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.

XIX. Literature Cited

AcosTA, JosE DE- 1590. Historia natural y moral de
las Indias en que se tratan las cosas notables del cielo,
elementos, metales, plantas y animales dellas y los
ritos y ceremonias, leyes y gobiemo y guerras de los
Indios. Juan de Leon, Seville, 535 + (36) pp. [not
seen].

. 1894. Historia natural y moral de las Indias.

Escrita por el P. Joseph de Acosta de la Compaiiia de
Jesus. Publicada en Sevilla en 1 590 y ahora fielmente

nELDL\NA: ZOOLOGY

reimpresa de la primera edicion. Ramon Angles, Ma-
drid, 2 vols.

Anghiera, see Martyr.

Anson, George. 1 748. A Voyage Round the World in
the Years MDCCXL, I, II, III, IV by George Anson
. . . Esq. Compiled and published by Richard Walter.
J. & P. Knapton, London.

AviLA Pikes, Fernando Dias de. 1 965. The type spec-
imens of Brazilian mammals collected by Prince
Maximilian zu Wied. American Museum Novitates,
no. 2209: 1-21.

. 1 974. CaracterizafSo zoogeografica da Provin-

cia Amazonica. I. Expedi^oes cientificas na Amazonia
Brasiliera. Anales Academia Brasiliera de Ciencias,
46(1): 133-158.

AzARA, Felix de. 1801. Essais sur I'histoire naturelle
des quadrupedes de la Province du Paraguay. Trans-
lated from the original Spanish by M. L. E. Moreau-
Saint-Mery. Charles Pougens, Paris, vol. 1, xxvii +
366 pp.; vol. 2, 499 pp.

. 1802. Apuntamientos para la historia natural

de los quadrupedos del Paraguay y Rio de la Plata. La
Viuda de Ibarra, Madrid, vol. 1, xiii + 318 pp.; vol.
2, (3) + 328 pp.

. 1809. Voyages dans I'Amerique Meridionale.

Contenant la description geographique politique et
civile du Paraguay et de la riviere de La Plata; I'histoire
de la decouverte et de la conquete de ces contrees; des
details nombreaux sur leur histoire naturelle, et sur les
peuples sauvages qui les habitent, le recit des moyens
employes par les Jesuits pour assujetir et civiliser les
indigenes, etc. Publics d'apres les manuscrits de Tau-
teur, avec une notice sur sa vie et ses ecrits. Translated
from the original Spanish and annotated by M. Son-
nini, with notes by G. Cuvier; edited by C. A. Walck-
enaer. Dentu, Paris, 4 vols, with atlas.

Bajon, M. 1777-1778. Memoires pour servir a I'his-
toire de Cayenne et de la Guiane Frangaise. Grange,
Paris, vol. 1, 1777; vol. 2, 1778.

Bancroft, Edward. 1769. An Essay on the Natural
History of Guiana in South America. T. Beckett and
P. A. DeHondt, London, (8) + 402 + (2) pp.

Barrere, Pierre. 1741. Essai sur I'histoire naturelle
de la France Equinoxiale ou de'nombrement des
plantes, des animaux, y des mineraux, que se trouvent
dans I'lsle de Cayenne, les Isles de Remire, sur les
cotes de la mer, & dans le continent de la Guyane.
Piget, Paris.

Bellin, Jaques Nicolas. 1763. Description geogra-
phique de la Guiane. Didot, Paris.

Bokermann. Werner C. A. 1957. Atualiza^o do iti-
nerario de viajem do Principe de Wied do Brasil (1815-
1817). Arquivos de 2^oologia (Rio de Janeiro), 10(3):

209-251.

Browne, Patrick. 1756. The Civil and Natural His-
tory of Jamaica. Privately printed, London.

BuFFON, George Louis Leclerc, Comte de. 1 753-1 789.
Histoire naturelle, generale et particuliere, avec la de-
scription du cabinet du roy. L'Imprimerie Royale, Paris,
44 vols., 1 750-1804; vols. 4-15, Quadrupedes par M.
de Buffon et M. Daubenton, 1753-1767; vol. 9, 1761;
supplement, vol. 107 par M. de Buffon, 1774-1789.

Catesby, Mark. 1731-1743. The Natural History of
Carolina, Florida and the Bahama Islands. Privately
printed, London, vol. 1,(2) + vii + (1) + xliv + 100 +
2 pp., 100 pis., fold, map; vol. 2, 2 + 120 + (6) + 2
pp., 120 pis.

Charlevoix, Pierre Francois Xavier DE. 1757. His-
toire du Paraguay. Oesaint, David, Durand, Paris, 6
vols.

Darwin, Chari,es Robert. 1839. Journal of Re-
searches into the Geology and Natural History of the
Various Countries Visited by H.M.S. Beagle. Under
the Command of Captain Fitzroy, R.N., from 1832
to 1836. Henry Colbum, London, xiv + 615 pp., 16
pis.

