosus-group ranges from Central America south to southwestern Ecuador in the
Pacific lowlands; its only Amazonian representative is P. oconnelli from east-central Colombia.
The longicaudatus-group ranges from southeastern Colombia through the western Amazon
Basin into the northern Parana Basin of Brazil and northern Paraguay. The goeldii-group ranges
throughout the Amazon Basin from eastern Peru to eastern Brazil. The guyannensis-group
occurs from the coastal Guianan region through the Rio Negro and eastern half of the Amazon
Basin in Brazil, with an isolate in Goias and Minas Gerais states. The cuvieri-group has a
similar distribution, but extends further up the Amazon into northern Peru, with one isolate
in east-central Peru. Finally, the trinitatus-gcoup is found from north-central Colombia eastward
across northern Venezuela to Trinidad.

Nueve grupos de especies de la rata espinosa subgenero Proechimys, familia Echimyidae, son
definidos primariamente en las bases de caracteres baculares que son soportados por razgos
cualitativos del craneo. El ultimo incluye la estructura del foramen incisivo y mesopterigoideo,
arista temporal y desarroUo del canal infraorbital y patron de contraplegamiento de los dientes
postcaninos.

Tres grupos son aparentemente monotipicos, incluyendo el grupo decumanus del suroccidente
del Ecuador y adyacente Peru, el grupo canicollis del nororiente de Colombia y adyacente
Venezuela, y el grupo simonsi del occidente de la cuenca amazonica desde Colombia hasta el
norte de Bolivia. Los seis grupos remanentes son politipicos, pero el numero de especies en
cada uno p>ermanece obscuro. El grupo semispinosus se extiende desde el sur de Centroamerica
hasta el suroccidente ecuatoriano en las tierras bajas del Pacifico; su solo representante de la
cuenca amazonica el P. oconnelli del centroriente colombiano. El grupo longicaudatus se dis-
tribuye desde el suroriente de Colombia a traves del occidente de la cuenca amazonica hasta
el norte de la cuenca del Parana en Brazil y norte de Paraguay. El grupo goeldii se distribuye
a traves de la cuenca amazonica desde el Peru oriental hasta el Brazil oriental. El grupo
guyannensis ocurre desde la region costera guayanesa a traves de rio Negro y la mitad oriental
de la cuenca amazonica en Brazil, con un aislamiento de poblaciones en los estados de Goias

From Museum of Vertebrate 2^ology, University of
California, Berkeley, CA 94720.

PATTON: SPECIES GROUPS OF PROECHIMYS 305

y Minas Gerais. El grupo cuvieri tiene una distribucion similar, pero se extiende mas arriba del
Amazonas en el interior del norte del Peru, con un aislamienlo de poblaciones en el centroriente
peruano. Finalmente, el grupo trinitatus es encontrado desde el centronorte de Colombia hacia
el oriente a traves del norte de Venezuela hasta Trinidad.

Nove grupos de especies de ratos-de-espinho, do subgenero Proechimys, familia Echimyidae,
sao definidos principalmente na base de carateres baculares que concordam tambem com
caracteres qualitativos do cranio. Estes incluem: a estrutura dos incisivos e do forame mesop-
terigoideo, o desenvolvimento das temporas e do canal infraorbital, e o padrao dos molares.

Tres grupos sao aparentemente monotipicos, incluindo o grupo decumanus do sudoeste do
Equador, o grupo canicollis do nordeste da Colombia e das areas adjacentes na Venezuela, e o
grupo simonsi da Bacia Amazonica ocidental, desde a Colombia ate o norte da Bolivia. Os seis
grupos restantes sao politipicos, mas o numero de especies em cada continua incerto. O grupo
semispinosus estende-se da America Central ao sudoeste do Equador, nas planicies do Pacifico.
Seu unico representante na Bacia Amazonica e P. oconnelli, do Centro-leste da Colombia. O
grupo longicaudatus estende-se do sudeste da Colombia, atraves da Amazonia ocidental, ate o
norte da Bacia do Parana no Brasil e no norte do Paraguai. O grupo goeldii ocorre na Bacia
Amazonica, do leste do Peru ao leste do Brasil. O grupo guyannensis ocorre da costa guianense,
ate o Rio Negro e a regiao oriental da Bacia Amazonica no Brasil, com uma especie isolada
nos estados de Goias e de Minas Gerais. A distribuifao do grupo cuvieri e parecida, mas este
ocorre tambem ate o norte do Peru, com uma especie isolada no centro-leste do Pais. Por final,
o grupo trinitatus e encontrado do centro-norte da Colombia, atraves da Venezuela, ate Trin-
idade.

