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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 

© 1987 Field Museum of Natural History 
Library of Congress Catalog Card Number: 87-82549 

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 

V 



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 



VI 



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. 



10 



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 

16 

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) 

38 

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

Monsieur Bajon (1763?) 40 

John Gabriel Stedman (1744-1797) 

40 

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 

87 

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) 

91 

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 



13 



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 



14 



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 



IS 



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 



16 



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 



17 



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) 



18 



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 



19 



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- 



20 



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 



21 













•c 



1 



2-* 

03 so 



<N ea 

O u 
Cl, a 



22 



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 



23 



' 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). 



24 



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 



25 





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). 



26 



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 



27 




^^ ^'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 



28 



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 



29 




d. 







c 5? 
11 

c 5 



1^ 

2 i" 
•S 



30 



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 



31 



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 



32 



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 



33 



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- 



34 



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 



35 



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 



36 



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 



37 



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- 



38 



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 


12 

45 


14 

28 


16 
50 


87% 
56% 


Edeimtata 










Genera 
Species 


10 
16 


10 
12 


12 
15 


83% 
75% 


Carnivora 










Genera 
Species 


11 
17 


10 
14 


14 

25 


71% 
56% 


PiNNIPEDIA 










Genera 
Species 










2 
2 


0% 
0% 


SlRENlA* 










Genera 
Species 


1 
1 


1 
1 


1 
2 


100% 
50% 


Perissodactyla 








Genera 
Species 


1 

1 


1 
1 


1 
1 


100% 
100% 


Artiodactyla 








Genera 
Species 


4 

7 


4 
6 


5 

7 


80% 

86% 


Lagomorpha 










Genera 
Species 


1 
1 


1 
1 


1 
1 


100% 
100% 


RODENTIA 










Sciuropmorpha 

Genera 1 
Species 5 


1 
3 


3 
6 


33% 
50% 


Myomorpha (Murinae 
Genera 3 
Species 1 7 


excluded) 

5 

17 


20 
45 


25% 
24% 


Caviomorpha 

Genera 
Species 


11 

24 


15 

22 


23 
47 


65% 
47% 


Cetacea* 










Genera 
Species 


2 
2 


2 
2 


2 
2 


100% 
100% 



Taxon 







Total 








cur- 








rently 








known 


Percent- 


Number 




for 


age of 


reported 


Current 


Brazil 


current 


by 


equiva- 


(esti- 


total col- 


Pelzeln 


lent 


mated, 


lected by 


(1883) 


number 


1984) 


Natterer 



Totals 










Genera 


69 


99 


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 



39 



(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). 



40 



HELDIANA: ZOOLOGY 




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u* 




JC^ # 




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H^'^^!^ %sj ■ 






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HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 



41 



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. 



42 



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 



43 



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 



44 



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 



45 



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): 



46 



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 



47 



. . . 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, \ 



48 



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 



49 



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- 



50 



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 



51 




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HELDIANA: ZOOLOGY 





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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 



53 



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- 



54 



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 



55 





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





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-. •»« 



^ 



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). 



56 



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 



13 



14 



13 



13 



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 



57 




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



58 



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 



59 



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. 



60 



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 



61 



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. 



62 



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 



63 



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 



64 



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 



65 







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66 



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 



67 



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 



17 



68 



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 



17 



\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 



17 



Cuy del monte 



HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 



69 



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 



17 



* 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 



70 



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 



71 




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72 



FIELDIANA: ZOOLOGY 





CAI.ITKIN M».pkn" '*• 



r M.I.ITIIKIN J..— i^M.. 



I. . ■■! .'■■■!■ 








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 



73 



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 

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 



19 



ARGENTINA: Rio Parana in 
Provinces Buenos Aires and 
Corrientes 



74 



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 



75 




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76 



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 



77 



lOV 




Coplapo'U 

a 

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 



T. 



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). 



78 



HELDIANA: ZOOLOGY 




u 



HERSHKOVITZ: HISTORY OF NEOTROPICAL MAMMALOGY 



79 



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 



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 



22 



80 



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 



81 



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- 



82 



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 



22 



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 



83 



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 



84 



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 



85 






"-(SI 



86 



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 



87 



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). 



88 



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 








11 


'CXXl/.fja td> 


1 




n 


\ 


1 




1,/ 


^ 


r 


1 




m 


1i- B^^Bl 


1 






^i»' 


Via '■ 




1 




, 
















\*«.ll m 


)<': fwfl 


1 






■^/ 




i '^fjS 


1 






'' ■'u t 






1 






r 

"*■■ ■ r 






1 




-,-: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 



89 



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. 



90 



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 



91 



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 



92 



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 


31 


24 (77%) 


0(0%) 


Camivora 


303 


58(19%) 


41 (13%) 


Pinnipedia 


39 


1 1 (28%) 


2(5%) 


Sirenia 


5 


2 (40%) 


1 (20%) 


Perissodactyla 


23 


2(9%) 


0(0%) 


Artiodactyla 


186 


1 1 (6%) 


12(6%) 


Lagomorpha 


52 


4(8%) 


14(27%) 


Rodentia 


563 


152(27%) 


104(18%) 


Cetacea 


4 


2 (50%) 


0(0%) 


Proboscidea 


2 


0(0%) 


0(0%) 


Hyracoidea 


5 


0(0%) 


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 



93 



narratives of travel provided much reliable data 
on geographic distribution, habitat, life histories, 
ecological backgrounds, itineraries and maps of 
the expeditionary routes, and stopping and col- 
lecting localities. Descriptions of the collected 
mammals, most of them by the naturalist-trav- 
elers themselves, were often based on skeletal, 
dental, and soft parts in addition to purely tegu- 
mentary characters. Their classifications were pu- 
tatively natural groupings on the ordinal, family, 
and, as a rule, the generic levels. The prevailing 
belief in the biblical version of creation and fixity 
of species, not confessed in writing, did not blind 
systematists of the period to evident relationships 
between species and their clusterings into supra- 
specific groups. Descriptions of species were, 
nevertheless, typological. Subspecies or geograph- 
ic races were, at best, vaguely conceived but de- 
scribed as species. The infrequent or rare use of 
trinomials was accidental or equivocal and not 
certainly intended for a clearly defined geographic 
race. The term usually used for deviates firom 
"types" was "variety." 

Controversies regarding origin of species or fau- 
nas centered on where, not how. Philosopher- 
chroniclers of the first era accepted Noah's ark 
literally as the one place of origin and disjiersal of 
the Recent fauna. Acosta may have been the first 
to suggest the former existence of intercontinental 
connections for passage of Old World animals into 
the New World. 

More and better knowledge of the world's fauna 
during the second era revealed the weaknesses or 
fallacy of the ark dogma. Staunch creationists such 
as Linnaeus pointed instead to a vaguely located 
region as the place from which all species dispersed 
to occupy predestined habitats for which they had 
been created. Other authorities like Buffon argued 
for multiple centers of origin, with sp)ecies origi- 
nating in the habitats for which they were adapted. 
Darwin also believed in multiple places of origin, 
or faunal regions separated by geographic barriers 
but with some trepidation. The belief in multiple 
creations was heretical. 

Inconsistencies between religious dogma and 
realities did not prevent Wied-Neuwied from rec- 
ognizing the geographic range of a species (or sub- 
species) as a property of that species. Another 
advance beyond scriptures was the concept of eco- 
logical life zones contributed by von Tschudi, who 
plotted them for Peru on the basis of plants, an- 
imals, and climate. 

The total of named Neotropical sjjecies of mam- 
mals counted ft-om 1758, the year of publication 



of the 10th edition of the Linnaean Systema Na- 
tura and starting date of zoological nomenclature, 
to mid- 1 9th century, exceeded by far that of the 
Nearctic region and any other equivalent area of 
the world. Neotropical mammals were also better 
known than those of other continents except west- 
em Europe. 

By mid- 1 9th century, about 90% of currently 
known Neotropical mammalian species larger than 
common tree squirrels had already been described, 
but no more than about 10% of the smaller forms. 

The great number and variety of Neotropical 
mammals (and animals generally) known to sci- 
ence by mid- 19th century and the accumulated 
knowledge gained from study of living and pre- 
served specimens in field and laboratory, much of 
it contributed by Charles Darwin, helped pscvt the 
way to the Darwinian revolution of the next half 
century. 



XYIII. Acknowledgments 

I am indebted to Benjamin W. Williams, As- 
sociate Librarian and Librarian of Rare Books, 
Field Museum of Natural History, for p)ermission 
to consult at pleasure in the Museum's Mary W. 
Runnells Rare Book Room the books needed for 
writing this article; and to Bruce D. Patterson, 
Robert M. Timm, Ronald H. Pine, Debra Mos- 
kovitz, and J. A. Gagliano for reviewing the manu- 
script. Map>s shown in Figures 11, 12, and 2 1 were 
prepared by the author with assistance of Mary 
Anne Rogers from accounts of the travelers cited 
and other sources. Photographic reproductions of 
the figures are by Field Museum of Natural His- 
tory Staff Photographer Ron Testa. Technical 
Assistants Barbara Brown and Mary Anne Rogers 
typed the manuscript and contributed in other ways 
toward its completion. 



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nELDL\NA: ZOOLOGY 



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368. 



98 



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 









B 






'C, 


alveolus 
alveolus 


m 

1 




m 

2 




m 


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 




B 



/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 




1.00 


0.96 






1.40 


1.20 


1.08 


1.08 


1.12 


0.96 


5.40 




1.00 


1.08 




1.00 


0.92 




1.20 


1.00 




0.96 


0.96 








1.00 


1.00 


1.04 


1.00 


0.92 




1.36 


1.08 


1.00 


0.96 


0.96 


0.92 


5.12 


1.40 


1.00 
1.08 




1.00 


1.00 
0.96 




4.80 




1.28 


1.16 




1.24 


1.08 


6.76 



2.80 



3.00 4.60 



2.20 4.40 



6.20 



1.80 
1.60 

1.80 



2.40 



to elongate or modify either the pj (Polydolopidae 
and Parabderitini caenolestids; see Marshall, 1980) 
or the m, (Abderitinae and Palaeothentinae caen- 
olestids), and (2) to reduce (e.g., Caenolestidae) or 
lose (Polydolopidae) the m4. All teeth except the 
c, are completely hypselodont and rootless. The 
cheekteeth are slightly curved, with the concavity 
forward (fig. IC-D). 

Incisor ("/J— Incompletely preserved in the ho- 
lotype (fig. 2A-B). The anterior end is broken, but 
the posterior end is unaltered, showing an open 
pulp cavity with no sign of root formation. It is 
as elongate and curved as in the most specialized 
caviomorphs (e.g., Ctenomyidae), although not 
upcurved posteriorly. The extra-alveolar part 
would have been nearly vertical, although not so 
much as in the Groeberiidae (cf. fig. 4B,D). The 
tooth extends along the ventral border lingually, 
first beneath the m, and then lingually to far below 
the mj, terminating where the ventral border be- 
comes an inflected flange (MACN CH-865, fig. 1 B) 
reminiscent of kangaroos; the base of the alveolus 
shapes a superficial prominence similar to that of 
hystricognathous rodents, although enveloped by 
the inflected ventral border. It is approximately 
oval in cross section, with the long axis oriented 
dorsoventrally and with a flatter medial surface. 
Apparently enamel covers the entire tooth, but a 
noticeably heavier layer extends as a ventral band. 

Canine fie J— None of the specimens in the hy- 
podigm include this tooth, but its alveolus (figs. 
1 C-D; 2C) is oblique, tapered, and relatively shal- 
low, indicating a closed-rooted tooth; its oblique 
orientation suggests that the occlusal apex was ap- 
pressed against the occlusal tip of i,. 

First Molar (m,)—The first molar is separated 
from the c, by a crested diastema (fig. 1 A) as long 
as m,. It is the largest cheektooth, almost rect- 
angular in cross section, with the longer lateral 



faces slightly concave; the lingual face sometimes 
bears a very shallow groove along the intra-alveo- 
lar portion. The anterior face is convex, occasion- 
ally somewhat pointed; the posterior face is almost 
flat, forming angles with the lateral faces slightly 
greater than 90°. 

Second Molar (^wj— The second molar is irreg- 
ularly quadrate, with the lateral faces slightly con- 
vex and the anterior and posterior ones flatter. The 
posterolingual angle is less than 90°, whereas the 
others are almost 90°. Its width is similar to that 
of m, (fig. lA). 

Third Molar (m^J—The third molar is the small- 
est cheektooth, being subtriangular in cross section 
rather than square. The anterior face is slightly 
convex, lingually flatter, and labially more strongly 
curved; the labial face converges posterolingually 
with the lingual face, forming a rounded pillar 
rather than a well-defined posterior face (fig. lA). 

Mandible— No nearly complete mandible is 
known, but parts of the horizontal ramus and base 
of the coronoid process are known. These parts 
indicate the mandible is extremely short and deep, 
like that of Groeberia minoprioi, although very 
different in other respects (cf. fig. 4B,D). The sym- 
physis is subvertical and unfused, with a nearly 
smooth symphyseal surface (i.e., normal in struc- 
ture instead of fused and forming the odd medial 
posterior projection peculiar to Groeberiidae). The 
depth of the mandible increases abruptly toward 
the mj. The deep masseteric fossa appears to be 
F)eculiarly confined to a dorsal position, as the 
masseteric crest is situated at a level between the 
alveolar rim and the lowest level of the rounded 
and inflected ventral border (figs. 1 B; 2 A). A sim- 
ilar condition is found in some Abderitinae caen- 
olestids (e.g., Parabderites bicrispatus; Marshall, 
1976, fig. 8a), although in P. bicrispatus the man- 
dibular body is not as deep and the alveolar border 



104 



FIELDIANA: ZOOLOGY 



not as extensively inflected. The coronoid process 
has its root beneath the m,, forming a strong, sa- 
lient lamina (known only by its root), so that a 
conspicuous diagonal valley is formed between the 
coronoid and the alveolar border behind m, (fig. 
lA); a similar structure is present in Groeberia 
minoprioi (see Patterson, 1952, p. 41); the valley 
is open labially and lingually limited by a prom- 
inence similar to that present in Australasian Po- 
toroinae. 

In many respects this strong, salient, ascending 
ramus and correlated features are reminiscent of 
highly fossorial caviomorphs, such as burrow-in- 
habiting Ctenomyidae. Although the mandibular 
angle is not preserved in any of the specimens, it 
probably was inflected, as suggested by the inflec- 
tion of the ventral border, beginning at the level 
of mj, which defines a lingual flanged crest (figs. 
IB; 2B) similar to that producing the extremely 
inflected angle in the Macropodidae. This lingual 
ventral flanged crest seems to be the lingual border 
of an expanded and relatively deep pterygoid fos- 
sa, resembling that of argyrolagids (see Simpson, 
1 970a). There is a relatively large alveolar foramen 
within the pterygoid fossa, level with the alveolar 
border and within a pit (fig. 3D), and a mental 
foramen beneath the anterior face of m, at the 
level of the alveolus of i, (fig. 2 A). 



Affinities 

As in the case of Groeberia (see Simpson, 1 970c), 
the conclusion that Patagonia is a marsupial rests 
on a combination of definite, negative, and indi- 
rect evidence. The most definite evidence for its 
being a marsupial is the inflected ventral border 
of the mandible and probably the related inflected 
angle. This evidence alone is inconclusive, as a 
few marsupials lack an inflected angle and a few 
placentals have one. However, no known placental 
has such an extended and upturned flange-shaped 
inflection, and even in marsupials it is rarely so 
well developed (e.g., Groeberiidae [Patterson, 
1952]. Argyrolagidae [Simpson, 1970a,b], and the 
Australasian Macropodidae). Unlike Groeberia, 
Patagonia has other characters supporting its mar- 
supial affinities, namely the lower procumbent in- 
cisor-like canine. In the Epidolopinae (Pascual &. 
Bond, 1981) there are three procumbent lower 
teeth, the third being unquestionably the canine. 
Within the more advanced Polydolopidae (Poly- 
dolopinae), there are one or three procumbent 
lower teeth; in the latter case, evidence suggests 



they consist of two incisors and a canine, the me- 
dial incisor being quite reduced and the canine 
well developed, single, and closed-rooted. 

As in Groeberia the negative evidence is that 
Patagonia has no features precluding its reference 
to the Marsupialia. It does exhibit characters mak- 
ing reference to any Eutheria highly improbable. 
Its habitus is rodent-like, but its two differentially 
procumbent lower teeth rule out reference to the 
Rodentia. While the incisor is rodent-like in shape, 
it is oriented differently than that in rodents, ex- 
tending along the ventral border of the horizontal 
ramus, first below the m,, then lingually to other 
molars, without curving upward. It apparently 
shapes the ventral border of the mandible. In ad- 
dition, the short diastema extends at the level of 
alveoli. Among known rodents, only Paramyidae 
and Ischyromyidae developed diastemas at the al- 
veolar level, but even in these groups, the incisor 
extends as in other rodents, not as in Patagonia. 
A more-or-less rodent-like habitus was also char- 
acteristic of some notoungulates, especially among 
Typotheria and Hegetotheria, but insofar as known 
not so extreme in development as in Patagonia. 
Neither the enlargement of the incisor nor the re- 
duction of the cheekteeth is known in any prim- 
itive Paleocene notoungulates or in other South 
American "ungulates." Even later rodent-like no- 
toungulates were much less specialized than the 
Oligocene Patagonia. South American marsupials 
diverged very early into unique evolutionary lin- 
eages (see Simpson, 1970a-c, 1971, 1 980; Pascual, 
1980a,b, 1981; Paula Couto, 1979; Reig, 1981). 

Patagonia peregrina is unquestionably a mar- 
supial because its unique and diagnostic combi- 
nation of characters are unknown in any eutherian. 
Nevertheless, it could be regarded as another of 
the extinct South American mammals considered 
by some as incertae sedis and by others as a tertium 
quid with regard to the eutherian-marsupial di- 
chotomy (McKenna, 1980; Reig, 1981). However, 
the marsupial affinities of other peculiar fossil 
mammals from South America remain unques- 
tioned, despite weaker support than that offered 
here for Patagonia. For example, the basis for con- 
sidering the Polydolopidae as marsupials is the 
combination of an inflected mandibular ramus, 
palatal vacuities, and a cheektooth formula of 

1-3 1-4 

P— -- and ^-r~z- These characters were formerly 

used to exclude the polydolopids from the Allothe- 
ria. But, as these characters are present in prim- 
itive therians outside South America, their diag- 
nosis of marsupials can be considered an "act of 



PASCUAL & CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 



105 






Fig. 4. Labial (1) and occlusal (2) outlines of mandibles, showing the diflFerent development of incisor. A, Ar- 
gyrolagus parodii Rusconi; B, Groeberia minoprioi Patterson; C, Proargyrolagus bolivianus Wolff, D, Patagonia 
peregrina gen. et sp. nov. Graphic scale = 2 mm. 



faith based on . . . geography and stratigraphic po- 
sition rather than on . . . biology" (McKenna, 1 980, 
pp. 58-59). We beheve that assignment of poly- 
dolopids to marsupials represents the most par- 
simonious conclusion. 

Like the newly described Proargyrolagus boli- 
vianus (Wolff, 1984), Patagonia peregrina is 
another peculiar marsupial that appears in the fos- 



sil record without known ancestors (see Simpson, 
1970c, p. 16) only to vanish again soon afterward: 
Groeberiidae (Divisaderan Age, Late Eocene); Pat- 
agoniidae (Colhuehuapian Age, Late Oligocene); 
Necrolesiidae (Santacrucian Age, Early Miocene); 
Argyrolagidae (Huayquerian to Uquian Ages, Late 
Miocene to Early Pleistocene). We believe there 
are cedent reasons to think of Proargyrolagus bo- 



106 



nELDL\NA: ZOOLOGY 



livianus Wolff, 1984, described as a Deseadan ar- 
gyrolagid, as possibly representing a distinct fam- 
ily of Argyrolagoidea. 

This raises the question of the position of Pat- 
agonia among the varied ranks of South American 
marsupials. The previous descriptions and illus- 
trations demonstrate that Patagonia peregrina has 
many peculiarities that are rare, differently devel- 
oped, or completely absent in other marsupials (cf 
fig. 4). The most striking of these are: 

1 . Mandible extremely short and deep, with un- 
fused subvertical symphysis, dorsally posi- 
tioned masseteric fossa, ventral border in- 
flected at level of the mj, enveloping there 
the alveolus of the incisor. 

2. Presence in each ramus of mandible of one 
rodent-like rootless incisor that extends lin- 
gually along ventral border of mandible to 
below the m,. 

3. Presence in each ramus of one procumbent 
canine, single- and closed-rooted, scarcely 
separated from the incisor and with the oc- 
clusal apex probably appressed to the inci- 
sive apex. 

4. Three rectangular and continuously growing 
cheekteeth arranged in close sequence. 

These and other less striking characters under- 
score the unique specializations of Patagonia per- 
egrina, leading to its assignment to a new family, 
Patagoniidae. But the distinctive combination of 
characters in the Patagoniidae identify it as a dis- 
tinct evolutionary group, that is, a different clado- 
genetic unit. Simpson (1945, 1970a, 1980) des- 
ignated natural evolutionary groups of marsupials 
as suF>erfamilies. Following this line of reasoning, 
Patagoniidae should be allocated to a new super- 
family, the Patagonioidea. 

What are the affinities of this new superfamily 
to other superfamilies within the superorder Mar- 
supialia? Any discussion of its affinities depends 
on the systematics of other taxa, many of which 
are problematic. The systematics of fossil and ex- 
tant South American marsupials, including the 
merits of recognizing Marsupialia as a superorder, 
are discussed by Simpson (1970a, 1971) and Pas- 
cual (1980b). 

The majority of South American marsupials 
represent the order Polyprotodonta; this is roughly 
equivalent to Ride's ( 1 964) Marsupicamivora, but 
also includes Ameghino's Paucituberculata (see 
Pascual, 1980b; contra Kirsch, 1977a,b; Reig, 
1981). There is as yet no compelling argument to 



include any South American families within the 
Australasian order Diprotodonta (Reig, 1981), de- 
spite some suggestions to the contrary (e.g., Pas- 
cual & Herrera, 1 973, 1 975). While the allocations 
of these groups seems unambiguous, the positions 
of most remaining groups (e.g., Argyrolagidae, 
Necrolestidae, and Groeberiidae) remain uncer- 
tain. With some reservation, Kirsch (1977b) in- 
cluded the Necrolestidae in the polyprotodont 
Borhyaenoidea (as did Patterson, 1958), and the 
Groeberioidea and Argyrolagoidea within the 
Paucituberculata. Independently, Clemens and 
Marshall ( 1 976) also treated these animals as mar- 
supials, recognizing each as superfamilies: Argy- 
rolagoidea, Necrolestoidea, and Groeberioidea. 
Like Simpson, they made these assignments with 
disclaimers that the interrelationships of these 
groups were far from clear. 

Reig (1981, p. 60) not only questioned whether 
the Argyrolagidae (his Microtragulidae) were mar- 
supials, as none of its known characters are ty- 
pologically diagnostic, but conjectured probable 
affinities to the Anagalida. Further, without rig- 
orous analysis, he suggested that the Argyrolagidae 
could be treated as an independent order, pro- 
posing the name Argyrolagida. He concluded that 
only more intensive study or additional records 
could substantiate allocation of this order to the 
Metatheria or the Eutheria. 

Remains of Patagoniidae exhibit a unique mo- 
saic of characters, some of which are absent or 
differently developed in Groeberiidae and Argy- 
rolagidae. Despite their similarities, each of these 
taxa apF)ears prima facie to represent indep)endent 
evolutionary trends. To assess their interrelation- 
ships, common and distinctive characters of each 
must be carefully weighed. Remains of Argyro- 
lagoidea obtained in the same horizon and locality 
as the hypodigm of Patagonia peregrina should 
be particularly useful in this regard and are now 
under study. Ordinal and subordinal allocation of 
the Patagonioidea await this more comprehensive 
analysis. Known representatives of this taxon are 
so highly derived, as is the case with other peculiar 
marsupials, that their relationships to other mar- 
supial groups are obscure and can only be clarified 
by an expanded record of earlier forms. 



Ecology and Historical Biogeography 

Biological inferences of Patagonia are necessar- 
ily limited to the mandibular fragments thus far 



PASCUAL & CARLINL NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 



107 



known. These demonstrate unique characters 
among marsupials, Hving or extinct, which are ob- 
viously related to a particular mode of life. No 
known eutherian possesses such mandibular fea- 
tures. Superficially it is similar to Groeberia, both 
being rodent-like marsupials: each has a short and 
deep mandible with a single enlarged, open-rooted 
incisor, deeply extended along the mandible, with 
the extra-alveolar part apparently nearly vertical. 
These represent functional not phylogenetic sim- 
ilarities, as similar states were attained by different 
routes: in Groeberia this tooth extended within an 
odd medial posterior projection of the symphysis, 
whereas in Patagonia the intra-alveolar portion is 
truly rodent-like, in being extended along the hor- 
izontal ramus (cf fig. 4B,D). No doubt both were 
powerfial gnawers as the lower incisor worked al- 
most vertically, much more so than in most ro- 
dents. The unknown face and snout of Patagonia 
was probably short and deep; whether it had two 
pairs of lagomorph-like upper incisors like Groe- 
beria remains unknown. Related to this gnawing 
specialization, both Groeberia and Patagonia show 
a short diastema near the alveolar level and a re- 
duced number of postincisive teeth, four in both; 
however, Patagonia has three cheekteeth, whereas 
Groeberia has four. The rodent-like habitus of Pat- 
agonia is especially advanced, because the three 
cheekteeth are truly hypselodont, rectangular- 
shaped in cross section, with at most only shallow 
lateral grooves representing the remnants of an- 
cestral bilobate cheekteeth. 

This combination of features suggests food was 
obtained by gnawing and prepared for swallowing 
by grinding. It represents extraordinary conver- 
gence on some desert-adapted and fossorial forms, 
such as the Octodontidae. The evolution of cheek- 
teeth toward a rectangular shape and numerically 
reduced sequence has been recognized as occurring 
within the Octodontoidea (from the Octodontidae 
to the Ctenomyidae; Pascual et al., 1965). The 
dental features of Patagonia are also convergent 
on those of the desert-dwelling African Bathyer- 
gidae, particularly to the sand rat Heterocephalus 
glaber, and to the North American Geomyidae. 
These convergent anatomical features suggest that 
Patagoniidae were probably fossorial marsupials. 

Anatomical convergence of Patagonia on des- 
ert-dwelling fossorial rodents is curious, because 
prevailing conditions in central Patagonia during 
the Colhuehuapian Age were not highly favorable 
to desert dwellers. The first record of platyrrhine 
monkeys in Patagonia occurs at the same locality 
and level (Fleagle & Sown, 1983) as Patagonia. 



Many other vertebrate remains recovered at this 
site (see Bordas, 1939; Donadio, 1983) suggest an 
environment of well-watered tropical woodlands. 
Conversely, however, both Argyrolagoidea and 
very advanced Cephalomyidae rodents from this 
site (currently under study) show dental features 
reminiscent of desert or at least drier environ- 
ments. Generally, the Colhuehuapian vertebrate 
fauna from central Patagonia (see Pascual, 1970; 
Pascual & Odreman Rivas, 1971; Marshall et al., 
1983) is composed of both forest and open-coun- 
try types, presumably brought together in a sub- 
tropical savanna. Thus, the Patagoniidae, Ceph- 
alomyidae, and Argyrolagoidea occurred in 
apparently inappropriate environments, probably 
restricted to xeric patches in the subtropical sa- 
vanna mosaic. Because the Colhuehuapian Pata- 
gonioidea were already highly specialized for xeric 
habitats, they probably evolved earlier in the Pa- 
leogene. It therefore seems likely that ancestral 
forms existed in the Deseadan (Early Oligocene). 
Another highly specialized group of marsupials, 
the Argyrolagoidea, suggests this hypothesis. For- 
merly believed present in the record fi-om the 
Huayquerian (Late Miocene) to the Uquian (Early 
Pleistocene; see Marshall et al., 1983), argyrola- 
goids have now been reported from the Deseadan 
of Bolivia (Wolff, 1984), and here from the Col- 
huehuapian beds of central Patagonia. 

The pre-Deseadan record contains no potential 
ancestor for either Argyrolagoidea or Patagonioi- 
dea. Simpson (1970c, p. 17) proposed that "these 
groups (including Groeberioidea) evolved in what 
are now (and quite likely were then) the tropics 
and are picked up in our record only when they 
spread rather briefly to what was for them a mar- 
ginal area." It seems quite probable that the en- 
vironments responsible for their initial divergence 
were poorly or not represented in the known fossil 
record. 

Global diastrophic movements in the Late 
Eocene, and apparently related climatic and en- 
vironmental changes, are thought to be responsi- 
ble for the cosmopolitan turnover in Early Oh- 
gocene mammal communities (Kurten, 1971). This 
turnover also occurred in South America (Pascual, 
1984). Mammal communities in the Deseadan 
(Early Oligocene) are substantially different from 
Eocene communities in composition (see Pascual 
et al., 1 985), apparently reflecting Stehlin's '^^grande 
coupure."" The apparently sudden occurrence of 
the Argyrolagoidea. and probably the Patagonioi- 
dea, in the Deseadan Age is probably another ex- 
ample of this global turnover. 



108 



nELDL\NA: ZOOLOGY 



It is remarkable that, to the numerous succes- 
sive parallel trends ("successive trends" or "iter- 
ation"; Simpson, 1953, pp. 248-259; 1961, p. 127) 
in the evolutionary history of South American 
mammals, especially from the Deseadan on, can 
be added the convergence of Oligocene patagoniid 
marsupials and Pliocene to Recent ctenomyid ro- 
dents on a common morphology. These conver- 
gences are products of similar responses to re- 
peated environmental conditions. The anatomical 
and functional similarities of Patagonia peregrina 
with the extant Ctenomys are so striking that we 
are tempted to call the former the "marsupial tuco- 
tuco." 



Acknowledgments 

All of the material studied here was discovered 
by 1983 and 1984 expeditions of the Museo Ar- 
gentino de Ciencias Naturales "Bernardino Ri- 
vadavia" (MACN), in which Lie. Oscar E. Don- 
adio and Lie. Miguel Soria (both of MACN) and 
the American paleontologists John G. Fleagle and 
Thomas M. Bown participated. Dr. Jose F. Bo- 
naparte, Chief of the Seccion Paleontologia Ver- 
tebrados, MACN, and responsible for these ex- 
peditions, generously put this and other marsupial 
material at our disposal. The x-ray plates were 
made by Dr. Roberto Guevara, Profesor de Odon- 
tologia, Universidad Nacional de La Plata, by the 
authority of the Dean, Dr. Oscar Barletta. We thank 
all of them very much. 



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tiembre 1981), Actas 3: 593-606. 

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de Noviembre de 1968), 2 (Paleontologia Sudameri- 
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. 1980a. Nuevos singulares tipos ecologicos de 

marsupiales extinguidos de America del Sur (Paieo- 
ceno tardio o Eocene temprano) del Noroeste Argen- 
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1 980b. Prepidolopidae, nueva familia de Mar- 



supialia Didelphoidea del Eoceno sudamericano. 
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. 1981. Adiciones al conocimiento de Bona- 



partherium hinakusijum (Marsupialia, Bonapartheri- 
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. 1983. Novedosos marsupiales paleogenos de 



la Formacion Pozuelos (Grupo Pastes Grandes) de la 
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. 1984. La sucesion de las Edades-mamifero, de 



los climas y del diastrofismo sudamericanos durante 
el Cenozoico: fenomenos concurrentes. Anales de la 
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PASCUAL 8c CARLINI: NEW SUPERFAMILY OF PALEOGENE MARSUPIALS 



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Pascual, R., and M. Bond. 1981. Epidolopinae 
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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 

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



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110 



FIELDIANA: ZOOLOGY 



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

Milton H. Gallardo and Bruce D. Patterson 



ABSTRACTS 

The karyotype of Rhyncholestes Osgood is described for the first time. The karyotype has 
2n = 14 and is similar in most respects to karyotypes of similar number found in other American 
and Australasian genera in several families. The karyotype of somatic (bone marrow) tissues 
from male Dromiciops Thomas is presented for the first time; surprisingly, it differs from the 
2n = 14 complement previously reported from female bone marrow and male gonads. The 
2n = 1 3 karyotype found in bone marrow of male Dromiciops lacks a minute element thought 
to be the Y chromosome. This instance of somatic chromosome elimination represents the first 
case reported for American marsupials and presents an interesting parallel to sex-chromosome 
mosaicism among Australasian Peramelidae and Petauridae. 



El cariotipo de Rhyncholestes Osgood es descrito por primera vez. El cariotipo consta de 
2n = 14 y es muy similar a cariotipos de igual mimero encontrados en otros generos americanos 
y australoasiaticos de varias familias. El cariotipo de tejidos somaticos (medula osea) de un 
Dromiciops Thomas macho es presentado por primera vez; sorprendentemente, difiere del 
complemento 2n = 14 reportado previamente de medula osea femenina y gonadas masculinas. 
El cariotipo 2n = 1 3 encontrado en medula osea del Dromiciops macho carece de un diminuto 
elemento que supuestamente corresponde al cromosoma Y. Este ejemplo de eliminacion so- 
matica de cromosomas representa el primer caso reportado en marsupiales americanos y pre- 
senta un interesante paralelo con el extenso mosaicismo de los cromosomas sexuales descrito 
entre las formas australoasiaticas. 



Descreve-se pela primeira vez, o cariotipo de Rhyncholestes Osgood. O cariotipo e de 2n = 
1 4, e, na maioria de seus aspectos, assemelha-se aos cariotipos de numeros similares encontrados 
em outros generos americanos e austral^sios. O cariotipo de tecidos somdticos (da medula ossea) 
de Dromiciops Thomas machos e descrito pela primeira vez. Supreendentemente, este cari- 
otipo difere do complemento de 2n = 1 4, previamente descrito para a medula ossea das femeas 
e para as gonadas dos machos. No cariotipo de 2n = 13, encontrado na medula ossea de 
Dromiciops machos, falta um elemento miudo, possivelmente o cromossomo Y. Este e o 
primeiro exemplo documentado da elimina9ao somatica de um cromossomo em marsupiais 
americanos, e apresenta um paralelo interessante ao mosaico frequentemente encontrado nos 
cromossomos sexuais de outras formas austral^sias. 



From the Institute de Ecologia y Evolucion, Univer- 
sidad Austral de Chile, Casilla 567, Valdivia, Chile; and 
Division of Mammals, Field Museum of Natural His- 
tory, Chicago, IL 60605-2496. 

GALLARDO & PATTERSON: KARYOTYPES OF MARSUPIALS 1 1 1 



Introduction 

Several unusual cytological features, including 
low diploid number (Hayman & Martin, 1969; 
Reig et al., 1977), paternally derived X inactiva- 
tion (Lyon, 1974a,b), multiple sex-chromosome 
systems (Hayman & Martin, 1969; Schneider, 
1977), somatic elimination of sex-chromosomes 
(Schneider, 1977; Close, 1984), and sperm con- 
jugation (Biggers & Creed, 1962; Biggers & De- 
Lamater, 1965), have made marsupials interesting 
subjects of cytological research. These studies have 
clarified fundamental cytological mechanisms. 
Additionally, results of the research have shed Ught 
upon directions of chromosomal evolution and 
upon interrelationships of lineages (Hayman & 
Martin, 1969; Reig et al., 1977; Sharman, 1982). 

A 14-chromosome karyotype occurs in several 
distinct lineages in all living American families: 
Didelphidae (Reig et al., 1 977), Microbiotheriidae 
(Spotomo & Fernandez, 1971; Reig et al., 1972), 
and Caenolestidae (Hayman et al., 1971). This 
karyotype also occurs in several Australian mar- 
supial lineages (Hayman & Martin, 1969) and is 
therefore considered the primitive chromosome 
number for Metatheria (Reig et al., 1977). Direc- 
tion of chromosome evolution in Metatheria has 
proceeded via centromeric dissociations— with 
pericentric inversions superimposed on the basic 
Robertsonian mechanism— to give rise to the re- 
maining 2n = 1 8 and 2n = 22 karyotypes known 
for American forms (Hayman & Martin, 1969; 
Reig et al., 1977). Extremes of karyotypic varia- 
tion in Australasian marsupials are 2n = 10 to 32 
(Schneider, 1977). 

Two autochthonous and endemic South Amer- 
ican genera, Rhyncholestes and Dromiciops, are 
especially interesting from an evolutionary view- 
point. Both are represented by a single species and 
occur only in the temperate Valdivian rainforests 
of southern Chile and Argentina. Rhyncholestes. 
one of three extant genera of Caenolestidae, is 
widely isolated from its relatives in the northern 
Andes and presents some striking morphological 
specializations. Dromiciops. thought by some to 
have special affinities with Australasian lineages 
(Sharman, 1982; Szalay, 1982), is the only hving 
genus of the otherwise extinct Microbiotheriidae 
(Marshall, 1982). Its affinities with other marsu- 
pial genera are currently uncertain. In this note we 
present the first somatic karyotypes of male Rhyn- 
cholestes raphanurus and Dromiciops australis. 
Additionally, we document the first instance of 



somatic sex-chromosome mosaicism in South 
American marsupials. 



Materials and Methods 

Seven specimens of D. australis (five males and 
two females) from Valdivia (39°32'S, 72°52'W), 
Osorno (4r06'S, 72°30'W), and Concepcion 
(37»26'S, 73°19'W) provinces, Chile, were ana- 
lyzed by the in vivo colchicine-hypotonic citrate 
technique using bone marrow as a source of mi- 
toses (Patton, 1967). Modifications of the same 
procedure were used for the one R. raphanurus 
collected at La Picada, Volcan Osorno (41*^6'S, 
72''30'W); incubation with colchicine lasted 2.5 
hours and a slightly more hypotonic solution of 
sodium citrate was used. A total of 4 1 9 mitotic 
plates was examined: 29 1 from male and 1 20 from 
female D. australis and 10 from R. raphanurus. 
Museimi specimens were deposited in the Collec- 
tion of Mammals, Instituto de Ecologia y Evolu- 
cion, Universidad Austral de Chile, and Field Mu- 
seum of Natural History. 



Results and Discussion 

Rhyncholestes raphanurus presents a 2n = 14 
complement, consisting of three pairs of large 
metacentric, one pair of medium-sized metacen- 
tric, and two pairs of small metacentric autosomes. 
The sex-chromosomes are an acrocentric X and a 
minute Y (fig. 1). This karyotype differs morpho- 
logically from the didelphid 2n = 1 4 in not show- 
ing a clear break between chromosome groups A 
and B. It also differs in arm ratios (table 1) from 
the other living caenolestids, Lestoros and Caeno- 
lestes (see Hayman et al., 1971). Moreover, the 
interstitial region of the short arm of pair two shows 
an achromatic area, resembling a secondary con- 
striction, not described in other caenolestids (but 
see discussion in Sharman, 1982). Nevertheless, a 
2n = 14 karyotype characterizes all three genera 
of Caenolestidae, which supports previous claims 
that this karyotype is primitive for Metatheria 
(Hayman & Martin, 1969; Hayman et al., 1971; 
Reig et al., 1 977) and reinforces the pattern of low 
karyotypic variation within marsupial families. 

Secondary constrictions can serve as chromo- 
some markers and are thus useful, in the absence 



112 



FIELDIANA: ZOOLOGY 



of banding data, for phylogenetic reconstruction. 
However, the secondary constriction evident in 
the karyotype of Rhyncholestes is unreported in 
other South American marsupials, although sec- 
ondary constrictions are widespread among Aus- 
tralasian marsupials (Hayman &. Martin, 1969). 
Considering commonality and in-group and out- 
group comparisons, we regard the secondary con- 
striction of Rhyncholestes as apomorphic. Thus, 
the similar structures of Australasian marsupials 
were apparently independently derived and can- 
not be traced back to some marsupicamivorous 
or other common ancestor. 

Chromosome counts from all four male D. aus- 
tralis consistently indicated 2n = 13 chromo- 
somes. The diploid number for females was 2n = 
14 as was previously reported (Spotomo & Fer- 
nandez, 1971; Reig et al., 1972). No differences 
among our specimens from geographically isolated 
localities were detected, nor were secondary con- 
strictions evident. 

Electron microscope studies of sex-chromo- 
somes in spermatocytes of D. australis and the 
didelphid Marmosa elegans demonstrate striking 
similarities (Fernandez et al., 1979). These simi- 
larities suggest that a 2n = 14 karyotype should 
be present in D. australis, its Y chromosome should 
resemble that of M elegans, and both genera should 
exhibit an XX/XY sex-chromosome system. 

We have consistently found 2n = 13 chromo- 
somes in somatic tissues of male Dromiciops and 
2n = 14 in female somatic tissue. The missing 
chromosome in males is dotlike and probably the 
Y chromosome (fig. 1). Translocation of the Y to 
an autosome is an unlikely mechanism for the dif- 
ferences between sexes because males have 2n = 
14 in germinal cells and because the sex vesicle 
appears normal (Fernandez et al., 1979). While it 
is possible that such a small chromosome might 
be overlooked in one or a few chromosomal 
spreads, its universal absence in all counted plates 
makes this alternative highly unlikely. Available 
data favor a somatic elimination of the Y chro- 
mosome. 

Previous studies have shown that both consti- 
tutive and facultative heterochromatin can be de- 
leted from marsupial cells in vivo without appar- 
ent deleterious effects on cell replication and 
survival (Hayman & Martin, 1969). Most exam- 
ples of somatic elimination of sex-chromosomes 
in marsupials involve the X chromosome in dos- 
age compensation (e.g., perameUds and petaurids; 
Close, 1984). Mitotic figures from the testes of 



B 



(( a (» 



A-1 

a 

B-1 

II 

C-1 



B-1 



A-2 A-3 



II •« 

C-2 XY 



D& Si! \l 

A-1 A-2 A-3 

U 



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. 
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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 



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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, 
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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- 
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. 1974b. Mechanisms and evolutionary origins 



of variable X-chromosome activity in mammals. Pro- 
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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. 