Edwards, George. 1743-1751. A Natural History of
Uncommon Birds, and of Some Other Rare and Un-
described Animals. Printed for the author at the Col-
lege of Physicians, London, 4 vols.

Fermin, Philippe. 1769. Description generale, histo-
rique, geographique et physique de la colonic de Su-
rinam. E. van Harrevelt, Amsterdam, 2 vols.

Ferreira Rodriguez, Alexandre, see Rodriguez Fer-
reira, Alexandre.

Gaffarel, Paul. 1907. See Martyr, Anghiera Pierre.

Gay, Claudio. 1 847. Historia fisica y politica de Chile.
Vol. 1, Zoologia. Museo de Historia Natural de San-
tiago [de Chile], Malde y Renow, Paris.

GuMiLLA, Jose. 1741. El Orinoco ilustrado, historia
natural, civil y geograhca, de este gran rio, y de sus
caudalosas vertientes; gobiemo, usos y costumbres de
los Indios, sus habitadores con nuevas, y utiles noticias
de animales, arboles, frutos, aceytes, resinas, yervas,
y singulares a nuestra santa fe, y casos de mucha edi-
licacion. M. Fernandez, Madrid.

. 1758. Histoire naturelle, civile et geographique

de rOrenoque. Translated from the Spanish. 2nd edi-
tion by M. Eidous. Veuve de F. Firard, Avignon, 3
vols.
Hernandez, Francisco. 1651. Nova plantarum, ani-
malium et mineralium mexicanorum historia. B.
Deuersini & Z. Masotti, typis V. Masacardi, Rome,
(16) + 950 + 90 pp.

. 1959. Historia natural de Nueva Espafia, vol.

2. Translated by Jose Rojo Navarro. Universidad Na-
cional de Mexico, D.F.

Herrera y Tordesillas, Antonio de. 1 60 1 . Historia
general de las Indias occidentalis: o, de los hechos de
los castellanos en las islas y tierra firme de mar oceano,
escrita por Antonio de Herrera, cronista mayor de Su
Magestad de las Indias y de Castilla. En la Emplen-
terea, 4 vols, [work not seen, cited from reviewer com-
ments].

. 1601-1615. Historia general de las Indias oc-
cidentalis: o, de los hechos de los castellanos en las
islas y tierra firme del mar oceano, escrita por Antonio
de Herrera, cronista mayor de Su Magestad de las
Indias y de Castilla. Madrid, 9 parts in 4 vols, [work
not seen, cited from reviewer comments].

1725-1726. The General History of the Vast

Continent and Islands of America, Commonly Called,
the West Indies, from the First Discovery Thereof:
With the Best Accounts the People Could Give of the
Antiquities. Collected from the original relations sent

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

to the Kings of Spain. Translated into English by Capt.
John Stevens. Batley, London, vol. 1, 1725; vols. 2-
6, 1726.
. 1728. Historia general de las Indias occiden-

talis: o, de los hechos de los castellanos en las islas y
tierra firme del mar oceano, escrita por Antonio de
Herrera, cronista mayor de Su Magestad de las Indias
y de Castilla. En ocho decadas. Sigue a la ultima de-
cada. la Description de las Indias, por el mismo autor.
J. B. Verdussen, Amberes [work not seen, cited from
Library of Congress catalogue].

Hershkovitz, Philip. 1972. The Recent mammals of
the Neotropical Region: A zoogeographic and ecolog-
ical 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, Al-
bany.

Humboldt. Alexander von. 1811-1812. Voyage de
Humboldt et Bonpland. Observations de zoologie et
d'anatomie comparee. Pt. 2, vol. 1 , Recueil d'obser-
vations de zoologie et d'anatomie comparee, faites
dans I'ocean Atlantique dans Tinterieur du nouveau
continent et dans la mer du sud pendant les annees
1799, 1800, 1801, 1802 et 1803. Levrault Schoell,
Paris, viii + 368 pp., 40 pis.

. 1838. Ueber den Manati des Orinoko. Trans-
lated from the original manuscript in French by the
editor. Wiegmann's Archiv fiir Naturgeschichte, 4(1):
1-10.

. 1884. Personal Narrative of Travels to the

Equinoctial Regions of America During the Years
1799-1804 by Alexander von Humboldt and Aime
Bonpland. Translated from the original French and
edited by Thomasina Ross. George Bell and Sons,
London, 3 vols.

Ihering, Herman von. 1 902. Natterer e Landsdorff.
Exploradores antigos do estado de Sao Paulo. Revista
do Museu Paulista (Sao Paulo), 5: 13-34.