Introduction

Spiny rats of the genus Proechimys represent one
of the most diverse groups of Neotropical rodents;
with the possible exception of tuco-tucos, Cteno-
mys, the number of taxa of spiny rats is unmatched
by any other caviomorph (Woods, 1984). The ge-
nus extends throughout lowland forests from Nic-
aragua to northern Paraguay and the coastal re-
gions of Brazil. Despite this diversity and large
geographic range, however, the group is taxonom-
ically one of the most poorly understood among
all of the Neotropical mammals. Only a few stud-
ies have succeeded in recognizing the number of
taxa sympatric at any single locality (e.g., Moojen,
1948; Patton & Gardner, 1972), and no study has
been able to follow geographic character trends
within a clearly defined taxon over any but the
shortest distances. Diagnosis of species and hence
definition of natural units in Proechimys have been
severely hampered by the often extreme level of
variability within and between populations for
most morphological characters that have been ex-
amined (Moojen, 1948; Hershkovitz, 1948; Pat-
ton &. Gardner, 1972). Even karyotypes, which
have proven useful in differentiating sympatric taxa
of spiny rats (Patton & Gardner, 1972), are often

highly variable geographically (Reig & Useche,
1976;Reigetal., 1 980; Gardner & Emmons, W84).

In this report I will challenge some of these pre-
cepts of character instability (see Thomas, 1928,
p. 262) by using the structure of the baculum and
specific characters of the cranium to define major
taxonomic groups of spiny rats. Taxa can be di-
agnosed, despite both within- and between-pop-
ulation variation, and the patterns of character
variation over geography are coherent, permitting
a consistent view of these taxa throughout their
range. Some of the more traditional characters that
have been used to make taxonomic decisions in
spiny rats (e.g., counterfold patterns on the cheek-
teeth) are not chaotic in their variation patterns,
but are quite helpful in defining units.

This paper will consider only members of the
subgenus Proechimys, excluding entirely the group
of species found along the Atlantic highlands of
Brazil that compose the subgenus Trinomys.

Species Groups of Proechimys

In the section below I provide the basic species
groups of spiny rats, subgenus Proechimys, listing

306

HELDIANA: ZOOLOGY

Fig. 1 . Geographic distribution of taxa of the simonsi- and guyannensis-^ovii>s. Type locaHties for each included
taxon are indicated by stars; dots represent other locahties hsted in the Appendix.

those named forms I consider as component parts.
In recognizing these groups and their membership
I make no conclusions here as to the specific, sub-
specific, or other status of these names. Because
of the plethora of names available and the con-
fusion with which each has been applied to the
genus over the past century, this synopsis is pro-
vided first to allow for coherent discussion; the
documentation upon which these decisions are
based follows.

I recognize nine species groups within the sub-
genus Proechimys. Five of these are widespread,
while the remaining ones are more restricted in
their ranges. Maps of the distribution of each group,
with localities of included holotypes, are presented
in Figures 1-4 (see Specimens Examined for lists
of localities). Unless otherwise stated, allocation
of any given holotype to a specific species-group
is based on my examination of that specimen. The

groups are defined by a combination of palatal
(particularly incisive foramina and mesopterygoid
fossa) characters, counterfold patterns of the
cheekteeth, temporal ridge development, infraor-
bital notch development, and bacular characters
(see below). In each case, the group name is taken
from the oldest assignable name for that unit.

guyannensis-%Tou^

Named forms in this group include:

guyannensis (E. Geoffroy, 1 803)
cherriei Thomas, 1 899
roberti Thomas, 1 90 1
vacillator Thomas, 1 903
oris Thomas, 1 904
warreni Thomas, 1 905

PATTON: SPECIES GROUPS OF PROECHIMYS

307

SNmi
(Mkm

Fig. 2. Geographic distribution of taxa of the goeldii-group (dots) and decumanus-group (triangles). Type localities
of taxa are indicated by stars.

boimensis Allen, 1916
arescens Osgood, 1 944
riparum Moojen, 1948
ar abupu Moo}tn, 1948

Comments— On the basis of septal patterns in
the bullae, Gardner and Emmons (1984) included
these taxa in their brevicauda-group, an all-inclu-
sive unit combining taxa that are here allocated
to six separate groups. As will be apparent below,
my guyannensis-group only shows close similarity
to the taxa included in the 5/wo/J5/-group. Mem-
bers of these two groups share virtually no char-
acters with the remaining taxa listed by Gardner
and Emmons (1984) in their brevicauda-group.

This group is confined in its distribution to the
Guianan region and southern Venezuela through
the central Amazon Basin of Brazil, with an isolate
(roberti) in Minas Gerais and Goias states in Brazil
(see map, fig. 1). It is sympatric with members of

the cuvieri-group in the Guianan region (see Petter,
1978) and with those of both the cuvieri- and goel-
dii-groups in the central Amazon Basin.

goeldii-group

Included are the following named forms:

goeldii Thomas, \ 90 5
steerei Goldman, 1911
kermiti M\Qn, 1915
pachita Thomas, 1923
hilda