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Reig, O. a., R. Fernandez, and O. A. Spotorno. 1 972. 
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mosomes in two species of Chilean didelphoid mar- 
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Reig, O. A., A. L. Gardner, N. O. Bianchi, and J. L. 
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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 


7 

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) 



6 


CVG M-30 BE 95 


617 


333.5 




29 


63.7 


61.0 


57.3 


30.0 


S 


USNM 397626 


630+ 


330 + 


52 


34 


64.5 


63.4 


59.1 


32.3 


6 


USNM 396 1 60 


590 


340 


51 


32 


63.6 


62.5 


57.3 


30.5 


S 


FMNH 60698 










63.4 


60.6 


56.5 


30.5 


9 


AMNH 208101 


570 


310 


47 


37 


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 


orbital 


orbital 


Zygo- 


brain- 


Length 


case 


Length 


Length 


lary 




dibular 


con- 


pro- 


con- 


matic 


case 


longer 


(incl. 


of 


mand. 


tooth- 


M'- 


tooth- 


striction 


cesses 


striction 


breadth 


width 


nasal 


bullae) 


mandible 


ramus 


row 


M' 


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 
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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, 
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Cabrera, A. 1958. Catalogo de los mamiferos de 
America del Sur. I. (Metatheria-Unguiculata-Camiv- 
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turales "Bernardino Rivadavia": 2k>ologia (1957), 4: 
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Collins, L. R. 1973. Monotremes and Marsupials. 
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Davis, J. A., Jr. 1965. Agreat year for rarities. Animal 
Kingdom, 68(5): 130-133. 

Emmons, L. H. 1 984. Geographic variation in densities 
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HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds. 
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Janson, C, J. Terborgh, and L. H. Emmons. 1981. 
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azonian forest. Biotropica, 12(Suppl.): 1-6. 

KiRSCH, J. A. W. 1977. The comparative serology of 
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de microbiotherinos vivientes en la fauna sudameri- 
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ings 48th Annual Conference American Association 
of Zoological Parks and Aquariums held in Portland, 
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. 1975. Observations on the water opossum or 

yapok Chironectes minimus in captivity. International 
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Sanborn, C. C 1951. Two new mammals from south- 
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SiMONETTA, A. M. 1979. First record of Ca/Mrow>'5;op5 cies of Cocha Cashu Biological Station, Manu Na- 

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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 


%F 


%V 


%F 


%V 


%F 


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 


%F 


% V 


%F 


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 


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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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 
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Streilein, K. E. 1982. Ecology of small mammals in 
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107. 

Taube, C. M. 1947. Food habits of Michigan opos- 
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Tyndale-Biscoe, C. H., and R. B. Mackenzie. 1976. 
Reproduction in Didelphis marsupialis and D. albi- 
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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 


Age 


eruption 


Wear 


class 


(months) 


dP' M' 







I 


3.0-3.5 


dP' M^ 







II 


4.5-5.0 


dP' M' 







III 


6.2-6.7 


P' M' 







IV 


7.9-8.7 


P' M* 







V 


10.9-11.7 


P' M- 


P^ 


M'-2 


VI 


12.8-14.1 


P' M* 


P' 


M^ 


VII 


> 16.1 



Source: G. A. Cordero (unpublished data). 



CORDERO & NICOLAS: FEEDING HABITS OF OPOSSUMS 



131 



Notes on Distribution of Some Bats 
from Southwestern Colombia 



Michael S. Alberico 



ABSTRACTS 

Noteworthy range extensions are presented for Noctilio albiventris, Rhinophylla alethina, 
Sturnira aratathomasi, and Lonchophylla handleyi, including the second Colombian report for 
the last. A previous report of Molossops brachymeles is clarified as representing M. abrasus. 



Se presentan algunas notables extensiones del rango de distribucion para las especies Noctilio 
albiventris, Rhinophylla alethina, Sturnira aratathomasi y Lonchophylla handleyi, este ultimo 
siendo el segundo reporte para Colombia. Un reporte anterior de Molossops brachymeles se 
clarifica como representativo de M. abrasus. 



Apresentam-se notaveis exten^oes mas distribui96es das especies Noctilio albiventris, Rhin- 
ophylla alethina, Sturnira aratathomasi, e Lonchophylla handleyi, esta ultima sendo apenas o 
segundo registro para a Colombia. Clarifica-se o registro anterior de Molossops brachymeles 
como representativo de M. abrasus. 



Introduction 

Despite considerable interest in Neotropical 
mammals, southwestern Colombia remains poor- 
ly understood in this respect. This is mainly a 
result of a lack of adequate collections caused by 
the inaccessible nature of much of the zone. Early 
collecting expeditions to which we owe much of 
our knowledge were undertaken around the turn 
of the century by personnel of the American Mu- 
seum of Natural History and summarized by Allen 
(1916). Bats were typically underrepresented in 
these early collections because of inadequate col- 
lecting techniques in use at the time. Now, with 
the aid of Japanese mist nets, we are able to obtain 
more complete samples of bat communities. In 
this report I present results of a continuing col- 
lecting effort during the past five years in this poor- 
ly known region, extending the known distribution 



From the Departamento de Biologia, Universidad del 
Valle, Call, Colombia. 



of Noctilio albiventris, Lonchophylla handleyi, 
Rhinophylla alethina, Sturnira aratathomasi, and 
Molossops abrasus. 

All specimens mentioned were collected in mist 
nets, prepared as standard study skins with skulls, 
and deposited in the mammal collection of the 
Departamento de Biologia, Universidad del Valle, 
Cali, Colombia (UV). 



Distribution 

Noctilio albiventris 

The lesser bulldog bat was recently reviewed by 
Davis (1976) and by Hood and Pitocchelli (1983). 
Both mapped the distribution as including eastern 
Colombia across the Llanos and Amazonas and 
the northern Caribbean coast. Davis (1976) re- 
ported the altitudinal range of the species as ex- 
tending up to 1 , 1 00 m. We have found this species 
to be common in the upper Cauca valley, between 



ALBERICO: DISTRIBUTION OF COLOMBIAN BATS 



133 



the Cordillera Central and the Cordillera Occi- 
dental of the Andes, where the elevation reaches 
this approximate limit. Fifteen specimens from 
the Departamento (= state) del Valle del Cauca 
and adjacent Departamento del Cauca were com- 
pared with the descriptions and measurements of 
all subspecies recognized by Davis (1976). This 
population is indistinguishable from N. a. minor 
in all characters examined and undoubtedly fol- 
lows the Rio Cauca south from the Caribbean low- 
lands. A similar southern extension is most prob- 
able in the valley of the Rio Magdalena to the 
Departamento de Huila, but has yet to be con- 
firmed by collections. 

Specimens Examined— Cauca: Rio Palo, 18 km 
S, 5 km E Puerto Tejada, 3°04'N, 76°22'W, 1,050 
m (3 92, UV3 1 3, 324, 325); Valle del Cauca: 2 km 
S, 4 km W Candelaria, 3°23'N, 76°23'W, 1,000 m 
(1 (5, UV676); Universidad del Valle (Melendez 
Campus), 8 km S Cali, 3°22'N, 76°32'W, 1,000 m 
(5 S6, UV2602, 2603, 2604, 2608, 2609; 2 99, 
UV2605, 2607); 13 km S, 1 km E Cali, 3°22'N, 
76°32'W, 1,000 m (2 66, UV2620, 2611; 1 9, 
UV2612). 



Lonchophylla handleyi 

This species was described on the basis of spec- 
imens from Peru and southern Ecuador by Hill 
(1980), who suggested that some individuals in 
existing collections might be misidentified as L. 
robusta. Lonchophylla handleyi was first reported 
for Colombia by Alberico and Orejuela (1982), 
who collected a single individual from near the 
Ecuadorian border at 850 m. A specimen recently 
collected from the Departamento del Valle del 
Cauca at 480 m provides the second record for 
Colombia. Both specimens are larger (greatest 
length of skull, 28.4 and 28.6 mm, respectively) 
than the largest L. robusta reported by Hill (1980) 
for Peru and Ecuador and are larger than any L. 
robusta in our collections from western Colombia. 
Both Colombian specimens of L. handleyi are from 
the lower slope Andean forests, probably one of 
the last habitats to be intensively sampled for 
mammals in this country. The presence of this 
species in a relatively narrow elevational band 
within this habitat type attests to the importance 
of continued collecting in the Pacific slope of the 
Andes in southwestern Colombia. 

Specimens Examined— Nariilo: 5 km E Junin, 
l''20'N, 78°08'W, 850 m (1 6, UV3007); Valle del 



Cauca: Rio Cajambre, approx. 60 km S Buena- 
ventura, 3°20'N, 77°00'W, 480 m (1 9, UV3694). 



Rhinophylla alethina 

This species was described based on specimens 
from western Colombia in the Departamento del 
Valle del Cauca (Handley, 1 966) and until recently 
was known only from the type locality. Albenco 
and Orejuela (1982) reported it from Narifio near 
the Ecuadorian border and suggested that it might 
have a broader geographic range than previously 
thought, which was confirmed by Baud (1982) who 
reported the species for Ecuador. Our collections 
show R. alethina to be relatively common in the 
Pacific lowlands and the adjacent lower slopes of 
the western Andes up to 850 m. That this species 
was only recently described and remains poorly 
known is undoubtedly due to insufficient collect- 
ing in the forests of this zone. 

Specimens Examined— Nariflo: 5 km E Junin, 
1°20'N, 78°08'W, 850 m (3 66, UV3029, 3033, 
3036; 5 99, UV3030, 3031, 3032, 3034, 3035). 
Valle del Cauca: Alto Anchicaya, 35 km S, 20 km 
E Buenaventura, 3°34'N, 76°54'W, 400 m (2 66, 
UV3166, 3167); Rio Azul, 5 km N, 25 km W 
Darien, 3°59'N, 76°44'W, 560 m (1 9, UV3391); 
Rio Cajambre, approx. 60 km S Buenaventura, 
3°20'N, 77°00'W, 480-520 m (l 6, UV3702; 1 9, 
UV3703); Rio Cahma, 13 km N, 14 km E Bue- 
naventura, 4°00'N, 76°59'W, 40 m (1 9, UV2809). 



Stumira aratathomasi 

In their description of this species, Peterson and 
Tamsitt (1968) reported three specimens, the ho- 
lotype from the Departamento del Valle del Cauca 
in western Colombia and two from an unknown 
locality in Ecuador. They stated that it might be 
restricted to the Pacific side of the Andes. Thomas 
and McMurray ( 1 974) provided measurements for 
the holotype and six individuals collected near the 
type locality and suggested that this species may 
be common at high elevations in the western An- 
des of Colombia. Our recent collections extend the 
known range some 150 km to the north in the 
Cordillera Occidental and, more importantly, re- 
cord the presence of S. aratathomasi in the Cor- 
dillera Central, where it was previously unknown. 
This species appears to inhabit medium to high 
elevation forests which are relatively continuous 



134 



FIELDIANA: ZOOLOGY 



in Colombia, and its occurrence both farther to 
the north and in the Cordillera Oriental is likely. 
Specimens Examined— Valle del Cauca: Cor- 
dillera Central: Hacienda "Los Alpes," 6 km S, 1 1 
km E Florida, 3°16'N, 76°09'W, 2,400 m (1 9, 
UV3482); Cordillera Occidental: Betania, 10 km 
N, 15 km W Bolivar, 4°26'N, 76''19'W, 1,800 m 
(1 9, UV3876); Parque Nacional "Los Farallones 
de Cali," 1 km S, 1 6 km W Cali, 3°22'N, 76°4 1 'W, 
2,600 m (1 9, UV3373); Paso de Galapagos, 8 km 
N, 4 km E El Cairo, 4°50'N, 76°12'W, 1,800 m 
{2 66, UV4131,4133). 



Molossops abrasus 

This species was reported for Colombia by Al- 
berico and Naranjo-H. (1982) as M. brachymeles, 
based on specimens from the Cauca valley in 
northern Valle del Cauca. Although often referred 
to by this latter specific epithet (see Cabrera, 1958; 
Freeman, 1981), the holotype of Dysopes abrasus 
from Brazil has been shown to represent this species 
(Husson, 1962; Carter & Dolan, 1978). The Co- 
lombian record extends the known distribution of 
M. abrasus in western South America from An- 
dean Peru some 1 ,600 km to the north. 

Specimens Examined— Valle del Cauca: 1 1 km 
S, 2 km W Cartago, 4''39'N, 75°56'W, 930 m (2 
66, UV2451, 2452; 1 9, 2453). 



Acknowledgments 

This report is the result of the combined efforts 
of many friends and students, too numerous to 
mention by name, who have collaborated either 
by accompanying the author in the field, by sharing 
specimens collected during other activities, or both. 
However, a few individuals have contributed more 
than could be expected in the normal turn of events, 
and their support in the field and out has been 
especially important in the present study: Eduardo 
Velasco, Gloria Giral, Alonso Gonzalez, Guiller- 
mo Cantillo, and Luz Marina Alberico. To these, 
the author is most appreciative. 



Literature Cited 

Alberico, M., and L. G. Naranjo-H. 1982. Primer 
registro de Molossops brachymeles (Chiroptera: Mo- 
lossidae) para Colombia. Cespedesia, II: 141-143. 

Alberico, M., AND J. E. Orejuela. 1982. Diversidad 
especifica de dos comunidades de murcielagos en Na- 
rino, Colombia. Cespedesia, Suplemento no. 3(4 1-42): 
31-40. 

Allen, J. A. 1916. List of mammals collected in Co- 
lombia by the American Museum of Natural History 
expeditions, 1910-1915. Bulletin of the American 
Museum of Natural History, 35: 191-238. 

Baud, F. J. 1982. Presence de Rhinophylla alethina 
(Mammalia, Chiroptera) en Equateur et repartition 
actuelle du genre en Amerique du Sud. Revue Suisse 
de Zoologie, 89: 8 1 5-82 1 . 

Cabrera, A. 1958. Catalogo de los mamiferos de 
America del Sur. Revista del Museo Argentino de 
Ciencias Naturales "Bernardino Rivadavia", Ciencias 
Zoologicas, 4: 1-307. 

Carter, D. C, and P. G. Dolan. 1978. Catalogue of 
type specimens of neotropical bats in selected Euro- 
pean museums. Special Publications, The Museum, 
Texas Tech University, 15: 1-136. 

Davis, W. B. 1976. Geographic variation in the lesser 
noctilio, Noctilio albiventris (Chiroptera). Journal of 
Mammalogy, 57: 681-107. 

Freeman, P. W. 1981. A multivariate study of the 
family Molossidae (Mammalia, Chiroptera): Mor- 
phology, ecology, evolution. Fieldiana: Zoology, n.s., 
7: 1-173. 

Handley, C. O., Jr. 1966. Descriptions of new bats 
{Choeroniscus and Rhinophylla) from Colombia. Pro- 
ceedings of the Biological Society of Washington, 79: 
83-88. 

Hill, J. E. 1 980. A note on Lonchophylla (Chiroptera: 
Phyllostomatidae) from Ecuador and Peru, with the 
description of a new species. Bulletin of the British 
Museum (Natural History), Zoology Series, 38: 233- 
236. 

Hood, C. S., and J. Pitocchelll 1983. Noctilio al- 
biventris. Mammalian Species, 197: 1-5. 

HussoN, A. M. 1962. The bats of Suriname. Rijks- 
museum van Natuurlijke Historic, Leiden, 58: 1-282. 

Peterson, R. L., and J. R. Tamsitt. 1968. A new 
species of bat of the genus Sturnira (family Phyllosto- 
matidae) from northwestern South America. Life Sci- 
ences Occasional Papers, Royal Ontario Museum, 12: 
1-8. 

Thomas, M. E., and D. N. McMurray. 1974. Ob- 
servations on Sturnira aratathomasi from Colombia. 
Journal of Mammalogy, 55: 834-836. 



ALBERICO: DISTRIBUTION OF COLOMBIAN BATS 



135 



Distributional Records of Bats 

from the Caribbean Lowlands of Belize 

and Adjacent Guatemala and Mexico 

Timothy J. McCarthy 



ABSTRACTS 

Thirty new species records are presented for the bat fauna of Belize, along with secondary 
records for eight bats that had been recorded previously from that country. Contiguous lowland 
localities in Guatemala provided new department records: nine for El Peten, five for Izabal, 
and two for Alta Verapaz. The El Peten records include the first confirmation of Vampyrum 
spectrum in Guatemala. One state record for Quintana Roo, Mexico, is reported. These species 
represent the genera Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, 
Micronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, 
Trachops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vam- 
pyressa, Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bau- 
erus, Eumops, and Molossus. Range extensions are acknowledged for Saccopteryx leptura, 
Diclidurus virgo, Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyl- 
lostomus discolor, Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vam- 
pyrodes caraccioli, Artibeus toltecus, and Bauerus dubiaquercus. A checklist of the bat fauna of 
Belize, which stands at 66 species, is presented. 



Se registran 30 especies que no habian sido citadas antes para la fauna de murcielago de 
Belice, con registros secundarios para ocho murcielagos ya conocidos de ese pais. En ciertas 
localidades contiguas de las tierras bajas de Guatemala, se obtuvieron nuevos registros depar- 
tamentales: nueve de El Peten, cinco de Izabal, y dos de Alta Verapaz. Los registros de El Peten 
incluyen la primera confirmacion de Vampyrum spectrum en Guatemala. Ademas, se presenta 
un nuevo registro estatal para Quintana Roo, Mexico. Las especies obtenidas estan segregadas 
en los generos Saccopteryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Mi- 
cronycteris, Lonchorhina, Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trach- 
ops, Chrotopterus, Vampyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa, 
Chiroderma, Artibeus, Centurio, Diphylla, Natalus, Myotis, Eptesicus, Lasiurus, Bauerus, Eu- 
mops, y Molossus. Para cada una de las siguientes especies de murcielagos se anota el alcance 
geografico de su distribucion conocida: Saccopteryx leptura, Diclidurus virgo, Noctilio leporinus, 
Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor, Vampyrum spec- 
trum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli, Artibeus tolte- 
cus, y Bauerus dubiaquercus. Se incluye una lista de 66 especies que representan la fauna de 
murcielagos de Belice. 

Apresenta-se records de 30 novas especies de morcegos para Belice, e de oito especies pouco 
conhecidas no pais. Areas adjacentes, na Guatemala, providenciaram novos records para: El 



From the Department of Mammalogy, American Mu- 
seum of Natural History, Central Park West at 79th 
Street, New York, NY 10024. 



MCCARTHY: DISTRIBUTION OF BATS 137 



Peten (nove especies), Izabal (cinco especies), e Alta Verapaz (dois especies). Os records de El 
Peten incluem as primeiras confirma^oes de Vampyrum spectrum na Guatemala. Um novo 
record para Quintana Roo, Mexico, e incluido. Estas especies representam os generos Saccop- 
teryx, Balantiopteryx, Diclidurus, Noctilio, Pteronotus, Mormoops, Micronycteris, Lonchorhina, 
Macrophyllum, Tonatia, Mimon, Phyllostomus, Phylloderma, Trachops, Chrotopterus. Vam- 
pyrum, Glossophaga, Uroderma, Vampyrops, Vampyrodes, Vampyressa, Chiroderma, Artibeus, 
Centurio. Diphylla, Natalus, Myotis, Eptesici4s, Lasiurus. Bauerus, Eumops, e Molossus. Re- 
conhece-se extensoes nas areas onde sao encontrados Saccopteryx leptura, Diclidurus virgo, 
Noctilio leporinus, Micronycteris nicefori, Macrophyllum macrophyllum, Phyllostomus discolor, 
Vampyrum spectrum, Glossophaga commissarisi, Uroderma bilobatum, Vampyrodes caraccioli, 
Artibeus toltecus. e Bauerus dubiaquercus. Apresenta-se uma lista da fauna de morcegos em 
Belice, que agora conta com 66 especies. 



Introduction 

Inventories of bat communities in Mexico and 
Central America have increased significantly dur- 
ing the last twenty-five years (Jones et al., 1977). 
Although the resulting data have enhanced our 
knowledge of the distributions and the zoogeo- 
graphical relationships of species, incomplete sur- 
veys exist for certain regions. The northern low- 
lands along the Caribbean coast of Honduras, 
Guatemala, Belize, and Quintana Roo, in Mexico, 
is one such region. Travel within this coastal ver- 
sant has improved with agricultural and settle- 
ment expansion. The isolation of Belize from its 
neighbors has been reduced with the construction 
of roads in Guatemala's frontier of El Peten and 
Mexico's former territory of Quintana Roo. A 
paved road from Izabal now connects El Peten 
and Belize with the Pan-American Highway in 
western Guatemala. Road development continues 
within Belize for all-weather travel. 

Belize is situated within the Caribbean lowland 
drainage of northern Central America. Contiguous 
with Belize on this eastern slope is the eastern 
portion of the department of El Peten to the west 
and, to the south, the department of Izabal, both 
of Guatemala. Southern Quintana Roo of penin- 
sular Mexico borders to the north (see fig. 1 and 
Gazetteer). The topography of these Caribbean 
lowlands extends from the lower ranges (600 m 
and below) of the eastern Sierra de Chama, the 
Sierras de las Minas, the Sierra de Santa Cruz, the 
Sierra del Meredon, and the Montarias del Mico 
in Alta Verapaz and Izabal, and the Maya Moun- 
tains of southern Belize and southeastern El Peten 
to the low undulating relief of southern Quintana 
Roo. The Maya Mountains represent a heavily 
eroded Paleozoic formation that now ranges at the 
top from 671 to 853 m in elevation, with the high- 



est peak at 1113m (Wright et al., 1959). Annual 
rainfall in portions of Izabal averages from 3,000 
to nearly 5,000 mm (Portig, 1976). Over 4,500 
mm of rainfall was reported from the most south- 
em coastal area of Belize. North and northwest- 
ward of the Maya Mountains, rainfall decreases 
appreciably to less than 1 ,500 mm in north-central 
El Peten and northern Belize, where less than 1 ,400 
mm was recorded near the Quintana Roo border 
(Walker, 1973). The severity of this northward 
reduction of rainfall is intensified by the increased 
lack of surface drainage into the Yucatan Penin- 
sula of Mexico. Because the limestone shelf of 
northern Belize has geological affinities with the 
Yucatan Peninsula (Wright et al., 1 959), the south- 
em limit of this peninsula can be considered the 
fault line extending from north of the Maya Moun- 
tains westward through the northem shore of Lake 
Peten-Itza, El Peten (Wadell, 1938; West, 1964). 
Effectively, the northem plain of Belize and north- 
em El Peten are portions of the Yucatan Peninsula. 
The northward shift from alluvial soils to shallow 
calcareous soils, along with the mentioned cli- 
matic changes, create edaphic conditions that af- 
fect the composition and the structure of the vege- 
tation that can be supported (Lundell, 1 934, 1 937; 
Standley & Record, 1936; Wright el al., 1959; Pen- 
nington & Samkhan, 1968). The potential effect 
of this transitional physiography on the distribu- 
tion and relative abundance of bats in this Carib- 
bean lowland region will require further inventory 
studies. 

This paper documents 30 new records for Belize. 
A checklist of the known bat fauna for this country 
is annotated in the Appendix. Sixty-six species are 
cited. Included here are also records from nearby 
localities for El Peten, Izabal, and Alta Verapaz, 
Guatemala, and Quintana Roo, Mexico. Nine 
species records from EI Peten, five records from 



138 



HELDIANA: ZOOLOGY 



Izabal, and two records from Alta Verapaz in- 
crease the number of reported species for these 
departments to 35, 31, and 40, respectively (see 
Jones, 1966; Carter etal., 1966; Rick, 1968; Smith, 
1972; LaVal, 1973a; Martinez R., 1980; Mc- 
Carthy, 1982). Jones et al. (1973) and Bimey et 
al. (1974) summarized the records for 31 species 
from Quintana Roo, and this paper provides one 
additional record. 



Materials and Methods 

The bats that I collected during the years 1974- 
1984 in Belize and El Peten (Parque Nacional Ti- 
kal), Guatemala, were obtained principally with 
mist nets set at ground level; aerial netting and the 
use of a bat trap were limited. Unless otherwise 
stated, mist netting was carried out during the first 
half of the night. A limited number of specimens 
were obtained with hand nets or plastic funnel 
traps at roost sites. Specimens were prepared as 
standard museum skins with skulls and/or skele- 
tons, or as fluid-preserved specimens. These 
vouchers are housed in Field Museum of Natural 
History, Chicago (FMNH); The Museum, Mich- 
igan State University, East Lansing (MSU); Car- 
negie Museum of Natural History, Pittsburgh 
(CM); and American Museum of Natural History, 
New York (AMNH). 

A survey of 45 museum collections in the United 
States, Canada, Mexico, Guatemala, and England 
resulted in additional specimens from Belize, El 
Peten, Izabal, Alta Verapaz, and Quintana Roo. 
Pertinent specimens (147) have been included in 
this report from the following institutions [collec- 
tors in brackets]: American Museum of Natural 
History, New York [N. Sullivan]; British Museum 
(Natural History), London, England (BM) [R. H. 
L. Disney; P. Williams; A. M. Hutson; R. E. Steb- 
bings]; Carnegie Museum of Natural History [N. 
A. Bitarj; Field Museum of Natural History [L. 
de la Torre]; Florida State Museum, University of 
Florida, Gainesville (FSM) [F. J. Bonaccorso]; 
Museum of Zoology, Louisiana State University, 
Baton Rouge (LSUMZ) [D. M. Uy]; Royal On- 
tario Museum, Toronto, Canada (ROM) [R. L. 
Peterson; J. Kamstra; J. Fragoso]; Texas Coop- 
erative Wildlife Collections, Texas A«&,M Uni- 
versity, College Station (TCWC) [D. C. Carter; M. 
D. Engstrom]; Texas Tech University, Lubbock 
(TTU) [P. Diamond]; and United States National 



Museum of Natural History, Washington, D.C. 
(USNM) [E. L. Tyson]. 

Systematic arrangement of species accounts and 
nomenclature, unless otherwise indicated, follow 
Jones et al. (1977) and Handley (1980). Disney 
(1968) did not provide data for the first records 
of Pteronotus davyi, Tonatia minuta, and Eptesi- 
cusfurinalis from Belize. Those data are presented 
in the respective accounts of this report, with ad- 
ditional records. Further secondary records from 
Belize of Mimon crenulatum, Trachops cirrhosus, 
Glossophaga commissarisi, Vampyressa pusilla, 
and Eumops auripendulus are also included. All 
of the species accounts are discussed in the context 
of their range and elevational distributions in 
Mexico and Central America. Hall (1981) was the 
primary reference for this unless cited otherwise. 

Forest types in Belize follow Wright et al. ( 1 959), 
whose classification was partially based on the sea- 
sonal formation series (Beard, 1 944), which refers 
to structural appearance. The correct political 
alignments between the states of the Yucatan Pen- 
insula are inconsistent among a number of pub- 
lished maps. The state boundary between Quin- 
tana Roo and Campeche on the map in Figure 1 
(see also Gazetteer) is based on a number of Gov- 
ernment of Mexico (Secretaria de Programacion y 
Presupuesto) maps, including "Carta Topografica, 
Merida" (1:1 ,000,000; 1 979 and 1 983) and "Mapa 
Geografica" (1:5,000,000; 1980). 



Species Accounts 
Family EMBALLONURIDAE 
Subfamily EMBALLONURINAE 
Saccopteryx leptura (Schreber, 1 774) 

Specimen Examined— BELIZE. Toledo: 2. 1 km 
NNE Salamanca Camp, Columbia Forest, 1 9 (cm). 

The known distribution of this small sac-winged 
bat north of Panama extends through Costa Rica 
and Nicaragua to Chiapas along the Pacific ver- 
sant. The presence of predominantly lowland Sac- 
copteryx leptura in southern Belize represents a 
country record and an extension of its distribution 
along the Caribbean side from southeastern Nic- 
aragua. 

Small bats were observed foraging up to heights 
of 13-13.5 m during the twilight period of the 
evening. Flight appeared to be concentrated within 



MCCARTHY: DISTRIBUTION OF BATS 



139 



a small, open area below the lower canopy of ev- 
ergreen seasonal forest. A short mist net was hand- 
hoisted to capture (24 March) this adult specimen. 
Saccopteryx bilineata was collected shortly after 
the capture of S. leptura. 



Balantiopteryx io Thomas, 1 904 

Specimens Examined— GUATEMALA. EI Pe- 
ten: Poptun, Finca Ixobel, \S 66, 1 8 99 (cm). 

The restricted distribution of Balantiopteryx io 
ranges from the Gulf lowlands of Veracruz, Oa- 
xaca, and Tabasco to the lowlands of Belize and 
eastern Guatemala. Kirkpatrick et al. ( 1 975), Cart- 
wright and Kirkpatrick (1977), and Sanborn (1936) 
represent the previous records for Belize and Iza- 
bal. The Poptun locality represents the first record 
for El Peten. 

The specimens reported here were collected ( 1 2 
June) by N. A. Bitar as they exited from a cave 
surrounded by secondary forest. The distribution 
of this colonial species may be restricted in part 
by the availability of adequate cave habitats as 
roosting sites. 



Subfamily DICLIDURINAE 

Diclidurus virgo Thomas, 1903 

Specimen Examined— BELIZE. Cayo: 1.5 km 
SSW Roaring Creek, 1 6 (fmnh). 

The white bat is represented by relatively few 
localities in Middle America, which extend from 
western (Nayarit) and eastern (southern Veracruz) 
Mexico through Central America. Specimens from 
southwestern El Peten were reported by Jones 
(1966). The single specimen from Belize repre- 
sents a northward range extension in the Carib- 
bean lowlands from northwest Honduras (Carter 
& Dolan, 1978) and a record for the country. 

The single bat apparently was roosting on the 
trunk of a fig tree (Ficus insipida) overhanging a 
pool along the Roaring Creek River. It was cap- 
tured (May) by C. Tzul after being observed on a 
number of occasions roosting near, but not among, 
a group of Rhynchonycteris naso. Jones ( 1 966), 
Starrett and Casebeer ( 1 968), and Handley ( 1 976) 
commented on the high foraging habits of Dicli- 
durus. Similar to the molossid bats, Diclidurus 
probably concentrates its foraging efforts at levels 
well above the tree canopy and beyond the reach 
of conventional collecting techniques, except fire- 



arms. This may explain why there are few speci- 
mens available in collections. 

Goodwin ( 1 969) considered Diclidurus virgo at 
best not more than subspecifically different from 
D. albus. Both species were recognized by Ojasti 
and Linares (1971), who questioned Goodwin 
( 1 969) because they believed that his South Amer- 
ican comparative material represented D. virgo and 
not D. albus. 



Family NOCTILIONIDAE 

Noctilio leporinus mastivus (Vahl, 1797) 

Specimens Examined— BELIZE. Cayo: Banana 
Bank, 1 <5, 1 9 (fmnh); Barton Creek, at Western 
Hwy., 1 9 (fmnh). Stann Creek: Melinda, Stann 
Creek River, 1 6 (fmnh). Toledo: 1.2 km E Agua- 
cate, Aguacate River, 2 66 (cm), 1 9 (bm); Big Fall, 
vicinity Rio Grande Bridge, 1 9 (fmnh); Salamanca 
Camp, 1 9 (bm). 

The fishing bat occurs along the riparian habi- 
tats of river systems, inland lakes, and coastlines 
in primarily lowland regions from northwestern 
(northern Sinaloa), eastern (southern Veracruz), 
and peninsular (Yucatan) Mexico throughout Cen- 
tral America (Davis, 1973;Hellebuycketal., 1985). 
Dickerman et al. (1981) reported a locality for 
Noctilio leporinus from Alta Verapaz as in the Ca- 
ribbean drainage when it was clearly in the Rio 
Usamacinta drainage of the Gulf lowlands. The 
Belizean localities extend northward the recorded 
occurrence of A^. leporinus from Izabal and north- 
western Honduras (Carter et al., 1966). 

All of the specimens were obtained (March, 
April, May, July, August) over rivers and a pond 
except for one individual, which was mist netted 
(28 August) low over a pasture adjacent to a flood- 
ed river. This bat was foraging primarily for insects 
since its feces contained the chitinous remains of 
these prey. Additional fishing bats from the lo- 
calities in Cayo and Stann Creek districts were 
captured, banded, and released. This bat was com- 
mon along the South Stann Creek drainage. Cocks- 
comb Basin. A specimen belonging to M. Craig, 
Belize Audubon Society, was collected at Indian 
Church (Lamanai), New River Lagoon, Orange 
Walk District. An old specimen of M leporinus in 
the collections of British Museum (Natural His- 
tory) was registered in 1 909 without pertinent field 
data. The two peninsular records from Campeche 
(Jones et al., 1973) and Yucatan (Bimey et al., 
1974) were obtained in coastline habitat along the 



140 



FIELDIANA: ZOOLOGY 



Gulf of Mexico. Although subsurface drainage 
predominates north of Belize into Quintana Roo, 
shallow inland "lagunas" are fairly common and 
probably support Noctilio populations. 



Family MORMOOPIDAE 
Pteronotus davyi fulvus (Thomas, 1 892) 

Specimens Examined— BELIZE. Cayo: Central 
Farm, 1 <5 (cm), 1 $ (fmnh); Ontario, 5.5 km W 
Teakettle, 1 S (fmnh); Unitedville, 9 km WSW 
Teakettle, 1 <5 (fmnh). Orange Walk: Tower Hill, 
B.S.I, compound, 3 99 (fmnh). Toledo: Aguacate, 
1 (3 (cm); 1.2 km E Aguacate, 1 $ (bm), 1 $ (cm); 
Rice Station, 2 6$ (fmnh); 0.4 km W Rice Station, 
1 $ (fmnh); San Antonio, 1 $ (fmnh). GUATE- 
MALA. El Peten: Parque Nacional Tikal, 1 $ (msu). 

Smith (1972) summarized the majority of the 
capture localities for this subspecies of naked- 
backed bat, which ranges from northwestern (So- 
nora), northeastern (Tamaulipas), and peninsular 
(Yucatan) Mexico southeastward to Honduras and 
El Salvador, but omitted the only record for Belize 
(Disney, 1968). Parque Nacional Tikal is the first 
record for El Peten, and the Belizean specimens 
provide additional records for Belize. 

Disney ( 1 968) did not present data for his single 
Pteronotus davyi specimen. This male was ob- 
tained ( 1 November) in Cayo District, at Listowel 
along the Belize River, and is housed in British 
Museum (Natural History). The subsequent spec- 
imens reported here were collected (October-De- 
cember, May, July, August) in open areas, bor- 
dering on vegetation and buildings, and over water. 
The specimen from El Peten was captured (25 
March) along a trail in upland deciduous forest. 
An additional P. davyi from Tikal was captured, 
banded, and released. 



Pteronotus personatus psilotis (Dobson, 1878) 

Specimens Examined— BELIZE. Toledo: 1 .2 km 

E Aguacate, Aguacate River, 1 9 (bm), 4 33, 1 9 
(CM); Big Fall, 1.5 km WSW Rio Grande Bridge, 
1 $ (fmnh); 0.8 km NW Blue Creek, 1 9 (fsm); 
Crique Jute, 1 $ (fmnh); Crique Lagarto, 1 km 
NW San Antonio, 1 9 (fmnh); Jacinto Creek, at 
Punta Gorda Road, 1 3 (msu); 0.4 km W Rice 
Station, 1 9 (fmnh); Salamanca Camp, 1 9 (usnm); 
San Antonio, 1 9 (fmnh); 0.9 km WNW San Pedro 
Columbia, 1 9 (fmnh). 



The distribution of Pteronotus personatus psi- 
lotis extends from western (southern Sinaloa) and 
eastern (Tamaulipas) Mexico southeastward to 
Honduras and El Salvador (Smith, 1972), with 
Caribbean lowland localities in Campeche (Jones 
et al., 1973), El Peten (Jones, 1966), and Alta Ver- 
apaz (Jones, 1966). Elevations range from 123 to 
984 m. These localities from southern Belize are 
the first records for the country. 

Fifty-three percent of the small moustache bats 
were collected (March, May, July) over open water; 
the remainder were foraging (January, April, Au- 
gust, December) in open areas adjacent to build- 
ings or corralled cattle. 



Mormoops megalophylla megalophylla 

Peters, 1864 

Specimens Examined— BELIZE. Belize: 6.6 km 

N Churchyard, 1 9 (cm). Cayo: 1.6 km NW Au- 
gustine, Rio Frio, 1 3 (ttu). Stann Creek: Melinda, 
1 3 (fmnh). Toledo: Forest Home, 1 3 (msu); Pueb- 
lo Viejo, 1 3 (fmnh). GUATEMALA. Izabal: 25 
km SSW Puerto Barrios, 1 3 (tcwc). 

The leaf-chinned bat has been reported through- 
out Mexico, Guatemala, El Salvador, and Hon- 
duras (Smith, 1972). Davis and Carter (1962), 
Jones (1966), and Taibel (1977) provided lowland 
records for El Peten and Alta Verapaz. Elevations 
range from near sea level to 2270 m. These lo- 
calities are the first records for Belize and Izabal. 

Except for one Belizean specimen, which was 
captured (9 June) in a cave, these leaf-chinned bats 
were associated (March, April, December) with 
open areas bordering on forest or orchard edges, 
including pine savanna. One Mormoops specimen, 
which was registered into the British Museum 
(Natural History) collections in 1892, may have 
been obtained in the vicinity of Belize City. The 
Guatemalan specimen was collected by D. C. Car- 
ter. 



Family PHYLLOSTOMIDAE 
Subfamily PHYLLOSTOMINAE 

Micronycteris brachyotis (Dobson, 1878) 

Specimens Examined— BELIZE. Cayo: 1 km 

NW Augustine, 2 $S (fmnh). Toledo: Crique Ne- 
gro, Columbia Forest, 1 3 (bm). 

The first Middle American specimen of Micro- 



McCARTHY: DISTRIBUTION OF BATS 



141 



nycteris brachyotis was initially reported from Nic- 
aragua as M. syhestris by Goodwin (1946), but 
was correctly identified by Sanborn (1949). Sub- 
sequent records are from the Gulf-Caribbean low- 
lands of southern Veracruz (Medellin L. et al., 
1983), Oaxaca (Schaldach, 1964), Chiapas (Davis 
et al., 1964), and El Peten (Jones, 1966; Rick, 
1 968; McCarthy, 1 982); and the Pacific-Caribbean 
versants of Costa Rica (Howell & Burch, 1974; 
Starrett, 1976; LaVal & Fitch, 1977) and Panama 
(Handley, 1966; Fleming etal., 1972; Bonaccorso, 
1979). Reported elevations range from 40 to 594 
m. The present specimens are the first records of 
the yellow-throated bat for Belize. 

Two specimens were captured (29 July) as they 
exited from a cave into a low deciduous seasonal 
forest, and a third bat was taken (28 May) along 
a path in an evergreen seasonal forest. The two 
specimens from Cayo, which I obtained while as- 
sisting a histoplasmosis survey, were listed as "M. 
bardyoiis" in a preliminary report (Quinones et 
al., 1978, p. 559) and no specific locality data were 
provided. 



Micronycteris megalotis mexicana Miller, 1898 

Specimens Examined— BELIZE. Corozal: San 

Antonio, 2 km NW Corozal, 1 6 (fmnh). Orange 
Walk: San Antonio, Rio Hondo, 2 55, 1 2 (fmnh). 
Toledo: Aguacate, 1 6 (cm); Big Fall, 2 km E Rio 
Grande Bridge, 1 2 (bm); Cuevas Creek Bridge, 10 
km NW Punta Gorda, 1 5, 1 2 (bm), 1 S (amnh), 
1 2 (msu); Nimli Punit, 1 2 (cm); Rocky Run Ranch, 
4.8 km NW Punta Gorda, 1 3, 1 2 (bm); Union 
Camp, 2 22 (bm); Vista Hermosa Ranch, 3.7 km 
WNW Punta Gorda, 1 2 (cm). GUATEMALA. El 
Peten: Parque Nacional Tikal, 1 <5 (fmnh). 

The distribution of this subspecies of big-eared 
bat extends from western (Jalisco), eastern (south- 
em Tamaulipas), and peninsular (Yucatan) Mex- 
ico, along the Pacific coastal and highland regions, 
to Costa Rica. Gardner et al. ( 1 970) suggested that 
the southern extent of Micronycteris megalotis 
mexicana is in the Cordillera Talamanca of Costa 
Rica. This species has been recorded most often 
at lowland-moderate elevations, up to 2870 m. 
Specimens from Isla Cozumel, Quintana Roo, rep- 
resent the only record for Quintana Roo (Jones et 
al., 1973). The records of A/, m. mexicana which 
are reported here are the first for Belize and El 
Peten. 

Belizean specimens were obtained (May, July, 
August, November) in diurnal roost sites (shallow 



caves and limestone chambers, bridge approach- 
ments, abandoned rum factory boiler) and col- 
lected in forest habitats (riparian marsh, evergreen 
and semi -evergreen, deciduous semi -evergreen, 
and deciduous seasonal). The Tikal specimen was 
captured (6 June) roosting in a passageway of an 
excavation tunnel within a ruin complex. A second 
juvenile male was captured, banded, and released 
(29 July) in escobal palm (Cryosophila argentea) 
forest, 1.9 km SE Tikal Reservoir. 



Micronycteris nicefori Sanborn, 1 949 

Specimen Examined— BELIZE. Toledo: 0.4 km 

NE Aguacate, 1 2 (fmnh). 

Handley ( 1 966) documented the first specimens 
of Micronycteris nicefori north of South America, 
from Panama. Subsequently, it has been reported 
from southeastern Nicaragua (Baker & Jones, 1975) 
and both the dry Pacific (Starrett, 1976) and wet 
Caribbean (LaVal, 1977) lowlands of Costa Rica. 
These Central American localities range from near 
sea level to over 100 m. This first record from 
Belize also represents a significant Central Amer- 
ican range extension along the Caribbean versant. 

The M. nicefori specimen reported here was mist 
netted on 1 5 December along a track in hilltop, 
evergreen seasonal forest. 



Micronycteris schmidtorum Sanborn, 1935 

Specimens Examined— BELIZE. Corozal: Pat- 
chakan, 2 22 (fmnh). Orange Walk: 1 .3 km W San 
Antonio, Rio Hondo, 1 6 (fmnh). Toledo: Big Fall, 
1 km E Rio Grande Bridge, 1 6 (cm). 

Micronycteris schmidtorum was described (San- 
bom, 1935) from specimens collected in the Ca- 
ribbean lowlands of Izabal. An additional Gua- 
temalan specimen was recorded in the Pacific 
piedmont (Dickerman et al., 1981). The remaining 
Central American records represent both the Pa- 
cific and Caribbean lowland slopes from Honduras 
(Sanbom, 1941), Nicaragua (Davis et al., 1964; 
Baker & Jones, 1 975), Costa Rica (Starrett & Case- 
beer, 1968; Fleming et al., 1972; Howell & Burch, 
1974; LaVal & Fitch, 1977), and Panama (Han- 
dley, 1 966). Specimens from Yucatan assigned to 
M. schmidtorum by Villa-R. (1966) were reiden- 
tified as M. megalotis by Jones et al. (1973). An 
identification of M. schmidtorum (Jones et al., 
1 973) for a specimen from Isla Cozumel, Quintana 
Roo, was questioned by Hall (1981) because this 



142 



FIELDIANA: ZOOLOGY 



specimen previously was identified as M. mega- 
lot is (Jones & Lawlor, 1 965). I examined this spec- 
imen (University of Kansas 9 1 539) and agree that 
it is M. schmidtorum. The northern distribution 
of this big-eared bat extends to the Caribbean coast 
of the Yucatan Peninsula. The specimens reported 
here are the first records for Belize. 