Kalm, Pehr. 1753-1761. En resa til Norra America.
Pa Kong. Svenska Vetenskapsacademiens befallning,
och publici Kostnad. Stockholm, 3 vols.

Kraft, Richard. 1983. Die von J. B. von Spix bes-
chriebenen neotropischen Primaten und Chiropteren.
Verzeichnis der in der Zoologischen Staatssammlung
Miinchen aufbewahrten Typusexemplare. Spixiana
Suppl., 9: 429-441.

KuHL, Heinrich. 1 820. Beitrage zur Zoologie und ver-
gleichenden Anatomic, Abt. 1. Frankfurt am Main,
1 52 + unnumbered pp.

Langguth, A. 1978. Revived application in the case
of the names for South American rodents published
by Brants ( 1 827). Z.N.(S.) 1 775. Bulletin of Zoological
Nomenclature, 35(2): 115-120.

Linnaeus, Carolus. 1758. Systema Naturae. Regnum
animate, ed. 10, vol. 1. Holmiae, 4 + 824 + 3 pp.

. 1 766. Systema Natura per regna tria naturae,

secundum classis, genera, species, cum characteribus,
differentius, synonymis locis . . . , ed. 12 reformata,
vol. 1. L. Salvii, Holmiae, 1 + 327 + 36 pp.

Marcgraf [or Marcgrav, or Marggraf] de Lieb-
STAD, Georg. 1648. Historiae rerum naturalium
Brasiliae, libro octo cum appendice de Tapuyis, et Chi-
lensibus. loannes de Laet, Antvverpianus, In ordinem

digessit & annotationes addidit multas, &. varia ab
auctore omissa supplevit & illustravit. In Piso, W.,
Historia Naturalis Brasilliae, asspicio et beneficio II-
lustras. I. Maritii Com. Nassau. Issius provincije et
Maris summi praefecti adomata in qua non tantum
plantx et animalia sed et indiginarum morbi, ingenia
et mores describuntur et iconibus scipta quingentas
illustrantus. Fransiscum Hackium, et Amstelodami,
Lud, Elizevirium, Leiden.

Martyr, Anghiera Pierre. 1907. De orbe novo. Les
huit decades. Translated from Latin with notes and
comments by Paul Gaffarel. Recueil de voyages et de
documents pour servir a I'histoire de la geographic
depuis le XIII' jusqu'a la fin du XVI"^ siecle. No. XXI.
Imprimerie Nouvelle Noel Texier, La Rochelle, France.

. 1912. De orbe novo. The Eight Decades of

Peter Martyr d'Anghiera. Translated from the Latin
with notes and introduction by Francis Augustus
MacNutt. G. P. Putnam's Sons, New York & London,
2 vols.

Miller, Gerrit S., Jr. 1929. Mammals eaten by In-
dians, owls, and Spaniards in the coast region of the
Dominican Republic. Smithsonian Miscellaneous
Collections, 82(5): 1-16.

Molina, Giovanni Ignazio. 1 782. Saggio sulla storia
naturale del Chili. S. Tommaso d'Aquino, Bologna,
367 + 1 pp.

Orbigny, Alcide Dessalines d'. 1835-1847. Voyage
dans I'Amerique Meridionale (le Bresil, la Republique
orientale de Uruguay, la Republique Argentine, la Pa-
tagonie, la Republique du Chile, la Republique de Bo-
livia, la Republique du Perou) execute pendant las
annees 1826, 1827, 1828, 1829, 1830, 1831, 1832, et
1833. Pi tois- Levrault et cie, Paris, Strasbourg, 9 vols.

Orbigny, Alcide d', and Paul Gervais. 1847. Mam-
miferes, vol. 4, pt. 2. In Orbigny, A. d'. Voyage dans
I'Amerique Meridionale (le Bresil, la Republique ori-
entale de I'Uruguay, la Republique Argentine, la Pa-
tagonie, la Republique du Chili, la Republique de Bo-
livia, la Republique du Perou) execute pendant les
annees 1826, 1827, 1828, 1829, 1830, 1831, 1832,
1833. Pitois-Levrault et cie, Paris, Strasbourg.

Osgood, Wilfred H. 1943. The mammals of Chile.
Publications of Field Museum of Natural History,
Zoological Series, 30: 1-268.

OviEDO Y Valdes, Gonzalo Fernandez de. 1 944- 1 945 .
Historia general y natural de las Indias, Islas y Tierra-
Firme del Mar Oceano. Prologo de J. Natalicio Gon-
zalez. Notas de Jose Amador de los Rios. Editorial
Guarania, Asuncion (Paraguay), 1 4 vols, [text and il-
lustrations said to be reproduced from the original
manuscript].

Pelzeln, August von. 1871. Ueber Omithologie
Brasiliens. Resultate von Johann Natterer's Reisen in
den Jahren 1817 bis 1835. A. Pichler's Witwe & Sohn,
Vienna.

. 1883. Brasilische Saugethiere. Resultate von

Johann Natterer's Reisen in den Jahren 1 8 1 7 bis 1 835.
Verhandlungen der kaiserlich-koniglichen zoologisch-
botanischen Gesellschaft in Wien, 33: 1-140.

PiLLERi, G., and Lucie Arvy. 1977. La expedicion de
Alcide d'Orbigny en Bolivia y el descubrimiento del
delphin boliviano de agua dulce Inia boliviensis (d'Or-

FIELDIANA: ZOOLOGY

bigny, 1834). Himanatomisches Institut, Bern, Swit-
zerland, 15 pp., 4 figs., 16 pis., fold. map.

PoEPPiG, Eduard. 1835-1836. Reise in Chile, Peru
und auf dem Amazonenstrome wahrend der Jahre
1827-1832. Friedrich Fleischer, Leipzig, vol. 1, xii +
466 pp.; vol. 2, V + 464 pp.

Rengger, J. R. 1830. Naturgeschichte der Saeuge-
thiere von Paraguay. Schweighauserschen Buchhand-
lung, Basel, Switzerland, 16 + 394 pp.

RoDRiGUES Ferreira, ALEXANDRE. 1971. Viagem fi-
losofica pelas capitanias do Grao-Para, Rio Negro,
Mato Grosso e Cuiaba. Iconografia. Vol. 2, Zoologia.
Conselho Federal de Cultura, Rio de Janeiro. Editora
Monumento S. A., Sao Paulo. Artes Graficas Gomes
de Souza S. A., Rio de Janeiro, 168 color plates.

. 1972. Viagem filosofica pelas capitanias do

mecophagusjubata). Proceedings of the Zoological So-
ciety. Annals of Natural History, ser. 1 , 4(23): 202.

1840. Information respecting botanical trav-

Grao-Para, Rio Negro, Mato Grosso e Cuiaba. Me-
morias. Zoologia, Botanica [nomenclatorial notes by
Luiz Carlos Souto; introduction by Jose Candido de
Melo Carvalho]. Conselho Federal de Cultura. De-
partamento de Imprenta Nacional, Rio de Janeiro,
246 pp.

Roth, Walter E. 1931. See Schomburgk, Richard,
and Schomburgk, Robert Hermann.

ScHiNZ, Heinrich R. 1821. Das Thierreich eingetheilt
nach dem Bau der Thiere als Grundlage ihrer Natur-
geschichte und der vergleichenden Anatomic von dem
Herm Ritter von Cuvier. Vol. 1 , Saugethiere und Vo-
gel. J. G. Cotta'schen Buchhandlung, Stuttgart and
Tiibingen.

. 1844-1845. Systematisches Verzeichniss aller

bis jetzt bekannten Saugethiere oder Synopsis Mam-
malium nach dem Cuvier'schen System. J. Gassmann,
Sohn, Solothum, Switzerland, 2 vols.

Schomburgk., Richard. 1847-1848. Reisen in Brit-
ish-Guiana in den Jahren 1840-1844. Im auftrag Sr.
Majestat des Konigs von Preussen. Nebst einer Fauna
und Flora Guiana's nach Vorlagen von Johannes Miil-
ler, Ehrenberg, Erichson, KJotzsch, Troschel, Cabanis
und Andem. Mit Abbildungen und einer Karte von
British-Guiana afgenommen von Sir Robert Schom-
burgk. Versuch einer Fauna und Flora von British-
Guiana ( 1 848). J. J. Weber, Leipzig, vol. 1 , 1 847; vols.
2-3, 1848.

. 1922-1923. Travels in British Guiana During

the Years 1840-1844 Carried out Under the Com-
mission of His Majesty the King of Prussia. Together
with a Fauna and Flora of Guiana According to the
Works of Johannes Miiller, Ehrenberg, Erichson,
KJotzsch, Troschel, Cabanis and Others. Including il-
lustrations and a map of British Guiana drawn by Sir
Robert Schomburgk. Translated and edited, with geo-
graphical and general indices and route maps, by Wal-
ter E. Roth. "Daily Chronicle," Georgetown, British
Guiana, vol. 1 (1922), 38 + 402 pp., 8 pis., 4 maps,
2 figs.; vol. 2 (1923), (12) + 443 pp., 8 pis., 7 maps.

Schomburgk, Robert Herman. 1 840. A Description
of British Guiana, Geographical and Statistical: Ex-
hibiting Its Resources and Capabilities, Together with
the Present and Future Condition and Prospects of the
Colony. Simpkin, Marshall, and Co., London.

. 1 840. Remarks on the greater ant-bear (Myr-

ellers. Recent expedition in Guiana. Annals of Natural
History, ser. 1, 4(24): 262-266; (25): 322-328; (26):
429-434; 5(28): 29-35; (31): 282-288; (32): 343-348;
(33): 399-403.