At Parque Nacional Tikal, one juvenile and two 
adult females, which were captured (30 July) in a 
hollow tree (Bursera semirouba) of an upland de- 
ciduous seasonal forest, were photographed, band- 
ed, and released. This site was revisited during the 
following March, but no Micronycteris were found. 
These individuals of M. schmidtorum were the 
first seen in El Peten. Similarly, Sanborn (1935) 
and Starrett and Casebeer (1968) reported indi- 
viduals from tree hollows. The Belizean specimens 
were captured (February, September, November) 
in the orchard vegetation of a village, along a sec- 
ondary forest edge, and in riparian secondary 
vegetation. 



Lonchorhina aurita aurita Tomes, 1863 



Macrophyllum macrophyllum (Shinz, 1821) 

Specimens Examined— BELIZE. Cayo: Sibun 
River at Indian Creek, 1 $ (fmnh). Toledo: Big 
Fall, 1.7 km NE Rio Grande Bridge, 1 $ (cm). 

Tabasco, Mexico, represents the northernmost 
occurrence for the long-legged bat, which is known 
from both the Caribbean and Pacific regions of 
Central America. Primarily a lowland inhabitant, 
Macrophyllum macrophyllum ranges from 40 to 
almost 600 m. These specimens represent a Ca- 
ribbean lowlands range extension from north- 
western Honduras (Valdez «& LaVal, 1971) and 
the first records for Belize. 

Harrison and Pendleton (1974), Gardner (1977), 
and Dickerman et al. (1981) indicated that long- 
legged bats may be closely associated with aquatic 
habitats. Similarly, the Belizean specimens were 
obtained ( 1 7 March, 1 April) from along the Sibun 
River, although not directly above water, and over 
the surface of the Rio Grande. The first bat was 
taken at approximately 0340 in a stand of shade 
trees, dominated by cohune palms (Orbignya co- 
hune), at the edge of an open pasture. 



Specimens Examined— BELIZE. Stann Creek: 

5.3 km WNW Quam Bank, Cockscomb Basin, 1 
9 (CM). Toledo: 0.8 km NW Blue Creek, 1 3, 1 2 
(amnh); Crique Jute Village, 1 9 (cm); Crique Ne- 
gro, Columbia Forest, 1 S (bm), 1 $ (usnm); 2. 1 km 
NNE Salamanca Camp, Columbia Forest, 3 66 
(CM). GUATEMALA. El Peten: Poptun, Finca 
Ixobel, 2 66 (cm). 

Lonchorhina aurita was first recorded in Middle 
America from Panama (Miller, 1912). Subsequent 
collecting has found this cave-dwelling bat north- 
ward through Central America to southeastern 
(southern Veracruz, Oaxaca, Tabasco) and pen- 
insular (Quintana Roo) Mexico. Predominately 
lowland, this distinctive leaf-nosed bat extends up 
to more than 1500 m in representative habitats. 
Jones et al. (1973) reported the only record from 
Quintana Roo, while specimens from Izabal (San- 
bom, 1936) are apparently the next Caribbean ver- 
sant record north of eastern Costa Rica (Nelson, 
1965); records from Nicaragua and Honduras are 
lacking. The specimens examined for this account 
are the first records from Belize and El Peten. 

All specimens from Belize were captured (March, 
April, May, August) in deciduous seasonal and 
evergreen seasonal forests. The Guatemalan bats 
were captured by N. A. Bitar as they exited from 
the cave discussed in the Balantiopteryx io ac- 
count. 



Tonatia bidens bidens (Spix, 1823) 

Specimens Examined— BELIZE. Cayo: Rio 

Frio, 1 .6 km W Augustine, 1 9 (cm). Toledo: Nimli 
Punit, 1 6 (cm); Orange Creek, 1.5 km SW Punta 
Gorda, 1 6 (msu); 2. 1 km NNE Salamanca Camp, 
Columbia Forest, 1 6 (cm); 2.2 km NNE Sala- 
manca Camp, Columbia Forest, 1 9 (cm). 

Goodwin (1946) first recorded Tonatia bidens 
in Central America from the Pacific lowlands of 
Costa Rica. Other humid lowland records include 
both the Caribbean and Pacific versants of Pan- 
ama, continuing along the Caribbean corridor of 
Nicaragua, Honduras, and Guatemala. The north- 
ernmost record is from eastern Chiapas (Medellin 
L., 1983). The Guatemalan records are from the 
Caribbean lowlands of El Peten (McCarthy, 1982) 
and Izabal (Carter et al., 1966). Elevations range 
from near sea level to around 660 m. The present 
specimens constitute the first records from Belize. 

Four adult males were taken (March, April) over 
a creek in a low transitional forest, in a high ev- 
ergreen seasonal forest, and in a deciduous sea- 
sonal forest. A subadult male was captured (24 
September) in the courtyard of a Mayan archae- 
ological site located in a high deciduous seasonal 
forest. 



MCCARTHY: DISTRIBUTION OF BATS 



143 



Tonatia evotis Davis and Carter, 1978 

Specimen Examined- GUATEMALA. El Pe- 
ten: Parque Nacional Tikal, 1 6 (fmnh). 

Davis and Carter ( 1 978) described Tonatia evo- 
tis on the basis of its smaller size in comparison 
to T. sylvicola; a female from Izabal was designated 
as the holotype. El Peten is part of a Gulf-Carib- 
bean distribution which extends from southern 
Veracruz, Tabasco, Chiapas, and Campeche to Be- 
lize, and continues along northern Honduras (Da- 
vis & Carter, 1978). Martinez R. (1980) recorded 
an additional eastern Guatemalan locality in Aha 
Verapaz. All recorded elevations are less than 100 
m. The T. evotis from Tikal represents the first 
record for El Peten. 

Two adult males and one pregnant female were 
mist netted (20 February, 29 and 25 March) in 
Tikal along the Uaxactun Road, at a permanent 
water pool in escobal palm forest, and in an upland 
deciduous seasonal forest. One male and the fe- 
male were banded and released. 



Tonatia minuta Goodwin, 1 942 

Specimens Examined— BELIZE. Cayo: 1.1 km 

W Augustine, 1 2 (fmnh); Central Farm, at Belize 
River, 1 9 (fmnh); 1.2 km E Macaw Bank, 1 2 
(fmnh). Toledo: Big Fall, 1.7 km NE Rio Grande 
Bridge, 1 2 (msu); San Lucas, 1 2 (msu). 

This small Tonatia was originally described from 
the Caribbean coast of Nicaragua as T. nicaraguae 
(Goodwin, 1942a). Its Middle American distri- 
bution is lowland ( 1 5 to 6 1 m) along Caribbean 
and Pacific versants, from southern Veracruz 
(Lackey, 1 970) to El Peten, Guatemala (McCarthy, 
1982) and Belize (Disney, 1968), continuing 
through Honduras (LaVal, 1969; Valdez & LaVal, 
1971; Greenbaum & Jones, 1978), Nicaragua 
(Jones et al., 1971; Greenbaum & Jones, 1978), 
and Costa Rica (Gardner et al., 1 970; LaVal, 1 977), 
to Panama (Davis et al., 1964; Handley, 1966). 
This account represents additional records for the 
small round-eared bat in Belize. 

Disney (1968) reported no data for the first To- 
natia minuta specimen from Belize, which was a 
female collected (25 November) in Cayo District, 
at Listowel, along the Belize River. This specimen 
was deposited in British Museum (Natural His- 
tory). The additional specimens reported here were 
captured (November, January, February, April, 
May) over rivers or in a deciduous seasonal forest. 

The name minuta is applied in accordance with 



the discussion by McCarthy ( 1 982). Gardner ( 1 976) 
referred to a personal communication with C. O. 
Handley, Jr., who suggested that all small Tonatia 
(including minuta) represent a single species, T. 
brasiliense. Because the taxonomy is poorly under- 
stood, a systematic review of this group would be 
useful. 



Mimon cozumelae Goldman, 1914 

Specimens Examined— BELIZE. Belize: 

Churchyard, Sibun River, 1 2 (fmnh). Cayo: 
"Mountain Pine Ridge", 2 33, 1 2 (bm); 0.8 km W 
Augustine, 1 6 (cm); 1 km NW Augustine, 2 $6 
(fmnh); Barton Creek, at Western Hwy., 2 $S, 3 
22 (fmnh). Toledo: vicinity Aguacate, 2 $S, 2 22 
(cm), 1 (5 (fmnh); Crique Negro, Columbia Forest, 
1 2 (bm); Pueblo Viejo, 1 3, 1 2 (fmnh); 2. 1 km 
NNE Salamanca Camp, Columbia Forest, 2 6S 
(cm); 2.2 km NNE Salamanca Camp, Columbia 
Forest, 1 <5 (cm); vicinity Union Camp, 2 5(5, 1 2 
(bm), 2 22 (cm). 

This spear-nosed bat ranges from southeastern 
(northern Oaxaca, southern Veracruz) and pen- 
insular (Yucatan, Quintana Roo) Mexico south- 
eastward along the humid Caribbean side of Cen- 
tral America. Specimens from Isla Cozumel, 
Quintana Roo, provided the original description 
for this species (Goldman, 1914). Recorded ele- 
vations extend to 495 m. The Belizean localities 
reported here are the first records for the country. 

Mimon cozumelae were collected (January, 
March, May, July, August, September, December) 
along the edge of deciduous and semi-evergreen 
seasonal forests bordered with pasture, on riparian 
flood plains, over rivers, along paths in high de- 
ciduous, semi-evergreen seasonal forests, and in 
caves. 

Schaldach (1964), Villa-R. (1966), and Hall 
(1981) considered cozumelae a subspecies of ben- 
nettii. I tentatively accept cozumelae at the specific 
level. 



Minion crenulatum keenani Handley, 1 960 

Specimens Examined— BELIZE. Cayo: Listow- 
el, Baking Pot, 1 S (fmnh). Toledo: Crique Negro, 
Columbia Forest, 1 6 (usnm). 

There are few records for Mimon crenulatum 
keenani from Middle America. The distribution 
of this distinctive spear-nosed bat extends along 
the Caribbean versant, from Panama (Handley, 



144 



FIELDIANA: ZOOLOGY 



1966; Bonaccorso, 1979), Costa Rica (Gardner et 
al., 1970; LaVal, 1977), Nicaragua (Greenbaum & 
Jones, 1978), Belize (Ruiz, 1983), El Peten 
(McCarthy, 1982), and Campeche (Jones, 1964) 
to the Gulf lowlands of eastern Chiapas (Medellin 
L., 1 983). All recorded elevations range below 265 
m. These specimens are the second and third rec- 
ords from Belize. The first record (Ruiz, 1 983) was 
obtained near Blue Hole, 14 km SE Belmopan, 
Cayo District. 

One Mimon crenulatum was captured (8 Oc- 
tober) in a house after it flew through an open 
window. The house was situated along the Belize 
River in an agricultural area. The second specimen 
was netted (29 March) along a path in evergreen 
seasonal forest. E. L. Tyson collected the specimen 
from Toledo District. 



Phyllostomus discolor verrucosus Elliot, 1905 

Specimens Examined— BELIZE. Toledo: Cri- 
que Lagarto, 1 km NW San Antonio, 1 S (fmnh); 
1 km NNE Salamanca Camp, Columbia Forest, 1 
(3 (cm). GUATEMALA. Alta Verapaz: Lanquin, 
Lanquin Cave, approx. 1 49 km WSW Puerto Bar- 
rios, 1 (5, 1 5 (fmnh). 

Records of Phyllostomus discolor extend from 
southern (Oaxaca, Veracruz) Mexico along both 
the Pacific and Caribbean corridors of Central 
America. Records are more common at lower el- 
evations, less than 600 m. The new records from 
southern Belize provide a limited range extension 
northward from eastern Izabal (Sanborn, 1936). 

An adult from Crique Lagarto was captured ( 1 
January) along the edge of low secondary forest 
bordering this settlement. The head of the bat was 
covered with yellow pollen. The second specimen 
was netted (21 March) in secondary vegetation, 
which resulted from slash-bum agriculture. Whit- 
ish pollen dusted the face, chest, and ventral wing 
surfaces. A male subadult Phyllostomus discolor 
that was taken ( 1 3 July) along a fenceline of sec- 
ondary vegetation between two pastures, 1 .9 km 
ENE Rio Grande Bridge, Big Fall, Toledo District, 
was photographed, banded, and released. L. de la 
Torre apparently captured (3 June) the two Phyl- 
lostomus from Alta Verapaz inside the entrance 
of Lanquin Cave. 

I tentatively follow Jones et al. (1977) in as- 
signing the specimens of Phyllostomus discolor 
from the Caribbean lowlands to the subspecies 
verrucosus. Sanborn (1936, p. 98) recognized ver- 
rucosus subspecifically, stating the "available mea- 



surements of rf/5co/or would place them much clos- 
er to verrucosus."'' He suggested the Panamanian 
P. d. discolor are assignable to verrucosus based on 
larger size. Felten (1956) and ^urt and Stirton 
(1961) concurred with his statement by referring 
a large series from El Salvador to verrucosus; with 
the availability of greater series of specimens, Da- 
vis and Carter (1962) indicated they could not 
recognize two subspecies of P. discolor in Central 
America and northern South America, acknowl- 
edging only P. d. verrucosus. Handley ( 1 966) ap- 
parently disagreed as he recognized the subspecies 
discolor in Panama. Multivariate analysis of mor- 
phological data (Power & Tamsitt, 1 973) suggested 
this species might be monotypic. 



Phylloderma stenops septentrionalis 

Goodwin, 1940 

Specimens Examined— BELIZE. Toledo: Cri- 
que Negro, Columbia Forest, 1 2 (usnm); 2. 1 km 
NNE Salamanca Camp, Columbia Forest, 2 $S 

(CM). 

This rarely encountered species has been re- 
corded north of Panama from the Caribbean coast 
of Costa Rica (LaVal, 1977), the highlands of Hon- 
duras (Goodwin, 1940), the Caribbean lowlands 
of Guatemala (McCarthy, 1982), and the Gulf 
lowlands of Chiapas (Carter et al., 1966). Limited 
elevational data are from lowland to approxi- 
mately 1320 m. The specimens oi Phylloderma 
stenops from Belize represent the eighth, ninth, 
and tenth specimens north of Panama and the first 
records from Belize. 

All specimens were mist netted (March, Decem- 
ber) in similar evergreen seasonal forest habitats. 
E. L. Tyson collected the specimen from Crique 
Negro. 

Handley ( 1 966) regarded the Panamanian spec- 
imens to be Phylloderma stenops stenops, and those 
from northward into Middle America were thought 
to be subspecifically different from the nominal 
species. LaVal (1977) did not designate a subspe- 
cies for his Costa Rican specimen. 



Trachops cirrhosus coflini Goldman, 1925 

Specimens Examined— BELIZE. Orange Walk: 

Richmond Hill (Goat Hill), 8.9 km SSW Orange 
Walk Town, 1 3, 1 $ (cm). Toledo: 2.2 km NNE 
Salamanca Camp, Columbia Forest, 1 $ (cm). 



MCCARTHY: DISTRIBUTION OF BATS 



145 



GUATEMALA. Izabal: 25 km SSW Puerto Bar- 
rios, 1 $ (tcwc). 

This lowland subspecies of the fringe-lipped bat 
is recognized from eastern (southern Veracruz) and 
southeastern (eastern Oaxaca) Mexico southeast- 
ward to Nicaragua. Recorded elevations are from 
near sea level to approximately 330 m. Jones 
(1966), Rick (1968), and McCarthy (1982) pro- 
vided records for El Peten. The description of this 
subspecies was based on specimens from eastern 
El Peten (Goldman, 1925). The first Belizean rec- 
ords were reported from Belize District in the vi- 
cinity of Belize City (Sanborn, 1941) and Rock- 
stone Pond (Pendergast, 1979). The specimen from 
Izabal is the first record for that Guatemalan de- 
partment. 

D. C. Carter obtained the single specimen from 
Izabal on 19 February. The additional Belizean 
specimens were mist netted (March, April) in de- 
ciduous marsh and evergreen forests. 



Chrotopterus auritus (Peters, 1856) 

Specimens Examined— BELIZE. Toledo: vicin- 
ity Crique Negro, Columbia Forest, 1 9 (fmnh); 
1.6 km NNE Salamanca Camp, Columbia Forest, 
1 9 (fmnh). 

Chrotopterus was first reported in Central Amer- 
ica from El Salvador (Burt & Stirton, 1961). Sub- 
sequently, this carnivorous bat has been recorded 
from southern (southern Veracruz, northern Oa- 
xaca, Chiapas) and peninsular (Yucatan, Quintana 
Roo) Mexico southeastward throughout Central 
America at lowland and upland elevations (40 to 
over 1 880 m). Chrotopterus auritus has been re- 
ported from Quintana Roo (Jones et al., 1 973) and 
El Peten (Rick, 1968; McCarthy, 1982). These 
specimens from southern Belize provide the first 
records for the country. 

The Belizean specimens were netted (10 April, 
28 July) in an evergreen seasonal forest at ground 
level along a path and at a height of about 13.7m 
over an intermittent stream bed. Both were active 
during the morning hours, 0418 and 0330, re- 
spectively. 

The subspecific name Chrotopterus auritus au- 
ritus has been applied to Middle American pop- 
ulations (Jones et al., 1971). Carter and Dolan 
(1978) stated the type specimen for Vampyrus au- 
ritus Peters, 1856, actually was collected in Santa 
Catarina, Brazil, not in Mexico. The discussion by 
Carter and Dolan (1978, p. 37) suggested that Pe- 



ters based his description on one or more speci- 
mens from Brazil and compared these with a spec- 
imen from an unrecorded locality in Mexico as 
the "verwandten Art aus Mexico." Handley ( 1 966) 
doubted that subspecies were recognizable. 



Vampyrum spectrum (Linnaeus, 1758) 

Specimen Examined— BELIZE. Toledo: Santa 
Elena, 1 S (fmnh). 

Two localities in southern Veracruz, Mexico 
(Goldman, 1917; Navarro L., 1 979) are the north- 
westernmost records of the false vampire bat's 
Middle American distribution, which continues in 
Nicaragua (Dobson, 1 878; Allen, 1910), Costa Rica 
(Casebeer et al., 1963; Armstrong, 1969; Gardner 
et al., 1970; Howell &. Burch, 1974; Vehrencamp 
et al., 1977; LaVal & Fitch, 1977), and Panama 
(Handley, 1966; Peterson & Kirmse, 1969; Bo- 
naccorso, 1979). Although primarily lowland in 
distribution, its highest recorded elevation was 
about 1815m. The occurrence of Vampyrum spec- 
trum in the Caribbean lowlands of Belize is doc- 
umented by this specimen. 

There appears to be no definite record of this 
carnivorous bat from Guatemala (Jones, 1966). 
Dobson (1878, p. 471) recorded "Guatemala" as 
part of the Central American range for Vampyrum, 
but did not list any examined specimens. Alston 
(1879-1882, p. 39) stated Dobson (pers. comm.) 
saw specimens from Guatemala, although Alston 
realized the collector, O. Salvin, had not obtained 
specimens of Vampyrum; hence, the identification 
of this species is doubtful. Five false vampire bats 
were mist netted on three separate dates in Parque 
Nacional Tikal, El Peten. Two females were cap- 
tured during the dry season (22 and 24 March) in 
an upland deciduous seasonal forest, in the vicin- 
ity of Central Plaza of the archaeological site, and 
at a permanent water pool in escobal palm forest, 
2.6 km SE Central Plaza. Two females and one 
male were netted during the wet season (22 July) 
at a location along an archaeological transect in 
escobal palm forest, 1 km SE Tikal Reservoir. All 
of these bats were released after being observed, 
measured, and/or photographed. These individ- 
uals provide the first record for Guatemala and, 
along with the specimen from Belize, bridge an 
intermittent distribution that now extends north- 
ward toward peninsular Mexico. 

The Vampyrum spectrum from Belize was cap- 
tured (8 April) during the early morning (0300) in 



146 



FIELDIANA: ZOOLOGY 



an open field. We were "trapping" Desmodus ro- 
tundus during a vampire bat control effort in the 
village. This large bat was captured after it made 
a number of low passes over horses and mules, 
which were encircled by mist nets. The bat died 
while enroute to captivity via an assistant. 

The Central American population of Vampy- 
rum was described as a distinct subspecies, V. s. 
nelsoni (Goldman, 1914), but Handley (1966) ar- 
gued that the species was monotypic. 



Subfamily GLOSSOPHAGINAE 

Glossophaga commissarisi commissarisi 

Gardner, 1962 

Specimens Examined— BELIZE. Belize: Rock- 
stone Pond, 2 SS, 3 99 (rom). Toledo: Aguacate, 1 
9 (fmnh), 1 9 (cm); Big Fall, 1 km SE Rio Grande 
Bridge, 2 $S (cm); Forest Home, 1 9 (fmnh); 2.8 
km NNW Punta Gorda, 1 9 (fmnh). GUATE- 
MALA. Izabal: 25 km SSW Puerto Barrios, 7 SS, 
6 99 (tcwc). 

Webster and Jones (1982) summarized the dis- 
tribution for this subspecies of nectivorous bat, 
which was documented from eastern (Veracruz) 
and southern (Oaxaca, Chiapas) Mexico and 
southern Belize southeastward throughout Central 
America. Hellebuyck et al. (1985) recently re- 
ported records from El Salvador. The specimens 
from Izabal are the first records from this Gua- 
temalan department. The specimens from Belize 
District extend northward the distribution of Glos- 
sophaga commissarisi along the Caribbean low- 
lands. 

According to D. C. Carter's field notes, the ma- 
jority of the Guatemalan Glossophaga commis- 
sarisi were mist netted (February, March) over a 
stream and in the adjacent undisturbed forest. 
Many of these nectivorous bats were captured in 
association with night-blooming "bat flowers" 
bordering on a stream. The Belizean specimens 
reported (Webster & Jones, 1982) from Lubaan- 
tun, Toledo District, were collected ( 1 8 April) in 
a disturbed semi-evergreen seasonal forest. Ad- 
ditional specimens were secured (January, July, 
September, December) in secondary and orchard 
vegetation of villages, in riparian secondary vege- 
tation, and from the hollow of a mamey tree (Pou- 
teria mammosa). 



Subfamily STENODERMATINAE 
Uroderma bilobatum molaris Davis, 1968 

Specimen Examined— MEXICO. Quintana Roo: 

2 km N, 8 km W Bacalar, 1 $ (tcwc). 

Davis (1968) recognized this subspecies of the 
tent-making bat from the Gulf-Caribbean versant 
of southern Veracruz, Tabasco, northeast Oaxaca, 
northern Chiapas, Belize, Honduras, Nicaragua, 
Costa Rica, and northwest Panama. Disney (1968) 
and Pendergast (1979) also reported the occur- 
rence of Uroderma bilobatum from Belize. The 
specimen reported here represents the first record 
for Quintana Roo and a marginal range extension 
into the Mexican peninsula of Yucatan. 

The above specimen was taken in a net on 6 
August by M. D. Engstrom along a path leading 
to an inland lagoon. 

Vampyrops helleri helleri Peters, 1866 

Specimens Examined— BELIZE. Cayo: Banana 
Bank, 5 99 (fmnh); 0.8 km W Macaw Bank, 1 6 
(fmnh). Toledo: Big Fall, 1.9 km ENE Rio Grande 
Bridge, 1 9 (amnh), 1 9 (cm), 1 $ (msu); Crique 
Negro, Columbia Forest, 1 $ (bm); Forest Home, 
1 (5 (fmnh), 1 (5 (msu); Salamanca Camp, 1 S (bm), 
1 (5 (fmnh), 1 9 (usnm); 1.8 km NNE Salamanca 
Camp, Columbia Forest, 1 9 (fmnh); vicinity Union 
Camp, 1 9 (bm), 2 99 (cm). 

The Middle American records of this fruit bat 
indicate a distribution from sea level to elevations 
of over 1 300 m and a range from southeastern 
Mexico (southern Veracruz, Oaxaca, Tabasco) 
throughout Central America. Lowland records 
have been reported from El Peten (Rick, 1968) 
and Izabal (Carter et al., 1966). This account con- 
stitutes the first records from Belize. 

Eighty-seven percent of the Vampyrops helleri 
specimens were captured along or in proximity to 
waterways. Eleven additional individuals were re- 
leased at Banana Bank, where a concentration of 
stenodermatines (Sturnira, Uroderma, Vampyres- 
sa, Chiroderma, Artibeus, and Vampyrops) was 
observed. The remaining localities were in upland 
evergreen seasonal forest and in disturbed village 
vegetation. A specimen in the collection of St. 
John's College, Belize City, was collected by E. L. 
Tyson in Columbia Forest. 

I follow Dickerman et al. (1981) for the taxo- 
nomic assignment of the subspecific epithet. 



MCCARTHY: DISTRIBUTION OF BATS 



147 



Vampyrodes caraccioli major G. M. Allen, 1908 

Specimens Examined— BELIZE. Toledo: Agua- 
cate, 1 (5 (CM); Big Fall, 1 .9 km ENE Rio Grande 
Bridge, 1 S (cm), 1 6 (fmnh); Big Fall, 2.1 km E 
Rio Grande Bridge, 1 S (bm); Crique Negro, Co- 
lumbia Forest, 1 S (bm), 1 5, 1 9 (msu); Salamanca 
Camp, 1 S (usnm); 1.6 km N Salamanca Camp, 
Columbia Forest, 1 S (fmnh); 2. 1 km NNE Sala- 
manca Camp, Columbia Forest, 4 66, I 9 (cm); San 
Antonio, 1 9 (fmnh). 

The published distribution of Vampyrodes car- 
accioli major northwestward of Costa Rica and 
Panama is confined to the Gulf-Caribbean low- 
lands as far as southern Mexico (Oaxaca, southern 
Veracruz, Chiapas); elevational data are less than 
300 m. The records from Belize extend the range 
of this stenodermatine north of Izabal (Sanborn, 
1936). 

The Belizean localities represent habitats of ri- 
parian lowland and upland evergreen seasonal for- 
ests and village secondary vegetation. The capture 
dates cover both the dry and wet seasons (March, 
April, May, July-September, December). 

I follow Carter and Dolan (1978) for the correct 
spelling of Vampyrodes caraccioli. 



Vampyressa pusilla thyone Thomas, 1 909 

Specimens Examined— BELIZE. Cayo: 1.6 km 

NW Augustine, 3 66, I 9 (cm); Banana Bank, 1 9 
(fmnh); Blancaneaux, 8.3 km NNE Augustine, 1 
9 (fsm). Toledo: vicinity Aguacate, 1 9 (bm), 3 99 
(cm); 1.2 km E Aguacate, 1 3, 1 9 (cm); Big Fall, 
1 km E Rio Grande Bridge, 1 9 (cm); Big Fall, 2. 1 
km E Rio Grande Bridge, 1 6 (cm); Big Fall, 1 .9 
km ENE Rio Grande Bridge, 1 5, 1 9 (cm), 1 9 
(fmnh); Crique Negro, Columbia Forest, 1 6 (msu), 
1 6 (usnm); Forest Home, 1 6 (msu); Pueblo Viejo, 
1 9 (fmnh); 1 .6 km NNE Salamanca Camp, Co- 
lumbia Forest, 1 3, 2 99 (fmnh). 

The general distribution of the little yellow-eared 
bat extends from southern (Oaxaca, southern Ve- 
racruz, Chiapas) and peninsular (Campeche) Mex- 
ico and continues southeastward along the Carib- 
bean slope to both the Pacific and Caribbean 
corridors of southern Nicaragua, Costa Rica, and 
Panama, into South America. Elevational data are 
primarily lowland, from sea level up to a recorded 
2200 m. Peterson (1966) reported the only record 
of Vampyressa pusilla in Belize, from Rockstone 
Pond, Belize District. There are also previous rec- 
ords from El Peten (Rick, 1 968) and southeastern 



Campeche (Jones et al., 1973). This account pro- 
vides additional records of this species. 

These specimens of Vampyressa pusilla were 
collected (February-May, July-September, De- 
cember) in moist habitats, the majority of which 
were associated directly with riparian vegetation 
or in village and pasture-edge vegetation situated 
near rivers. Evergreen seasonal forest provided an 
upland habitat. 



Chiroderma villosum jesupi J. A. Allen, 1900 

Specimens Examined— BELIZE. Cayo: Banana 

Bank, 1 3, 5 99 (fmnh). Corozal: Chan Chen, 1 6 
(fmnh). Toledo: Big Fall, vicinity Rio Grande 
Bridge, 1 6 (fmnh); Big Fall, 1 .7 km NE Rio Grande 
Bridge, 1 9 (msu); Big Fall, 1 .9 km ENE Rio Grande 
Bridge, 1 3, 1 9 (cm); San Antonio, 1 6 (fmnh); 1 
km WNW San Pedro Columbia, 1 9 (fmnh). GUA- 
TEMALA. EI Peten: Parque Nacional Tikal, 1 6 
(fmnh). 

The Middle American occurrence of Chiroder- 
ma villosum has been documented in southern 
(Oaxaca, southern Veracruz, Chiapas) and pen- 
insular (Campeche, Quintana Roo) Mexico, Gua- 
temala, Nicaragua, Costa Rica, and Panama. Hel- 
lebuyck et al. (1985) recently reported this fruit 
bat from El Salvador. Locality records reach from 
the coastal lowlands to upland habitats at 1 300 m. 
Southeastern Campeche (Jones et al., 1973) and 
northern Quintana Roo (Bimey et al., 1974) are 
previous Caribbean lowland localities, in addition 
to these first records from Belize and El Peten. 

All but one of the Belizean Chiroderma were 
associated either directly with or in the vicinity of 
riparian evergreen or semi-evergreen seasonal for- 
ests (April, May, August, September, December). 
One individual was captured (15 November) in 
village orchard vegetation. Five additional indi- 
viduals were released at Banana Bank. The Tikal 
specimen was captured (24 March) along the per- 
manent water pool mentioned in the Tonatia ev- 
otis account. 



Artibeus toltecus toltecus (Saussure, 1 860) 

Specimens Examined— BELIZE. Cayo: vicinity 
Augustine, 2 66, 4 99 (fsm); 1 .6 km NW Augustine, 
Rio Frio, 1 3, 1 9 (fmnh), 5 66 (ttu), 4 66 (cm); 
"Rio On," ? km N Augustine, 1 9 (ttu); 1.1 km 
S Baldy Beacon, Bald Hills, 3 99 (cm); vicinity San 
Luis, 7.1 km SSW Augustine, 1 9 (ttu). Toledo: 



148 



FIELDIANA: ZOOLOGY 



Orange Point, 1 2 (fmnh); Pueblo Viejo, 3 9$ 
(fmnh); Union Camp, 5 S6, 4 92 (cm). 

In his revision of the small Artibeus of Middle 
America, Davis (1969) recognized the range of 
Artibeus toltecus toltecus from southern Tamau- 
lipas, Mexico, southeastward along the mountain- 
ous region of the Gulf versant, upland of southern 
Mexico, Guatemala, Honduras, Nicaragua, and 
Costa Rica. He did not examine Panamanian spec- 
imens. Handley (1966) summarized the Pana- 
manian localities for /I. toltecus. This bat primarily 
occurs at elevations between 328 and 1640 m, 
although elevations near sea level were recorded 
(Davis, 1969). Consequently, the occurrence of ^. 
toltecus in the Maya Mountain range of southern 
Belize and southeastern El Peten was not unex- 
pected. These Belizean localities represent the first 
northern Caribbean lowland records. 

The Belizean localities range in elevation from 
near sea level to approximately 720 m. Artibeus 
toltecus is more common at the higher elevations. 
These dark-colored Artibeus were captured (De- 
cember-February, April, June, September) in hab- 
itats of deciduous seasonal forest, semi-evergreen 
seasonal forest, transitional forest, and pine forest- 
savanna. 

The subspecies toltecus is applied, based on the 
proximity of Belize to its distribution as defined 
by Davis (1969). 



Centurio senex senex Gray, 1 842 

Specimens Examined— BELIZE. Belize: 1 .4 km 
S San Pedro, Ambergris Caye, 1 3, 1 9 (fmnh). 
Cayo: 1.6 km NW Augustine, Rio Frio, 1 <5 (ttu); 
vicinity Augustine, Rio On, 1 9 (ttu); Blanca- 
neaux, 8.3 km NNE Augustine, 1 9 (fsm); Central 
Farm, 1 3, 1 9 (fmnh); Teakettle, Young Gal Road 
at Belize River, 1 3, 1 9 (fmnh); Xunantunich, 1 
$ (fmnh). Corozal: 1.2 km E, 1.6 km N Corozal, 
1 (5 (LSUMZ). Orange Walk: 1.6 km NW San An- 
tonio, Rio Hondo, 1 9 (fmnh). Toledo: Big Fall, 
1 .9 km ENE Rio Grande Bridge, 1 S (cm); Crique 
Negro, Columbia Forest, 1 5, 1 9 (usnm); Forest 
Home, 1 9 (amnh); vicinity Union Camp, 2 99 
(BM), 1 9 (CM). GUATEMALA. Alta Verapaz: Lan- 
quin, vicinity Lanquin Cave, approx. 149 km WSW 
Puerto Barrios, 1 $ (amnh). Izabal: 25 km SSW 
Puerto Barrios, 1 3, 5 99 (tcwc). 

The recorded distribution of the wrinkle-faced 
bat extends from western (southern Sinaloa), 
northeastern (southern Tamaulipas), and penin- 
sular (Campeche and Quintana Roo) Mexico and 



continues southeastward through Central America 
at principally lower to upland elevations (sea level 
to 1882 m). The records given here are the first 
for Belize, Alta Verapaz, and Izabal. 

The distribution of this unusual bat in Belize 
reflects apparent ecological flexibility. Centurio se- 
nex has been captured in low littoral forest and 
mangrove swamp edge on the coastal sand strip 
of Ambergris Caye, to about 720 m in evergreen 
and semi-evergreen seasonal forest on the south- 
ern slope of the Maya Mountains. Evergreen sea- 
sonal and transitional forests, secondary forest, and 
agriculturally disturbed areas provide additional 
habitats. This bat was captured throughout the 
year. Two males and one female were mist netted 
and released at Orange Point, Toledo District. 
Brother N. Sullivan collected (15-17 January) the 
specimen from Alta Verapaz, but I assume the bat 
was captured outside of Lanquin Cave. The spec- 
imens from Izabal were obtained (February, 
March) by D. C. Carter and field party. Field data 
are limited, but four Centurio were captured over 
a stream. 



Diphylla ecaudata Spix, 1823 

Specimens Examined— BELIZE. Cayo: vicinity 
Augustine, 1 S (rom); San Antonio, 1 6 (fmnh). 
Toledo: Crique Jute, 1 $ (amnh); San Antonio, 1 
9 (fmnh); Santa Elena, 1 9 (fmnh). 

The distribution of Diphylla ecaudata appears 
primarily restricted along the Gulf side and in the 
Yucatan Peninsula of Mexico southeastward 
throughout Central America, where this bat occurs 
from the coastal lowlands up into the mountainous 
highlands (1880 m). The hairy-legged vampire bat 
has been recorded from El Peten (McCarthy, 1 982) 
and Quintana Roo (Jones et al., 1973). The spec- 
imens reported here are the first records from Be- 
lize. 

Four of the localities represent village environ- 
ments where Diphylla was captured (April, July, 
August, December) along with Desmodus rotundus 
during vampire bat control activities. Mist netting 
was carried out in direct immediacy to domestic 
livestock and homes. The feeding activities of £>/- 
phylla in these villages were not documented, al- 
though one blood meal was obtained for analysis. 
P. Boreham, Imperial College Field Station, En- 
gland, reported (in litt.) a weak precipitin reaction 
for a mammal host from the blood meal sample 
without a response for bird or reptile. It is not 
known if this blood meal was obtained in the vil- 



McCARTHY: DISTRIBUTION OF BATS 



149 



lage (Santa Elena). Gardner (1977) summarized 
the sanguivorous preference ofDiphylla as for pri- 
marily avian hosts. The hairy-legged vampire from 
Augustine was apparently taken (22 February) in 
a deciduous seasonal forest. 



Family NATALIDAE 

Natalus stramineus saturatus 

Dalquestand Hall, 1949 

Specimens Examined— BELIZE. Cayo: 1.6 km 

NW Augustine, Rio Frio, 2S6,2 9i (fsm); 0.8 km 
W Augustine, 2 66, 3 22 (cm); 1.5 km N Augustine, 
5 22 (cm); Sibun Camp, Hummingbird Hwy. at 
Silver Creek, 1 2 (fmnh). Orange Walk: Richmond 
Hill (Goat Hill), 8.9 km SSW Orange Walk Town, 
1 2 (cm). Stann Creek: Kendal, 1 6 (fmnh). Toledo: 
vicinity Aguacate, 1 3, 3 22 (cm); 1.2 km E Agua- 
cate, 1 2 (cm); Vista Hermosa, 3.7 km WNW Punta 
Gorda, 8 66, 6 22 (fmnh). 

The northern range of Natalus stramineus sa- 
turatus extends from both northwestern (Sinaloa) 
and northeastern (Nuevo Leon) Mexico, including 
the Yucatan Peninsula, southeastward through 
Central America where the number of records for 
this species is noticeably reduced beyond Guate- 
mala to Panama. Although predominately a low- 
land species, elevations were recorded as high as 
2400 m. The presence of the funnel-eared bat in 
Belize was anticipated, as it appears to be well 
reported throughout the Gulf-Caribbean versant. 

Those specimens obtained (April, August, Sep- 
tember) at roost sites in Belize were from caves. 
Other capture localities include low riparian forest 
and open areas bordering on forest, in orchard 
habitats, and alongside a building. 



Family VESPERTILIONIDAE 

Subfamily VESPERTILIONINAE 

Myotis elegans Hall, 1962 

Specimens Examined— BELIZE. Belize: Belize 
City, Landivar, 1 2 (amnh), 1 3, 1 2 (fmnh), 1 6 
(msu); Mussel Creek, 7.5 km W Burrell Boom, 1 

6, 1 2 (fmnh). 

LaVal (1973a) summarized the lowland distri- 
bution of Myotis elegans. ranging from the Gulf 
(eastern San Luis Potosi, Veracruz), Pacific coastal 
(Chiapas), and peninsular (southeastern Cam- 



peche) regions of Mexico to Honduras, Nicaragua, 
and northeastern Costa Rica. Subsequent records 
were reported from the Pacific side of Costa Rica 
and the Caribbean lowlands of El Peten (LaVal, 
1977; McCarthy, 1982). The majority of eleva- 
tions are less than 1 20 m, ranging to 750 m. These 
additional Caribbean lowland localities are the first 
records from Belize. 

Two elegant Myotis were netted (1 July) along 
a tractor track, in low riparian vegetation domi- 
nated by bamboo and thistle palms. Four indi- 
viduals were obtained (January, February, May, 
December) at a coastal locality in low vegetation 
bordering on disturbed mangrove (Rhizophora 
mangle, Avicennia germinans) habitat. 



Eptesicus furinalis gaumeri 

(J. A. Allen, 1897) 

Specimens Examined— BELIZE. Belize: Belize 
City, Landivar, 1 2 (cm). Cayo: Central Farm, 2 
22 (cm), 5 66, 16 22 (fmnh), 2 66,19 (TTu); Little 
Vaquero Creek, 9.3 km NNW Augustine, 1 5, 1 2 
(fsm); Ontario, 5.5 km W Teakettle, 1 2 (fmnh); 
Teakettle, 1 6 (fmnh). Corozal: Estero Lagoon, 4 
km W Patchakan, 1 3, 1 2 (fmnh); Santa Clara, 1 
2 (fmnh). Orange Walk: Honey Camp Lagoon, 1 
6, 2 22 (fmnh); Tower Hill, B.S.I, compound, 3 66 
(CM), 1 5, 4 22 (fmnh); 2 km SSW Tower Hill 
Bridge, 1 2 (cm). Stann Creek: Melinda, 3 22 (fmnh); 
Dangriga (Stann Creek), 1 <5 (usnm). Toledo: Or- 
ange Creek, 1.5 km S Punta Gorda, 1 2 (msu); 
Punta Gorda, 1 2 (msu). 

The Mexican distribution of Eptesicus furinalis 
gaumeri ranges from the western (Jalisco) and the 
eastern (San Luis Potosi) versants southeastward 
to South America. Davis (1965), Disney (1968), 
and Starrett and Casebeer (1968) reported records 
from all of the Central American countries except 
El Salvador. Lowland elevations range from near 
sea level to 1800 m, the majority being below 500 
m. This tropical brown bat has been reported from 
El Peten (Rick, 1968; McCarthy, 1982) and Quin- 
tana Roo (Jones et al., 1973). The localities here 
are additional records for Belize. 

Disney ( 1 968) did not present locality data for 
his two specimens of Eptesicus furinalis. Both were 
males, captured (16 November, 29 December) in 
Cayo District, near Central Farm and Esperanza 
(4.5 km W Central Farm). These are located in 
British Museum (Natural History). An additional 
1 96 individuals were captured from three of the 
localities reported here; the majority of these were 



ISO 



FIELDIANA: ZOOLOGY 



banded and released during a behavioral study. 
The majority was found in direct association with 
buildings, utilizing the infrastructure of the walls 
or floors and the space behind window shutters as 
roost sites. Individuals have been taken over water 
(creeks and a swimming pool) at three localities 
and in riparian vegetation along two lagoons. 



Lasiurus borealis (Muller, 1776) 

Speomens ExAMnsfED— BELIZE. Orange Walk: 
Tower Hill, B.S.I, compound, 1 9 (fmnh). Stann 
Creek: 5.3 km NNW Quam Bank, Cockscomb 
Basin, 1 9 (cm). GUATEMALA. El Peten: Parque 
Nacional Tikal, 1 9 (fmnh). 