. 1841. Reisen in Guiana und am Orinoko wah-

rend der Jahre 1835-1839. Nach seinen Berichten und
Mitteilungen an die geographische Gesellschaft in
London. Edited by O. A. Schomburgk, with a foreword
by Alexander von Humboldt. Georg Wigand, Leipzig.

. 1931. Travels in Guiana and on the Orinoco

During the Years 1835-1839. According to his Re-
ports and Communications to the Geographical So-
ciety of London. Edited by O. A. Schomburgk, with
a preface by Alexander von Humboldt. English trans-
lation by Walter E. Roth. "The Argosy" Company,
Limited, Georgetown, British Guiana.

Seba, Albertius. 1734-1765. Locuplestissimi rerum
naturralium thesauri accurata descripti et iconibus ar-
tificiosissimisexpressio, per universam historiam. . . .
Jansson-Waesberg, Amsterdam, 4 vols., 449 color pis.

Spix, Jean de. 1 823. Simiarum et vespertilionum bra-
siliensium species novae: ou histoire naturelle des es-
pecies nouvelles de singes et de chauve-souris obser-
vees et recueillies pendant le voyage dans I'interieur
du Bresil. Typis Francisci Serephici Hiibschmanni,
Monaco, 16 + 72 pp., 38 color pis.

Spix, Johann Baptist von, and Carl Friedrich von
Martius. 1823-1831. Reise in Brasilien auf Befehl
Sr. Majestat Maximilian Joseph 1., Konigs von Baiem,
in den Jahren 1817 bis 1820. M. Lindauer, Munich,
vol. 1, 1823; vol. 2, 1828; vol. 3, 1831.

. 1824. Travels in Brazil in the Years 1817-

1820, Undertaken by Command of His Majesty the
King of Bavaria. Translated from the original German
by H. E. Lloyd. Longman, Hurst, Rees, Orme, Brown,
and Green, London, vol. 1, 24 + 327 pp., 4 pis.; vol.
2, 8 + 298 pp., 5 pis.
Stedman, John Gabriel. 1 796. Narrative of a Five
Years' Expedition Against the Revolted Negroes of
Surinam in Guiana on the Wild Coast of South Amer-
ica; from the Year 1772 to 1777: Elucidating the His-
tory of that Country, and Describing Its Productions,
viz. Quadruf>eds, Birds, Fishes, Reptiles, Trees, Shrubs,
Fruits, & Roots; with an Account of the Indians of
Guiana, & Negroes of Guinea. J. Johnson and J. Ed-
wards, London, vol. 1,18 + 423 pp., 18 pis., 3 maps;
vol. 2, 9 + 419 + (5) pp., 38 pis., 2 maps.

Stevens, John. 1726. See Herrera y Tordisillas, An-
tonio de.

Tamayo H., Manuel, and C. Daniel Frassinetti. 1 980.
Catalogo de los mamiferos fosiles y vivientes de Chile.
Boletin Museo Nacional de Historia Natural de Chile,
37: 323-399.

Thomas, Oldfield. 1908. On certain African and S.-
American otters. Annals and Magazine of Natural
History, ser. 8, 1: 387-395.

. 1911. The mammals of the tenth edition of

Linnaeus; an attempt to fix the type of the genera and
the exact bases and localities of the species. Proceed-

HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY

ings of the Zoological Society of London, 1911:
158.

120-

TiEFENBACHER, L., ED. 1983. Fcstschrift zu Ehren von
Dr. Johann Baptist Ritter von Spix. Spixiana, Suppl. 9.

TscHUDi, Johann Jacob von. 1844a. Mammalium
conspectus quae in Republica Peruana reperiuntur et
pleraque observata vel collecta sunt in itinerere. Ar-
chiv fiir Naturgeschichte, 10(1): 244-255.

. 1844b. Untersuchungen iiber die Fauna Pe-
ruana. Physiognomik von Peru (1844); Therologie
(1844b); Omithologie [with J. Cabanis] (1845-1846);
Herpetologie (1845 [1846]). Scheitlin und ZoUikofer,
St. Gallen, Switzerland.

. 1 846. Peru. Reiseskizzen aus den Jahren 1 838-

1842. Scheitlin und Zollikofer, St. Gallen, Switzer-
land.

1847. Travels in Peru During the Years 1838-

1 842. On the Coast, in the Sierra, Across the Cordi-
lleras and the Andes into the Primeval Forests. English
translation from the original German by Thomasina
Ross. Wiley and Putnam, New York.
Vazquez DE EspiNOSA, Antonio. 1942. Compendium
and description of the West Indies. Translated by
Charles Upson Clark. Smithsonian Miscellaneous
Collections, 102: i-xii; 1-862.

. 1948. Compendio y descripcion de las Indias

Occidentals. Transcrito del manuscrito original por
Charles Upson Clark. Smithsonian Miscellaneous
Collections, 108: i-xii; 1-801.

Wagner, Johann Andreas. 1840-1855. Die Sauge-
thiere in Abbildungen nach der Natur mit Beschrei-
bungen von Dr. Johann Christian Daniel von Schre-
ber, Supplementenbanden 1-5. Erlangen, Germany.

. 1 842. Diagnosen neuer Arten brasilischer Sau-

gethiere. Archiv fur Naturgeschichte, 8(1): 356-362.

. 1843. Diagnosen neuer Arten brasilischer

. 1844-1846. Die geographische Verbreitung der

Saugethiere. Abhandlungen der Mathem.-physika-
lischen Classe der Koniglich Bayerischen Akademie
der Wissenschaften, 4(1): 1-116 (1844); (2): 39-108
(1845); (3): 1-114 (1845); (3): 241-244, explanation
of maps, plates I-IX (1846).

1850. Beitrage zur Kenntniss der Saugethiere

America's. Abhandlungen der Mathem.-physikali-
schen Classe der Koniglich Bayerischen Akademie der
Wissenschaften (Miinchen), 5: Erste Abt. [Marsupi-
alia, Chiroptera], 1 19-208 (1847); Zweiten Abt. [Ro-
dentia], 269-332 (1847); Dritte Abt. Vierte Ordnung
Affen, 405-480(1848).

Waterhouse, George R. 1838-1839. Mammalia. /«
Darwin, C, The Zoology of the Voyage of H.M.S.
Beagle, Under the Command of Captain Fitzroy, Dur-
ing the Years 1832-1836. With Notes by Charles Dar-
win, part II. Smith, Elder and Co., London, no. 1,
1838; no. 2, 1838; no. 3, 1838; no. 4, 1839.

Wied-Neuwied, Maximilian Alexander Philipp, Prinz
von. 1820. Ueber ein noch unbeschriebenes Sau-
gethier aus der Familie der Nager. Isis (Oken's), 1: 43.

. 1820-1821. Reise nach Brasilien in den Jahren

1815 bis 1817. Heinrich Ludwig Bronner, Frankfurt
A.M., vol. 1, 1820; vol. 2, 1821.

. 1821. Ueber die Brasilianischen Hirschart. Isis

(Oken's), 2: 649-651.

. 1822-1831. Abbildungen zur Naturgeschichte

Brasiliens. (Recueil de Planches Coloriees d'Animaux
du Bresil.) Weimar, Germany, 90 color pis. with de-
scriptive text issued in 1 5 parts with 6 pis. each (30
pis. of mammals in parts 1-5, 1822-1824).

. 1826. Beitrage zur Naturgeschichte von

Brasilien, vol. 2. Gr. H. S. priv. Landes-Industrie-
Comtpoirs, Weimar, Germany.

Handflugler. Archiv fiir Naturgeschichte, 9(1): 365-
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.

Literature Cited

BoRDAS, A. F. 1939. Diagnosis sobre algunos mami-
feros de las capas con Colpodon del valle del Rio Chu-
but. Physis, 14:413-433.

Clemens, W. A., and L. G. Marshall. 1976. Amer-
ican and European Marsupialia. Pars 123. Marsupi-
alia. In Westphal, F., ed., Fossilium Catalogus. I: An-
imalia. W. Junk, The Hague, 1 14 pp.

DoNADio, O. E. 1983. Los lacertilios del Colhuehua-
pense de la provincia del Chubut, Argentina. Circular
Informativa de la Asociacion Paleontologica Argen-
tina, 11: 5-6.

Fleagle, J. G., and T. M. Bown. 1983. New Primate
fossil from Late Oligocene (Colhuehuapian) localities
of Chubut Province, Argentina. Folia Primatologica,
41: 240-266.

Gould, S. J. 1 983. Nature's Great Era of experiments.
Natural History, 7: 12-21.

KiRSCH, J. A. W. 1977a. The classification of marsu-
pials, pp. 1-48. In Hunsaker II, D., The Biology of
Marsupials. Academic Press, New York, San Francis-
co, London, 537 pp.

. 1977b. The comparative serology of Marsu-
pialia, and a classification of marsupials. Australian
Journal of Zoology, Supplementary Series, 52: 1-152.

KuRTfeN, B. 1971. The Age of Mammals. Columbia
University Press, New York, 250 pp.

Marshall, L. G. 1976. Revision of the South Amer-
ican fossil marsupial subfamily Abderitinae (Mam-
malia, Caenolestidae). Publicaciones Museo Munici-
pal de Ciencias Naturales de Mar del Plata "Lorenzo
Scaglia," 2(3): 57-90.