The subspecies teliotis ranges southward from 
both the western and eastern regions of Mexico to 
Oaxaca and the northern Yucatan Peninsula. 
Specimens of Lasiurus borealis from the Guate- 
malan central highlands were assigned by Jones 
(1966) to the Central American subspeciesyra«/z/7, 
based on Handley (1960). Carter et al. (1966) as- 
signed specimens from both lowland and highland 
localities in Chiapas Xofrantzii, suggesting that the 
region of the Isthmus of Tehuantepec represents 
the break heXwecn front zii and teliotis. Hall (1981) 
concurred with this arrangement. Similarly, Jones 
et al. (1973) suggested that southern Mexico, in- 
cluding the Yucatan Peninsula, may represent a 
zone of intergradation between frantzii and teli- 
otis. Few specimens of L. borealis are available 
from El Salvador (Burt & Stirton, 1961), Honduras 
(Goodwin, 1942b), Nicaragua (Davis & Carter, 
1962— as L. b. teliotis), Costa Rica (Goodwin, 
1 946; Gardner et al., 1 970) and Panama (Handley, 
1 966). Recorded elevations (near sea level to about 
2540 m) are primarily low or moderate (< 1 155 
m). Koopman (1959) reported the only record from 
Quintana Roo. This account represents the first 
records for Belize and eastern Guatemala from El 
Peten. 

The red bats captured in Belize (April, May) 
were netted over a stream and a swimming pool. 
The Tikal specimen was taken (30 July) while it 
was flying in an open area near a large man-made 
reservoir. 

I hesitate to assign a subspecific designation be- 
cause I see no practical purpose in doing so until 
adequate series of specimens from throughout the 
range of Lasiurus borealis become available. Han- 
dley (1960) had fewer specimens of L. borealis at 
hand for a proper evaluation of subspecific vari- 



ation. Consequently, the limits of the distributions 
for the recognized subspecies remain unresolved. 



Lasiurus ega (Gervais, 1855) 

Specimens Examined— BELIZE. Belize: Trop- 
ical Park, Mi. 14.5 Western Hwy., 1 S (fmnh). 
Orange Walk: Tower Hill, B.S.I, compound, 2 6i 
(FMNH), 1 (3 (cm). Stann Creek: 5.3 km WNW Quam 
Bank, Cockscomb Basin, 2 66, I 9 (cm). Toledo: 
Big Fall, 1.7 km NE Rio Grande Bridge, 1 9 (cm); 
Orange Creek, 1 .5 km SW Punta Gorda, 1 6 (msu). 

Similar to Lasiurus borealis, the distribution for 
the two recognized subspecies of the yellowish bat 
is not well understood. While L. e. panamensis 
was recognized along the Pacific versant of Chia- 
pas (Baker &. Patton, 1 967) and Guatemala (Dolan 
& Carter, 1979;Dickermanetal., 1981), Goodwin 
( 1 969) identified panamensis from the moderate 
elevations of the Gulf drainage in northern Oaxaca 
and suspected L. e. xanthinus may occur in the 
drier Pacific portion of that state. Baker et al. ( 1 97 1 ) 
determined the variation in karyotypes and pelage 
color of L. ega from near Brownsville, Texas, re- 
sembled those from eastern coastal and southern 
Mexico and referred the Texas specimens to L. e. 
panamensis. Meanwhile, L. e. xanthinus was rec- 
ognized in the Yucatan Peninsula (Jones et al., 
1973; Bimey et al., 1 974). The yellow bat is poorly 
represented from the remainder of Central Amer- 
ica, which includes Honduras (Goodwin, 1942b; 
LaVal, 1969; Greenbaum &. Jones, 1978), Costa 
Rica (Goodwin, 1946; Starrett & Casebeer, 1968; 
Gardner et al., 1970; LaVal & Fitch, 1977), and 
Panama (Handley, 1966). Where designated, the 
subspecies panamensis has been applied to these 
preceding Central American localities, although 
Hall (1981) did not acknowledge panamensis north 
of Costa Rica. Elevational data are similar to those 
for L. borealis. Ingles (1958) reported two L. ega 
from Quintana Roo. Jones et al. (1973, p. 23) 
translated Ingles's locality from Spanish as "Puer- 
to Morelos" when it was actually a collection site 
only 16 km east of the state border with Yucatan, 
along the highway from Valladolid (Yucatan) to 
Puerto Morelos (Quintana Roo). Alvarez and Ra- 
mirez-P. (1972) cited an additional Caribbean 
lowland record from southeastern Campeche. This 
account provides the first L. ega records from Be- 
lize. 

Eight yellowish bats were captured (April, May) 
over streams, a river, and a swimming pool. 
Another was netted ( 1 8 August) at about 5 m above 



MCCARTHY: DISTRIBUTION OF BATS 



151 



the ground while circhng a building located in grass- 
sedge savanna. 



Lasiurus intermedius intermedius 

(H. Allen, 1862) 

Specimen Examined— BELIZE. Toledo: Crique 
Jute, 1 9 (cm). 

The range of this subspecies of the large yellow 
bat extends southeastward from Mexico to Hon- 
duras (Handley, 1960; Carter et al., 1966), El Sal- 
vador (Hellebuyck et al., 1985), and Guatemala 
(Carter et al., 1 966). Lasiurus intermedius has been 
recorded in Mexico from the northern Yucatan 
Peninsula and Chiapas northwestward to Texas 
along the eastern coast and to Sinaloa on the Pa- 
cific side. Recorded elevations range from lowland 
to highland (1620 m) habitats. A single specimen 
of L. intermedius from northern Quintana Roo 
(Bimey et al., 1974) provided the only record for 
that Mexican state. This Belizean specimen rep- 
resents the first record for the country. 

The above specimen was obtained on 30 March 
over the stream Crique Jute surrounded by sec- 
ondary vegetation. 



Bauerus dubiaquercus (Van Gelder, 1959) 

Specimens Examined— BELIZE. Cayo: 1.6 km 

NW Augustine, Rio Frio, 1 2 (rom). Toledo: 2. 1 
km NNE Salamanca Camp, Columbia Forest, 1 6 

(CM). 

The published localities of the rarely encoun- 
tered Bauerus dubiaquercus are scattered from the 
Islas Tres Marias (Nayarit), Jalisco, and southern 
Veracruz in Mexico to eastern Honduras and Cos- 
ta Rica (Engstrom & Wilson, 1981; Dinerstein, 
1985). Mainland elevations range from approxi- 
mately 460 to 1450 m and appear to represent 
mid-elevation and montane forest habitats (Pine, 
1 966; Pine et al., 1 97 1 ; Engstrom & Wilson, 1981; 
Dinerstein, 1985). These first occurrences of Bau- 
erus in Belize extend northward a scattered dis- 
tribution along the northern Caribbean lowlands 
in Central America. 

J. Kamslra and J. Fragoso collected (8 July) one 
specimen inside the main Rio Frio cave, located 
in a deciduous seasonal forest at approximately 
410m. The second Bauerus was netted (26 March) 
along an open forestry track in an evergreen forest 
at about 180 m. 



Engstrom and Wilson (1981) and Martin and 
Schmidly (1982) evaluated the taxonomic status 
of Antrozous (Bauerus) dubiaquercus and con- 
cluded the chromosomal, cranial, postcranial, and 
phallic differences between this bat and Antrozous 
(Antrozous) pallidus were sufficient to recognize 
Bauerus as a distinct genus. I follow their conclu- 
sions and agree that the species is monotypic since 
the mainland sample size that previously was as- 
signed to A. d. meyeri Pine, 1971, was limited to 
a total of five specimens representing both sexes. 



Family MOLOSSIDAE 

Eumops auripendulus auripendulus (Shaw, 1 800) 

Specimens Examined— BELIZE. Orange Walk: 

Orange Walk Town, 1 ? (cm); Tower Hill, B.S.I, 
compound, 1 9 (fmnh). 

The recorded distribution of Eumops auripen- 
dulus auripendulus includes both moist uplands 
and drier lowland coastal and plateau areas, rang- 
ing from eastern Oaxaca, Tabasco, Quintana Roo, 
and Belize, through Guatemala, western Hondu- 
ras, El Salvador, western Nicaragua, Costa Rica, 
and Panama, into South America (Eger, 1974; 
Greenbaum & Jones, 1978). Villa-R. (1956) and 
Eger (1974), respectively, reported this free-tailed 
bat from Quintana Roo and Belize (Belize District: 
Rockstone Pond). This account provides the sec- 
ond and third records for Belize. 

The Orange Walk specimen consists of a man- 
dible and partial skull, which were recovered from 
an owl {Tyto alba) roost in a church tower. The 
second specimen was discovered (July) alive by L. 
G. Hoevers, after it apparently was attacked by a 
bird. 



Eumops bonariensis nanus (Miller, 1900) 

Specimens Examined— BELIZE. Orange Walk: 

Orange Walk Town, 2 ?? (cm). 

Eger (1977) summarized the few available Mid- 
dle American localities for this small mastiff bat, 
which are limited to southeastern Mexico (south- 
em Veracruz, Tabasco, Yucatan), eastern Hon- 
duras, and Panama. These and additional locali- 
ties in Panama (Dolan & Carter, 1979) and 
Nicaragua (Hall, 1981) are restricted to coastal 
lowland environments. This is the first recording 
of Eumops bonariensis for Belize. 



152 



HELDIANA: ZOOLOGY 



Entire specimens oi Eumops bonariensis as yet 
are unavailable from Belize. Documentation is 
based on two fragmented sets of maxillary tooth- 
rows, which were sifted from regurgitated rubble 
beneath an owl {Tyto alba) roost in a church tower. 



Moiossus ater nigricans (Miller, 1 902) 

Specimens Examined— GUATEMALA. Izabal: 

25 km SSW Puerto Barrios, 2 33, 4 99 (tcwc). 

The black mastiff bat is a common inhabitant 
of roof spaces throughout its lowland Middle 
American range, from western (Sinaloa) and east- 
em (Tamaulipas) Mexico southeastward into South 
America. This species has been reported from the 
Caribbean lowlands of Quintana Roo (Jones et al., 
1973) and Belize (Murie, 1935; Pendergast, 1979). 
These Moiossus ater from Izabal are the first rec- 
ord for that department. Apparently, all of these 
specimens were collected on 1 5 February over a 
stream by D. C. Carter and his field party. 



Addendum 

While this paper was in press, other papers and 
additional information concerning bats in Belize 
came to my attention. Two recent papers provide 
new country records of the glossophagines Li- 
chonycteris obscura (Hill, 1985) and Hylonycteris 
underwoodi (McCarthy & Blake, 1987), which 
increase the known bat fauna to 68 species. Both 
records are from Toledo District. McCarthy and 
Blake (1987) also reported the occurrence of the 
following bats: Rhynchonycteris naso, Saccopteryx 
bilineata, Balantiopteryx io, Noctilio leporinus, 
Pteronotus parnellii, Micronyctehs megalotis, M. 
nicefori, M. schmidtorum, Tonatia evotis, Mimon 
cozumelae, Phyllostomus discolor, Trachops cir- 
rhosus, Chrotopterus auritus, Vampyrum spec- 
trum, Glossophaga soricina, Carollia brevicauda, 
C. perspicillata, Sturnira lilium, Uroderma bilo- 
batum, Vampyressa pusilla, Artibeus jamaicensis, 
A. lituratus, A. phaeotis, A. watsoni, Centurio se- 
nex, Natalus stramineus, Rhogeessa tumida, Bau- 
erus dubiaquercus, and Eumops underwoodi. 

Hill, J. E. 1 985. The status of Lichonycteris degener 
Miller, 1931 (Chiroptera: Phyllostomidae). Mam- 
malia, 49(4): 579-582. 

McCarthy, T. J., and M. Blake. 1 987. Noteworthy 



bat records from the Maya Mountains Forest Re- 
serve, Belize. Mammalia, 51(1): 161-164. 

A bat specimen from Stann Creek was reported 
(Miller & Allen, 1928, p. 180) as Myotis nigricans 
nigricans. I examined this specimen at USNM and 
found it to be an immature Eptesicus furinalis. It 
is included in this paper under the species account 
for the latter species. 

Miller, G. S., Jr., and G. M. Allen. 1928. The 
American bats of the genera Myotis and Pizonyx. 
Bulletin of the United States National Museum, 144: 
1-218. 

Silva-Taboada and Koopman (1964, p. 3) re- 
ported specimens of Tadarida laticaudata (= Nyc- 
tinomops laticaudatus) from Corozal District. Most 
of the bat species discussed in an unpublished dis- 
sertation by A. M. Cartwright were also reported 
by Kirkpatrick et al. (1975) and Cartwright and 
Kirkpatrick (1977). The remaining identifications 
(Cartwright, 1977, pp. 240, 242-246, 250, 251), 
which were from Belize District, included Rhyn- 
chonycteris naso, Saccopteryx bilineata, Carollia 
brevicauda, C. perspicillata, Sturnira lilium, Arti- 
beus lituratus, A. phaeotis, Des modus rotundus, 
Eptesicus furinalis, Rhogeessa tumida, and Mo- 
iossus moiossus. Baker et al. (1985, p. 236) re- 
ported cytogenetic data from specimens of Rho- 
geessa tumida that I collected in Belize District. 

Baker, R. J., J. W. Bickham, and M. L. Arnold. 
1985. Chromosomal evolution in Rhogeessa (Chi- 
roptera: Vespertilionidae): Possible speciation by 
centric fusions. Evolution, 39(2): 233-243. 

Cartwright, A. M. F. 1977. Patterns of Neotrop- 
ical chiropteran reproduction including histological 
and ecological aspects of bats collected in Belize. 
Ed.D. diss.. Ball State University, Muncie, Ind., 278 
pp. 

Silva-Taboada, G., and K. F. Koopman. 1964. 
Notes on the occurrence and ecology of Tadarida 
laticaudata yucatanica in eastern Cuba. American 
Museum Novitates, 2175: 1-6. 

A mammal checklist was included in a resource 
paper on Belize (Hartshorn et al., 1984). The list 
of bats supposedly was a compilation of known 
and expected species. The result is inaccurate and 
undocumented. The reader is referred to the 
checklist in the Appendix as correct. 



Hartshorn, G., et al. 1984. Belize. Country En- 
vironmental Profile. A Field Study. United States 



MCCARTHY: DISTRIBUTION OF BATS 



153 



Agency for International I>evelopment, San Jose, 
Costa Rica. 151 pp. 



Additional specimens of Micronycteris mega- 
lotis (2: Belize District, Cayo District), Artibeiis 
toltecus (11: Cayo District), Centurio senex (2: Cayo 
District), and Diphylla ecaudata (3: Cayo District) 
were found in the mammal collection of Royal 
Ontario Museum. D. J. Tallman collected speci- 
mens of M. megalotis ( 1 ) and Mimon cozumelae 
(2) from Orange Walk District, which were de- 
posited in Bell Museum of Natural History, Uni- 
versity of Minnesota. I secured further voucher 
specimens (AMNH) of Mimon cozumelae (1), 
Vampyrodes caraccioli ( 1 ), Vampyressa pusilla ( 1 ), 
and Bauerus dubiaquercus (1) from Toledo Dis- 
trict. 

Certain specimens (Sturnira lilium and Rho- 
geessa tumidd) that were catalogued by Dobson 
(1878, pp. 540, 246) were listed as collected in 
"Honduras." These were obtained by D. Dyson 
and H. Cuming between November 1844 and late 
1845. During that time, "Honduras" correspond- 
ed to the present region that extends from southern 
Quintana Roo, Mexico, southeastward to northern 
Honduras. Many early collectors did not differ- 
entiate between the area of Belize ("British settle- 
ment in Honduras") and that of the Republic of 
Honduras ("Spanish Honduras"), but recorded 
only "Honduras" or "Bay of Honduras" without 
further locality data. The above specimens did not 
originate from present day Honduras, but were 
collected in Belize. Additional specimens of Mi- 
cronycteris megalotis (Dobson, 1 878, p. 479) from 
the "Bay of Honduras" and Rhynchonycteris naso 
(Dobson, 1878, p. 368) from "Honduras" remain 
orphaned records of the historical literature. 

Uroderma bilobatum was reported (Sanchez-H. 
et al., 1986) from southern Quintana Roo while 
this volume was delayed. Ten specimens were col- 
lected at Ruinas de Kohunlich (18°23'N; 88''42'W), 
about 16 km W Estevez on the Belizean border. 
Four other species {Pteronotus davyi, Mormoops 
megalophylla, Tonatia evotis, T. minuta) were 
documented for the first time from Quintana Roo, 
from localities within 35 km of the northern bor- 
der of Belize. The known bat fauna of Quintana 
Roo is now represented by 36 species. 



Sanchez-H., O., G. Tell£z-G., R. A. Medelun, and 
G. Urbano-V. 1986. New records of mammals 
from Quintana Roo. Mexico. Mammalia, 50(2): 275- 
278. 



Acknowledgments 

Grateful acknowledgments are due H. Flowers, 
O. Rosado, R. Belisle, and E. O. Bradley, E>e- 
partment of Forestry, Belize, and H. Topsey, W. 
Branche, E. A. Graham, and J. Palacio, Depart- 
ment of Archaeology, Belize, for the necessary per- 
mission to conduct fieldwork. The Department of 
Agriculture, Belize, provided opportunities and 
support while I headed the Vampire Bat Education 
and Control Program (1975-1978). Division of 
Mammals, Field Museum of Natural History, pro- 
vided partial equipment needs (1974) and partial 
funding ( 1979). Mellon North American Mammal 
Research Institute, O'Neil Fund, and Section of 
Mammals, Carnegie Museum of Natural History, 
supported further fieldwork (1982, 1984). Nu- 
merous colleagues and friends have been of direct 
assistance to my needs in Belize, and the following 
are but a few: E. Awe, P. Boreham, H. Bums, W. 
D. Burton, J. Cab, G. Cal, G. C. Canto, J. Chun, 
M. Craig, L. de la Torre, L. Dieckman, R. Foster, 
P. W. Freeman, P. Gamble, E. Garcia, J. P. and 
M. Garcia, H. H. Genoways, E. Gomez, L. Hen- 
derson, R. W. Henderson, L. G. and E. Hoevers, 
A. M. Hutson, R. J. Izor, T. Juring, E. King, K. 
Leacock, D. Owen-Lewis, N. MacKenzie, G. T. 
McCarthy, H. Pastor, R. H. Pine, A. Rabinowitz, 
M. L. Reed, C. J. Rushin, K. and T. Salisbury, R. 
Schmidt, B. M. Silva, M. F. Valentine, L. Waight, 
L. Wilkins, and A. C. S. Wright. Various regiments 
and the Royal Air Force of Her Majesty's British 
Forces provided certain logistical support over the 
years. 

My fieldwork in Parque Nacional Tikal, El Pe- 
ten, Guatemala, was made possible while assisting 
D. J. Howell and J. G. Cant during their respective 
visits to the park. M. Dary-R., Universidad de 
San Carlos, and F. Polo-Sifontes, Instituto de An- 
tropologia y Historia, graciously assisted and per- 
mitted my work in 1979. J. R. Martinez R. kindly 
provided a copy of his thesis. 

The courtesies rendered to me by the personnel 
of the following institutions while providing either 
information, the loan of specimens, or assistance 
during visits are greatly appreciated: American 
Museum of Natural History; British Museum 
(Natural History); Carnegie Museum of Natural 
History; Field Museum of Natural History; Flor- 
ida State Museum, University of Florida; Museum 
of Natural History, University of Kansas; Mu- 
seum of Zoology, Louisiana State University; The 
Museum, Michigan State University; Royal On- 
tario Museum; Texas Cooperative Wildlife Col- 



154 



HELDIANA: ZOOLOGY 



lection, Texas A «& M University; The Museum, 
Texas Tech University; United States National 
Museum of Natural History. I am grateful to N. 
A. Bitar, F. J. Bonaccorso, D. C. Carter, M. D. 
Engstrom, A. M. Hutson, J. Kamstra, R. L. Pe- 
terson, and E. L. Tyson for sharing information 
concerning their field efforts in either Belize, El 
Peten, Izabal, or Quintana Roo. 

Besides J. R. Choate, J. K. Jones, Jr., K. F. 
Koopman, and R. M. Timm, two anonymous re- 
viewers enhanced earlier manuscripts with their 
comments and editorial skills. E. Mendez kindly 
reviewed my Spanish abstract. Verification of the 
state boundaries in the Yucatan Peninsula was 
provided by T. Dachtera, National Geographic 
Society, and G. de la Torre M., Secretaria de Pro- 
gramacion y Presupuesto, Mexico. M. A. Schmidt, 
R. A. Rollin, V. Risoli, and S. Cramer typed var- 
ious versions of the manuscript. S. M. Velez guid- 
ed the manuscript during my absence. This report 
is a contribution of the Mammals of Belize Pro- 
gram. 



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Jones, J. K., Jr., P. Swanepoel, and D. C. Carter. 
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KooPMAN, K. F. 1959. The zoogeographical limits of 
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Lackey, J. A. 1970. Distributional records of bats from 
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Martin, C. O., AND D. J. ScHMiDLY. 1982. Taxonomic 
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Martinez R., J. R. 1980. Deteccion de virus rabico 
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McCarthy, T. J. 1982. Bat records from the Carib- 
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MCCARTHY: DISTRIBUTION OF BATS 



157 



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Gazetteer 

The numbers for localities are plotted in Figure 1 . 



MEXICO 

QUINTANA ROO 



GUATEMALA 

El PETfeN 



1. Bacalar 18°43'N; 88°22'W 



2. Parque Nacional Tikal 1 7°20'N; 89°39'W 

3. Poptun 16°2rN; 89''26'W 



158 



FIELDIANA: ZOOLOGY 




Fig. 1. Distribution of localities in the districts of Belize; Quintana Roo, Mexico; and El Peten and Izabal, 
Guatemala. The numbers refer to those listed in the Gazetteer. This map does not display the Caribbean lowlands 
of Guatemala and Mexico in their entirety. 



MCCARTHY: DISTRIBUTION OF BATS 



159 



IZABAL 

4. Puerto Barrios 15°43'N; 88°36'W 

BELIZE 
CoROZAL District 

5. Chan Chen 18°26'N; 88»27'W 

6. Corozal 18°24'N; 88°24'W 

7. Patchakan 18°24'N; 88''29'W 

8. Santa Clara 18°18'N; 88°30'W 

Orange Walk District 

9. Honey Camp Lagoon 18°03'N; 88°27'W 

10. Orange Walk Town 18°05'N; 88°34'W 

11. San Antonio, Rio Hondo 18°1 1'N; 88°39'W 

12. Tower Hill, Belize Sugar Industries (B.S.I.) 

18°02'N; 88''34'W 

Belize District 

13. Belize City 17°30'N; 88°12'W 

14. Churchyard 17°18'N; 88°33'W 

15. Mussel Creek 17°39'N; 88°24'W 

16. Rockstone Pond (Altun Ha) 17°46'N; 

88°22'W 

17. San Pedro, Ambergris Caye 17°55'N; 

87°58'W 

18. Tropical Park 17°28'N; 88''23'W 



16°47'N: 



30. Teakettle 17°13'N; 88°5rw 

31. Xunantunich 17«t)5'N; 89°08'W 



Stann Creek District 

32. Kendal 16°49'N; 88°22'W 

33. Melinda 17°00'N; 88°18'W 

34. Quam Bank, Cockscomb Basin 

88°28'W 



Toledo District 

35. Aguacate 16°10'N; 89°06'W 

36. Big Fall 16''15'N; 88°53'W 

37. Blue Creek 16°12'N; 89°03'W 

38. Crique Jute and Salamanca Camp (Forestry 

Camp) 16°16'N; 89°0rw 

39. Crique Negro, Columbia Forest 16°17'N; 

89°02'W 

40. Cuevas Creek and Jacinto Creek Bridges, at 

Punta Gorda Road 16°09'N; 88°53'W 

41. Forest Home 16'^8'N; 88°50'W 

42. Nimli Punit 16°20'N; 88''48'W 

43. Orange Point 16°04'N; 88°49'W 

44. Pueblo Viejo 16°13'N; 89°09'W 

45. Punta Gorda 16°07'N; 88°48'W 

46. Rice Station (Agricultural Station) 16°08'N; 

88°51'W 

47. San Antonio 16°15'N; 88°02'W 

48. San Lucas (deserted) 16°05'N; 89°06'W 

49. San Pedro Columbia 16°17'N; 88°58'W 

50. Santa Elena 16°14'N; 89'X)6'W 

51. Union Camp 16°24'N; 89°08'W 



Cayo District 

19. Augustine 16°58'N; 88°59'W 

20. Baldy Beacon, Bald Hills 17°0rN; 88°47'W 

21. Banana Bank 17°15'N; 88''48'W 

22. Barton Creek at Western Hwy. 17°13'N; 

89''57'W 

23. Central Farm and Listowel 17°irN; 

89°00'W 

24. C.I.T.A.,Sibun River at Indian Creek 17''16'N; 

88''34'W 

25. Macaw Bank 17°05'N; 89'^4'W 

26. Roaring Creek 17°15'N; 88''47'W 

27. San Antonio 17°05'N; 89°0rW 

28. San Luis 16°54'N; 89°00'W 

29. SibunCamp 17°05'N; 88°39'W 



Appendix 

This district checklist of the bat fauna of Belize 
is based on published accounts. The citations refer 
to the initial taxonomic treatments of specimens. 
Districts are arranged from north (left) to south 
(right). Abbreviations are as follows: Cz = Coro- 
zal; OW = Orange Walk; Bz = Belize; Cy = Cayo; 
SC = Stann Creek; Td = Toledo. 

In order to give an accurate list of bats, it is 
necessary to present certain discrepancies that have 
appeared in the literature. Dobson (1878) referred 
to certain early sp>ecimens that may have origi- 
nated from Belize. One reference (Dobson, 1878, 



160 



FIELDIANA: ZOOLOGY 



p. 520) to "Half-Moon Key, Honduras" for a spec- 
imen of Artibeus perspicillatus (= jamaicensis) be- 
longs to Belize, since this specimen was collected 
by O. Salvin (see Salvin, 1864). Sanderson (1941, 
p. 228) recalled "Anoura sp." in his descriptive 
narrative of a visit to then British Honduras. This 
species was not identified (Hershkovitz, 1951) in 
the Sanderson bat collection. Diaemus youngi was 
cited from Belize (Villa-R., 1966, p. 340), but R. 
L. Peterson (pers. comm.) stated that the specimen 
in question was actually from Guyana. Specimens 
of 17 species of bats were listed in Disney (1968) 



without locality data. These specimens, which are 
housed in British Museum (Natural History) and 
Royal Ontario Museum, are all from Cayo Dis- 
trict. J. L. Eger (pers. comm.) identified the ques- 
tionable specimen of Molossus bondae in Disney 
( 1 968, p. 7) as M. molossus. Quinones et al. ( 1 978, 
p. 559) reported six species, which I collected and 
identified, without the exact locality information 
other than "the Maya Mountains region." This 
locality is 1 km NW Augustine, Cayo District. 
Sixty-six bat species are recognized in Belize. 



Species 


Cz OW 


Bz 


Cy 


sc 


Td 


References 


Rhynchonycteris naso 






X 






Murie, 1935, pp. 17-18; Disney, 1968, p. 7 


Saccopteryx bilineata 


X 


X 


X 


X 




Sanborn, 1937, p. 331; Hershkovitz, 1951, 
p. 553; Disney, 1968, p. 7; Pendergast, 
1979, p. 10 


Saccopteryx leptura 










X 


This paper 


Peropteryx kappleri 






X 






Cartwright & Kirkpatrick, 1977, p. 466 


Peropteryx macrotis 








X 




Hershkovitz, 1951, p. 553 


Centronycteris maximilliani 








X 




Sanborn, 1941, p. 372 


Balantiopteryx io 




X 


X 






Kirkpatrick et al., 1975, p. 330; Cartwright 
& Kirkpatrick, 1977, p. 466 


Diclidurus virgo 






X 






This paper 


Noctilio leporinus 


X 




X 


X 


X 


This paper 


Pteronotus davyi 


X 




X 




X 


Disney, 1968, p. 7; this paper 


Pteronotus parnellii 




X 


X 


X 




Hershkovitz, 1951, pp. 553-554; Smith, 
1972, p. 74; Cartwright & Kirkpatrick, 
1977, p. 466; Quinones et al., 1978, p. 
559; Pendergast, 1979, p. 10 


Pteronotus personatus 










X 


This paper 


Mormoops megalophylla 




X 


X 


X 


X 


This paper 


Micronycteris brachyotis 






X 




X 


This paper 


Micronycteris megalotis 


X X 








X 


This paper 


Micronycteris nicefori 










X 


This paper 


Micronycteris schmidtorum 


X X 








X 


This paper 


Lonchorhina aurita 








X 


X 


This paper 


Macrophyllum macrophyllum 






X 




X 


This paper 


Tonatia bidens 




... 


X 




X 


This paper 


Tonatia evotis 






X 


X 




Sanborn, 1941, pp. 372-373; Disney, 1968, 

p. 7 
Disney, 1968, p. 7; this paper 


Tonatia minuta 






X 




X 


Mimon cozumelae 




X 


X 




X 


This paper 


Mimon crenulatum 






X 




X 


Ruiz, 1983, p. 374; this paper 


Phyllostomus discolor 










X 


This paper 


Phylloderma stenops 










X 


This paper 


Trachops cirrhosus 


X 


X 






X 


Sanborn, 1941, p. 374; Pendergast, 1979, 
p. 10; this paper 


Chrotopterus auritns 










X 


This paper 


Vampyrum spectrum 










X 


This paper 


Glossophaga commissarisi 




X 






X 


Webster & Jones, 1982, p. 5; this paper 


Glossophaga soricina 




X 


X 






Villa-R., 1966, p. 231; Disney, 1968, p. 7; 
Cartwright & Kirkpatrick, 1977, p. 466; 
Howell, 1977, p. 510; Quinones el al., 
1978, p. 559; Pendergast, 1979, p. 10 



Continued on next page 



MCCARTHY: DISTRIBUTION OF BATS 



161 



Species 



Cz OW Bz Cy SC Td 



References 



CaroUia brevicauda 



Carollia perspicillata 



Sturnira lilium 
Uroderma bilobatum 

Vampyrops helleri 
Vampyrodes caraccioli 
Vampyressa pusilla 
Chiroderma villosum 
Artibeus intermedius* 
Artibeus jamaicensis* 



Artibeus lituratus* 



Artibeus phaeotis 


X 


X 


X 


X 






Artibeus toltecus 
Artibeus watsoni 




X 


X 


X 




X 


Centurio senex 
Desmodus rotundus 


X 


X 


X 
X 


X 
X 


X 


X 


Diphylla ecaudata 
Natalus stramineus 
Thyroptera tricolor 
Myotis elegans 
Myotis keaysi 




X 


X 


X 
X 

X 


X 
X 

X 


X 
X 


Eptesicus furinalis 
Lasiurus horealis 
Lasiurus ega 
Lasiurus intermedius 
Rhogeessa tumida 


X 


X 
X 
X 


X 
X 
X 


X 
X 


X 
X 
X 


X 

X 
X 


Bauerus dubiaquercus 
Nyctinomops laticaudatus 
Eumops auripendulus 
Eumops bonariensis 
Eumops glaucinus 
Eumops underwoodi 
Molossus ater 


X 


X 
X 


X 
X 


X 
X 

X 
X 




X 


Molossus molossus 
Molossus sinaloae 






X 


X 
X 


X 





Hershkovitz, 1951, pp. 555-556; Villa-R., 
1966, p. 270; Disney, 1968, p. 7; Pine, 
1972, p. 42; Quinones et al., 1978, p. 
559; Pendergast, 1979, p. 10 

Hershkovitz, 1951, p. 555; Disney, 1968, 
p. 7; Pine, 1972, p. 42; Quinones et al., 

1978, p. 559; Pendergast, 1979, p. 10 
Disney, 1968, p. 7; Pendergast, 1979, p. 10 
Disney, 1968, p. 7; Davis, 1968, p. 697; 

Pendergast, 1979, p. 10 

This paper 

This paper 

Peterson, 1966, p. 676; this paper 

This paper 

Davis, 1984, p. 14 

Dobson, 1878, p. 520; Andersen, 1908, p. 
263; Gaumer. 1917, p. 298; Disney, 
1968, p. 7; Davis, 1970b, p. 118; Pen- 
dergast, 1971, p. 102 

Murie, 1935, p. 19; Hershkovitz, 1951, pp. 
556-557; Disney, 1968, p. 7; Pendergast, 

1979, p. 10 

Disney 1968, p. 7; Davis, 1970a, p. 398; 

Pendergast, 1979, p. 10 
This paper 
Davis, 1970a, p. 393; Pendergast, 1979, p. 

10 
This paper 
Hershkovitz, 1951, p. 557; Disney, 1968, 

p. 7; Cartwright & Kirkpatrick, 1977, p. 

466; Quinones et al., 1978, p. 559; Pen- 
dergast, 1979, p. 10 
This paper 
This paj)er 

Sanborn, 1941, p. 382 
This paper 
Hershkovitz, 1951, pp. 557-558; LaVal, 

1973a, p. 25; Quinones et al., 1978, p. 

559 
Disney, 1968, p. 7; this paper 
This paper 
This paper 
This paper 
LaVal, 1973b, p. 29; Kirkpatrick et al., 

1975, p. 331 
This paper 
Murie, 1935, p. 19 
Eger, 1974, p. 5; this paper 
This paper 
Eger, 1977, p. 42 
Eger, 1977, p. 55 
Murie, 1935, p. 19; Pendergast, 1979, p. 

10 
Murie, 1935, p. 19 
Hershkovitz, 1951, p. 559; Disney. 1968, 

p. 7 



* Davis (1984) examined the Artibeus "lituratus" complex in Middle America and restored Artibeus intermedius 
J. A. Allen to specific status. Specimens cited in the publications listed for A. jamaicensis and A. lituratus should be 
reevaluated. 



162 



HELDIANA: ZOOLOGY 



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



Charles O. Handley, Jr. 



ABSTRACTS 

The larger species of Artibeus of the Amazon Basin are defined, and a new giant species is 
named and described from Venezuela and Colombia. Artibeus fallax, A. Hercules, and A. pla- 
nirostris are regarded as subspecies of Artibeus jamaicensis, by far the most variable of the 
larger Artibeus of the region. 

The smaller Artibeus are keyed and arranged in six species groups. A new dwarf species is 
described from Brazil, Ecuador, Guyana, Peru, and Venezuela. Distribution and diversity of 
the smaller species are discussed. Artibeus cinereus, once thought to range throughout Central 
America and much of South America and to include all of the smaller taxa except A. concolor 
and A. hartii, is restricted to include only the nominate form and A. quadrivittatus of the lower 
Amazon Basin and adjacent coastal areas. 

With these additions and changes in status, at least nine species of Artibeus now are known 
to occur in northeastern South America. 



Las especies de gran tamaiio de Artibeus de la Cuenca del Rio Amazonas son definidas y una 
nueva especie gigante de Venezuela y Colombia es nombrada y descrita. Artibeus fallax, A. 
hercules, y A. planirostris son consideradas como subespecies de Artibeus jamaicensis, que es 
el mas variable de los grandes Artibeus de la region. 

Una clave es preparada para las especies de Artibeus menores, y las especies son arregladas 
en seis grupos. Una nueva especie enana de Brasil, Ecuador, Guyana, Peru, y Venezuela es 
descrita. La distribucion y la diversidad de las especies menores son discutidas. Artibeus cinereus, 
que antes se penso estaba distribuida en Centro America y una gran parte de Sudamerica, y 
que incluyera todas las taxa mas pequeiias (a excepcion de A. concolor y A. hartii), es ahora 
restringuida para incluir solamente la especie nominal y A. quadrivittatus a la Cuenca baja del 
Rio Amazonas y a las areas costeras adyacentes. 

Con estas adiciones y cambios de "status," por lo menos nueve especies de Artibeus ya son 
conocidas y se encuentran en el nordeste de Sudamerica. 



Sao definidas as especies maiores de Artibeus que ocorrem na Bacia Amazonica, e uma especie 
nova, gigante, e descrita. Artibeus fallax, A. hercules, e A. planirostris sao consideradas subes- 
pecies de Artibeus jamaicensis, certamente a especie mais variavel dos Artibeus maiores da 
regiao. 

Uma chave para os Artibeus menores, os quais foram designados a seis grupos de especies, 
e fomecida. Uma especie nova aiia e descrita do Brasil, Equador, Guiana, Peru, e Venezuela. 
A diversidade, e as distribui^oes geograficas destas especies, sao discutidas. Artibeus cinereus, 
o qual acreditava-se abranger toda America Central e grande parte da America do Sul, alem 
de incluir todos taxa menores com excessao de A. concolor e A. hartii, e reduzido a um unico 
taxon, restrito ao sul da Bacia Amazonica e as suas areas adjacentes. 

Incluindo as adi9oes e mudan^as de status propostas neste trabalho, sao reconhecidas, atual- 
mente, ao menos nove especies de Artibeus na regiao nordeste da America do Sul. 



From the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. 

HANDLEY: NEW SPECIES OF ARTIBEUS 163 



Introduction 

Mammals and their ectoparasites were collected 
in Venezuela between 1965 and 1968 by the 
Smithsonian Venezuelan Project (SVP), supported 
in part by a contract (DA-49-MD-2788) of the 
Medical Research and Development Command, 
Office of the Surgeon General, U.S. Army. Nu- 
merous papers have described the ectoparasites 
and mammals of the Project. Throughout these 
papers undescribed species of mammals have been 
referred to by alphabetical designations. Some of 
these have been named subsequently by Handley 
and Ferris (1972), Handley and Gordon (1980), 
and Handley (1984). This paper deals with fruit- 
eating bats of the genus Artibeus Leach. 

The cranial measurements reported here were 
taken as outlined by Handley (1959, p. 98). Hind 
foot, tibia, calcar, and forearm were measured on 
dry museum si)ecimens or on specimens preserved 
in alcohol; all other external dimensions were 
measured on fresh specimens in the field. All mea- 
surements are in millimeters. Coloration was de- 
termined under Examolites (Macbeth Corp., New- 
burg, NY 12533) with natural light excluded. 



A New Giant Artibeus 

It is now generally agreed that in and around 
the Amazon Basin there are three large species of 
Artibeus. Handley (1976) recognized them as: (1) 
/1./m//^;>jo51« Gray— smaller, molars 3/3, rostrum 
arched, postorbital process poorly developed, fur 
long, coloration blackish, facial stripes faint or ab- 
sent, interfemoral membrane (IM) naked; (2) A. 
jamaicensis Leach — larger, molars 3/3, rostrum 
arched, postorbital process poorly developed, fur 
short, coloration gray-brown, facial stripes present 
but not sharply defined, IM naked; and (3) A. li- 
turatus Olfers— larger, molars 2/3, rostrum flat- 
tish, postorbital process well developed, fur short, 
coloration chocolate brown, facial stripes promi- 
nent and well defined, IM hairy. However, as shown 
by Koopman (1978) and Honacki et al. (1982), 
there is no consensus on the delimitation of these 
species. 

The difficulty in defining the species arises pri- 
marily from the fact that Artibeus jamaicensis is 
unusually variable geographically in morphology. 
The other species show very little variation in this 
region. Artibeus jamaicensis is large in the Ama- 



zon Basin, so large in fact that the subspecies there, 
A. j. fallax Peters and A. j. Hercules Rehn, until 
recently have been aligned by most authors with 
the universally large A. lituratus, although they 
differ from it in many characteristics other than 
size. To the southeast of the Amazon Basin {A. j. 
planirostris Spix) and to the north of it (A. j. trin- 
itatis Andersen), A. jamaicensis is dramatically 
smaller, in fact similar in size to A. fuliginosus. 
Everywhere east of the Andes A. jamaicensis has 
3/3 molars; west of the Andes and in Central 
America it has 2/3 molars. 

Specimens in the SVP collection show that the 
large Artibeus jamaicensis fallax and small A. j. 
trinitatis apparently intergrade in the Llanos of 
Venezuela where the habitat is marginal for A. 
jamaicensis and where it is an uncommon bat. 
Furthermore, intergradation between the small, 
1 2-molar A. j. trinitatis and the slightly larger, 10- 
molar A. j. aequatorialis Andersen of the north- 
west coast of South America can be seen in spec- 
imens from northern Colombia. 

These two zones of intergradation are of crucial 
importance in the nomenclature oi Artibeus, for 
they serve to link "/I. jamaicensis^'' of the West 
Indies and Central America and "A. planirostris" 
of eastern South America. They are especially im- 
portant in the present context because of the dis- 
covery of a fourth large Artibeus, superficially sim- 
ilar to but larger than A. j. fallax, occurring together 
with it in southern Venezuela and with the small 
A. j. trinitatis in western Venezuela and northern 
Colombia. It can be recognized as follows: 



Artibeus amplus new SF>ecies 

HoLOTYPE— USNM 440932, adult female with 
suckling young, skin and skull, collected 1 5 April 
1968 by Norman E. Peterson, F. P. Brown, Jr., 
and J. O. Matson at Kasmera, 2 1 km SW Machi- 
ques, Estado Zulia, Venezuela, 270 m, in a damp 
cave in a cliff across the Rio Yasa from the Kas- 
mera Biological Station, eastern foothills of the 
Sierra de Perija. Original number, svp 22086. 

Etymology— Latin amplus, large, referring to 
the large size of this bat, one of the largest Artibeus. 

Distribution— Northern foothills of the Cen- 
tral Andes in Colombia; lower eastern slopes of 
the Sierra de Perija and the Venezuelan Andes in 
western Venezuela; and the vicinity of Cerro Dui- 
da and the low mountains of southeastern Bolivar 
in southern Venezuela. It probably occurs in ad- 



164 



HELDIANA: ZOOLOGY 



jacent parts of Guyana and Brazil as well. The SVP 
collectors found A. arnplus near streams and in 
other moist areas (98%); in evergreen forest (90%) 
and in forest openings such as yards and orchards 
(10%). Most specimens were mist netted (86%), 
but some ( 1 4%) were found roosting in the twilight 
zone of caves. Elevational range, 24-1200 m. 
Holdridge life zones (Ewel & Madriz, 1 968): Trop- 
ical humid forest (10%), Tropical very humid for- 
est (22%), Premontane humid forest (12%), Pre- 
montane very humid forest (2%), Premontane rain 
forest (4%), Lower montane very humid forest 
(10%), and Lower montane rain forest (40%). Ridge 
slopes and valley floor in the area where the ho- 
lotype was collected were clothed with second 
growth evergreen forest, while lawns, shrubbery, 
banana and papaya plants, and scattered grapefruit 
trees characterized the grounds of the biological 
station. 

Description— Size large (forearm 70.0, greatest 
length of skull 31.3, maxillary toothrow 11.2 — 
averages of Venezuelan specimens). Coloration of 
fur as in sympatric Artibeus jamaicensis (dorsum 
blackish brown to brown; facial stripes present but 
obscure; underparts blackish brown, usually frost- 
ed with white; underarms with abundant long, 
usually whitish hairs); ears dark fuscous to black, 
paler basally; lips and noseleaf blackish; mem- 
branes blackish; wing tips undifferentiated or gray- 
ish, never white. Horseshoe of noseleaf bound 
down mediobasally; legs and interfemoral mem- 
brane slightly hairy, the latter particularly medio- 
ventrally, where hairs extend as a short fringe be- 
yond edge of membrane; forearm long. 