. 1980. Systematicsofthe South American mar-
supial family Caenolestidae. Fieldiana: Geology, n.s.,
5: i-vii, 1-145.

Marshall, L. G., R. Hoffstetter, and R. Pascual.
1983. Mammals and stratigraphy: Geochronology of
the continental mammal-bearing Tertiary of South
America. Palaeovertebrata, Memoire Extraordinaire,
Montpellier, 1983: 1-93.

McKenna, M. C. 1980. Early history and biogeogra-
phy of South America's extinct land mammals. In
Ciochon, R. L., and A. B. Chiarelli, eds.. Evolutionary
Biology of the New World Monkeys and Continental
Drift. Plenum Publishing Corp., New York, xvi + 528
pp.

Mendia, J. E., AND A. Bayarsky. 1981. Estratigrafia
del Terciario en el valle inferior del rio Chubut. VII
Congreso Geologico Argentino, San Luis (20-26 Sep-
tiembre 1981), Actas 3: 593-606.

Pascual, R. 1970. Evolucion de comunidades, cam-
bios faunisticos e integraciones biocenoticas de los
vertebrados cenozoicos de Argentina. Actas IV Con-
gresso Latinoamericano de Zoologia (Caracas, 10-16
de Noviembre de 1968), 2 (Paleontologia Sudameri-
cana): 991-1088.

. 1980a. Nuevos singulares tipos ecologicos de

marsupiales extinguidos de America del Sur (Paieo-
ceno tardio o Eocene temprano) del Noroeste Argen-
tino. Actas II Congreso Argentino de Paleontologia y
Bioestratigrafia y I Congreso Latinoamericano de Pa-
leontologia (Buenos Aires, 2-6 de Abril 1 978), 2: 151-
173.

1 980b. Prepidolopidae, nueva familia de Mar-

supialia Didelphoidea del Eoceno sudamericano.
Ameghiniana, 17(3): 216-242.

. 1981. Adiciones al conocimiento de Bona-

partherium hinakusijum (Marsupialia, Bonapartheri-
idae) del Eoceno temprano del Noroeste Argentino.
Anais II Congresso Latino-Americano de Paleonto-
logia (26 a 30 de Abril, 1981, Porto Alegre-Brasil), 2:
507-520.
. 1983. Novedosos marsupiales paleogenos de

la Formacion Pozuelos (Grupo Pastes Grandes) de la
Puna, Salta, Argentina. Ameghiniana, 20(3-4): 265-
280.
. 1984. La sucesion de las Edades-mamifero, de

los climas y del diastrofismo sudamericanos durante
el Cenozoico: fenomenos concurrentes. Anales de la
Academia Nacional de Ciencias Exactas, Fisicas y
Naturales, 36: 1 5-37.

PASCUAL 8c CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS

109

Pascual, R., and M. Bond. 1981. Epidolopinae
subfam. nov. de los Polydolopidae (Marsupialia, Po-
lydolopoidea). Anais II Congresso Latino- Americano
de Paleontologia (26 a 30 de Abril, 1981, Porto Alegre-
Brasil), 2: 479-488.

Pascual, R., AND H. E. Herrera. 1973. Adiciones al
conocimiento de Pliolestes tripotamicus Reig, 1955
(Mammalia, Marsupialia, Caenolestidae) del Plioceno
superior de la Argentina. Ameghiniana, 10(1): 36-50.

. 1975. Stilotherium Ameghino, 1887, el mas

primitivo Caenolestidae conocido. Consideraciones
sobre la transicion Didelphidae-Caenolestinae (Mar-
supialia). Actas I Congreso Argentine de Paleontologia
y Bioestratigrafia (12-16 Agosto, 1 974, Tucuman, Ar-
gentina), 2: 417-430.

Pascual, R., and O. E. Odreman Rivas. 1971. Ev-
olucion de las comunidades de los vertebrados del
Terciario argentine. Los aspectos paleozoogeograficos
y paleoclimaticos relacionados. Ameghiniana, 8(3-4):
372-412.

Pascual, R., J. PiSANO, and E. J. Ortega. 1965. Un
nuevo Octodontidae (Rodentia, Caviomorpha) de la
Formacion Epecuen (Plioceno medio) de Hidalgo
(Provincia de La Pampa). Ameghiniana, 4(1): 19-30.

Pascual, R., M. G. Vucetich, G. J. Scillato-Yane,
AND M.Bond. 1985. Main pathways of mammalian
diversification in South America, pp. 219-247. In
Stehli, F., and S. D. Webb, eds.. The Great American
Biotic Interchange. Plenum Publishing Corp., New
York.

Patterson, B. 1952. Un nuevo y extraordinario mar-
supial deseadiano. Revista del Museo Municipal de
Ciencias Naturales, Mar del Plata, 1: 39-44.