Skull superficially like that of Artibeus jamai- 
censis, but relatively longer and narrower; rostrum 
long and flattish; supraorbital ledges subparallel 
and together with postorbital processes often poor- 
ly develojDed or even ill-defined; zygomata not very 
flared from skull, usually subparallel to one another, 
and in side view, thin and fragile; posterolateral 
angle of skull not particularly flared; palate rela- 
tively narrow and toothrows ovoid in outline; 
postpalatal extension usually long, narrow, and 
parallel sided; dentition as in A. jamaicensis, ex- 
cept I' only weakly bilobed; dental formula 2/2- 
1/1-2/2-3/3 X 2 = 32. This bat is the only known 
host of Strebla paramirabilis Wenzel and Tri- 
chobius assimilis Wenzel (Diptera: Streblidae), so 
it can be distinguished from other Artibeus by its 
parasites as well as its morphology. 

Measurements of the holotype, an adult female: 
total length 101, tail vertebrae 0, hind foot (dry) 
17, ear from notch 25, forearm 69.2, tibia 24.1, 



calcar 6.2, weight 70.4 g. Greatest length of skull 
31.9, zygomatic breadth 18.3, postorbital breadth 
7.7, breadth of braincase 13.3, depth of braincase 
1 1.6, length of maxillary toothrow 1 1.2, postpal- 
atal length 9.8, palatal breadth outside of M^ 12.9, 
rostral breadth at base of canines 8.3. See Table 
1 for additional measurements. 

Comparisons— Four large species of Artibeus 
occur in Venezuela, all of them together in the 
southern part of the country. Among these, Arti- 
beus amplus and A. jamaicensis are most alike; 
but despite the superficial resemblance, the two 
can be distinguished by many characters, both ex- 
ternal and cranial. All A. amplus examined have 
the lower edge of the noseleaf horseshoe bound 
down, while about 95% of A. jamaicensis from the 
same localities have it free; all A. amplus have the 
interfemoral membrane slightly hairy and fringed 
medially, hwX A. jamaicensis never does; and while 
A. jamaicensis often has the wings white-tipped, 
A. amplus never does. Cranially, A. amplus differs 
from A. jamaicensis in having a longer, narrower 
skull; longer, somewhat more ffattened rostrum 
(most easily seen in dimensions of rostral shield); 
less arched nasals; margins of supraorbital nearly 
parallel, rather than converging posteriorly, and 
usually not as well developed; zygomata thinner 
and more fragile and usually subparallel rather 
than diverging markedly posteriorly; posterolater- 
al angle of skull not so flaring; palate narrower and 
toothrows usually less nearly circular in outline; 
and postpalatal extension usually longer and nar- 
rower, parallel sided (not flaring posteriorward). 
The two species are hosts of different species of 
parasitic streblid ffies. 

Specimens Examined— Total 55. COLOMBIA. 
Antioquia: La Tirana, 33 km SW Zaragoza, 520 
m (2 usnm). VENEZUELA. Apure: Nulita, Selvas 
de San Camilo, 29 km SSW Santo Domingo, 24 
m (2 usnm). Bolivar: 21 to 33 km NE Icabani, 
775-851 m (6 usnm); Km 125, 85 km SSE El 
Dorado, 826-1165 m (5 usnm). T.F. Amazonas: 
Belen, Rio Cunucunuma, 56 km NNW Esmeralda, 
1 50 m (9 usnm); Cabecera del Caiio Culebra, Cerro 
Duida, 40 km NNW Esmeralda, 1 1 40-1 200 m (2 1 
usnm); Caiio Culebra, Cerro Duida, 50 km NNW 
Esmeralda, 800 m (2 usnm); Tamatama, Rio Ori- 
noco, 2 km above Boca del Casiquiare, 135 m (2 
usnm). Zulia: Kasmera, 21 km SW Machiques, 
270 m (3 USNM, 1 ucv); 15 km W Machiques (1 
amnh); Novito, 19 km WSW Machiques, 1 135 m 
(1 usnm). 

Remarks— In previous publications of SVP, Ar- 
tibeus amplus has been known as ""Artibeus sp. D". 



HANDLEY: NEW SPECIES OF ARTIBEUS 



165 



Table 1 . Measurements of adult Artibeus ampins and A. jamaicensis. For each measurement, line 1 includes the 
mean plus or minus two standard errors, line 2 the extremes, and line 3, in parentheses, the number of specimens 
measured. All specimens are from Venezuela unless otherwise stated. 



Total length 



100.4 ± 3.88 
93-104 
(5) 

89.9 ± 2.22 

80-100 

(21) 

86.4 ± 2.90 

77-95 

(19) 

83.8 ± 4.14 

73-91 

(10) 

88.1 ± 1.62 

80-93 

(17) 

86.3 ± 1.60 

82-90 

(8) 



Hind foot 
(dr>) 



Ear 



Forearm 



Greatest 
length 



Zygomatic 
breadth 



15.9 ± 0.40 
14-18 
(19) 

15.4 ± 0.32 
15-16 
(10) 

17.8 ± 0.32 
16-19 
(17) 



18.3 ± 0.32 

18-19 

(8) 



16.9 ± 0.30 

16.0-17.7 

(12) 

17.0 ± 0.28 
16.6-17.6 

(6) 

19.1 ± 0.14 
18.7-19.6 

(13) 

19.3 ± 0.32 

18.6-20.2 

(8) 



Postorbital 
breadth 



Artibeus amplus, males and females, Zulia and Colombia 
18.4 ± 0.60 23.7 ± 1.28 70.8 ±1.78 31.4 ± 0.26 18.6 ± 0.20 
17-19 22-26 68.6-75.3 31.0-31.9 18.1-18.8 

(7) (7) (7) (7) (7) 

A. amplus, males and females, T.F. Amazonas and Bolivar 
18.3 ± 0.30 23.0 ± 0.98 69.1 ± 0.90 31.2 ± 0.24 18.4 ± 0.14 
17-20 18-26 65.0-73.2 30.3-32.8 17.4-19.1 

(22) (21) (22) (29) (30) 

A. jamaicensis, females, Zulia 
22.6 ± 0.96 61.1 ± 0.64 27.7 ± 0.24 
17-25 58.9-64.2 26.7-28.5 

(19) (19) (19) 

A. jamaicensis, males, Zulia 
22.2 ± 0.98 59.3 ± 0.92 27.4 ± 0.14 
20-25 56.2-61.4 27.1-27.7 

(10) (10) (10) 

A. jamaicensis, females, T.F. Amazonas 
24.6 ± 0.46 66.8 ±1.12 30.7 ± 0.28 
23-26 62.1-70.1 29.4-31.3 

(17) (17) (14) 

A. jamaicensis. males, T.F. Amazonas 
24.6 ± 0.52 65.4 ± 1.42 30.7 ± 0.26 
24-26 62.4-68.6 30.2-31.4 

(8) (8) (8) 



7.9 ± 0.20 

7.6-8.3 

(7) 

7.8 ± O.IO 

7.3-8.4 

(31) 

6.8 ± 0.08 

6.5-7.1 

(19) 

6.8 ± 0.14 

6.3-7.0 

(10) 

7.5 ± 0.10 
7.1-7.7 

(14) 

7.6 ± 0.16 

7.2-7.9 
(8) 



A New Dwarf Artibeus 

The taxonomy of the smaller Artibeus is in a 
state of flux. As recently as 35 years ago all of the 
smaller species except A. concolor Peters and A. 
hart a Thomas were believed to be variants of .-i. 
cinerens Gervais. Since then, first one and then 
another of the supposed subspecies of .4. cinereus 
has been shown to be independent species. Today 
only A. bogotensis Andersen, A. glaucus Thomas, 
A. pumilio Thomas. A. quadrivittatus Peters. A. 
rosenbergi Thomas, and A. watsoni Thomas re- 
main associated with A. cinereus (Honacki et al., 
1982). However, except for^. quadrivittatus, these 
do not properly belong with A. cinereus either. 

Artibeus glaucus and A. bogotensis intergrade in 
Ecuador and form an Andean-northern South 
American species sympatric with A. cinereus in 
southern Venezuela. Artibeus glaucus thus has two 
subspecies, the nominate form and A. g. bogoten- 
sis. Artibeus watsoni Thomas of northwestern South 
Ameinca and Central America is closely related, 
but intergradation with A. g. glaucus or A. g. bo- 
gotensis has not been obsei^ed. 



Artibeus pumilio is an enigmatic taxon. Many 
museum specimens bear the name A. pumilio, but 
perhaps the only specimen properly associated with 
the name is the holotype. This specimen may be 
only an odd variant of one of the other species, 
but not of the species described here. For the pres- 
ent, A. pumilio must be regarded as unplaceable. 
The same can be said for A. rosenbergi, charac- 
teiized by a curiously long, narrow skull such as 
can be found occasionally in large samples of most 
species o{ Artibeus. Because of their equivocal sta- 
tus, neither A. pumilio nor A. rosenbergi is includ- 
ed in the appended list of species and key. The 
characteii sties and status of these taxa will be the 
subject of another paper. 

Thus, .A. cinereus now has been shorn of all of 
its supposed subspecies except A. c. quadrivittatus. 
Its supposed range has been reduced from encom- 
passing most of Central Ameiica and tropical South 
America to occupying only the Amazon Basin 
(possibly only the lower basin) and adjacent coast- 
al areas. Sympatric with A. cinereus in much of its 
range is a distinctive dwarf species which can be 
known as: 



166 



HELDIANA: ZOOLOGY 



Table 1. Continued. 



Braincase 
breadth 


Braincase 
depth 


Maxillary Postpalatal 
toothrow length 


Width at 
molars 


Width at 
canines 


Tibia 


13.5 ± 0.20 

13.2-14.0 

(7) 


Artibeus amplus. males and females, 
1 1.0 ± 0.26 1 1.2 ± 0.16 9.7 ± 0.20 
10.6-11.6 11.1-11.5 9.3-10.0 
(7) (7) (7) 


Zulia and Colombia 

13.2 ± 0.26 8.4 ±0.16 
12.7-13.5 8.2-8.8 
(7) (7) 


25.9 ± 1.28 

24.1-28.2 

(6) 


13.3 ± 0.12 

12.9-14.0 

(30) 


A. amplus, males and females, T.F. Amazonas and Bolivar 
11.2 ±0.06 11.2 ±0.10 9.8 ±0.14 13.3 ± 0.10 8.6 ± 0.08 
10.7-11.5 10.7-11.8 9.1-10.6 12.8-13.9 8.3-8.9 
(30) (31) (28) (30) (30) 


24.8 ± 0.38 

23.1-26.2 

(22) 


12.3 ± 0.10 

12.0-12.8 

(19) 


10.2 ± 0.16 

9.5-10.9 

(19) 


A. jamaicensis. females 
10.0 ±0.12 8.6 ±0.14 
9.4-10.5 8.2-9.3 
(17) (18) 


, Zulia 

12.1 ± 0.16 

11.2-12.7 

(19) 


7.6 ± 0.10 

7.2-8.0 

(19) 


22.4 ± 0.50 

20.1-23.8 

(19) 


12.1 ± 0.14 

11.8-12.4 

(10) 


10.3 ± 0.18 

10.0-10.9 

(10) 


A. jamaicensis. males, 
10.1 ± 0.16 8.4 ± 0.12 
9.7-10.4 8.2-8.8 
(8) (10) 


Zulia 

12.2 ± 0.18 

11.8-12.6 

(10) 


7.7 ± 0.12 

7.4-7.9 

(9) 


21.7 ± 0.52 

20.1-23.0 

(10) 


13.2 ± 0.12 

12.9-13.5 

(14) 


10.8 ± 0.18 

10.2-11.5 

(14) 


A. jamaicensis. females, T.F 
11.4 ±0.20 9.4 ±0.18 
11.0-12.0 8.9-10.0 
(13) (14) 


. Amazonas 
13.7 ± 0.22 
13.2-14.4 
(13) 


8.6 ± 0.12 

8.3-8.9 

(13) 


24.1 ± 0.46 

22.3-25.4 

(17) 


13.4 ± 0.22 

12.8-13.7 

(8) 


11.0 ± 0.18 

10.7-11.4 

(8) 


A. jamaicensis, males, T.F. 
11.4 ±0.18 9.3 ± 0.24 
11.1-11.8 8.7-9.8 
(8) (8) 


Amazonas 
13.8 ± 0.30 
13.0-14.3 
(8) 


8.8 ± 0.12 

8.6-9.1 

(8) 


23.3 ± 0.52 

22.0-24.4 

(8) 



Artibeus gnomus new species 

HoLOTYPE— USNM 387534, adult female, skin 
and skull, collected 14 June 1966 by A. L. and M. 
D. Tuttle at El Manaco (= Km 74), 59 km SE El 
Dorado, Bolivar, Venezuela, 150 m, in a mist net 
in an orchard. Original number, svp 9298. 

Etymology— Latin gnomus, diminutive fabled 
being, dwarf, alluding to the small size of this 
species, one of the smallest Artibeus. 

Distribution— The Amazon Basin and bor- 
dering regions; from northern Amazonas Territory 
(14 km SSE Pto. Ayacucho) and northern Bolivar 
State (28 km SE El Manteco) in Venezuela and 
northern Guyana, to Para (Belem) and Mato Gros- 
so (Serra do Roncador), Brazil, and Loreto (Santa 
Rosa), Peru. SVP collectors netted A. gnomus 
mostly in moist sites (92%) in evergreen forest 
(52%) or openings such as savannas (25%) and 
yards and orchards (23%). Elevations range 1 19- 
161 m in Venezuela, sea level to 530 m in Brazil. 
Holdridge life zones: Tropical dry forest (22%), 
Tropical humid forest (67%), Tropical very humid 
forest (2%), and Premontane humid forest (9%). 



Description— Body size small (forearm aver- 
ages 36-38, greatest length of skull 18.5-18.7, and 
maxillary toothrow 5.7-6.0). Dorsal coloration 
gray-brown to brown; underparts paler; facial 
stripes very white and sharply defined. Soft parts 
coloration in life (usnm 36 1 742, male, Belem, Bra- 
zil): ear narrowly edged with yellow, brightest to- 
ward base; antitragus entirely yellow; tragus yel- 
low, brightest distally and on posterior basal lobe; 
noseleaf and horseshoe gray-brown medially, 
cream color laterally; lips and chin gray-brown; 
iris brown; forearm and fingers brownish flesh col- 
or; wings blackish, except membrane between fin- 
gers II and III transparent, grayish; interfemoral 
membrane sooty brown; legs and feet dark brown; 
claws horn color. Face short; shape and propor- 
tions of ears, noseleaf, horseshoe, lips, chin, and 
interfemoral membrane as in Artibeus cinereus; 
noseleaf minutely hirsute; lower edge of horse- 
shoe free; basal part of forearm hairy; hind ex- 
tremities (except for short hairs on feet) appear 
naked. 

Skull small, short, and broad; zygomata sub- 
parallel; rostrum narrow, very short, moderately 



HANDLEY: NEW SPECIES OF ARTIBEUS 



167 



Table 2. Measurements of adult male and female (combined) Artibeus gnomus and A. glaucus bogotensis. For 
each measurement, line 1 includes the mean plus or minus two standard errors, line 2 the extremes, and line 3, in 
parentheses, the number of specimens measured. All specimens are from Venezuela. 



Total length 



47.5 ± 1.40 

44-54 

(13) 

52.2 ± 0.86 

49-56 

(19) 



Hind foot 
(dry) 



Ear 



Forearm 



Greatest 
length 



Zygomatic 
breadth 



A. glaucus bogotensis. Km 125, 85 km SSE El Dorado 
10.6 ± 0.22 17.4 ±0.38 39.6 ± 0.60 20.3 ± 0.10 11.6 ±0.10 
10-11 16-19 36.8-41.9 19.4-21.2 10.8-12.1 

(19) (19) (19) (50) (44) 



Postorbital 
breadth 



Artibeus gnomus, Rio Supamo, Los Patos, and EI Manaco 
9.5 ± 0.28 16.9 ± 0.62 36.7 ± 0.54 18.5 ± 0.18 11.0 ±0.18 
9-10 14-19 34.0-38.3 17.9-19.1 10.4-11.2 

(13) (13) (13) (14) (8) 



4.9 ± 0.10 

4.5-5.2 

(14) 

5.0 ± 0.04 

4.6-5.3 

(51) 



deep and arched, and much swollen posterolater- 
ally (part on rostral shield, part within orbit, above 
eye); excavation for orbital nerve large and deep; 
braincase short and deep, with swelling at pos- 
terodorsal apex interrupting junction of sagittal 
and lambdoidal crests; postpalatal extension rel- 
atively short; internal edge of pterygoid fossa 
strongly ridged, narrowing mesopterygoid fossa and 
cupping pterygoid fossa which opens straight back; 
vomerine ridge visible in mesopterygoid fossa; va- 
cuities in roof of posterior nares much anterior to 
mesopterygoid fossa and not easily seen; outline 
of maxillarv toothrows nearly circular; upper ca- 
nine small (especially in basal diameter); M' with 
accessory internal ridge on lateral cusps, and with 
relatively wide talon; m, present (75 of 79 speci- 
mens examined). 

Measurements of the holotype, an adult female: 
total length 47, tail vertebrae 0, hind foot (dry) 9, 
ear from notch 18, forearm 36.5, tibia 12.6, calcar 
4.9, weight 10.5 g. Greatest length of skull 18.2, 
zygomatic breadth 10.8, postorbital breadth 4.8, 
breadth of braincase 8.5, depth of braincase 7.2, 
length of maxillary toolhrow 5.5, postpalatal length 
6.5, palatal breadth outside of M' 7.1, rostral 
breadth at base of canines 4.6. See Table 2 for 
additional measurements. 

Comparisons— ^r//Z)e«5 gnomus differs from A. 
concolor and A. hartii in many ways, but most 
significantly in lack of M\ From all other small 
Anibeus {A. anderseni, A. cinereus, and A. glaucus 
bogotensis) that occur within its range, A. gnomus 
can be distinguished by its possession of mj. Among 
the specimens examined, m, is consistently absent 
in these other taxa while it is consistently present 
in A. gnomus (except in southern Venezuela, where 
it is absent from both mandibles in four of 53 
specimens and from one mandible only in two 
others). In addition, A. gnomus differs from all of 



the sympatric taxa in its more prominent white 
facial stripes; more colorful ears, noseleaf, and lips; 
average browner, less grayish coloration of pelage; 
shorter face and rostrum (except when compared 
with A. concolor); more swollen supraorbital re- 
gion; average larger and deeper orbital nerve ex- 
cavation (sometimes equally large and deep in A. 
g. bogotensis); and more cupped pterygoid fossa, 
with internal ridge so enlarged as to significantly 
narrow the mesopterygoid fossa. 

Artibeus gnomus differs from the sympatric taxa 
individually in several other ways. It is much 
smaller than A. concolor (forearm averages 36-38 
vs. 46-48). In contrast to A. hartii it has notched 
inner upper incisors, brownish rather than dark 
chocolate coloration, and a wide, unfringed inter- 
femoral membrane. Compared with A. aruierseni 
(including the holotype, fmnh 21331), ^. gnomus 
is similar in size (slightly larger than Rio Madeira 
A. anderseni); has rostrum much deeper, more 
arched, narrower, and shorter; face not dished; 
orbit larger; zygomata more nearly parallel; and 
vacuities in roof of p)OSterior nares far forward of 
mesopterygoid fossa, rather than opening in it or 
close to it. 

At Belem, Brazil, both Artibeus gnomus and A. 
cinereus were numerous and were often taken in 
the same nets. There, fresh specimens of the two 
species were compared. Artibeus gnomus is smaller 
in size, and has a smaller head and shorter face; 
facial stripes much brighter, more sharply defined, 
and more prominent; ears, noseleaf, and lips more 
brownish, less grayish; ear edgings, antitragus, and 
tragus bright yellow, rather than cream; and nose- 
leaf edged with cream, rather than plain gray- 
brown. Furthermore, it has zygomata more nearly 
parallel; rostrum deeper and shorter; supraorbital 
area much swollen and its edges nearly parallel; 
and smaller teeth. 



168 



HELDIANA: ZOOLCX}Y 



Table 2. Continued. 



Braincase 
breadth 



Braincase 
depth 



Maxillary 
toothrow 



Postpalatal 
length 



Width at 
molars 



Width at 
canines 



Tibia 



8.5 ± 0.14 

8.1-9.0 

(14) 

9.0 ± 0.06 

8.5-9.5 

(48) 



Artibeus gnomus, Rio Supamo, Los Patos, and El Manaco 



7.4 ± 0.14 



5.7 ± 0.06 



6.3 ± 0.10 



7.5 ± 0.12 



4.9 ± 0.08 13.2 ± 0.46 



7.1-8.0 5.5-5.9 6.0-6.7 7.1-7.9 

(14) (14) (14) (14) 

A. glaucus bogotensis, Km 125, 85 km SSE El Dorado 
7.9 ± 0.06 6.5 ± 0.04 7.0 ± 0.08 8.0 ± 0.06 
7.2-8.3 6.0-6.8 6.5-7.5 7.5-8.7 

(48) (51) (47) (50) 



4.6-5.1 
(14) 

5.1 ± 0.06 

4.9-5.6 

(50) 



11.2-14.4 
(13) 

13.6 ± 0.36 

12.3-15.8 

(19) 



In southern Venezuela Artibeus gnomus is sym- 
patric with A. glaucus bogotensis. Compared with 
Venezuelan specimens and with the holotype (bm 
99.11.4.35) of this taxon, A. gnomus is much 
smaller and shorter faced; has a deeper, shorter 
rostrum; disproportionately wider zygomatic 
spread; and smaller teeth. 

In addition to comparisons of /I. gnomus with 
sympatric species, two other small Artibeus need 
to be considered: 

1. Artibeus g. glaucus —This species occurs 
nearby in the Andes. It (including the holotype, 
BM 94.8.6.13) possesses m„ and its skull has the 
basic shape of /i. gnomus. However, it is much 
larger and darker in color, has the hind extremities 
much hairier, the supraorbital region usually less 
swollen, and the pterygoid fossa much less cupped 
and opening to the mesopterygoid fossa. 

2. Artibeus fvatsoni— West of the Andes and ex- 
tending into Central America is another small 
species, A. watsoni Thomas, which like A. gnomus 
possesses m,. It (including its holotype, bm 
0.7. 1 1 . 1 9) is larger than A. gnomus; has larger teeth; 
longer rostrum, with reduced supraorbital swell- 
ing; shallower and less well-defined orbital nerve 
excavation; and like A. glaucus has the pterygoid 
fossa not cupped and opening into the mesopter- 
ygoid fossa (which consequently is not narrowed 
by the inner pterygoid ridge). 

Discussion— The ten small species of Artibeus 
recognized here can be associated in six species 
groups: 

1. Artibeus concolor Group— Amazon and up- 
per Orinoco basins and Guianas. Includes only 
Artibeus concolor. 



2. Artibeus hartii Group— Mexico and Central 
America, across northern South America to Trin- 
idad, and south to Peru east of the Andes and to 
Ecuador west of the Andes. Includes only Artibeus 
hartii. 

3. Artibeus glaucus Group— Mexico, Central 
America, and South America to Mato Grosso and 
Peru. Includes Artibeus glaucus (with two subspe- 
cies, A. g. bogotensis and A. g. glaucus), A. gnomus, 
and A. watsoni. 

4. Artibeus toltecus Group— Mexico and Cen- 
tral America. Includes Artibeus aztecus Andersen 
and Artibeus toltecus Saussure, each with several 
subspecies. 

5. Artibeus cinereus Group— Guiana region, 
coastal Brazil, and lower Amazon Basin (dubious- 
ly also upper Amazon Basin). Includes only Ar- 
tibeus cinereus, with A. c. quadrivittatus as a sub- 
species. 

6. Artibeus phaeotis Group — Mexico, Central 
America, and South America to upper Amazon 
Basin and western Ecuador. Includes Artibeus an- 
derseni Osgood and Artibeus phaeotis Miller, ' with 
several subspecies. 

Diversity in the small Artibeus is greatest in east- 
ern South America, where representatives of five 
of the six groups occur and where three of the 
groups are endemic. Altogether six species occur 
in and around the Amazon Basin, while only one 
is known with certainty in the central portion of 
the Basin; there are three in the lower Amazon 



' Until recently (Koopman, p. 152, in Honacki et al., 
1982) it has not been generally recognized that Artibeus 
phaeotis and A. ravus are conspecific. They inlergrade in 
eastern Panama and western Colombia. Both names date 
from Miller (1902). Although A. ravus was named first, 
on an earlier page, A. phaeotis became embedded in the 
literature as the name of this species. 



HANDLEY: NEW SPECIES OF ARTIBEUS 



169 



and on the southern fringes in Brazil and Bohvia, 
four or five in southern Venezuela, and five in east- 
em Peru, Ecuador, and Colombia. In contrast, only 
three of the species groups occur in Central Amer- 
ica, and only one of them is endemic there. 

Several distributional patterns are represented 
in the complex of Amazonian species. Artibeus 
concolor is found throughout the Basin but scarce- 
ly beyond it; A. cinereus occurs in the lower Am- 
azon and along the coast for some distance north 
and south of the river; A. anderseni is known from 
the upper Amazon and an isolated area in northern 
Colombia; A. glaucus is higher up, in the Andes, 
and eastward around the northern edge of the Ba- 
sin in Venezuela; the range of A. hartii resembles 
that of A. glaucus, but extends on into Central 
America; and the dwarf /I. gnomus has a p)eculiar 
circular range, completely ringing the Amazon Ba- 
sin but apparently not extending into its interior. 

Key to the Smaller Species of Artibeus 

1. Molars 3/3 (mj large) 2 

r. Molars 2/3 (mj minute) or 2/2 3 

2. V notched; facial stripes absent; coloration 
pale brown; interfemoral membrane broad 
and naked; forearm 43-52 mm 

Artibeus concolor 

2'. I' not notched; facial strip>es present; color- 
ation dark chocolate brown; interfemoral 
membrane narrow and fringed; forearm 36- 
42 mm Artibeus hartii 

3. Supraorbital region much swollen; molars 
2/3 (2/2 m A. g. bogotensis and occasionally 
in the others) . . . Artibeus glaucus Group, 4 

3'. Supraorbital region little, or not at all, swol- 
len; molars 2/2 7 

4. Rostrum short and moderately arched; pter- 
ygoid fossa cupijed and opening back, causing 
mesopterygoid fossa to be narrowed; forearm 
34-38 mm Artibeus gnomus 

4'. Rostrum long and much or only moderately 

arched; pterygoid fossa not cupped, opening 

into and not narrowing mesopterygoid fossa 

5 

5. Rostrum much arched; orbital nerve exca- 
vation shallow and often ill-defined; dorsum 
pale brownish; ears pale; forearm 35-41 mm 

Artibeus watsoni 

5'. Rostrum moderately arched; orbital nerve 
excavation deep and well defined 6 

6. Molars usually 2/3; dorsum dark grayish or 
blackish; ears dark; forearm 38-42 mm . . . 

Artibeus glaucus glaucus 



6'. Molars 2/2; dorsum pale brownish or grayish; 

ears pale; forearm 37-41 mm 

Artibeus glaucus bogotensis 

7. Interfemoral membrane narrow and fringed; 
coloration blackish 

Artibeus toltecus Group, 8 

7'. Interfemoral membrane broad, naked; col- 
oration brownish 9 

8. Larger, forearm 42-48 mm 

Artibeus aztecus 

8'. Smaller, forearm 37-41 mm 

Artibeus toltecus 

9. Rostrum deep and arched; palate long and 
moderately wide Artibeus cinereus 

9'. Rostrum shallow and flattened; palate short 

and very wide 

Artibeus phaeotis Group, 10 

10. Maxillary toothrow 5.2-6.2 mm; rostrum 

often tilted up anteriorly 

Artibeus anderseni 

10'. Maxillary toothrow 6.7-7.1 mm; rostrum 

usually not tilted up anteriorly^ 

Artibeus phaeotis 

Specimens Examined— /irf/Aeiii anderseni — 
BRAZIL. Amazonas: Borba, Rio Madeira (1 
amnh). Rondonia: Porto Velho (2 amnh, 2 fmnh, 
including holotype of ^. anderseni); Sto. Antonio 
do Hauayara (4 amnh). COLOMBIA. Bolivar: Ca- 
tival. Upper Rio San Jorge, 120 m (16 fmnh). 
Antioquia: Aljibos, 26 km S and 22 km W Zara- 
goza, 630 m (2 usnm); nr. La Tirana, 24 km S and 
22 km W Zaragoza, 520 m (2 usnm). ECUAIX)R. 
Napo: Rio Suno (Abajo) (4 amnh). Pastaza: Mon- 
talvo, Rio Bobonaza ( 1 fmnh); Rio Pindo Yacu 
(1 fmnh); Rio Yana Rumi (1 fmnh). PERU. Hua- 
nuco: Monte Alegre (1 amnh). Loreto: Boca Rio 
Curaray (1 amnh); Boca Rio Peruate, Rio Ama- 
zonas, 90 m (1 FMNH); Lagarto, Alto Ucayali (1 
amnh); Mazan (1 amnh); 59 km W Pucallpa (1 
usnm); Puerto Indiana, Rio Amazonas (2 amnh); 
Rio Morona (Quebr. Pushaga), Alto Amazonas, 
220 m (2 FMNH); Rio Yavari Mirim (Quebr. Es- 
peranza), 200 m (2 fmnh); Santa Cecilia, Rio Man- 
iti, Iquitos, 110 m (3 fmnh); Santa Luisa, Rio 
Nanay, Iquitos, 160 m (1 fmnh); Sarayacu, Rio 
Ucayali (1 amnh). Pasco: San Juan, Oxapampa, 



^ Couplet 10 will separate Artibeus anderseni and A. 
phaeotis in South America and in southern Central 
America, but it will not distinguish A. anderseni from 
Mexican A. phaeotis nanus. In such a comparison, A. 
anderseni can be recognized by its relatively broader 
skull. 



170 



FIELDIANA: ZOOLOGY j 



274 m (3 usnm). Departamento (?): Yuhucumayo, 
1200 ft [= Puno: Yahuaramayo, 366 m?] (1 mcz). 

Artibeus cinereus c/iiere«5— BRAZIL. Amazo- 
nas: Sta. Clara, Vila Bela Imperatriz [nr. Parintins] 
(1 amnh). Para: Fordlandia, Rio Tapajos (2 amnh); 
Maracano, Rio Jamunda [= Nhamunda?], Faro (5 
amnh); Rio Yumunda, Faro ( 1 bm). 

Artibeus cinereus quadrivittatus — BRAZIL,. 
Maranhao: Juryassu [= Turia9u?] (1 bm). Para: 
Belem (10 usnm); Benevides (1 bm); Para [= Be- 
lem] (1 bm); Ilha do Taiuna, Rio Tocantins (3 
amnh). Pemambuco: Pemambuco [= Recife] (2 
bm). Rio Grande do Norte: Natal (1 usnm). SUR- 
IN AME. Surinam ( 1 bm). VENEZUELA. Bolivar: 
Hato San Felipe, Serrania de Nuria (1 ucv); Hato 
San Jose, 20 km W La Paragua, 300-324 m (2 
usnm). 

Artibeus glaucus bogotensis — COLOMBIA. 
Cundinamarca: Bogota (2 bm): nr. Bogota ( 1 bm); 
Curiche, nr. Bogota (2 bm, including holotype of 
A. bogotensis); Fomeque (1 amnh); Fusagasuga (2 
mcz); Rio Negro, nr. Bogota (2 bm). GUYANA. 
Kanuku Mts. (3 bm). VENEZUELA. Bolivar: El 
Manaco, 59 km SE El Dorado, 150 m (3 usnm); 
Hato San Jose, 20 km W La Paragua, 300-324 m 
(3 USNM); 23 to 45 km NE Icabaru, 824-851 m (3 
USNM); Km 125, 85 km SSE El Dorado, 826-1 165 
m ( 1 20 usnm); Rio Supamo, 50 km SE El Manteco, 
1 50 m (2 usnm). T.F. Amazonas: Belen, Rio Cu- 
nucunuma, 56 km NNW Esmeralda, 150 m (1 
usnm); Caiio Culebra, Cerro Duida, 50 km NNW 
Esmeralda, 800 m (3 usnm). 

Artibeus glaucus glaucus— BOIAW A. Santa 
Cruz: Buenavista, 400 m (1 fmnh). ECUADOR. 
Napo: Baeza (1 bm). PERU. Cuzco: Collpa de San 
Lorenzo, Quincemil, 700 m (1 1 fmnh); Hda. Ca- 
dena, Quincemil, 1000 m (9 fmnh). Junin: Chan- 
chamayo, 1000 m (2 bm, including holotype oi A. 
glaucus); Huacapistana ( 1 f>4Nh). Puno: Rio Inam- 
bari, 670 m (3 amnh); Santo Domingo (1 amnh); 
Yahuaramayo, 366 m (1 bm, 1 usnm). 

Artibeus gnomus-ToXdA 104. BRAZIL. Mate 
Grosso: Serra do Roncador, 264 km N (by road) 
Xavantina, 533 m (17 usnm). Para: Belem, Sta. 
A, IPEAN (7 usnm); Belem, Utinga (5 usnm); Be- 
lem, Benevides (2 usnm). ECUADOR. Pastaza: 
Canelos, upper Rio Bobonaza (1 amnh). GUY- 
ANA. E. Berbice District: Wikki River (3 usnm). 
Mazaruni-Potaro District: Issano Road, 1 2 mi W 
of Bartica-Potaro Road (1 usnm). PERU. Loreto: 
59 km SW Pucallpa (1 usnm); Santa Rosa, Alto 
Ucayali (10°42'S, 73''50'W) (2 amnh). VENE- 
ZUELA. Bolivar: El Manaco, 59 km SE El Do- 
rado, 150 m (12 usnm); Km 38, SE El Dorado, 



100 m (1 ucv); Los Patos, 28 km SE El Manteco, 
1 50 m (4 usnm); Rio Supamo, 50 km SE El Man- 
teco, 150 m (1 usnm); Salto Chalimaha, Rio Pa- 
ramichi, Rio Paragua (1 ucv); Salto Ichun, Rio 
Paragua (2 ucv). T.F. Amazonas: Belen, Rio Cu- 
nucunuma, 56 km NNW Esmeralda, 150 m (2 
usnm); Boca Mavaca, 84 km SSE Esmeralda, 1 38 
m (1 usnm); Caiio Leon, Cerro Duida, 325 m (1 
amnh); Capibara, Brazo Casiquiare, 106 km SW 
Esmeralda ( 1 usnm); Esmeralda, Cerro Duida, 325 
m (3 amnh); 14 to 65 km S, SSE, and SSW Pto. 
Ayacucho, 119-161 m (16 usnm); Rio Mavaca, 
1 08 km SSE Esmeralda, 1 40 m (7 usnm); San Juan, 
Rio Manapiare, 163 km ESE Pto. Ayacucho, 155 
m (6 usnm); Tamatama, Rio Orinoco, 135 m (7 
usnm). 

Artibeus phaeot is— HoXoXyTpcs o{A. phaeotis and 
A. ravus, plus hundreds of other specimens from 
Mexico, Central America, and NW South Amer- 
ica. 

Artibeus pumilio— PERU. Loreto: Masisea, 
Tushemo, Rio Ucayali, 328 m (1 bm, holotype of 
A. pumilio). 

Artibeus watsoni— PANAMA. Chiriqui: Boga- 
va, 250 m (5 bm, including holotype of A. watsoni); 
Progreso (34 usnm); Puerto Armuelles (2 usnm). 

Remarks— In previous publications of SVP, 
Artibeus gnomus has been known as '''' Artibeus sp. 
A". 



Acknowledgments 

Among the persons who helped me put together 
this paper I am especially grateful to Sally DeMott, 
who measured the SVP skulls; Linda Gordon, who 
compiled the tables and worked with me on the 
comparisons; and Jane Ailes Small, who read and 
criticized the manuscript and did the word pro- 
cessing. Curators of several collections kindly per- 
mitted me to study specimens under their care in 
the preparation of these descriptions: American 
Museum of Natural History (AMNH), British 
Museum (Natural History) (BM), Field Museum 
of Natural History (FMNH), Museum of Com- 
parative Zoology, Harvard University (MCZ), and 
Universidad Central de Venezuela (UCV). The 
SVP collection is in United States National Mu- 
seum of Natural History (USNM); a portion of its 
specimens have been returned to Venezuela. 



HANDLEY: NEW SPECIES OF ARTIBEUS 



171 



Literature Cited 



EwEL, J. J., AND A. Madriz. 1968. Zonas de Vida de 
Venezuela. Ministerio de Agricultura y Cria, Caracas, 
265 pp., map. 

Handley, C. O., Jr. 1959. A revision of American 
bats of the genera Euderma and Plecotus. Proceedings 
of the United States National Museum, 110: 95-246. 

. 1976. Mammals of the Smithsonian Venezue- 
lan Project. Brigham Young University Science Bul- 
letin, Biological Series, 20(5): 1-91. 

. 1984. Newspeciesof mammals from northern 

South America: A long-tongued bat, genus Anoura 
Gray. Proceedings of the Biological Society of Wash- 
ington. 97: 513-521. 

Handley, C. O., Jr., and K. C. Ferris. 1972. De- 
scriptions of new bats of the genus Vampyrops. Pro- 



ceedings of the Biological Society of Washington, 84: 
519-523. 

Handley, C. O., Jr., and L. K. Gordon. 1980. New 
species of mammals from northern South America. 
Mouse possums, genus Marmosa Gray, pp. 65-72. In 
Eisenberg, J. F., ed.. Vertebrate Ecology in the North- 
em Neotropics. Smithsonian Institution Press, Wash- 
ington, D.C. 

HoNACKJ, J. H., K. E. Kjnman, and J. W. Koeppl, eds. 
1982. Mammal Species of the World. Allen Press, 
Inc., and the Association of Systematic Collections, 
Lawrence, Kansas, 694 pp. 

Koopman, K. F. 1978. Zoogeography of Peruvian bats 
with special emphasis on the role of the Andes. Amer- 
ican Museum Novitates, 2651: 1-33. 

Miller, G. S., Jr. 1902. Twenty new American bats. 
Proceedings of the Academy of Natural Sciences of 
Philadelphia, 54: 389-412. 



172 



HELDIANA: ZOOLOGY 



Seasonality of Reproduction 
in Peruvian Bats 

Gary L. Graham 



ABSTRACTS 

The reproductive conditions of 3,489 specimens were used to determine seasonal patterns 
of pregnancy and parturition in Peruvian bats. More species that are trophic generalists yield 
birth records for the dry season than do trophic specialists. Relatively more highland than 
lowland species have births recorded for both seasons (dry and wet). Presumably, trophic 
generalists and highland species experience less seasonal variation in food supplies compared 
to the other groups. A larger percentage of nectarivorous species than frugivores have birth 
records for the dry season; the reverse is true for the wet season. These patterns are associated 
with greater floral resource abundance during the dry season and a greater abundance of fruit 
resources during the wet season. 



Las condiciones reproductivas de 3489 especimenes fueron usadas para describir patrones 
estacionales de embarazo y alumbramiento en murcielagos peruanos. Mas especies de gener- 
alistas alimenticios tienen registros de nacimiento durante los dos estaciones (seca y mojada) 
que tienen especialistas. Relativamente mas especies desde tierras altas que desde tierras bajas 
tienen registros de nacimiento durante la estacion seca. Possiblamente, generalistas alimenticios 
y especies de tierras altas sufrir menos variacion estacional de provisiones alimentos que los 
otros grupos. Un mayor porcentaje de especies nectivoras que frugivoras tienen registros de 
nacimiento durante la estacion seca pero, el opuesto exista para la estacion mojada. Estos 
patrones son asociados con un mayor abundancia de recursos de flores durante la estacion seca 
y con un mayor abundancia de recursos de frutas durante la estacion mojada. 



As condi96es reprodutivas de 3489 especimes de morcegos p)eruanos foram usadas para 
descrever seus padroes de parturi^ao. Os morcegos de habitos alimentares generalizados pos- 
suem uma propor9ao maior de especies que possuem registros de partos durante a epoca da 
seca, do que os morcegos com habitos alimentares especializados. Proporcionalmente mais 
especis de morcegos de areas elevadas, do que das planicies, possuem registros de partos durante 
as duas epocas (seca e chuvosa). Morcegos generalizados e os de areas elevadas provavelmente 
experimentam menos varia^ao epocal em a quantidade da comida, do que os morcegos das 
planicies ou morcegos com habitos alimentares especializados. Uma porcentagem maior das 
especies nectarivoras, do que das especies frugivoras, possuem registros de partos durante a 
epoca da seca, sao a regra ao inves da exce9ao entre os morcegos que consumen fruta. Estes 
padroes podem ser interpretados como adpata96es de individuos a recursos alimentares que 
sao regularmente, ou ocasionalmente, obteniveis durante a epoca da seca. 



From the Department of Biology, University of New 
Mexico, Albuquerque, New Mexico 87131. 

GRAHAM: PERUVIAN BATS 173 



Introduction 

Although the timing of reproductive events is 
an important Hfe history adaptation, there are few 
studies of the reproductive phenology of South 
American bats (Racey, 1982). Most of these stud- 
ies are of individual species and cover only part 
of the year. Gestation, parturition, lactation, and 
weaning should be timed to correspond with vari- 
ations in the abundance and diversity of food sup- 
plies (Fleming et al., 1972; Bradbury & Vehren- 
camp, 1977; Bonaccorso, 1979; Wilson, 1979; 
August & Baker, 1 982). These variations are known 
to be seasonal in most of the Neotropics (Janzen, 
1967, 1973; Foster, 1982; Wolda, 1982; Smythe, 
1982; Terborgh, 1983). Which reproductive event 
is actually synchronized with peak food supplies 
is likely to be determined by the relative cost of 
each event. Lactation is the most costly period for 
most females, but weaning and dispersal pose the 
greatest survival problem for young bats (Wilson, 
1979; Racey, 1 982). These authors, and Tuttle and 
Stevenson (1982), agree that the weaning of young 
is the most critical period for individuals of most 
bat populations. 

Most species of Neotropical bats that have been 
studied are polyestrous (Fleming et al., 1972 
Thomas, 1972; Bradbury & Vehrencamp, 1977 
Myers, 1977; Wilson, 1979; Bonaccorso, 1979 
Humphrey & Bonaccorso, 1979). Each year, fe- 
male bats confront the problem of timing two 
(rarely three in a few vespertilionids) periods of 
lactation and weaning with variations in food 
availability. Individuals of most species handle 
this problem by producing their young so that the 
first is weaned at the beginning of the wet season 
as fruit supplies are reaching their peak, and the 
second later in the wet season when fruit avail- 
ability reaches a second peak or remains relatively 
high (Wilson, 1979; Tuttle «& Stevenson, 1982). 
Exceptions to this general pattern of seasonal poly- 
estry have been observed in most studies (Fleming 
et al., 1972; Thomas, 1972; Myers, 1977; Myers 
& Wetzel, 1983; Bonaccorso, 1979; see Mares & 
Wilson, 1 97 1 ; Bradbury & Vehrencamp, 1 977; and 
August & Baker, 1982 for good discussions of dry 
season birth periods). 

The purpose of this study is to examine some 
of the details of dry season reproduction in Pe- 
ruvian bats. I demonstrate that the different feed- 
ing assemblages and the faunas of different zoo- 
geographical regions differ in the proportion of 
species with birth records during the dry season. 



I also offer suggestions as to how dry season births 
may be adaptive. 



Methods 

The bat specimens from Peru housed in the col- 
lections of the American Museum of Natural His- 
tory, Field Museum of Natural History, Louisiana 
State University Museum of Zoology, National 
Museum of Natural History, and the Texas Co- 
operative Wildlife Collection were examined in 
late 1977 and early 1978 for information on re- 
production. Data were taken directly from Tuttle 
(1970) and Bowles et al. (1979) for specimens not 
in the above collections but included in their re- 
ports. Information on reproductive condition of 
bats was also obtained from the notes of A. L. 
Gardner and J. L. Patton. Fieldwork at several 
localities in Peru, conducted from June through 
August 1977 and from June through November 
1978, enabled me to collect and record the repro- 
ductive data for many of the bats now in the Lou- 
isiana State University collection. Diet informa- 
tion was taken from the literature (Heithaus et al., 
1975; Paradiso, 1975; Gardner, 1977) and is given 
in Graham ( 1 983). Nomenclature follows Graham 
(1983) except for Artibeus glaucus, which I now 
consider a distinct species. 

Recorded data consisted of species identifica- 
tion, locality, elevation, sex, age, and macroscopic 
reproductive condition. Individuals with incom- 
pletely ossified phalangeal epiphyses were classi- 
fied as juveniles. Females were designated as preg- 
nant or lactating if this information was included 
on specimen tags or indicated by dissection of fluid 
preserved females. The crown-rump lengths of 
embryos (including extra-embryonic membranes) 
were either measured (in mm) or taken from spec- 
imen tags. Length of testes was also noted when 
it was recorded on sp>ecimen tags, but was not used 
to establish reproductive patterns because of un- 
certainties in the relationship between testes size 
and sexual activity (Taddei, 1976; Thomas, 1972). 
Monthly pregnancy frequencies are based only on 
those female SF>ecimens for which I am confident 
that the presence (or absence) of embryos had been 
properly recorded. This limitation excludes most 
of the specimens collected prior to 1960. 

I classified each species with adequate data (those 
with evidence of at least one birth period) into 
those that can give birth in the dry (May-Septem- 
ber) and/or wet (October-April) seasons. I as- 



174 



FIELDIANA: ZOOLOGY 



sumed that a birth between 1 May and 3 1 August 
also indicates weaning of the young during the dry 
season. Species with births recorded only for the 
month of September (usually a dry month) were 
not included in either breeding season, since these 
young could be weaned either as the dry season 
ended or as the wet season began. The above clas- 
sifications were accomplished by determining the 
distribution of births across all months, as indi- 
cated by the distribution of juveniles, lactating fe- 
males, and embryo sizes. If a single birth was in- 
dicated for a given season, then that species was 
identified as having the potential to produce young 
during that season. The presence of juveniles or 
lactating females was assumed to indicate partu- 
rition during the month prior to capture. If a fe- 
male was carrying a large, near-term embryo, par- 
turition was assumed to take place in that month. 
On the other hand, if the embryo was very small 
relative to the adult body length, birth was as- 
sumed to take place four months later for most 
species, three months later for small insectivorous 
species (Findley & Wilson, 1 974), and five months 
later for emballonurids (Bradbury & Vehrencamp, 
1977) and Desmodus rotundus (Wilson, 1979). 

Classification of a species as one that can give 
birth during the dry season does not necessarily 
mean that it does so each year. I was unable with 
this method to determine the actual frequency of 
births during either season. A dry season classi- 
fication simply means that individuals can at least 
occasionally produce young during the dry season. 

The lack of birth records for a given species for 
either season can be the result of an actual absence 
of birthing during that season, or it may be due to 
an inadequate sample from that season. If I as- 
sume that the lack of a breeding record for a given 
season is not due to inadequate sampling, then I 
can use the G test (Sokal & Rohlf, 1981) to assess 
the significance of the differences between the 
groups of species compared. This is a safe as- 
sumption, since most of the specimens were col- 
lected during the Peruvian dry season (the season 
with which this study is principally concerned) and 
because the groups of bats predicted to have the 
greatest proportion of dry season births are the 
groups most poorly sampled (see Appendix and 
below). 



Regions and Climate 

I follow Koopman's ( 1978) division of Peru into 
three zoogeographical regions. He lists SF)ecies as 



lowland if they were collected below 1000 m east 
of the Andes and highland if collected above this 
elevation. Pacific coastal species are those col- 
lected either along the arid coast or adjacent An- 
dean foothills (including the mesic areas of north- 
western Peru). 

The Pacific coastal plain is characterized by low 
(< 50 mm) annual rainfall, most of which falls 
between December and April. The mesic areas on 
the northwestern Andean slopes experience great- 
er rainfall over an extended period. Rainfall east 
of the Andes (fig. 1 ) is seasonal and abundant, with 
the greatest amounts deposited between the ele- 
vations of 1 000 and 3000 m (Bowman, 1916). The 
wet season begins in October and continues through 
April and the dry season begins in May and ex- 
tends through September. 

Most investigators agree that food resources are 
primarily modulated by rainfall seasonality (Jan- 
zen, 1967, 1973; Frankie et al., 1974; Ricklefs, 
1975; Buskirk & Buskirk, 1976; Wolda, 1978a,b, 
1982; Foster, 1982; Smythe, 1982; Terborgh, 
1983). Many of the following generalizations on 
seasonal changes in food supplies for bats of mid- 
dle and higher elevations are based on the as- 
sumption that Peruvian plants and insects respond 
to environmental conditions in a manner similar 
to closely related organisms in Costa Rica, Pan- 
ama, or Puerto Rico (as described by the above 
authors). Terborgh's (1983) study of the changes 
in fruit, flower, and insect supplies in Manu Na- 
tional Park indicates that in the lowlands of south- 
eastern Peru, fruit abundance and diversity usually 
increase in October with the onset of the wet sea- 
son, peak in November and December and again 
in March, and decrease to the lowest level in May 
and June. Not only is the first peak greater but it 
also includes a greater proportion of plants in the 
family Moraceae (Terborgh, pers. comm.), which 
have fruit favored by bats (Gardner, 1 977). Flower 
abundance peaks in the dry season, but some flow- 
ers are present throughout the year (Terborgh, 
1983). At middle to higher elevations, the phe- 
nology of fruit and flower species is probably sim- 
ilar to lowland species, but less seasonal (Ter- 
borgh, 1977; Nevling, 1971). Changes in insect 
resources are more difficult to generalize because 
the species in different size classes (Smythe, 1982; 
Wolda, 1982) and in different habitats (Bradbury 
& Vehrencamp, 1976; Terborgh, 1983; Janzen, 
1983) have different periods of peak abundance 
and diversity. If the supply across size classes and 
habitats is considered, then it is probably rather 
high throughout the year. I will not discuss the 



GRAHAM: PERUVIAN BATS 



175 



600. 



500. 



400. 



5 300. 



I 200_ 



100. 




Month 



Fig. 1. Average rainfall amounts at 
Iquitos (lowland) and Yurac (middle el- 
evation), Department of Loreto, Peru. 
Data are taken from a map published in 
1971 by the Servicio Nacional de Me- 
teorologia e Hidrologia in Peru and are 
averaged from 1 years of records. 



seasonal changes in food levels west of the Andes 
in Peru because less is known of this region. 



Results 

More than 3,400 female specimens divided 
among 109 species were included in this study. 
Twins were recorded for only one female, a Car- 
ollia perspicillata, collected in November bearing 
two well-developed embryos (24 and 29 mm). 
Specimens were collected almost exclusively be- 
tween May and December, with June through Au- 
gust having the largest samples (see sample sizes 
in Appendix). The reproductive records are also 
unevenly distributed among the species (Appen- 
dix). These sampling problems made it difficult to 
identify the reproductive patterns for most species. 
If, however, the percentages of all the females that 
were pregnant are determined for each species for 
each month, many species have data that fit the 
pattern of seasonal polyestry. This pattern is well 
illustrated by the pregnancy curve ofCarollia per- 
spicillata (fig. 2). Pregnancy levels peak in Septem- 
ber and December and are followed by periods of 



parturition, as suggested by the October drop in 
the frequency of pregnant females and by the ju- 
veniles collected in January (Appendix). The be- 
ginning of the second reproductive cycle of the 
season is also indicated by the females of Carollia 
castanea (l), Vampyrops dorsalis (7), and Artibeus 
planirostris (1) that were simultaneously lactating 
and pregnant in November and December (Ap- 
pendix). On the other hand, the pregnancy curve 
of Myotis (fig. 2) and the records of juveniles and 
lactating females of Carollia (fig. 2) and Artibeus 
(Appendix) for May-September indicate that births 
can occur outside of the wet season. Individuals 
of some species (i.e., Desmodus rotundus, Myotis 
nigricans, and perhaps Glossophaga soricina and 
Artibeus planirostris; see Appendix) may be able 
to produce young during any month of the year. 
Table 1 lists the number of species in each fam- 
ily or subfamily with births recorded during the 
seasons. I was able to identify birth periods for 79 
(72.5%) of the 109 species listed in the Appendix. 
Most species (63 of the 79. 79.7%) have birth rec- 
ords for the wet season but a surprisingly large 
proportion of all of the species (46 out of 79, 58.2%) 
have records of dry season parturition. The ab- 
sence of birth records during the wet season 



176 



HELDIANA: ZOOLOGY 



Carollia perspiclllata 

60-1 3 3 24 38 100 18 15 23 15 





Fig. 2. Changes in pregnancy levels of Carollia perspiclllata and Myotis nigricans. The numbers across the top 
refer to the sample sizes for each month. The ordinate represents the percentage of all females for each month that 
were recorded as pregnant. 



(Mormoopidae) and the dry season (Furipteridae 
and Thyropteridae) may represent real periods of 
no births but may also be the result of inadequate 
sampling. 

Are there groups of bats that have a greater ten- 
dency than other groups toward parturition during 
the dry season? If seasonal fluctuations in food 
supplies are less in the highlands than in the low- 
lands, then proportionately more highland than 
lowland species could be expected to produce and 
wean young during the dry season and, if most 
species are polyestrous, during both seasons. Thir- 
ty-one of the 58 lowland species (53.4%) and 17 
of the 27 highland species (63.0%) have records 
of dry season births (table 2). Although the 9.6% 
difference is in the predicted direction, it is not 
significant {P > 0.25). The highland region has a 
greater proportion of species with birth records 
from both seasons (51.9% vs. 34.5%, P < 0.05). 
The highland Sturnira bogotensis that were lac- 



tating and pregnant in June and August (Appen- 
dix) provide evidence that some species with dry 
season birth records are polyestrous, becoming 
pregnant again during the dry season. Bats of the 
Pacific coastal and slope region also show a strong 
tendency toward dry season births. 

If trophic generalists (those that consume more 
than one type of food, such as fruit and insects) 
can switch to another resource when one type be- 
comes scarce, they should be less vulnerable to 
seasonal fluctuations in their food supply than are 
trophic specialists (those species that use only one 
major type of food). This reduced seasonality of 
food resources should be reflected by a tendency 
for births to occur during the dry season and, if 
polyestry is common, for both seasons. Nine of 
the 1 1 generalist species (81.8%) and 23 of the 68 
specialists (33.8%, table 2) have records of indi- 
viduals that have given birth during the dry sea- 
son, and relatively more generalists (63.6%) than 



GRAHAM: PERUVIAN BATS 



177 



Table 1. Number of species of Peruvian bats with 
births recorded for each season. 



Family 


Dry 


Season* 
Wet 


Both 


Total 
spe- 

ciest 


Emballonuridae 


1 


3 




4 


Noctilionidae 


2 


1 


1 


2 


Mormoopidae 
Phyllostomidae 
Phyllostominae 
Glossophaginae 
Carolliinae 


2 

6 
6 

2 


6 
4 
5 


3 
3 

2 


2 

9 

7 

5 


Stumirinae 


4 


6 


4 


6 


Stenodermatinae 


10 


22 


10 


22 


Desmodontinae 


1 


2 


1 


2 


Furipteridae 
Thyropteridae 
Vespertilionidae 
Molossidae 


7 
5 


1 
1 
7 
5 


4 

2 


1 

1 

10 

8 


Totals 


46 


63 


30 


79 



* The number of species with records of birth in either 
the dry or the wet season or in both. A species is listed 
in all three columns if it has birth records for both sea- 
sons. 

t The total number of species that have data indicating 
birth periods. The sum of the dry and wet season records 
does not equal the total, because some species have rec- 
ords for both seasons. 



specialists (29.7%) have records for births during 
both seasons (table 2). These large proportional 
differences are significant (P < 0.01 and P < 0.05, 
respectively). 

Since floral resources are greatest in the dry sea- 
son when fruit resources are lowest, nectarivorous 
species should show a greater tendency for indi- 
viduals to produce and wean young during the dry 
season compared to frugivores. Six of the seven 
(85.8%) bats that are at least partially nectarivo- 
rous have birth periods in the dry season com- 
pared with less than half (1 7 of 40, 42.5%) of those 
that are partially frugivorous (table 2). This large 
relative difference is significant (P < 0.05). Dry 
season production and weaning of young is prob- 
ably the common pattern for most bats that eat 
nectar. This is indicated by the large number of 
juveniles (13) and lactating females (35) recorded 
for this group from May through August. Since 
the resource abundances are reversed during the 
wet season, the pattern of frugivorous sp>ecies hav- 
ing relatively more birth records (39 of the 40 
species that are partially frugivorous, 97.5%) than 
nectarivores (3 of the 7 species that are partially 
nectarivorous, 42.9%) was expected {P < 0.001). 
Percentages of the species in these trophic groups 
that have birth records for both seasons are similar 



(40.0% and 42.9% for frugivores and nectarivores, 
resp)ectively, P > 0.50). 

The number of insectivorous species that have 
birth records for each season is almost equal (17 
for the dry, 1 8 for the wet). This is the expected 
pattern if insect abundance and diversity remain 
high throughout the year. 

If a species is seasonally polyestrous, the prob- 
ability of recording a birth for the wet season for 
that species is greater, since both young are pro- 
duced during that season. If most species are sea- 
sonally polyestrous, then the majority of species 
should have wet season birth records. All of the 
ecological groups mentioned above (except the 
nectarivores) show considerable evidence of par- 
turition during the wet season (table 2). 



Discussion 

Seasonal polyestry is the dominant pattern in 
other studies of Neotropical bat reproduction 
(Hemingetal., 1972; Thomas, 1972;Taddei, 1976; 
LaVal & Fitch, 1977; Myers, 1977; Bonaccorso, 
1979; Wilson, 1979; August & Baker, 1982). The 
majority of bats in Peru, especially frugivores, 
give birth during the wet season. Although this 
pattern is expected if most species are seasonally 
polyestrous most of the time, this conclusion is 
compromised by the possibility that some species 
may be monestrous, producing and weaning their 
young during the wet season. I was unable, for 
most species, to distinguish between these two re- 
productive patterns. However, the females col- 
lected from November and December that were 
simultaneously lactating and pregnant provide 
evidence for seasonal polyestry. Females that fol- 
low this pattern wean their first young concur- 
rently with the onset of the heavy rains, as fruit 
supplies are reaching their peak, and wean their 
second young near the middle of the wet season 
when food resources remain plentiful. 

In Peru, as in many of the other Neotropical 
areas discussed in the above studies, young bats 
are at least occasionally bom and weaned in the 
dry season. These bats are members of zoogeo- 
graphical and ecological groups whose tendencies 
toward dry season births can be predicted. If food 
availability for a particular group of bats does not 
fluctuate greatly throughout the year relative to a 
second group, then the first group should include 
a larger percentage of species with birth records 
for the dry season and, if the species are polyes- 



178 



HELDIANA: ZOOLOGY 



Table 2. Number of species of Peruvian bats with births recorded for each season, by geographical region and 
ecological group. 



Season* 


Region 


Dry 


Wet 


Both 


Total 


and group 


% (N) 


% (N) 


% (N) 


species* 


Geographical Regions 










Lowlands 


53.4(31) 


79.3 (46) 


34.5(19) 


58 


Highlands 


63.0(17) 


88.8 (24) 


51.9(14) 


27 


Pacific coastal 


80.0 (8) 


60.0 (6) 


30.0 (3) 


10 


Trophic Groups 










Insectivores 


58.6(17) 


62.1 (18) 


22.2 (6) 


29 


Frugivores 


38.7(12) 


100 (33) 


38.7(12) 


33 


Nectarivores 


66.7 (2) 


33.3(1) 


... 


3 


Piscivores 


100 (1) 




... 


1 


Vampires 


50.0(1) 


100 (2) 


50.0(1) 


2 


Frugivore/insectivores 


60.0 (3) 


80.0 (4) 


40.0 (2) 


5 


Nectarivore/insectivores 


100 (3) 


66.7 (2) 


66.7 (2) 


3 


Piscivores/insectivores 


100 (1) 


100 (1) 


100 (1) 


I 


Omnivorest 


100 (2) 


100 (2) 


100 (2) 


2 


Trophic Specialists:}: 


33.8 (23) 


79.4 (54) 


29.7(19) 


68 


Trophic Generalists§ 


81.8(9) 


90.9(10) 


63.6 (7) 


11 



* See notes to Table 1 . N = Number of species; % = % of total species. 
t Includes species that eat more than two major types of items, 
t Includes insectivores through vampires (see text, p. 1 77). 
§ Includes frugivores/insectivores through omnivores. 



trous, for both seasons. Two groups of Peruvian 
bats, highland species and trophic generalists, pos- 
sibly experience less seasonal fluctuation in food 
supplies relative to lowland species and trophic 
specialists. However, the reasons for the damp- 
ened fluctuations differ with each group. 

At high elevations in Peru, even during the dry 
season, clouds cover the forests most of the time 
(Terborgh, 1977; pers. obs.), providing moisture 
that is largely unavailable to lowland plants. This 
yearlong availability of moisture presumably keeps 
plant productivity and ultimately food availability 
from fluctuating as greatly as in the lowlands. This 
reduced fluctuation in food supplies is reflected by 
a weak trend toward relatively more highland birth 
records for the dry season and by a stronger high- 
land trend for birth records for both seasons. 

Species with generalized feeding habits are 
thought to be able to switch resources when one 
becomes reduced. This dampens the effect of sea- 
sonal changes in food availability and explains 
their larger percentages of birth records for the dry 
season and for both seasons. Fleming et al. ( 1 972), 
Heithaus et al. ( 1 975), and Bonaccorso ( 1 979) pro- 
vide data from Costa Rica and Panama that il- 
lustrate seasonal switches made by several of the 
same species included in my study as feeding gen- 
eralists (i.e., Phyllostomus discolor and Glossoph- 
aga soricina). 



Relatively more nectarivorous than frugivorous 
species have individuals with dry season birth and 
weaning records, but the records for both seasons 
are not different. These were the expected patterns, 
since both groups presumably experience seasonal 
variations in their food supplies; floral resources 
are greatest and fruit resources lowest during the 
dry season (Terborgh, 1983). Hence, dry season 
births may actually be the rule for bats that con- 
sume nectar but the exception for frugivores. The 
reverse is true for the wet season. These seasonality 
differences explain the almost equal proportions 
of species with birth records for both seasons. 

Thus, for trophic generalists, highland, and nec- 
tarivorous bat species, many females produce their 
first young in the dry season and become pregnant 
again, producing their second young during the 
wet season. Obviously, many more data are need- 
ed to clarify these patterns and to address the pos- 
sibility of differential survival of young between 
the two seasons. 

In Peru, the onset and termination of the rainy 
season can vary annually (Terborgh, 1983). Food 
supplies track these variations (Terborgh, 1983; 
Foster, 1982), and so may bat reproduction. I do 
not believe that climatic variability is the principal 
factor responsible for the births recorded in Peru 
between May and August because the ecological 
groups that I compared (frugivores and nectari- 



GRAHAM: PERUVIAN BATS 



179 



vores, trophic specialists and generalists) showed 
different seasonal birth patterns. If unusual dry 
season rains were the principal factor, then equal 
proportions of the different ecological groups 
should have responded by producing young during 
the normally dry season. 

My data support the hypothesis that dry season 
births are adaptations for producing and perhaps 
weaning young when food resources are available 
predictably or occasionally during this time. These 
patterns need to be confirmed by long term studies 
of actual bat communities, and of the resources 
used by individuals at both highland and lowland 
localities and during both seasons. 



Acknowledgments 

I am grateful to the many people who assisted 
me in Peru and in the United States. Antonio Brack 
E., Eric Cardich Briceno, Richard Bustamante M., 
and Susana Moller-H. of the Direccion General 
Forestal y de Fauna, Ministerio de Agricultura, 
Lima, provided the necessary collecting permits. 
Hernando de Macedo R. and Ramon Ferreyra of 
the Museo de Historia Natural "Javier Prado" 
were also helpful. I thank Arturo Koenig R., Man- 
uel A. Plenge, and Gustavo del Solar for their 
hospitality and help in making this study possible. 
I appreciate the assistance provided by Faucet and 
Aero Peru. Without the reliable assistance of my 
Peruvian associates, Manuel Sanchez, Klaus Wehr, 
and Reyes Rivera, the fieldwork would have been 
much more difficult. 

Karl F. Koopman (American Museum of Nat- 
ural History), Don E. Wilson (National Museum 
of Natural History), David J. Schmidly and Wil- 
liam B. Davis (Texas Cooperative Wildlife Col- 
lection), and Patricia W. Freeman (formerly of 
Field Museum of Natural History) were very help- 
ful during trips to their museums. Koopman and 
James L. Patton (Museum of Vertebrate Zoology) 
loaned fluid-preserved bats. James B. Cope (Jo- 
seph Moore Museum), Alfred L. Gardner (Na- 
tional Museum of Natural History), and Patton 
were kind enough to send me unpublished manu- 
scripts and field notes. For valuable assistance in 
the field I thank Linda J. Barkley, J. William Eley, 
Gary R. Graves, John P. O'Neill, Theodore A. 
Parker, III, Thomas S. Schulenberg, and Morris 
D. Williams. The excellent suggestions on the 
manuscript by Philip Myers were appreciated. I 



thank Susan T. Graham for her support during the 
several revisions of this work. 

1 gratefully acknowledge the financial support 
of the LSUMZ Peruvian fieldwork by John S. 
Mcllhenny, Irving and Laura R. Schweppe, E. W. 
Mudge, and the late Babette M. Odom. Travel to 
other museums was made possible in part by an 
LSU Foundation-Graduate Student Travel Award. 



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WoLDA, H. 1978a. Fluctuations in rainfall, food and 
abundance oftropical insects. Journal of Animal Ecol- 
ogy, 47: 369-381. 

. 1978b. Fluctuations in abundance of tropical 

insects. American Naturalist, 112: 1017-1045. 

. 1982. Seasonality of Homoptera on Barro Col- 
orado Island, pp. 319-330. In Leigh, E. G., Jr., A. S. 
Rand, and D. M. Windsor, eds., The Ecology of a 
Tropical Forest. Smithsonian Institution Press, Wash- 
ington, D.C., 468 pp. 



GRAHAM: PERUVIAN BATS 



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FIELDIANA: ZOOLOGY 



Tent Construction by Bats 

of the Genera Artibeus and Uroderma 



Robert M . Timm 



ABSTRACTS 

Herein, I describe new styles of tents cut and utilized by Artibeus anderseni, A. glaucus, A. 
gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, and U. magnirostrum; review 
and summarize the literature on tent use by Artibeus and Uroderma; and discuss the effectiveness 
of tents as diurnal roosts. Artibeus anderseni alters the shape of Heliconia leaves by cutting the 
lateral nerves and interconnected tissue extending out from the midrib. Artibeus glaucus cuts 
the basal lateral nerves in Xanthosoma, causing the two sides of the leaf to collapse downward 
around the midrib. Artibeus phaeotis cuts the lateral nerves and interconnected tissues in both 
banana and Heliconia imbricata; the basal cuts veer out from the midrib such that a distinctive 
V-shaped enclosure is formed by the hanging leaf Artibeus toltecus cuts the basal nerves on 
Anthurium, causing the sides of the leaf to fold down around the midrib to form a pyramid- 
shaped tent. Artibeus watsoni was found to make four distinctive styles of tents, including simple 
V-shaped cuts on bifurcated palms, cuts of a few side veins on aroids to produce a rounded 
pyramid, elongate J-shaped cuts on banana and Heliconia, and polygonal cuts on Carludovica 
pal mat a. Artibeus watsoni has the greatest repertoire in tent styles, and uses the most diverse 
array of plant species and leaf shapes. Two styles of tents constructed by Uroderma bilobatum 
are reported for the first time, one on the large pinnately leafed palm Scheelea rostrata and the 
second on banana. The common denominator between the Uroderma bilobatum tents reported 
herein and those previously described is that all are on large, broad leaves and all have a 
distinctive V-shaped pattern cut by the bats. Uroderma magnirostrum also creates an inverted 
elongate V-shaped tent on pinnately leafed palms. 

All New World tent-makers described to date are tropical members of the phyllostomid 
subfamily Stenoderminae. Each species of tent-making bat has one or more distinctive style of 
tent. Bats select leaves of specific shapes, sizes, and angles for tent construction. Most species 
appear to be obligate tent-roosters. Tents provide bats with a cryptic diurnal roost site, in 
addition to providing shelter from both the sun and rain and an early warning to the approach 
of predators. 



Aqui yo describo nuevos estilos de carpas cortadas y utilizadas por Artibeus anderseni, A. 
glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, y U. magnirostrum 
reviso y hago un sumario de la literatura acerca del uso de carpas por filostomidos; y discuto 
la efectividad de las carpas como perchas diumas. Artibeus anderseni altera la forma de las 
hojas de Heliconia cortando las nervaduras centrales y tejido interconectado que se extiende 



From the Division of Mammals, Field Museum of 
Natural History, Chicago, Illinois 60605-2496. The au- 
thor's present address is Museum of Natural History and 
Department of Systematics and Ecology, University of 
Kansas, Lawrence, Kansas 66045. 



TIMM: ARTIBEUS AND URODERMA 187 



exteriormente desde la nervadura central. Artibeus glaucus corta la base de las nervaduras 
laterales en Xanthosoma, causando asi que los dos lados de la hoja colapsen hacia abajo al 
rededor de la nervadura central. Artibeus phaeotis corta las nervaduras laterales y tejidos in- 
terconectados en banana y Heliconia imbricata; los cortes basales viran hacia afuera desde la 
nervadura central de tal modo que una distintiva cavidad en forma de V es formada por la 
hoja colgante. Artibeus toltecus corta las nervaduras basales de Anthurium, adi que los lados 
de la hoja doblen en redor de la nervadura central de tal modo que una cavidad es formada 
en forma de una piramide. Artibeus watsoni fue encontrada haciendo cuatro distintos estilos 
de carpas, incluyendo simples cortes en forma de V en palmas bifurcadas, cortes en unas pocas 
nervaduras laterales en araceas para producir una piramide redondeada, cortes alargados en 
forma de J en banana y Heliconia. y cortes poligonales en Carludovica pal mat a. Artibeus watsoni 
tiene el mas grande repertorio en estilos de carpas y usa la mas diversa serie de plantas y formas 
de hojas. Dos estilos de carpas construidas por Uroderma bilobatum son reportadas por primera 
vez; una en la larga y pinnada hoja de palma Scheelea rostrata y la segunda en banana. El 
comun denominador entre carpas de Uroderma bilobatum repnirtadas aqui y aquellas previa- 
mente descritas es que todas usan hojas grandes y anchas y todas tienen un distintivo patron 
en forma de V cortado por los murcielagos. Uroderma magnirostrum tambien corta una carpa 
en forma de V-invertido en hojas pinnadas de palmas. 

Todos los filostomidos cortadores y utilizadores de carpas descritos del Nuevo Mundo son 
membros tropicales de la subfamilia Stenoderminae. Cada especie tiene uno o mas estilos 
distintos de carpas. Los murcielagos escogen hojas de formas especificas, y constroen sus carpas 
en angulos peculiares. La mayor parte de las especies parecen utilizar las carpas obligatoriamente. 
Las carpas oferecen una percha oculta durante el dia, asi como un abrigo del sol, de la lluvia, 
y de predadores. 



Neste trabalho, (1) descrevo novos estilos de tendas cortadas e utilizadas por Artibeus an- 
derseni, A. glaucus, A. gnomus, A. phaeotis, A. toltecus, A. watsoni, Uroderma bilobatum, e U. 
magnirostrum; (2) reviso e resumo a literatura sobre o uso de tendas pelos morcegos da familia 
Phyllostomidae, e (3) discuto a eficiencia de tendas como alojamentos diumos. Artibeus an- 
derseni altera a forma das folhas de Heliconia, cortando as veias laterais e os tecidos interligados 
que estendem da veia central. Artibeus glaucus corta as veias basilares laterais em Xanthosoma, 
causando com que os dois lados da folha caiam contra o centro. Artibeus phaeotis corta as veias 
laterais e os tecidos interligados nas folhas de bananas e de Heliconia imbricata. Os cortes 
basilares partem da veia central, formando um abrigo distinto em forma de "V". Artibeus 
toltecus corta as veias basilares de Anthurium, causando com que os lados da folha dobrem em 
volta da veia central, criando um abrigo em forma de piramide. Artibeus watsoni constroi 
quatro tipos diferentes de tendas, incluindo simples cortes em forma de "V" em folhas de 
palmeiras bifurcadas; cortes em algumas das veias laterais em folhas de trepadeiras (resultando 
em piramides redondas); cortes alongados, em forma de "J", em folhas de bananas e de 
Heliconia; e cortes poligonos em fdlhas de Carludovica palmata. Artibeus watsoni possue o 
maior repertorio de estilos de tendas, e usa o conjunto mais di verso de especies de plantas e 
de configura96es de folhas. Dois estilos de tendas construidas por Uroderma bilobatum sao 
descritos pela primeira vez; um nas fdlhas grandes da palmeira Scheelea rostrata, e outro nas 
folhas de bananas. Fatores comuns entre as tendas construidas por Uroderma bilobatum aqui 
descritas, e as descritas previamente, sao a forma distinta em "V" cortada pelos morcegos, e 
o uso de folhas grandes e largas para a construfao das tendas. Uroderma magnirostrum tambem 
constroe tendas em forma de "V" invertido nas folhas de palmeiras. 

Todos morcegos construidores de tendas no Novo Mundo p)ertencem a subfamilia Steno- 
derminae (familia Phyllostomidae), e cada especie exibe um ou mais estilos caracteristicos de 
constru9ao. A maioria destas especies de morcegos parecem alojar-se obrigatoriamente em 
tendas, as quais oferecem nao so um abrigo diumo camuflado, mas tambem prote9ao contra 
sol, chuva, e predadores. 

188 HELDIANA: ZOOLOGY 



Introduction 

The use of cut leaves for diurnal roosting sites 
by bats was first described by Thomas Barbour 
(1932), who reported on diurnal roosts of Uro- 
derma bilobatum near the Panama Canal. He found 
these bats roosting under the leaves of two culti- 
vated palms identified as Livistona chinensis and 
Prichardia pacifica. Not only were these bats found 
roosting under palm fronds, but they had also al- 
tered the leaf to produce a diurnal roosting struc- 
ture. Barbour (1932, p. 307) stated that "by nip- 
ping the ridges of the plications on the under side 
the leaf is weakened and as the bitten spots are 
skillfully and serially distributed the leaf finally is 
sufficiently weakened so that the distal portion 
droops sharply downward." Chapman (1932, p. 
555) discovered Artibeus watsoni roosting under 
the cut veins of a bifurcated palm, Geonoma cu- 
neata (reported as G. decurrens), on Barro Colo- 
rado Island, Panama, and first called these mod- 
ified leaves "tents." Ingles (1953) also reported A. 
watsoni constructing tents in the palm Geonoma 
oxycarpa (reported as G. binervia) on Barro Col- 
orado Island. Goodwin and Greenhall (1961, p. 
262) found Artibeus cinereus roosting "under cut 
leaves of palm trees and on the under side of ba- 
nana leaves" and Uroderma bilobatum roosting 
under cut leaves of the carat palm, Sabal glau- 
cescens, and coconut palm, Cocos nucifera, on 
Trinidad. Ectophylla alba was reported by Timm 
and Mortimer (1976) to alter the leaves of five 
species of Heliconia in Costa Rica; the bats se- 
lected specific leaves for both size and angle to the 
ground. Artibeus jamaicensis was found by Foster 
and Timm (1976) roosting under the cut leaflets 
on a pinnately leafed palm, Scheelea rostrata, in 
a tropical dry forest in Costa Rica. Recently, Timm 
( 1 984) reported tent construction by another phyl- 
lostomid, Vampyressa pusilla, in Costa Rica, and 
Koepcke (1984) found Mesophylla macconnelli 
utilizing similar tents in Peru. Only one Old World 
bat, Cynopterus sphinx (Pteropodidae), has been 
reported to alter the shape of palm leaves to pro- 
duce a diurnal roosting structure (Goodwin, 1 979). 
Reviews of roosting site selection by bats were 
provided by Tuttle (1976) and Kunz (1982). 

To date, seven species of phyllostomids {Arti- 
beus cinereus, A. jamaicensis, A. watsoni, Ecto- 
phylla alba, Mesophylla macconnelli, Uroderma 
bilobatum, and Vampyressa pusilla) have been re- 
ported to modify leaves of plants to produce diur- 
nal roosting sites herein referred to as tents. All 



are tropical members of the phyllostomid subfam- 
ily Stenodermatinae. 

The phyllostomid genus Artibeus, which in- 
cludes some 1 5 species, is widespread in the Noo- 
tropics from northern Mexico southward to Ar- 
gentina and Chile. These bats range in size from 
10 g (A. anderseni and A. watsoni) to 70 g (A. 
lituratus). Uroderma, a closely related genus of 
medium-sized stenodermatines that includes only 
two species, is found from southern Mexico through 
the Amazon Basin of South America. The better 
known of the two species, U bilobatum, weighs 
from 13 to 21 g. 

Herein I describe tent construction and utili- 
zation by Artibeus anderseni, A. glaucus, A. gno- 
mus, A. phaeotis, A. toltecus, A. watsoni, Uroderma 
bilobatum, and U. magnirostrum; describe several 
new styles of tents; review and summarize the lit- 
erature on tent use by Artibeus and Uroderma; 
discuss the effectiveness of tents as diurnal roosts; 
and suggest directions for future research. 



Methods 

Descriptions of Study Areas 

Costa Rica— Bosque Brrancia, also known lo- 
cally as Bosque Blanco, is located 0.8 km west of 
Cuarto Cruces on the south side of the Pan Amer- 
ican Highway (Route 1) in Guanacaste Province, 
in the Pacific lowlands of western Costa Rica. Bos- 
que Brrancia is a restricted area of undisturbed 
lowland forest classified as Tropical Moist Forest; 
the dominant vegetation includes A nacardium ex- 
celsum and Scheelea rostrata. This stand of forest 
probably has not been logged previously, and rep- 
resents a close approximation to the original (pre- 
Columbian) forests of this part of Guanacaste. 
Further descriptions of this forest can be found in 
Janzen (1971) and Wilson and Janzen (1972). 

Parque Nacional de Corcovado is located on 
the Osa Peninsula of southwestern Costa Rica, 
Puntarenas Province (between 08°27'N and 
08°39'N, and 83°25'W and 83°45'W); the eleva- 
tion ranges from sea level to 400 m. Corcovado 
lies within the Tropical Wet Forest Life 21one 
(Holdridge, 1967), with lowland evergreen forest 
being the dominant forest type. Mean annual rain- 
fall is 3,800 + mm and the wettest months are from 
August through November; mean monthly tem- 
peratures range from 25.0°C to 26.5°C. Vegetation 
and habitat types at Corcovado have been de- 



TIMM: ARTIBEUS AND URODERMA 



189 



scribed by Herwitz (1981) and Hartshorn (1983). 
Areas surveyed included both primary forest and 
secondary scrub along the coast. 

The La Selva Biological Station is the field sta- 
tion of the Organization for Tropical Studies lo- 
cated 1 km SW of Puerto Viejo de Sarapiqui, 
Heredia Province in the Caribbean lowlands of 
northeastern Costa Rica (10°27'N,84°00'W); ele- 
vation ranges from 29 to 100 m. Mean annual 
rainfall is 3,990 mm, with the wettest months being 
November, December, and February; mean 
monthly temperatures range from 24.5°C (Decem- 
ber) to 26.1°C (April). La Selva lies within the 
Tropical Wet Forest Life Zone, with lowland ev- 
ergreen forest being the dominant forest type. 
Vegetation and habitat types of La Selva have been 
described by Slud (1960), Holdridge et al. (1971), 
Sawyer and Lindsey (1971), and Hartshorn ( 1 983). 
One unusual feature of the subcanopy of the La 
Selva forest is the diversity and abundance of dwarf 
palms (Hartshorn, 1983), especially the broad- 
leaved species, Geonoma congesta and Asterogyne 
martiana. These species are regularly utilized by 
Artibeus watsoni for tent construction. The tran- 
sect surveys, which included all Artibeus tents ob- 
served, were restricted to primary forest. The Uro- 
derma tents described from this site were restricted 
to an open banana patch. 

Palo Verde (Refugio Nacional de Fauna Sil- 
vestre Dr. Rafael Lucas Rodriguez Caballero) is a 
wildlife refuge located 2 km S and 12 km E of 
Bolson, in the Pacific lowlands of Guanacaste 
Province, northwestern Costa Rica (10''30'N, 
85°20'W); the elevational range is from 3 to 183 
m. Palo Verde lies within the Tropical Dry Forest 
Life Zone, with lowland deciduous forest and riv- 
erine swamp forest being the dominant forest types. 
The vegetation and habitat types of Palo Verde 
have been described by Slud ( 1 980) and Hartshorn 
(1983). Mean annual rainfall is 1,700+ mm, with 
the wettest months being April, May, September, 
and October; mean monthly temperatures range 
from 26.0°C (November) to 29.7°C (April). The 
immediate vicinity of the survey at this site was 
in a mosaic of primary forest and secondary scrub 
that included considerable bananas scattered 
throughout. 

The newly expanded Parque Nacional Braulio 
Carrillo is located in northeastern Costa Rica (be- 
tween 10°05'N and 10°25'N, and 83°54'W and 
84'^5'W); the elevation of the park ranges from 
100 to 2600 m. Braulio Carrillo is located on the 
eastern, Caribbean slope of Volcan Barba in He- 



redia Province. The elevational range at which Ar- 
tibeus toltecus and associated tents were observed 
was from 700 to 1400 m, within the Premontane 
Rain Forest Zone with midelevational evergreen 
forest and tall palms being the dominant forest 
types. Mean annual rainfall at this elevational range 
is perhaps as much as 5,000 mm, although no 
exact measurements are yet available. The vege- 
tation and habitat types at the midelevational 
ranges also have yet to be described. On 14-15 
April 1986, 3 km of trail ranging from 700 to 1 100 
m were surveyed for bat tents. Additionally, an 
intensive netting effort with Richard K. LaVal and 
Don E. Wilson was conducted in this area over a 
7-day period to determine what species of bats 
were present and their relative abundance. 

At Finca Las Cruces (2 km S of San Vito, Pun- 
tarenas Province, 08°45'N, 82°58'W, 1200 m) in 
the Premontane Wet Forest-Rain Forest transition 
area, approximately 2 km of trail leading down to 
the Rio Jaba was surveyed for tents on 1 3 August 
1982. 

Ecuador— Cascada San Rafael lies 17 km (by 
road) west of the village of Reventador, Napo 
Province, in northeastern Ecuador (00°5.8'S, 
77°34.4'W). Rainfall averages 1,500 to 2,000 mm; 
average temperatures range from 18°C to 22°C. 
The elevation is 1200 m. Reventador lies within 
the Humid Subtropical Life Zone. 

Lagarto Cocha and Zancudo Cocha are military 
encampments along the Rio Aguarico of Ama- 
zonian Ecuador named for prominent lagoons. 
Both areas are undisturbed primary lowland rain 
forests classified as Moist Forest, with an annual 
rainfall averaging from 2,000 to 3,000 mm; the 
elevation is approximately 200 m. 

Peru— Hacienda Amazonia lies just north of 
the confluence of the Alto Rio Madre de Dios and 
the Rio Pinipini in the department of Madre de 
Dios, southeastern Peru (12°58'S, 71'>09'W), just 
north of Atalaya. The Hacienda is located just east 
of Parque Nacional del Manu in the Upper Trop- 
ical Zone on the eastern foothills of the Andes. On 
25 July 1985 Barbara L. Clauson searched the 
ridge above the Hacienda for bat tents at an ele- 
vation of 825 m in primary rain forest that had 
received some selective timber harvest; she re- 
turned to this site again on 3 November 1985. 

Cerro de PantiacoUa lies above the Rio Palotoa, 
10-15 km NNW of Shintuya, in the department 
of Madre de Dios, southeastern Peru (12°35'N, 
71''18'W). On 15 November 1985 Clauson 
searched a steep sloping ridge at 600 m. The sur- 



190 



HELDIANA: ZOOLOGY 



Table 1 . Individual measurements (in cm or degrees) collected from four tents constructed by Artibeus anderseni 
on Heliconia in Ecuador. 





Blade 




Petiole 


Basal 
height 




Length 


Width 


Angle 


Length 


Angle 


Remarks 


176 
158 
154 
140 


34 
38 
36 

27 


40 
15 
50 


80 
75 
88 
60 


80 
55 
80 
70 


200 
220 
260 
140 


1 adult male A. anderseni 
3 male A. anderseni 



rounding forest was primary rain forest in the Up- 
per Tropical Zone on the eastern Andean foothills. 
Reference specimens of the bats have been de- 
posited at Field Museum of Natural History, Chi- 
cago; Escuela Politecnica Nacional, Quito, Ecua- 
dor; and Universidad Nacional de Costa Rica and 
Servicio de Parques Nacionales, San Jose, Costa 
Rica. Voucher specimens of plants have been de- 
posited in the herbaria at Field Museum, Duke 
University, Missouri Botanical Garden, and/or 
University of Wisconsin. 