. 1958. Affinities of the Patagonian fossil mam-
mal Necrolestes. Breviora (Museum of Comparative
Zoology), 49: 1-14.

Paula Couto, C. de. 1952. Fossil mammals from the
beginning of the Cenozoic in Brasil. Marsupialia: Poly-
dolopidae and Borhyaenidae. American Museum
Novitates, 1559: 1-27.

. 1961. Marsupiais fosseis do Paleoceno do Bra-
sil. Anais Academia Brasileira Ciencias, Rio de Ja-
neiro, 33(3/4): 321-333.

. 1979. Tratado de Paleomastozoologia. Aca-

demia Brasileira de Ciencias, Rio de Janeiro, 590 pp.

Reig, O. A. 1981. Teoria del origen y desarrollo de la
fauna de mamiferos de America del Sur. Monogra-
phiae Naturae (Publicadas por el Museo Municipal de
Ciencias Naturales "Lorenzo Scaglia"), Mar del Plata,
1: 1-162.

Ride, W. D. L. 1964. A review of Australian fossil
marsupials. Journal of the Royal Society of Western
Australia, 47(4): 91-131.

Simpson, G. G. 1945. The principles of classification
and a classification of mammals. Bulletin of the Amer-
ican Museum of Natural History, 85: i-xxvi, 1-350.

. 1953. The Major Features of Evolution. Co-
lumbia University Press, New York, 248 pp.

1 96 1 . Life of the Past. Yale Paper Bound, Yale

University Press, New Haven, Conn.

1970a. The Argyrolagidae, extinct South

American marsupials. Bulletin of the Museum of
Comparative Zoology, 139: 1-86.

1970b. Additions to knowledge of the Argy-

rolagidae (Mammalia, Marsupialia) from the Late Ce-
nozoic of Argentina. Breviora (Museum of Compar-
ative Zoology), 361: 1-9.

1970c. Addition to knowledge of Groeberia

(Mammalia, Marsupialia) from the Mid-Cenozoic of
Argentina. Breviora (Museum of Comparative Zool-
ogy), 362: 1-17.

1971. The evolution of marsupials in South

America. Anais Academia Brasileira de Ciencias, Su-
plemento, 43: 103-118.

1 980. Splendid Isolation. The Curious History

of South American Mammals. Yale University Press,
New Haven and London, IX + 266 pp.
Wolff, R. G. 1984. A new Early Oligocene Argyro-
lagid (Mammalia: Marsupialia) from Salla, Bolivia.
Journal of Vertebrate Paleontology, 4(1): 108-1 13.

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

Archer, M. 1982. A review of Miocene thylacinids
(Thylacinidae, Marsupialia), the phylogenetic position
of the Thylacinidae and the problem of apriorisms in
character analysis, pp. 445-476. In Archer, M., ed.,
Carnivorous Marsupials. Royal Society of New South
Wales.

BiGGERS, J. D. 1 966. Reproduction in male marsupials,
pp. 251-280. In Rowlands, I. W.. ed., Symposia of
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,
New York, 8 unnumbered pp. + 304 pp.

Cabrera, A. 1958. Catalogo de los mamiferos de
America del Sur. I. (Metatheria-Unguiculata-Camiv-
ora). Revista del Museo Argentino de Ciencias Na-
turales "Bernardino Rivadavia": 2k>ologia (1957), 4:
1-307.

Collins, L. R. 1973. Monotremes and Marsupials.
Smithsonian Publication 4888, Smithsonian Institu-
tion, Washington, D.C., 323 pp.

Davis, J. A., Jr. 1965. Agreat year for rarities. Animal
Kingdom, 68(5): 130-133.

Emmons, L. H. 1 984. Geographic variation in densities
and diversities of non-flying mammals in Amazonia.
Biotropica, 16: 210-222.

HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds.
1 982. Mammal Species of the World. Allen Press and
Association of Systematics Collections, Lawrence,
Kansas, 694 pp.

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
Marsupialia, and a classification of marsupials. Aus-
tralian Journal of Zoology, supp. sen, 52: 1-152.

Marshall, L. G. 1982. Evolution of South American
Marsupialia, pp. 251-272. In Mares, M. A., and H.
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
Natural History, Zoological Series, 14: 1-156.

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
Zoo Yearbook, 15: 4-6.

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-
ciops. Acta Anatomica, 72: 489-50 1 .

IZOR & PINE: CALUROMYSIOPS IRRUPTA

123

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-

logeny and classification, pp. 621-640. //J Archer, M., tralian species of Anlechinus (Dasyuridae: Marsupi-

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

Terborgh, J. W., J. W. FiTZPATRiCK, AND L. Emmons. South Walcs.
1984. Annotated checklist of bird and mammal spe-

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