Accounts of Species 

Artibeus 

Artibeus anderseni Osgood, 1916 

Artibeus anderseni occupies an extensive range 
in western Amazonia; however, little is known of 
its biology. This species has long been considered 
a junior synonym of A. cinereus. In resurrecting 
A. anderseni as a distinct species, Koopman (1978, 
p. 14) stated, "Besides its shorter face and more 
abrupt forehead, A. anderseni apparently always 
lacks the last lower molars, which A. cinereus in 
western Amazonia almost always has." I concur 
with Koopman in recognizing A. anderseni as a 
distinct species. 

In late October and early November of 1983, 
the trails and forest surrounding the military en- 
campments were searched at Lagarto Cocha and 
Zancudo Cocha in eastern Ecuador. Artibeus an- 
derseni was found to alter the shape of leaves of 
several small, forest Heliconia species to produce 
diurnal roosting structures. To create a tent from 
a Heliconia leaf, the bat severs the lateral nerves 
and interconnecting veins that extend along both 
sides of the midrib. The cuts ran along the central 



90% of the leaf from near (0 to 14 cm) the base 
to near (10 to 20 cm) the tip. Nerves and inter- 
connected tissues were severed, but not complete- 
ly, so that they did afford some support for the 
sides of the leaf. Cuts ran parallel to the midrib 
for most of its length, but did flare outward slightly 
toward the base. The lateral nerves were cut from 
3 to 8 mm from the midrib; the midrib was not 
cut. Claw marks where the bats roosted started 50 
cm from the base in one tent and ran for 1 6 cm 
distally; in another they started at 70 cm and ran 
for 10 cm distally. Measurements of the blade 
length, blade width, blade angle, petiole length, 
petiole angle, and basal height of four tents are 
provided in Table 1 . 

At Zancudo Cocha one Heliconia tent was un- 
occupied for two days in succession, then on the 
third day was occupied by an adult male with 
enlarged testes, an adult lactating female, and a 
juvenile male Artibeus anderseni (fig. 1). Another 
Heliconia tent was unoccupied. 

At Lagarto Cocha 13 tents were found in Hel- 
iconia. One was occupied by three subadult males 
not in breeding condition; a second tei t contained 
a single adult male with enlarged testes. 



Artibeus cinereus (Gervais, 1856) 

Artibeus cinereus, Gervais's fruit-eating bat, is 
found on the islands of Grenada, Trinidad, and 
Tobago, and throughout the Amazon Basin and 
adjacent coastal areas. Surprisingly little has been 
published on roosting behavior or ecology of this 
widely distributed species. On Trinidad, Goodwin 
and Greenhall (1961, p. 262) stated, "It roosts in 
small colonies of a few individuals under the cut 
leaves of palm trees and on the under side of ba- 
nana leaves." On Tobago, Husson (1954, p. 64) 
reported a single male Artibeus cinereus "hanging 
in a banana tree in cultivated country near the 
shore." 



TIMM: ARTIBEUS AND URODERMA 



191 




Fig. 1. Photograph of an adult male, adult female, and juvenile Artibeus anderseni roosting in a Heliconia leaf 
tent. Details of the cut side nerves can be seen along the midrib of the leaf 



Artibeus glaucus Thomas, 1 893 

Artibeus glaucus is found at midelevalions along 
the eastern slopes of the Andes from Venezuela to 
Peru. The status of the name glaucus has long been 
in a state of flux; it often has been considered a 
subspecies of A. cinereus. I concur with Handley 
(1987) in regarding .4. glaucus as a distinct species. 

A single old adult male Artibeus glaucus (fmnh 
124844) was observed roosting under a cut Xan- 
thosoma leaf at Cascada San Rafael, Ecuador, on 
21 September 1983. Four cut leaves on separate 
plants were observed in close proximity to each 
other; a fifth cut leaf was observed approximately 
10 to 1 5 m to the south. Only the one cut leaf was 
occupied by the single bat. An adult female A. 
glaucus was netted in the vicinity that evening. All 
tent leaves were cut down at the time and four 
were measured. On 26 November, two more cut 
leaves were found in this Xanthosoma population; 
one contained three A. glaucus. The Xanthosoma 
in which the bats were roosting were part of a 



population of Xanthosoma that occupied approx- 
imately 1 ha on a steep east-facing hillside. 

The basal lateral nerves from 2 to 5 (usually 3) 
of the Xanthosoma leaves were severed near the 
midrib, causing the two sides of the leaf to collapse 
downward around the midrib. The midrib was not 
cut. Four of the five cut leaves were measured in 
September; one found in November was measured 
(table 2). The leaves that were selected by Artibeus 
glaucus for tents all had the midrib running ap- 
proximately parallel to the ground, whereas the 
majority of unaltered leaves in the p>opulation stood 
at more vertical angles. 



Artibeus gnomus (Handley, 1987) 

Artibeus gnomus, the dwarf fruit-eating bat, is 
found in a peculiar circular range ringing the west- 
em edge of the Amazon Basin. Although this 
species has a wide distribution from Venezuela 
and Guyana to Peru, it was only recently recog- 



192 



HELDIANA: ZOOLOGY 



Table 2. Individual measurements (in cm or degrees) collected from five tents constructed by Artibeus glaucus 
on Xanthosoma in Ecuador. 



Blade 



Length Width Angle 



Petiole 
length 



Basal 
height 



Remarks 



57 68 60 140 230 3 basal cut veins on each side, 2-3 cm from midrib 

53 • • • 40 • • • 230 3 basal cut veins on each side, 1.5-3.0 cm from midrib 

55 62 40 • •■ 230 2 veins cut left, 4 cut right side, 2-3 cm from midrib 

64 36 45 137 300 3 basal veins cut on each side, 1.5-3.0 cm from midrib; 

adult male A. glaucus hanging 16 cm from base 
61 66 20 145 275 5 basal veins cut on each side, 3.5-4.0 cm from midrib; 3 ^. 

glaucus 



nized as a distinct species, and little biological in- 
formation is available (Handley, 1987). 

On 15 November 1985 Barbara L. Clauson 
found a single adult male Artibeus gnomus roost- 
ing under a cut Monstera lechleriana leaf. The 
Monstera was growing as an epiphyte approxi- 
mately 10 m off the ground on a tree on a sloping 
hillside at 600 m elevation at Cerro de Pantiacolla, 
southeastern Peru. The single cut leaf was green 
and healthy and hung horizontally. No other cut 
leaves were observed in the immediate vicinity. 

The altered leaf was 70 cm long and 38.5 cm at 
its widest point. All lateral nerves along the basal 
nearly two-thirds of the leaf were severed imme- 
diately adjacent to the thick midrib; this included 
the basal 12 nerves on one side and 14 on the 
other. The midrib was severed at 44.5 cm from 
the base which caused the apical third of the leaf 
to droop downward perpendicular to the midrib. 
All nerves proximal to the midrib cut were sev- 
ered. The lateral nerves along the apical, drooping 
25.5 cm were unaltered. 

The tent resulting from these cuts was quite en- 
closed, being formed by the sides of the leaf col- 
lapsing downward around the midrib and the dis- 
tal third of the leaf folding down, perpendicularly 
to the midrib. The lone Artibeus gnomus was 
roosting 9 cm toward the base from the severed 
midrib of the leaf. 



Artibeus jamaicensis Leach, 1821 

Artibeus jamaicensis, the Jamaican fruit-eating 
bat, is found throughout much of tropical Central 
America, the northern half of South America, and 
the Greater and Lesser Antilles. At many localities 
this is one of the most common species of bats 
encountered and consequently has received more 
study than any other phyllostomid. 



Artibeus jamaicensis was reported roosting un- 
der the cut leaflets of Scheelea rostrata in Costa 
Rica by Foster and Timm (1976). Scheelea ros- 
trata is a large, pinnately leafed palm with the 
leaflets extending out at right angles from the hor- 
izontal rachis. Leaflets within the middle 1.3 m 
region of the frond were cut at varying distances 
that increased going up to the center of the cut 
area, then decreased. "As a result . . . the distal 
parts of the leaflets folded perpendicularly, hung 
vertically below the frond, and formed a broadly 
lanceolate tent" (Foster & Timm, 1976, p. 266). 

Although several Artibeus jamaicensis occupied 
the roost, only two males were captured, one an 
adult with enlarged testes, and the second a smaller 
male not in breeding condition. Artibeus jamai- 
censis apparently has a harem mating system, in 
which a single breeding male defends a roost used 
by several females and their offspring; nonbreed- 
ing males may be found either singly or in small 
groups (Morrison, 1979; Kunz et al., 1983). Ar- 
tibeus jamaicensis has been found roosting in a 
wide variety of situations, including caves, hollow 
trees, buildings, and under unaltered leaves (Tut- 
tle, 1976), and thus is certainly not an obligate 
tent-roosting species, as apparently are the smaller 
species oi Artibeus. 



Artibeus phaeotis (Miller, 1 902) 

Artibeus phaeotis, the pygmy fruit-eating bat, is 
found from central Mexico to northern South 
America (Timm, 1985). Most accounts of habitat 
for pygmy fruit-eating bats mention their being 
netted in close proximity to stands of bananas, 
Musa X para^/5/aca (Ramirez-Pulidoetal., 1977; 
Watkins et al., 1972). Davis (1970) suggested that 
they might roost under the leaves of bananas. 

During the summer of 1982, Artibeus phaeotis 



TIMM: ARTIBEUS AND URODERMA 



193 





Fig. 2. Top, Dorsal view of a banana leaf {Midsa x paradisiacd) showing the cut nerves running aloQg the midrib 
and flaring out toward the base; bottom, tent of Art ibeus phaeotis made from a banana leaf 



was observed roosting under the leaves of banana 
and Heliconia imbricata at Palo Verde and La 
Selva, respectively, in Costa Rica. In all cases the 
leaves had been altered to produce tents. 

Art ibeus phaeotis constructs roosts in both ba- 
nanas and Heliconia imbricata by biting the lateral 
nerves and interconnecting tissue that extend at 
right angles from the midrib, causing the blade to 
fold over in a V-shaped enclosure. The two sides 
of the leaf collapse downward, hanging beneath 
the midrib (fig. 2). Nerves and interconnected tis- 
sues are not completely severed, thus the sides of 
the leaf provide some support for the entire length 
of the cut. The cuts ran from the base of the leaf 
to near the tip (table 3). Near the base, the cuts 
flared out from the midrib to the sides to form an 
elongate J-shaped pattern. The uncut tip and basal 
portion of the leaf provide additional strength. The 
undersides of roost leaves are obscured from view 
from almost all angles except from directly be- 
neath the tent. 

To characterize Musa clumps, each of which 



presumably represented an individual plant, the 
number of stalks (ramets) per clump was counted; 
the height of each ramet was estimated to the near- 
est half meter. This was done for all clumps of 
Musa in the patches that could be located, both 
those with tents and without bat tents. When a bat 
tent was located, the following measurements were 
taken in centimeters: the angle of the petiole (mid- 
way from stalk to blade), angle of blade (midway 
on blade), petiole length, blade length, blade width, 
height of base of blade, height of tip of blade, 
height of roost, isolation distance (distance from 
nearest solid object on the same vertical level), 
length of uncut basal portion of blade, and length 
of uncut distal portion. I also noted whether the 
roosts were in direct sunlight or in shade (table 3). 
Also for all tents the number of stalks on that 
column was counted, and the age of the leaf rel- 
ative to other expanded leaves was recorded. The 
oldest (lowest leaf) on a plant was assigned the 
number 1 , then the rest counted up from there. 
I scored 232 individual stalks of Musa, which 



194 



HELDIANA: ZOOLOGY 



Table 3. Individual measurements (in cm or degrees) collected from 19 tents constructed by Artibeus phaeotis 
on leaves of banana, Musa x paradisiaca, in Costa Rica. 



Petiole 


Length 


Blade 
Angle 


Width 


Baiwl . 
height 


Height 
Tip Roost 


Isolation 


Uncut 




Length 


Angle 


Base 


Tip 


43 


40 


110 


-27 


20 


196 


156 


182 


81 





15 


S3 


38 


136 


-14 


19 


230 


146 


225 


120 


10 


45 


43 


42 


146 


-40 


22 


213 


170 


228 


125 


1.5 


48 


49 


12 


120 


-55 


23 


250 


180 


240 


53 


5 


8 


43 


S2 


100 


-16 


18 


205 




200 


70 





30 


SO 


55 


120 


-3 


21 


200 


165 


210 


25 


3 


8 


S4 


50 


142 


-36 


20 


235 


160 


230 


115 


1 


44 


40 


75 


80 


+ 15 


19 


190 


195 


206 


80 


0,25 


20 


36 


68 


175 


-12 


26 


320 


260 


320 


>300 





1.5 


60 


70 


195 


+25 


30 


360 


400 


400 


> 300 


0,35 





44 


66 


130 


+23 


25 


250 


260 


290 


120 





40 


41 


50 


94 


+5 


20 


140 


155 


170 


40 


0,40 


25 


34 


45 


97 


-22 


20 


205 


175 


200 


100 








SO 


68 


130 





23 


180 


220 


230 


60 





2 


S6 


60 


147 


-37 


27 


260 


180 


240 


35 





28 


33 


55 


85 


-12 


21 


205 


215 


220 


18 





5 


16 


52 


60 


+28 


13 


130 




140 


35 





12 


46 


50 


129 


+ 15 


21 


250 


220 


260 


120 





65 


40 


54 


88 


+ 15 


14 


170 




180 


220 





3 



were in 4 1 clumps scattered in 1 00 m of second 
growth forest. The mean number of ramets per 
clump was 5.4; the range from 1 to 23. I located 
26 leaves that were cut by bats, a ratio of roughly 
one tent per 8. 1 ramets. Fourteen clumps had cut 
leaves; if a clump had cut leaves it had a mean of 
1.9 cut leaves (range 1 to 2). Of the 26 altered 
leaves, complete data were taken on 19. The re- 
maining tents were decomposing; in a few cases 
the bats had not completed the tents. On the par- 
tially completed tents, only one side of the midrib 
was cut along one-quarter to one-third the distance 
of the blade, but did not cause the side to collapse. 
Only 2 of the 1 9 complete tents were located di- 
rectly in the sun. Bat tents were found in clump 
sizes ranging from 1 to 23 (mean = 6.2). The av- 
erage height of all plants (N = 232) was 2.3 m. 
The average height of plants with tents was 2.0 m 
(N = 16). 

One adult male with enlarged testes and six 
pregnant females were watched over a 3 -day p)e- 
riod in one of the roosts. Additionally, two solitary 
nonpregnant female Artibeus phaeotis roosted un- 
der separate leaves (fig. 3). 

Artibeus phaeotis appears to select banana leaves 
with specific characteristics. Usually these are the 
oldest fully expanded leaves, just over 2 m above 
the ground, with the center of the midrib nearly 
horizontal to the ground and positioned far enough 
from nearby stems and branches to limit access 
by predators. Roost sites generally are located in 



the shade of surrounding forest trees where ap- 
parently they are protected by the forest overstory 
from wind, blowing rain, and sunlight. 

Although bananas are not native to the New 
World, they are now common throughout the range 
of Artibeus phaeotis and probably provide roost 
sites in other localities. Heliconia, Calathea, and 
broad-leafed palms are uncommon at Palo Verde, 
hence are not readily available to A. phaeotis for 
tent sites there. Artibeus phaeotis used only banana 
leaves for tent making at Palo Verde, but con- 
structed similar tents in Heliconia imbricata at La 
Selva. Heliconia imbricata is the largest species of 
Heliconia in Costa Rica, and its leaves are similar 
in size and shape to banana. The tents in Heliconia 
were similar in all respects to those in banana 
leaves. An adult male and adult female were found 
roosting together under a single Heliconia leaf in 
late June. 

Villa-R. ( 1 967) found a single specimen roosting 
near the mouth of a small cave in Mexico. 



Artibeus toltecus (Saussure, 1 860) 

Artibeus toltecus, the lowland fruit-eating bat, is 
found along the coasts of eastern and western Mex- 
ico from Nuevo Leon and Sinaloa south through 
Central America and perhaps to extreme north- 
western Colombia. This species appears to be re- 
stricted to midelevational slopes, and in Costa Rica 



TIMM: ARTIBEUS AND URODERMA 



195 




Fig. 3. Photograph of an adult female Artibeus phaeotis roosting in a banana leaf tent. Details of the cut side 
nerves can be seen along the midrib of the leaf. 



I have found it only from 650 to 1 500 m in ele- 
vation. 

In April 1986 I observed a single Artibeus tol- 
tecus roosting under a cut leaf of Anthurium ca- 
peratum in Braulio Carrillo National Park, north- 
eastern Costa Rica. The Anthurium was growing 
as an epiphyte on a small tree at 800 m (IVi km 
S, 11 km E of San Miguel, 10°17'N, 84'>05'W). 
One leaf on the plant was altered; it was IVi m off 
the ground and the midrib hung parallel to the 
ground. Four or five lateral nerves were cut basally 
on each side, causing the sides of the leaf to fold 
down around the midrib. A break of the midrib 
at its midpoint caused the distal half of the leaf to 
droop down (fig. 4). Seven additional tents of this 
style were observed on Anthurium in this area, 
ranging in elevation from 700 to 1400 m. It is 
assumed that they were made by A. toltecus, the 
only small species of Artibeus we netted there, al- 
though these tents were not occupied. Six tents 
were observed in a 3-km transect ranging from 
700 to 1 100 m in elevation. 

Davis (1944) reported that Artibeus toltecus 



roosts imder banana leaves, although he did not 
indicate that the bats were modifying the leaves. 
Davis (1944, p. 378) stated: 

. . . they had regularly established roosts tin- 
der the large, drooping leaves of the banana 
trees, each one easily recognized by the man- 
ner in which the vane of the leaf hung limply 
suspended from the midrib. The closely ap- 
pressed vanes of the leaf, plus the natural 
darkness within the depths of the grove, af- 
forded good concealment. These bats, too, 
were wary and that feature coupled with the 
nature of their retreat caused considerable 
difficulty in procuring specimens. 

In light of Davis's description of the roost sites of 
A. toltecus in banana leaves and my own obser- 
vations on A. toltecus, I suspect that this species 
was creating tents similar to those I observed for 
A. phaeotis in Costa Rica. The tents formed by A. 
phaeotis in Musa (see fig. 2) are similar in ap- 



196 



HELDIANA: ZOOLOGY 




Fig. 4. Top, Ventral view of an Anthurium caperatum leaf showing the cut nerves along the base of the leaf and 
the broken midrib; bottom, tent of Art ibeus toltecus. 



TIMM: ARTIBEUS AND URO DERMA 



197 



pearance to those described by Davis (1944) for 
A. toll ecus. 

Artibeus toltecus has also been reported in caves 
(Davis et al., 1964; Jones, 1966; Jones & Alvarez, 
1 964), and Goodwin ( 1 934, p. 1 2) reported a single 
specimen collected "in one of the buildings 
(church?) at San Lucas . . . [the] rest of the con- 
gregation seemed to be Glossophaga." 



Artibeus watsoni Thomas, 1 90 1 

Artibeus watsoni, Thomas's fruit-eating bat, is 
one of the smaller members of the genus Artibeus 
and found from southern Veracruz south through 
Central America to northern South America. It 
appears to be restricted to lowland and midele- 
vation humid forests. 

During the summers of 1974, 1982, 1984, and 
1 986, numerous individuals of Artibeus watsoni 
were seen roosting under 19 different sj)ecies of 
broadleafed plants at several localities in Costa 
Rica. At Parque Nacional de Corcovado, Costa 
Rica, trails were surveyed on three separate oc- 
casions for the presence of tents made by Artibeus 
watsoni, in June and August 1982 and again in 
August 1984. In mid-June 1982, the following 
groupings of Artibeus watsoni were observed: two 
(both adults, a male and pregnant female), two 
(pregnant female and one not captured), two (not 
captured), and six hanging singly (of which two 
were captured and found to be adult males). Ad- 
ditionally, several tents on banana and Heliconia 
were marked for relocation later in the summer. 
A tent marked on Heliconia imbricata was relo- 
cated 60 days later. The tent was still intact, al- 
though it was beginning to break down; a single 
A. watsoni v/zs using it. All other marked tents had 
decomposed. 

On 10 August 1982 I found 90 tents constructed 
by Artibeus watsoni along the trail through Cor- 
covado's "Monkey Woods." These tents were made 
from the following species of plants: Musa x par- 
adisiaca (49, 54%), Anthurium ravenii (13, 14%), 
unidentified aroid (11, 12%), Heliconia imbricata 
(9, 10%), Heliconia latispatha (1, 1%), Heliconia 
sp. (3, 3%), and Calathea insignis (4, 4%). Tents 
located on Anthurium ravenii were most often 
found clumped, with an average of 2.6 tents per 
plant, whereas in the other species of plants it was 
uncommon to find more than one tent per indi- 
vidual plant. Bats were found singly (five) or in 



three groupings of four, three, and two individuals. 
Four of the single bats were all adult males. 

A trail running up to a ridge top was surveyed 
from 9 through 1 1 August 1982, with the following 
results: 25 tents found of which 16 were on Hel- 
iconia imbricata (64%), 8 on Calathea insignis 
(32%), and 1 on Carludovica palmata (4%). Three 
tents were occupied by two (sexes unknown), one 
male, and one female. Near the mouth of the Rio 
Llorona on 8 and 9 August I counted the following 
groups of bats: eight (three adult females, three 
young, and two not caught), two (adult female with 
volant young), two (sexes unknown), and three 
singles (one a nonreproductive adult female). Ad- 
ditional tents were observed in banana, coconut 
palm (Cocos nucifera), Calathea insignis, and Car- 
ludovica cf. drudei. 

In August 1 984, 1 found 63 tents constructed by 
Artibeus watsoni. These were distributed on the 
following plants: Anthurium ravenii (36, 57%), 
Heliconia sp. (14, 22%), Musa x paradisiaca (7, 
1 1%), Calathea insignis (3, 5%), Carludovica pal- 
mata (\, 1.6%), IVelfia georgii (1 , 1.6%),and(/eo/i- 
oma sp. (1, 1.6%). Only 3 of the 63 tents were 
occupied; one had two bats and two each had sin- 
gle bats. As I noted in 1982, tents on Anthurium 
ravenii were often clumped on the same plant with 
an average of 2.5 tents per plant. 

At La Selva in July 1982, 43 Artibeus watsoni 
tents were located over a 5 -day period in the fol- 
lowing species of plants: Asterogyne martiana (33, 
77%), Geonoma congest a (6, 14%), Geonoma cu- 
neata (2, 5%), and an unidentified species of Cy- 
clanthaceae (2, 5%). One adult male A. watsoni 
was found under an Asterogyne martiana tent on 
the first day. On the fifth day an adult female with 
young was found under another A. martiana tent 
that had been unoccupied for the previous fovu" 
days, as was a third adult (not captured) under 
another A. martiana tent. All other tents were un- 
occupied. 

In 1974 1 surveyed approximately 10 km of trails 
at La Selva and found 29 tents on the following 
species of palms: Asterogyne martiana (19, 66%), 
Bactris wendlandiana ( 1 , 3%), Geonoma congesta 
(2, 7%) and Geonoma cuneata (7, 24%); all were 
unoccupied. Foster and Timm (1976) reported 
tents in these palms, although they were not able 
to associate bats with the tents. My recent studies 
at La Selva have confirmed that these tents were 
made by A. watsoni. 

At Finca Las Cruces in mid-August 1982, 13 
tents constructed by Artibeus watsoni were located; 



198 



HELDIANA: ZOOLOGY 





2 



'^ 



e 
o 

•a 

00 



Si 






eo| 
C * 

(/J c* 
. k» 

a T 
•2 o 

I a 

c g 
. u 

C f 
3 ■-> 
P « 

« to 

x: CO 

♦^ 

o S 

li 

-3 e 



o V 



TIMM: ARTIBEUS AND URODERMA 



199 




Fig. 6. Top, Dorsal view of the cyclanth, Carludovica palmata, showing the polygonal cuts; bottom, tent of Artibeus 
waisoni on C. palmata. 



200 



HELDIANA: ZOOLOGY 




Fig. 7. Photograph of adult female Artibeus watsoni and her subadult offspring roosting in a Carludovica leaf tent. 
Details of the polygonal cuts and folds can be seen in the background. 



one was occupied by an adult male. The tents were 
distributed on the following species of cyclanths: 
Asplundia euryspatha (6, 46%), Carludovica drudei 
(4, 31%), and Cyclanthus bipartitus (3, 23%). 

Artibeus watsoni uses a variety of species of plants 
and a wide array of leaf shapes for diurnal roosts. 
I have found four distinct styles of tents at a single 
locality (Corcovado). These styles include the sim- 
ple V-shaped cuts on bifurcated palms (fig. 5), cut- 
ting a few side veins on aroids to produce a round- 
ed pyramid, the elongated J-shaped cuts on banana 
and Heliconia leaves, and the polygonal cuts on 
Carludovica (figs. 6-7). For each distinct leaf shape, 
the cuts create a well-concealed diurnal roost. Ar- 
tibeus watsoni probably is an obligate tent-rooster, 
as it has only been found roosting under cut leaves. 

On several instances a bat occupied the same 
tent, or tents in close proximity, for two to three 
days in succession. Those tents might then remain 
unoccupied for several days in succession. Dis- 
turbed bats generally flew directly to another tent 
from 20 to 50 m away, or attempted to return to 
the tent where originally found. 



Tents generally are found clumped, both on a 
single plant if leaf morphology and age are appro- 
priate, and in restricted areas. Up to five tents have 
been found on a single Anthurium ravenii, and 
when present the mean number of tents was 2.5. 

At Parque Nacional de Corcovado, Choe and 
Timm (1985) found that Artibeus watsoni showed 
strong preference for Anthurium ravenii leaves that 
were medium sized, low within the plant, and grew 
closer to the ground than average A. ravenii leaves. 
Also at this site, Boinski and Timm (1985) doc- 
umented that squirrel monkeys {Saimiri oerstedi) 
were major predators on A. watsoni, with the adult 
male monkeys being the most successful at cap- 
turing bats. Additionally, double-toothed kites 
{Harpagus bidentatus) followed troops of foraging 
squirrel monkeys, using them as "beaters." When 
tent-making bats were flushed by the monkeys and 
escap)ed, they were routinely captured and con- 
sumed by the attending double-toothed kites. 

Artibeus watsoni has long been known to cut 
palm tents for diurnal roosts, although prior to 
this study little had been published on roosting 



TIMM: ARTIBEUS AND URODERMA 



201 




Fig. 8. Dorsal view of Scheelea rostrata showing the leaflets cut by Uroderma bilobatum to form a tent 



202 



HELDIANA: ZOOLOGY 



behavior of this species. The elongate J-shaped 
cuts made on bifurcated palms were first described 
and illustrated by Chapman (1932, p. 555). He 
stated that "both vanes of the leaf whence the bat 
flew were cut diagonally to the midrib of the leaf, 
so that their terminal portions drooped downward 
to form a tentlike shelter." Chapman appropri- 
ately termed these three-sided diurnal roosts 
"tents," and I have expanded the use of the word 
tents to include all modified leaves by bats. 

Barbour's (1932) original description of Uro- 
derma bilobatum cutting palm leaves for roosts 
also provides a secondhand report (p. 308) by H. 
C. Clark stating that "Clark has just found for the 
first time a youngish coconut palm, a single leaf 
of which was being cut by bats of the genus Uro- 
derma in a very similar way." The common use 
of young coconut palms {Cocos nucifera) by Ar- 
tibeus watsoni in Costa Rica, coupled with the total 
lack of evidence that Uroderma bilobatum uses 
juveniles of this palm, leaves that small, or roosts 
that close to the ground, suggests that the tents 
seen by Clark were in fact made by A. watsoni. 
Artibeus watsoni also is abundant on Barro Col- 
orado Island, Panama. Apparently no voucher 
specimens of the bats were preserved at the time. 
Allen (1939, p. 69) reported "a specimen of A. 
watsoni that was hanging by day from the under- 
side of a banana leaf" Perhaps the natural-looking 
folds caused by the cuts running parallel to the 
midrib were not noticed at the time. Ingles (1953) 
reported on tents of A. watsoni in two species of 
Geonoma on Barro Colorado Island; one tent was 
occupied by three individuals. Thomas's fruit-eat- 
ing bat has been found roosting in an artificial tent, 
an inverted hanging box. Wilson (1970) reported 
that several females raised young in the comer of 
a suspended box on Barro Colorado Island. 



Uroderma 

Uroderma bilobatum Peters, 1865 

Uroderma bilobatum has been given the dis- 
tinctive "common" name of Peters's tent-making 
bat. Tents constructed by Uroderma bilobatum 
were seen at three separate localities in Costa Rica 
during the summers of 1982, 1984, and again in 
1986, the first at Bosque Brrancia near Cuarto 
Cruces in the Pacific lowlands of northwestern 
Costa Rica, the second at Corcovado on the Osa 
Peninsula, and the third at La Selva in the Carib- 
bean lowlands. 



On 25 August 1982 a colony of Uroderma bi- 
lobatum was roosting under a modified frond of 
the palm Scheelea rostrata at Bosque Brrancia. 
The colony included an adult male with enlarged 
testes and four adult females. 

The Scheelea rostrata frond in which the colony 
of Uroderma bilobatum roosted was a mature leaf, 
approximately 6.5 m in length. The bats were 
hanging approximately 4.5 m off"the ground; most 
were clustered together, although a few were spread 
out over 50 cm of the frond. The cut leaflets started 
at about 3.5 m off" the ground and proceeded up 
the frond for the next 2.5 m (fig. 8). The general 
pattern of the cut leaflets was a tapering effect, with 
the cuts on the lowermost leaflets being farthest 
from the midrib. Leaflets along the proximal 2 m 
and the distal 50 cm were unmodified. Only the 
midrib of the leaflets was cut. Each leaflet had a 
distinctive V-shaped fold at its base where it was 
attached to the midrib. The bats were hanging 
from the leaflets rather than the midrib. 

From the dorsal aspect of the leaf, the proximal 
portion of the tent (cut leaflets) extended 50 cm 
further down on the right side than on the left to 
include 10 basal leaflets whose opposites on the 
left were unaltered. The basalmost cut leaflet was 
cut 34 cm from the midrib. Proceeding distally, 
the length of the unmodified basal portion of each 
leaflet decreases. The basalmost cut leaflet on the 
left was cut 19 cm from the midrib. The overall 
appearance of the tent was a sharp, convergent 
taper for the next meter. Following this section, 
there was a 7 5 -cm section in which the cuts were 
close to the midrib (within 3 cm). On the distal- 
most 30 cm of the tent, the leaflets were cut closer 
to the midrib on the left side than on the right. 
Similar tents, each housing a colony of Uroderma 
bilobatum, were found in a large stand of Scheelea 
rostrata at Corcovado in 1984, and William A. 
Haber (pers. comm.) informed me that he has seen 
similarly cut leaves in the same species of palm at 
Cahuita (09°44'N, 82°49'W) in the Caribbean low- 
lands of southeastern Costa Rica in 1 984. 

In June of 1982 and again in March of 1986, I 
found numerous banana leaves cut by Uroderma 
bilobatum just to the north of the field station at 
La Selva. The midrib on vertical leaves was cut 
to the extent that the distal portion of the leaf 
collapsed downward to form a two-sided tent (fig. 
9). Severing the midrib on vertical leaves had the 
effect of folding the leaf back upon itself creating 
a tight, dark crevice at the fold where the bats 
roosted (fig. 1 0). In addition to severing the mid- 
rib, the bat cut a large V-shaped pattern running 



TIMM: ARTIBEUS AND URODERMA 



203 





Fig. 9. Left, Dorsal view of a banana leaf showing the cut midrib and the large V-shaped cuts through the side 
nerves; right, tent of Uroderma bilobatum made from a banana leaf. 



from the midrib to the base of the leaf. The side 
veins and interconnected tissues were partly sev- 
ered. However, because the leaf stood nearly ver- 
tical, these V-shaped cuts did not cause further 
folding of the leaf The only cut creating the tent 
was that of the midrib. 



In 1982 five tents in widely separated banana 
leaves were located (table 4). One was occupied 
by eight Uroderma bilobatum, which included one 
adult male with enlarged testes and seven females. 
In 1986 eight tents were observed in the same 
banana patch. On this occasion eight U. bilobatum 



204 



FIELDIANA: ZOOLOGY 




TIMM: ARTIBEUS AND URODERMA 



205 



tents were clustered in three clumps of bananas. 
Only one tent was occupied; it contained 1 3 bats. 

This folded broad-leaved style of tent is un- 
doubtedly the tent style illustrated by Walker ( 1 960, 
p. 30) in his photograph of roosting Uroderma 
bilobatum, although he did not describe it nor 
mention where it was observed. Interestingly, I 
have searched several dozen banana and larger 
Heliconia groves throughout Costa Rica and Ec- 
uador specifically looking for this style of tent, and 
none were observed. Uroderma bilobatum is an 
abundant and widespread species in the lowlands, 
but employment of this particular style of tent 
appears spotty as I have not observed it elsewhere. 

In Panama, Barbour (1932) found that colony 
size under a single cut leaf of Prichardia pacifica 
varied from a few bats to 56. Prior to Barbour's 
discovery that Uroderma altered leaves, Goldman 
(1920, p. 199) stated of these bats in Panama. 

In the forest near Gatun Uroderma biloba- 
tum was located several times, a few in a 
place, clinging during the day in clusters to 
the midribs on the under sides of large palm 
leaves. They usually choose darkened spots 
where the leaf was folded over, or over- 
hanging pinnae shut out much of the light. 

Burt and Stirton (1961) reported U. bilobatum in 
El Salvador roosting beneath the leaves of bananas 
and coconut palms. I strongly suspect that the bats 
had altered these leaves and that these authors had 
failed to notice it or failed to associate the hanging 
bats with the damage to the leaves. Bloedel (1955, 
p. 234) stated of Uroderma bilobatum in Panama 
that: 

I observed these bats only in their palm-leaf 
tents. ... In the latter part of March most 
females have nursing young, and are roost- 
ing in clusters of 20 to 40, while the males 
are separated from them, usually solitary or 
in small groups of from 2-5. 

In Trinidad, Goodwin and Greenhall (1961, p. 
254) found them 

on the under side of the fan-shaped leaves 
of certain palm trees, especially the carat 
palm (Sabal glaucescens). . . . The bat makes 
a series of cuts across the pleated surface of 
a leaf, causing half of the leaf to bend at an 
angle to form a protected retreat. 



Table 4. Individual measurements (in cm or de- 
grees) collected from four tents constructed by Uroderma 
bilobatum on banana, Musa x paradisiaca, in Costa 
Rica. 



Blade 


Petiole 


Cut 
midrib 




Length Width 


angle 


from base 


Remarks 


210 95 


40 


117 


Not occupied 


220 90 


70 


90 


8 U. bilobatum 




80 




1 U. bilobatum 




80 




Not occupied 


210 95 






Not occupied 



They were found roosting in colonies of 1 or more 
individuals. In Suriname, Husson (1962, p. 161; 
1978, p. 143) collected three pregnant female U. 
bilobatum "in a plantation where they were found 
hanging on the imder side of a leaf of [a] so-called 
'paloeloe,' Ravenala guyariensis." In Nicaragua, 
Jones ( 1 964, p. 507) collected four female U. bi- 
lobatum that "hung together about 10 feet above 
the ground in the 'tent' formed by a cut palm frond. 
Each was pregnant with a single embryo." In Gua- 
temala, Dickerman et al. (1981, p. 409) reported, 
"Palm leaf tents were frequently found occupied 
by one to seven individuals, but nursing females 
were usually found alone or with juveniles." 

Leaves selected by Uroderma bilobatum for tents 
are all large, and of a variety of shapes. The large 
V-shaped pattern cut into the leaves is a charac- 
teristic of Uroderma tents. Artibeus watsoni also 
uses a variety of leaf shapes for tents; however, 
the nature of the cuts and style of tent created 
vary with leaf shape. Uroderma bilobatum, on 
the other hand, makes patterned cuts that appear 
to be an innate response to large leaves, regardless 
of the shape. As noted in the banana tents, a single 
cut across the midrib creates the tent, and the 
V-shaped pattern had no effect upon tent shape; 
this perhaps represents wasted effort by the bats. 

There are a few records of Uroderma bilobatum 
being found roosting in a hollow tree and one "un- 
der the eave of house" (Davis, 1968, p. 695). In 
all cases these have been of single individuals, and 
I suspect represent either recently dispersed young 
that have yet to join a breeding colony or bachelor 
males. Uroderma bilobatum roosting under a cut 
Prichardia leaf was illustrated by Kunz (1982). A 
colony of U. bilobatum roosting under a banana 
leaf tent was illustrated by Keller ( 1 986). Mac- 
donald's ( 1 984, p. 806) photograph of two tent- 
making bats under a Heliconia tent is erroneously 
labeled Uroderma bilobatum. These bats are ac- 



206 



HELDIANA: ZOOLOGY 



Fig. 11. Tent of Vroderma 
magnirostrum on the pinnately 
leafed palm Astrocaryum muru- 
muru. 




tually a small species of Artibeus, probably A. 
phaeotis; the tent style and size and coloration of 
the bats are typical oi A. phaeotis. 



Uroderma magnirostrum Davis, 1968 

Although Vroderma magnirostrum is a widely 
distributed bat found from Mexico to Bolivia, it 
was not recognized as a species distinct from U. 
bilobatum until 1968 (Davis, 1968), and few spec- 
imens are represented in collections. When W. B. 
Davis described this new species he commented 



that although it was widely distributed only seven 
specimens had been collected prior to the wide- 
spread use of mist nets in the 1960s, and that all 
specimens available to him had come from local- 
ities less than 1000 feet in elevation. He stated 
that "These facts strongly suggest basic differences 
in the habits of the two species and that those bats 
with a deep rostrum are not 'tent-makers' as are 
members of the species Uroderma bilobatum" 
(Davis, 1968, p. 678). There have been no reports 
on the behavior or ecology of U. magnirostrum. 
On 25 July 1985 Barbara L. Clauson discovered 
a colony of two male and three female Uroderma 



TIMM: ARTIBEUS AND URODERMA 



207 



magnirostrum roosting under the cut leaflets of the 
pinnately leafed palm, Astrocaryum murumuru. 
The single occupied tent found was on the ridge 
above Hacienda Amazonia at 825 m in south- 
eastern Peru. The entire colony was collected by 
John W. Fitzpatrick. On 3 November Clauson re- 
turned to the site to measure ,»ie leaf and noted 
an additional cut leaf in the same plant. 

When first observed, four bats were hanging to- 
gether and one was hanging several centimeters 
away. When observed an hour later, all five were 
hanging together in a tight cluster approximately 
7.5 m off the ground. The bats were hanging from 
the leaflets rather than the midrib, approximately 
200 cm from the tip of the leaf The colony in- 
cluded one adult male with enlarged testes, two 
adult females, and two subadults, one female and 
one male. 

The roosting structure of Uroderma magniros- 
trum was in a pinnately leafed palm (fig. 1 1 ). The 
bats severed the leaflets along the upper two-thirds 
of the leaf; those along the lower third were un- 
altered, as were the leaflets at the very tip. As the 
leaflets proceeded up the tent they were severed 
closer to the midrib forming an elongate, conver- 
gently tapering tent (fig. 1 1). The general appear- 
ance of the U. magnirostrum tent is similar to that 
described herein for U. bilobatum on the pinnately 
leafed palm Scheelea rostrata. 

The Astrocaryum frond in which the colony of 
Uroderma magnirostrum roosted was a mature leaf, 
approximately 6.1 m in length and 1.9 m in width 
at the widest point, with the petiole 1.1m long. 
The leaf left the trunk (d.b.h. .4 m) at 3 m from 
the ground and hung at an angle of approximately 
50°. The bats were hanging approximately 7.5 m 
off" the ground. The cut leaflets started at 1.5 m 
from the lowest leaflet and proceeded up the frond 
for the next 2.9 m to nearly the tip (fig. 1 1). The 
cuts on the lowermost leaflets were furthest from 
the midrib. The lowest severed leaflets were cut 
up to 34 cm from the midrib, whereas the distal 
leaflets were severed only 2 cm from the midrib 
of the leaf The midribs of the leaflets were cut 
causing the distal portion of the leaflets to fold 
downward. Leaflets along the proximal 1.5 m and 
the distal .5 m were unmodified. The trunk, pet- 
iole, and midrib of this palm were covered with 
sharp, penetrating spines several centimeters in 
length. After 14'/2 weeks this tent was still alive 
and green, most of the leaflets appearing as fresh 
in November as they did in July. 

This Astrocaryum contained a second cut leaf 
that was unoccupied. This roost was also in a ma- 



ture leaf which was an older leaf than the occupied 
tent, with many broken, yellowed, and brown leaf- 
lets. The leaf was approximately 6 m in length and 
2 m in width at the widest point, with a petiole .8 
m long. The leaf left the trunk at 2.9 m from the 
ground and hung at an angle of approximately 60°. 
Cuts were distributed asymmetrically along the 
length of the leaf The cut leaflets on the left side 
of the leaf started 3.05 m from the lowest leaflet 
and proceeded up the frond for the next 1 .3 m, to 
.88 m from the tip. The cut leaflets on the right 
side of the leaf started 3.25 m from the lowest 
leaflet and proceeded up the frond for the next 
1.49 m, to .49 m from the tip. The cuts on the 
lower leaflets were farthest from the midrib. The 
lowest severed leaflets were cut up to 35 cm from 
the midrib, whereas the distal leaflets were severed 
as close as 1.5 cm from the midrib. Leaflets along 
the proximal 3.05 m and 3.25 m and the distal 
.88 m and .49 m were unmodified. 

I propose the common name of Davis's tent- 
making bat for this species. 



Conclusions 

A review of the literature on tent-making bats 
contains some 32 primary references covering the 
55-year period from 1932 through 1986. Surpris- 
ingly, we actually know very little about the bi- 
ology of these bats. 

As late as 1975 Eisentraut was yet doubting th^t 
bats were cutting leaves to make tents, stating: 

. . . observers maintain that the bats form 
these tentlike structures themselves, by 
making a series of holes running across the 
middle of a large palm leaf The bats then 
supposedly bend the outer half of the leaf 
around, so they can then rest inside this 
'tent'. . . . On the basis of personal obser- 
vations in tropical regions in Africa, I tend 
to believe instead that these holes were made 
by insect larvae while the leaves were still 
rolled up. A storm can then easily break the 
leaf along the line of holes and form the tent 
roof which is so convenient for the bats (Ei- 
sentraut, 1975, p. 142). 

Eisentraut, by his own admission, had never seen 
a bat tent. I believe that if he had, he would have 
come to the same conclusion Thomas Barbour did 



208 



HELDIANA: ZOOLOGY 



nearly a half century earlier, that the bats and not 
insects were making the cuts. 

Although we have yet to actually observe bats 
cutting leaves to form the roosting structures de- 
scribed herein, I hope the volume of data pre- 
sented here and in my other works establishes for 
a fact that many species of small and medium sized 
stenodermines are indeed tent-makers. The ob- 
servations presented represent data collected from 
several hundred tents located over a 1 5 -year pe- 
riod. Several facts consistently emerge between my 
observations and those independently corrobo- 
rated by others. Bats of the genera Artibeus and 
Uroderma (as well as Ectophylla, Mesophylla, and 
Vampyressa) roost under cut leaves. These leaves 
may be on a wide variety of species of plants, but 
generally the shape of the leaves is similar. The 
shape of the cuts is very characteristic for each 
species of bat and the patterns and styles of tents 
created by the bat species are consistent. 

The behavioral repertoire associated with tent- 
making in bats certainly evolved more than once, 
as evidenced by the patchy distribution of tent- 
making species with the chiropteran suborders 
Megachiroptera and Microchiroptera. Within the 
Megachiroptera, a single species of tent-maker is 
known, Cynopterus sphinx. Within the Microchi- 
roptera, tent-makers are known only from one 
subfamily of the Phyllostomidae, the Stenoder- 
minae. The Stenoderminae constitute an ex- 
tremely speciose and diverse group of bats, with 
more than 30 species currently recognized. Tent 
construction within stenodermines may be a trait 
that evolved once, twice, or as many as three times. 
The Artibeus- Uroderma group are sister genera and 
form one clade of the tent- making repertoire. Sec- 
ondly, the Mesophylla- Vampyressa group are sis- 
ter genera (and perhaps should be considered con- 
generic) and would constitute the second clade. 
Finally, Ectophylla would constitute a third lin- 
eage. The relationship between these three lineages 
is uncertain and warrants further investigation. 

Knowing that bats modify the leaves of several 
species of plants to produce diurnal roosting struc- 
tures led to the following questions: (1) Are bats 
selecting specific species of plants for tents? (2) 
What styles of tents are cut by bats and do these 
differ between species? (3) Do bats select for a 
particular angle, size, or shape of leaf for diurnal 
tents? (4) Are leaves selected preferably in larger 
clumps or smaller clumps? (5) Are older or youn- 
ger leaves selected? (6) Are leaves of a particular 
height class selected? (7) Are leaves that are not 
adjacent to solid objects selected? (8) What do 



typical tents look like? (9) How and why did tent 
construction evolve? 

On occasion I have found "cheaters," species of 
bats roosting in a tent made by another species. 
Is cheating an evolved strategy of roost site selec- 
tion of some bats? 

Bats of the genera Artibeus and Uroderma ac- 
tively modify leaves to produce diurnal roosting 
structures, but by biting the tissue between veins 
along the midrib and leaving the midrib and most 
veins intact, do not kill the leaves. The resulting 
tent is available for use as a roost for an extended 
period of time; one was observed in use for more 
than 60 days. Bats select for specific sizes and 
shapes of leaves. Tents provide concealment from 
predators and protection from the rain, wind, and 
sun. This type of roost offers the additional ad- 
vantage that the bats are warned about the ap- 
proach of a potential predator, because even slight 
movements of the leaf stem or the leaf itself are 
transmitted as magnified vibrations to the roosting 
bat. Tents may provide bats with suitable roosting 
sites that would not otherwise be available in close 
proximity to prime food resources. 

One of the most productive areas for future re- 
search will be exploring aspects of the biology of 
these bats from an evolutionary perspective. Fu- 
ture subjects I will be addressing include the role 
of tent roosting in controlling ectoparasites and 
the correlation between complexity of tents and 
social systems in these bats. I believe that tent- 
making originated as an antipredation strategy and 
has since, secondarily, evolved to play a major 
role in controlling ectoparasites and in social be- 
havior. 

Many factors influence the choice of roost site 
selection by bats. Included among these are vul- 
nerability to predation, physical stability of the 
site, proximity to food sources, and general ap- 
propriateness of the nest microenvironment for 
the rearing of young. It seems likely that tent con- 
struction requires considerable time and energy 
expenditure by bats, attesting to intense selection 
pressures involved. 



Acknowledgments 

I thank Eduardo Lopez Pizarro and El Depar- 
tamento de Vida Silvestre and Fernando Cortes 
and Servicio de Parques Nacionales of Costa Rica 
for making this study possible. The Organization 
for Tropical Studies (OTS), Rebecca Butterfield, 



TIMM: ARTIBEUS AND URODERMA 



209 



William A. Haber, Gary Hartshorn, Charles E. 
Schnell, and Joe M. Wunderle are gratefully ac- 
knowledged for assistance with logistics. Robert J. 
Izor assisted with field logistics in Costa Rica and 
Peru, and John W. Fitzpatrick assisted in Peru. In 
Ecuador I thank the Comandancia General del 
Ejercito Ecuatoriano, the Corporacion Estatal Pe- 
trolera Ecuatoriana, and the Ministerio de Agri- 
cultura y Ganaderia for making our studies there 
possible. Luis Albuja, Ramiro Barriga, Angelitos 
Garrett, Myriam Ibarra, Gustavo Orces, and Don- 
ald J. Stewart provided logistic assistance in Ec- 
uador. Kerry A. Barringer, William C. Burger, 
Thomas B. Croat, Robin B. Foster, Barry Ham- 
mel, and Timothy Plowman provided identifica- 
tions or confirmed identifications of the plants. 
Barbara L. Clauson, Alfred L. Gardner, Lawrence 
R. Heaney, Karl F. Koopman, Thomas H. Kunz, 
Bruce D. Patterson, and Timothy Plowman pro- 
vided valuable suggestions on earlier drafts of the 
manuscript. My wife, Barbara, provided superb 
assistance with all aspects of this project, including 
providing several of the photographs used and all 
data on Uroderma magnirostrum. Rosanne Mie- 
zio prepared the illustrations. Nina Cummings, 
Ron Testa, and Diane White expeditiously and 
cheerfully executed my photography requests. This 
project was funded in part by grants from the Rice 
Foundation of Chicago, the National Science 
Foundation [INT-8303 194], National Geographic 
Society, and Field Museum of Natural History. I 
especially thank Mr. and Mrs. Arthur A. Nolan, 
Jr. for their continuing support of my research. 

This paper is dedicated to Philip Hershkovitz 
in recognition of his contributions to Neotropical 
mammalogy, and most especially for the friend- 
ship he has shown me. 



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Appendix 

List of scientific names of plants mentioned in the text and used by Neotropical bats for tent con- 
struction. 



Anacardiaceae 

Anacardium excelsum (Bertero & Balbis) Skeels 



Araceae (cont'd.) 
Monstera lechleriana Schott 
Xanthosoma sp. 



Araceae 



Cyclanthaceae 



Anthurium caperatum Croat &. Baker 
Anthurium ravenii Croat & Baker 



Asplundia euryspatha Hari. 
Carludovica dmdei Masters 



TIMM: ARTIBEUS AND URODERMA 



211 



Cyclanthaceae (cont'd.) 
Carludovica palmata R. & P. 
Cyclanthus bipartitus Poit. 



Marantaceae 

Calathea insignis Petersen 

Musaceae 

Heliconia imbricata (Kuntze) Baker 
Heliconia latispatha Benth. 
Musa X paradisiaca L. 

Plenakospermum guyanense Endl. (syn. Ravenala 
guyanensis Petersen) 



Palmae 

Asterogyne martiana (H. Wendl.) H. Wendl. ex 

Hemsley 
Astrocaryum murumuru Mart. 
Bactris wendlandiana Burret 
Cocos nucifera L. 

Geonoma congesta H. Wendl. ex Spruce 
Geonoma cuneata H. Wendl. ex Spruce (syn. G. 

decurrens H. Wendl.) 
Geonoma oxycarpa Martius (syn. G. binervia 

Oerst.) 
Livistona chinensis (Jacq.) R. Br. ex. Mart. 
Prichardia pacifica Seem. & H. Wendl. 
Sabal mauritiiformis (Karsten) Griseb. &. H. 

Wendl. ex Griseb. (syn. S. glaucescens Lodd. ex 

H. E. Moore) 
Scheelea rostrata (Oersted) Burret 
Welfia georgii H. Wendl. ex Burret 



212 



FIELDIANA: ZOOLCX5Y 



Comparative Ultrastructure and Evolutionary Patterns 

of Acinar Secretory Product 

of Parotid Salivary Glands in Neotropical Bats 

Carleton J. Phillips, Toshikazu Nagato, and Bernard Tandler 



ABSTRACTS 

Secretory products produced by acinar cells of the parotid salivary glands of 1 5 species of 
Neotropical bats {Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, T. 
sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium, 
Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis, 
and Molossus molossus) were compared by transmission electron microscopy. Extensive inter- 
and intrageneric differences were found in the ultrastructure of the mature acinar secretory 
granules. This variation in secretory cell product exceeded any previously reported intraordinal 
phenotypic variation at the cellular level, but was in keeping with previously reported bio- 
chemical data on salivary protein polymorphism in primates and rodents. Data from molecular 
biology and systematics lend support to the hypothesis that the microscopic variations are 
directly representative of genie differences among species. It also is postulated that intrageneric 
microscopic differences at least partly are due to neutral (nonfunctional) differences in molecular 
structure or charge (or both) rather than evolutionary selection. Among the phyllostomids, a 
general trend in parotid acinar cell product was found in Artibeus and Ariteus, in which a 
decrease in enzymatic content of the product could be correlated with ultrastructural differences. 
The secretory product in Artibeus and Ariteus also differed significantly from that of the genus 
Sturnira. and it is proposed that the phenotypic differences between Sturnira and the other two 
stenodermatines represent a major genetic difference of systematic importance. The ultrastruc- 
tural appearance and substructure of the parotid acinar secretory granules could not be con- 
sistently correlated with diet alone, although insectivorous-animalivorous SF)ecies have enzyme- 
rich, mostly electron-dense granules, whereas two fruit bats, Artibeus and Ariteus, have pale, 
enzyme-poor parotid granules. 



Productos de secrecion producidos por celulas acinares de las glandulas salivales de 1 5 esF>ecies 
de murcielagos neotTopicales (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia 
bidens, T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira 
lilium, Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida bra- 
siliensis y Molossus molossus) fueron comparados mediante microscopio electronico de trans- 
mision. Extensivas diferencias inter- e intragenericas fueron encontradas en la estructura de 
granulos glandulares acinares maduros. Esta variacion en productos de celulas secretoras excedio 



From the Department of Biology, Hofstra University, 
Hempstead, NY 11550 (Phillips); and Department of 
Oral Biology, School of Dentistry, Case Western Reserve 
University, Cleveland, OH 44106 (Nagato and Tandler). 
Dr. Nagato's present address is Department of Oral and 
Maxillofacial Surgery, Ehime University School of Med- 
icine, Shizukawa, Shigenobu, Onsen-gun, Eshime 79 1 -02, 
Japan. 

PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 213 



cualquier variacion fenotipica intraordinal previamente reportada a nivel celular, pero estuvo 
en armonia con datos bioquimicos previamente reporlados de polimorfismo en proteinas sa- 
livales de primates y roedores. Datos de biologia molecular y sistematica proveen apoyo a la 
hipotesis de que las variaciones microscopicas son direclamente representativas de diferencias 
geneticas entre especies. Es lambien postulado que diferencias microscopicas intragenericas, al 
menos parcialmente, son debidas a diferencias neulrales (no funcionales) en estructura molecular 
o carga (o ambas) mas bien que a seleccion evolutiva. Entre los filostomidos, una tendencia 
general en el producto parotido de las celulas acinares fue encontrado en Artibeus y Ariteus, en 
los cuales una disminucion en contenido enzimatico del producto p>odria estar relacionado con 
diferencias ultraestructurales. El producto secretado en Artibeus y Ariteus tambien se diferencio 
significativamente de aquel del genero Sturnira y es propuesto que las diferencias fenotipicas 
entre Sturnira y los otros dos stenodermatinos representan una mayor diferencia genetica de 
importancia sistematica. La apariencia ultraestructural y subestructura de los granulos secre- 
torios parotidos acinares podria no estar consistentemente correlacionada solo con la dieta, 
aunque especies insectivoras-animalivoras tienen granulos ricos en enzimas mayormente densos 
en electrones, mientras que los murcielagos frugivoros, Artibeus y Ariteus, tienen granulos 
parotidos palidos y pobres en enzimas. 



Produtos secretorios produzidos por acinos das glandulas parotidas de 15 especies de mor-' 
cegos neotropicos (Pteronotus parnellii, Phyllostomus elongatus, P. latifolius, Tonatia bidens, 
T. sylvicola, Trachops cirrhosus, Glossophaga soricina, Leptonycteris sanborni, Sturnira lilium. 
Artibeus jamaicensis, Ariteus flavescens, Eptesicus lynni, E. brasiliensis, Tadarida brasiliensis, 
e Molossus molossus) foram comparados atraves da microscopia de transmissao eletronica. 
Vastas diferengas intra- e intergenericas foram encontradas na ultra-estrutura dos granulos 
maduros dos acinos secretorios. Esta variafao no produto das celulas secretorias supera qualquer 
varia^ao fenotipica intraordinal previamente relatada para o nivel celular, porem concorda 
com as rela96es publicadas sobre dados bioquimicos do polimorfismo de proteinas salivares 
em primatas e em roedores. Dados sistematicos, e de biologia molecular, apoiam a hipotese 
que variances microscopicas sao diretamente representativas das diferen^as geneticas entre 
especies. Propoese tambem, que estas diferen^as microscopicas intragenericas sao ao menos 
parcialmente causadas por diferen9as neutras (i.e., naofuncionais) nas estruturas moleculares 
ou nas suas cargas eletricas (ou ambas), ao inves de serem consequencias da sele^ao evolutiva. 
Entre os morcegos da familia Phillostomidae, foi encontrado em Artibeus e em Ariteus um 
padrao geral nos produtos dos acinos parotideos, onde uma redu9ao do conteudo enzimatico' 
e correlacionado a diferencas nas ultraestruturas dos granulos produzidos. Os produtos secre- 
torios em Artibeus e em Ariteus sao significantemente diferentes dos produtos do genero Sturnira. 
e prop6e-se que as diferen9as fenotipicas entre Sturnira e os outros dois stenodermatinos 
representam uma grande diferen9a genetica, de importancia sistematica. A aparencia ultraes- 
trutural, e a subestrutura dos granulos secretorios dos acinos parotideos, nao se correlacionam 
consistentemente com a dieta por si, apesar de que as especies insetivoras-animalivoras possuem 
granulos ricos em enzimas e densos em eletrons, equanto que dois morcegos frugivoros, Artibeus 
e Ariteus, (wssuem granulos parotideos palidos e com poucas enzimas. 



Introduction 

Neotropical bats are extremely diversified; ex- 
isting species represent perhaps the most outstand- 
ing mammalian example of ecomorphological ad- 
aptation. Dentitions, jaw morphology, brains, 
kidneys, tongues, and digestive tracts are only a 
few examples among the anatomical features that 
have been investigated in recent years (e.g., Phil- 
lips, 1971; Forman, 1972; Phillips et al., 1977, 



1984; Freeman, 1979, 1981; Eisenberg «t Wilson, 
1978; Studier et al., 1983). The major salivary 
glands are yet another system that has attracted 
attention, primarily because histological, ultra- 
structural, and histochemical investigations have 
consistently revealed striking interspecific differ- 
ences and unusual histological and secretory fea- 
tures (Wimsatt, 1956; DiSanto, 1960; Junqueira 
& Fava de Moraes, 1965; Junqueira et al., 1967, 
1973; Phillips et al., 1977; Mineda, 1977, 1978; 



214 



HELDIANA: ZOOLOGY 



Pinkstaff et al., 1982; Tandler & Cohan, 1984; 
Nagatoetal., 1 984; Tandler & Phillips, 1985; Phil- 
lips & Tandler, 1985, 1987; Tandler et al., in press). 

Mammalian salivary glands are highly complex 
organs that not only secrete digestive enzymes but 
also can secrete hormones, antibodies, and toxins, 
to name but a few known products (e.g., Tandler, 
1972; Hand, 1980b). Data are available on the 
biochemistry of saliva in humans and common 
laboratory species; but very little is known about 
the specific biochemistry of the parotid acinar se- 
cretory granules except for laboratory rats in which 
some of the proteins have been characterized (Ro- 
binovitch & Sreebny, 1969; Ball, 1974; Wallach 
et al., 1975). The complex structure and function 
of salivary glands is underscored by data from 
studies of Neotropical bats, which recently have 
been analyzed by both transmission electron mi- 
croscopy and histochemistry. For example, the ac- 
cessory submandibular gland of Trachops cirrho- 
sus has been shown to differ histologically from 
any known mammalian salivary gland, with the 
exception of the same gland in Megaderma lyra 
and M. spasma. Megaderma lyra is an Old World 
ecological equivalent of Trachops (Phillips &. 
Tandler, 1985, 1987; Phillips et al., 1987). Both 
of these unrelated species feed on frogs (Lekagul 
& McNeely, 1977; Tuttle & Ryan, 1981), which 
possibly has been a factor in the convergent evo- 
lution of their submandibular glands. A previously 
unknown cellular organelle has been described in 
another Neotropical bat, Tonatia sylvicola (Na- 
gato et al., 1984). In this species, the presence of 
xmique crystalloid smooth endoplasmic reticulum 
in seromucous acinar cells is sex-linked (being 
found only in submandibular acinar cells in males); 
a steroid product produced by this organelle pos- 
sibly serves as a species-isolating mechanism or 
as part of a chemo-behavioral system, or both. 
Lastly, a comparative investigation of the secre- 
tory product in seromucous acinar cells in sub- 
mandibular glands of five species ofArtibeus has 
revealed that the ultrastructural characteristics of 
secretory products can have systematic implica- 
tions (Tandler et al., 1983, 1986). This study was 
of particular interest because the salivary gland 
data matched genie data independently derived by 
Koop and Baker (1983). 

For the present investigation we surveyed par- 
otid acinar cell secretory products in a selected 
group of 1 5 species of Neotropical bats. Our group 
comprised four families (Mormoopidae, Phyllo- 
stomidae, Vespertillionidae, and Molossidae) and 
included a group of species in which dietary habits 



ranged from insectivory and animalivory to om- 
nivory and frugivory. This investigation is the first 
comprehensive interspecific survey of secretory 
product ultrastructure, and addresses the follow- 
ing questions: ( 1 ) what is the range of variation in 
secretory products; (2) what are the evolutionary 
patterns in parotid secretory product; and (3) what 
systematic conclusions can be reached by com- 
parative ultrastructural analysis? 



Materials and Methods 

Numbers and sex of specimens used in the pres- 
ent study are given in the Appendix. All of these 
bats were collected during fieldwork in Mexico, 
Jamaica, and Suriname. Voucher specimens for 
all species and collecting localities are deposited 
in the mammal collections of either the Carnegie 
Museum of Natural History or The Museum, Tex- 
as Tech University. Bats typically were collected 
at night with mist nets, and were kept overnight 
without food until they were killed between 0900 
and 1 200 the following morning. The animals were 
anesthetized with 0.25 ml of sodium pentobarbital 
(50 mg/ml, intraperitoneally) and the salivary 
glands removed, placed on dental wax, flooded 
with freshly mixed fixative, and diced into pieces 
measuring approximately 1 mm'. 

Two fixation protocols were used at different 
times during the project. Specimens collected in 
Mexico and Jamaica were fixed in 2% glutaral- 
dehyde in 0. 1 M phosphate buffer and then stored 
unrefrigerated in fresh fixative. The specimens col- 
lected in Suriname were fixed in a modified tri- 
aldehyde-dimethylsulfoxide (dmso) mixture, first 
described by Kalt and Tandler (1971) and later 
modified slightly for fieldwork (Phillips, 1 985). The 
trialdehyde fixative, consisting of 3% glutaralde- 
hyde, 1% paraformaldehyde, 0.5% acrolein, 2.5% 
DMSO, and 1 mM CaClj in a 0.05 M cacodylate 
buffer and sucrose at pH 7.2, proved superior to 
the simple glutaraldehyde fixative in that (1) a 
higher percentage of tissues proved to be ade- 
quately fixed for study, and (2) the mitochondria 
tended to remain intact instead of being disrupted. 
Although the composition of the fixatives was dif- 
ferent, we have not found any evidence that these 
differences introduced undesirable artifacts that 
would influence our analysis. Additional details 
about the techniques can be found in both Phillips 
(1985) and Nagato et al. (1984). 

To remove unbound aldehydes, the tissue blocks 
were subjected to prolonged washing in phos- 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



215 



acinar lumen. 




acinar cell 



secretory 
graauLej 



striated Lntercalated acinus 

duct duct 

Fig. I . Schematic diagram of a secretory unit in a typical parotid gland. A cluster of secretory cells, arranged 
around a lumen to form an acinus, drains into an intercalated duct that in turn empties into a striated duct. In reality, 
more than one acinus may be associated with an intercalated duct, which itself may be branched. The acinar cells 
elaborate the primary saliva, which has plasma-like concentrations of electrolytes, and add their organic secrelory 
products to it. The precise role of the intercalated duct cells has not been established, but they probably play a role 
in secretion and transport. The principal function of the striated ducts is resorption of electrolytes, mainly sodium, 
from the primary saliva, rendering it hypotonic. 



phate-buffered sucrose. The blocks were postfixed 
for two hours in phosphate-buffered 2% OSO4, 
rinsed in distilled water, soaked overnight in cold 
aqueous 0.25 uranyl acetate, rinsed again in dis- 
tilled water, dehydrated in ascending concentra- 
tions of ethanol, passed through propylene oxide, 
and embedded in Epon-Maraglas (Tandler & Wal- 
ter, 1977). Thin sections were doubly stained with 
methanolic uranyl acetate (Stempak & Ward, 1 964) 
and lead citrate (Venable & Coggeshall, 1965). All 
sections were examined in a Siemens Elmiskop la 
transmission electron microscope (tem). Semithin 
sections ( 1 nm) used for orientation were stained 
with toluidine blue (Bjorkman, 1962) and exam- 
ined in a Zeiss Ultraphot. 

Tissue samples used for comparisons were se- 
lected carefully from among available tissue blocks. 
We generally avoided edges of tissues where me- 
chanical trauma often affects not only the cellular 
ultrastructure but also the microscopic appearance 
of the secretory granules. In selecting representa- 
tive "mature" granules we took into account the 
full array of inter- and intracellular variability as 
well as the often complex substructural geometry 
of the granules. Selection of the most representa- 
tive granules admittedly was subjective, but based 
on our experience, four principal criteria were 
applied: (1) cells selected for analysis showed no 
signs of shrinkage or swelling, and sensitive or- 
ganelles, such as mitochondria, were not distorted; 



(2) the appearance of the granule was not altered 
in any meaningful way by variations in fixation; 

(3) the appearance of the secretory granules had 
to be unrelated to their location within the spec- 
imen block; and (4) the development of the gran- 
ules could be traced (from Golgi complex to apical 
cytoplasm) without any major breaks in devel- 
opmental sequence. 



Results ' 

The basic histology of the parotid salivary gland 
was similar in all 15 sp)ecies examined; in each 
species the acinus was formed by a cluster of cells 
connected to the striated duct by intercalated ducts 
of varying lengths (fig. 1). At the tem level, acinar 
cell secretory products were found to differ in all 
species examined (figs. 2-5). These differences 
could not be related to granule ontogeny by com- 
paring "immature" Golgi-GERL (Golgi-endoplas- 
mic reticulum-lysosomes)-associated granules with 
"mature" granules in the apical cytoplasm. In our 
samples that consisted of both males and females, 
we found no evidence of sexual dimorphism in 
secretory granule substructure, and in our largest 
samples (5 to 1 specimens) we found no examples 
of individual or geographic variation that could 
not be attributed to typical inter- or intracellular 
variation or to granule geometry. Although vari- 



216 



FIELDIANA: ZOOLOGY 




PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



217 



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218 



FIELDIANA: ZOOLOGY 



ation may occur at the individual level, we found 
no unequivocal evidence that it is detectable by 
transmission electron microscopy. Last, the ex- 
amples selected to illustrate each species (figs. 2- 
5) should be regarded as "average" or represen- 
tative of the most common, well-fixed granule 
morphology. 

Pteronotus parnellii— The secretory granules 
consisted of a moderately dense, homogeneous, 
variably thick outer zone and a slightly lighter in- 
terior in which were embedded a collection of lin- 
ear densities, usually randomly disposed but 
sometimes showing a degree of orderliness. Each 
of these densities was outlined by a thin, barely 
discernible, lucent layer (fig. 2). 

Phvllostomus elongatus— The matrix of the 
secretory granules was uniformly dense and con- 
tained a series of irregular shell-like densities, which 
in thin section appeared as ribbons consisting of 
a dense line flanked by thin lucent plies. In certain 
granules, the ribbons were aligned in concentric 
fashion, imparting a layered appearance to the pe- 
riphery of these cell structures. 

Phvllostomus latifolius— These granules 
bore some resemblance to those in P. elongatus in 
that some had one or two ribbon-like layers im- 
mediately subjacent to their limiting membrane, 
but many granules lacked these layers. Instead, the 
prevailing inclusion was a lucent, coreless strand 
disposed in a complex skeinlike or twisted pretzel 
conformation (fig. 2). 

Ton ATI A BiDENS— These secretory granules con- 
tained several groups of hexagonally-packed tu- 
bules just beneath the limiting membrane. In fa- 
vorable transverse sections, such packets appeared 
as honeycomb structures, whereas in longitudinal 
section they appeared as parallel linear densities. 
The dense matrix shows a vermiculate pattern (fig. 
2). 

ToNATiA SYLVicoLA— The secretory granules in 
this species differed from those in T. bidens. The 
granules consisted of light and dark material that 
displayed two principal patterns. Either the dark 
material was centrally placed, where it often con- 
tained one or several lucent spaces, or the light 
and dark material each occupied one hemisphere 
of the granule. In either case, crystalloid tubules 
of the sort found in T. bidens never were present 
(fig. 2). 

Trachops cirrhosus— In this species, the gran- 
ules consisted of a large, central, very dense spher- 
ule surrounded by a lucent cortex. The cortical 
material contained linearly arranged punctate den- 
sities that, at low to moderate magnifications, gave 



the appearance of several continuous dense layers 
in the outer portion of the granules (fig. 2). 

Glossophaga soricina— The bulk of each 
granule was occupied by homogeneously dense 
material, which formed a smooth interface with 
the remainder of the granule. The other portion 
of each granule was pale and contained minute, 
dense specks (fig. 3). The lighter material occa- 
sionally intruded into the dense zone in the form 
of a tortuous ribbon. 

Leptonycteris sanborni— The secretory gran- 
ules contained an amorphous, dense, central in- 
clusion in a paler matrix. A small bundle of short 
filaments was occasionally present in the matrix 
(fig. 3). 

Sturnira liuum— The secretory granules in this 
species resembled those in Trachops cirrhosus. 
They consisted of a large, central, dense core sur- 
rounded by a rim of lucent material in which were 
embedded distinct punctate densities often dis- 
posed in layers (fig. 3). 

Artibeus jamaicensis— The granules were large 
and pale and contained some faintly discernible, 
twisted fibrils. A few granules possessed a small, 
dense, usually peripheral inclusion (fig. 3). 

Ariteus FLA vescens— These secretory granules 
had a pale matrix in which were numerous, flat, 
narrow, moderately dense lamellae in a random 
arrangement. Viewed on edge, the lamellae ap- 
peared as dense lines; face on, as gray, irregular 
structures (as seen in fig. 3). In a few granules, 
there was a dense polygonal plate that lacked pe- 
riodicity. 

Eptesicus lynni— These granules had a com- 
plex substructure. The limiting membrane was un- 
derlaid by a layer of moderately dense material 
that made incursions, arcuate and anfractuous, into 
a dense matrix, producing a variety of patterns. 
These extensions of the F>eripheral light material 
had a layered structure (fig. 4, top). 

Eptesicus brasiliensis— As in E. lynni. the lim- 
iting membrane was subtended by layered mate- 
rial, usually in several plies, that extended into the 
dense matrix. These extensions were less tortuous 
than those in E. lynni and appeared to subdivide 
the granule interior (fig. 4, bottom). 

Tadarida brasiliensis— The secretory granules 
in this species exhibited a spectrum of patterns. 
The most common was relatively simple, with 
either short, flat, dense prisms or dense dots being 
suspended in a slightly less dense matrix. A few 
granules contained dense hollow spheres, whereas 
others had a mazelike configuration based on light 
and dark laminations (fig. 5, top). 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



219 




Fig. 4. Higher magnification of mature secretory granules in parotid acinar cells in Eptesicus lynni (top) and E. 
hrasiliensis (bottom). Note the subtle but consistent differences in the granule substructure. Eptesicus lynni, x 63,000; 
E. brasiliensis, x 76,800. 



220 



HELDIANA: ZOOLOGY 




Fig. 5. Parotid acinar cell granules in Tadarida brasiliensis (top) are compared to those found in Molossus molossus 
(bottom). Note the range of variation in granule substructure in T. brasiliensis. Tadarida brasiliensis, x 26,000; 
Molossus molossus, x 24,000. 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



221 



MoLOSSUs MOLOSSUS— The variable morphol- 
ogy of the granules in this species appeared to 
depend on their stage of maturation. Early gran- 
ules were small, with a moderately dense matrix 
in which were some prominent dense particles. As 
the granules matured (based on size, density, and 
spatial relationship to the Golgi complex), they 
enlarged and the particles decreased somewhat in 
density. The mature granules had a farinaceous 
matrix probably resulting from comminution of 
the dense particles (fig. 5, bottom). 



Discussion 

The idea of using comparative ultrastructural 
analysis of homologous, morphologically differ- 
entiated cells to study evolutionary pathways and 
to explore systematic relationships is new to the 
study of mammalian orders, but has been used in 
broader studies of vertebrates and invertebrates 
(Eakin, 1968; Rieger & Tyler, 1979; PhiUips & 
Tandler, 1987). Nevertheless, the potential value 
of ultrastructural comparisons has been demon- 
strated recently by studies of gastric mucosa, ret- 
ina, and submandibular salivary glands (Phillips 
et al., 1984; Feldman & Phillips, 1984; Tandler et 
al., 1983, 1986; Phillips & Tandler, 1987). The 
discovery of patterns of intersF>ecific differences in 
cellular architecture and cellular secretory prod- 
ucts is in keeping with previously successful use 
of histology in evolutionary and systematic mam- 
malogy (e.g.. Quay, 1954; Forman, 1972; Phillips 
& Oxberry, 1972; Sands et al., 1977; Naumova, 
1981; Hood & Smith, 1982, 1983). 

Secretory cells, such as the parotid acinar cells 
used in the present study, seem to hold special 
promise for comparative investigation. Firstly, the 
entire secretory process— from nuclear dna to syn- 
thesis of proteins and complex carbohydrates and 
packaging of materials into secretory granules and 
their subsequent discharge— has been studied in- 
tensively over the past several decades (e.g., 
Jamieson & Palade, 1971; Castle et al., 1975; Pa- 
lade, 1975). Secondly, the secretory process seems 
to be relatively conservative— in the sense that 
basic pathways are the same in virtually all secre- 
tory cells— and, therefore, interspecific compari- 
sons are facilitated and interpretation is somewhat 
simplified (Phillips & Tandler, 1987). 

In the present study we limited our descriptive 
comparisons to "mature" secretory granules. These 
are the secretory granules that accumulate within 
the apical cytoplasm of the acinar cell; this product 



either is discharged from the cell into the acinar 
lumen (thus becoming part of the formative saliva) 
or, after an unknown storage interval, is broken 
down through autophagy and recycled within the 
cell. The glycoprotein components of the acinar 
secretory product are elaborated by the rough en- 
doplasmic reticulum acting in concert with the 
Golgi complex and are perhaps the most interest- 
ing feature from a comparative point of view. Syn- 
thesis of these glycoproteins begins with transcrip- 
tion of a very small segment of the genome into 
mRNA. The exportable proteins of the secretory 
granules could be regarded as providing a "win- 
dow" on the genome because, although all somatic 
cells have the same genome (Briggs & King, 1 952, 
1957), only a small portion actually is operational 
in any given fully differentiated cell. A protein 
synthesized for export thus directly reflects a por- 
tion of the operational genome. Any polysaccha- 
ride components of the secretory granules prob- 
ably are at least one more step removed from the 
genome because synthesis of complex carbohy- 
drates is enzymatically determined and usually 
takes place within saccules in the Golgi complex 
(Tandler, 1978). Even so, the polysaccharides also 
have considerable potential for comparative anal- 
ysis. 

Production of secretory granules can be consid- 
ered in terms of ontogeny. The first "granules" 
typically are seen in direct proximity to the Golgi 
complex, where carbohydrates are linked to pro- 
teins and the membrane that will encase the gran- 
ule is synthesized in a process involving both the 
GERL and the Golgi complex itself (Tandler, 1978; 
Hand, 1 980a; Hand & Oliver, 1 984). Newly formed 
("immature") granules differ greatly from "ma- 
ture" granules (see Castle et al., 1975) and were 
not used by us in describing the product for each 
of our species. Nevertheless, in view of species 
differences found by us, it is interesting to ask 
whether or not immature granules of one species 
might resemble mature granules in another species. 
Such similarities might be expected if hetero- 
chronic differences accounted for differences in 
mature product. However, because no such cross- 
species similarities were found by us between 
immature and mature secretory granules, hetero- 
chrony does not seem to account for sp>ecies dif- 
ferences, at least among the 1 5 bats examined here. 

No two Neotropical sfiecies examined here ex- 
hibited the same mature secretory granules in their 
parotid gland acinar cells, although in a few cases 
there was a degree of resemblance. This extreme 
variability is easily the greatest ever reported for 



222 



HELDIANA: ZOOLOGY 



homologous cells within an order or, as in the case 
of the Phyllostomidae, within a family of mam- 
mals (or any other vertebrates). To what can we 
attribute this striking finding? 

One consideration is fixation, which certainly 
affects the appearance of any cellular feature as 
viewed with transmission electron microscopy. In- 
deed, it can be said that the appearance of cells 
and their products essentially is the consequence 
of their intrinsic chemistry combined with the 
chemistry of the fixative at the moment in time 
when fixation occurred. Different fixatives and tis- 
sue processing can have profound effects on the 
microscopic appearance of secretory granules in 
salivary glands (Simson et al., 1978). However, we 
used consistent processing techniques and two 
similar fixatives. Our specimens of Ariteus and 
Leptonycteris both were fixed with 2% glutaral- 
dehyde, whereas all of our other specimens were 
fixed with a trialdehyde-DMSO fixative (Kalt & 
Tandler, 1971; Phillips, 1985). We were able to 
eliminate fixation as a source of variation because 
we also have examined specimens of Artibeus that 
had been fixed in both solutions, coincidentally 
with Leptonycteris, Ariteus, and all of the other 
species examined (see A. phaeotis parotid in Phil- 
lips et al., 1977). 

Generally speaking, given consistent prepara- 
tion techniques, microscopic differences in secre- 
tory granule substructure can be ascribed to bio- 
chemical differences among the granules. 
Microscopically detectable sequestration of indi- 
vidual types of macromolecules within secretory 
granules has been demonstrated only rarely (Ra- 
vazzola & Orci, 1980; Kousvelari et al., 1982) but 
clearly is the best available explanation of intra- 
granule substructure. Based on fundamental prin- 
ciples of biochemistry, it thus can be concluded 
that macromolecules packaged in the secretory 
granules most likely sort themselves out according 
to charge and steric effects, as well as chemical 
interactions, to yield a characteristic pattern for 
each species. However, it also should be noted that 
a homogeneous appearance of intragranular sub- 
stance does not in itself preclude sequestration of 
different enzymes within the granule. Separate lo- 
calization of different enzymes (a-amylase and 
chymotrypsinogen A or a-amylase and trypsino- 
gen) within pancreatic cell zymogen granules has 
been demonstrated with a combination of hrp 
(horseradish-p)eroxidase)-labeled and ferritin-la- 
beled antibodies (Ono et al., 1980). 

At present it is impossible to correlate exactly 
secretory granule substructure with particular 



chemical components such as certain enzymes or 
mucosubstances, so from microscopic images alone 
we cannot say precisely how the granules in our 
species differ chemically from one another. How- 
ever, some conclusions can be inferred from the 
literature. For example, an electron-dense image 
(see, for example, fig. 2), labeled classically as "se- 
rous," can be associated with granules rich in en- 
zymes. Such an image is typical in species such as 
laboratory rodents and primates, for which some 
data are available on the biochemistry of parotid 
saliva (e.g., Jacobsen «Sc Hensten-Pettersen, 1974). 
On the other hand, the presence of electron-dense 
"serous" granules does not preclude the presence 
of mucosubstances within the secretory granules. 
Pinkstaff'et al. (1982) reported that, although the 
parotid product in the little brown bat, Myotis 
lucifugus, was "serous" with standard histological 
techniques, both neutral and acidic mucosub- 
stances could be demonstrated histochemically. 
The parotid granules in Artibeus and Ariteus are 
interesting in this regard because electron-dense 
material is scarce (especially in Artibeus, fig. 3) and 
enzyme production is extremely low, at least in 
Artibeus (Junqueira et al., 1973). By way of con- 
trast, Sturnira lilium has largely electron-dense 
granules (fig. 3) and thus differs considerably from 
the other two stenodermatines; Sturnira produces 
saliva rich in enzymes (Junqueira et al., 1973). 

Although the parotid granules in Artibeus (and, 
by extension, Ariteus) are low in enzyme content, 
to what can we attribute their tem image? This is 
an intriguing question because Wimsatt (1956) re- 
ported that the gland was negative for mucosub- 
stances, whereas Radtke (1972) reported the pres- 
ence of sialomucins in parotid acinar cells. In part, 
this apparent disagreement is the result of differ- 
ences in techniques that cannot be resolved by 
transmission electron microscopy. 

What can be determined about the parotid sal- 
ivary glands in Neotropical bats that relates to 
their diets, evolutionary history, or systematic re- 
lationships? Clearly the ultrastructure of parotid 
secretory granules has systematic significance, be- 
cause no two genera (or species either) are exactly 
alike. This finding is in keeping with our studies 
of the submandibular gland in five species of Ar- 
tibeus (Phillips et al., 1977; Tandler et al., 1983, 
1986). The seromucous cells in this gland were 
found to contain granules that allowed for three 
groupings of Artibeus species {A. cinereus-A. 
phaeotis, A. jamaicensis-A. lituratus, and A. con- 
color) that matched the independently derived 
genie data (from isozyme analysis) reported by 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



223 



Koop and Baker ( 1 983). The differences in parotid 
acinar product in Eptesicus lynni and E. brasi- 
liensis (fig. 4) also are interesting from this per- 
spective because independent genie data from 1 9 
presumptive loci show that E. lynni probably orig- 
inated from the E. fuscus species complex, sepa- 
rate from the origin of E. brasiliensis (Arnold et 
al., 1980). Eptesicus lynni was found to share only 
62% of analyzed alleles with E. brasiliensis. 

It might seem surprising that the microscopic 
images of salivary gland secretory products are 
generically (and often specifically) distinctive, but 
variation of such a fine resolution actually is in 
keeping with numerous genetic studies of mam- 
malian saliva. Several genetic markers have been 
found in human saliva (Azen & Opp)enheim, 1 973; 
Ashton & Balakrishinan, 1974; Tan & Ashton, 
1976), and salivary proteins in particular tend to 
be polymorphic (Azen, 1972, 1973). Additionally, 
both sex and strain differences in salivary proteins 
have been reported in laboratory mice (Ikemoto 
& Matsushima, 1984). Although none of these dif- 
ferences has been demonstrated microscopically 
(all are based on biochemical analysis alone), their 
occurrence nevertheless is significant to compar- 
isons among mammalian species. 

Although the sometimes subtle but consistent 
microscopic differences between species within the 
genera Phyllostomus, Tonatia, and Eptesicus (figs. 
2, 4) are in keeping with our hypothesis of the 
sensitivity of secretory product to genie differ- 
ences, they cannot readily be related to any known 
ecological differences between species within each 
genus. It is altogether possible that such species 
differences do not represent direct evolutionary 
selection. Instead, the differing images might rep- 
resent species-specific protein polymorphisms of 
a type that would not significantly affect "perfor- 
mance" of the saliva even though differences in 
primary molecular structure or surface charges (or 
both) could be indirectly detected by transmission 
electron microscopy (Phillips & Tandler, 1987). 
The existence of such "nonfunctional" (and pre- 
sumably nonselected, i.e., "neutral") interspecific 
variation in a protein molecule has been demon- 
strated previously in the otherwise conservative 
hemoglobin molecule (Perutz, 1983). In this ex- 
ample the tertiary and quaternary structures ap- 
parently are conserved regardless of large numbers 
of functionally neutral amino acid substitutions in 
the primary structure. Although these substitu- 
tions in themselves most often have no effect on 
the functional capacity of the molecule, they 
nevertheless are known sometimes to change mo- 



lecular surface charges (Perutz, 1983). For the 
present we postulate that similar nonselected, non- 
functional variations could account for the micro- 
scopically detectable intrageneric differences in 
salivary proteins in Tonatia bidens and T. sylvi- 
cola, Phyllostomus latifolius and P. elongatus, and 
Eptesicus lynni and E. brasiliensis. This explana- 
tion seems more parsimonious than the alterna- 
tive, which would be to assume that the acinar cell 
component of the parotid saliva is "functionally" 
different in closely related, ecologically similar 
species. The extent or degree of microscopic dif- 
ferences in the mature acinar cell product ulti- 
mately might tell us more about relative times of 
divergence than about ecological differences among 
congeneric species of bats. 

Microscopic comparisons of salivary gland se- 
cretory products could be valuable to cladistic 
studies of chiropteran families, at least to judge 
from our data. The phyllostomines generally are 
regarded as the most primitive (least derived) of 
the Phyllostomidae (Smith, 1976), and their par- 
otid acinar products thus would qualify as plesio- 
morphic (following Henning, 1 966). The products 
in all five species examined (Phyllostomus elon- 
gatus, P. latifolius, Tonatia bidens, T. sylvicola, 
and Trachops cirrhosus) contained large amounts 
of electron-dense material (fig. 2), as did the se- 
cretory product in Pteronotus parnellii (a closely 
related mormoopid), Eptesicus lynni and E. bra- 
siliensis (Vespertilionidae), and Tadarida brasi- 
liensis and Molossus molossus (Molossidae), all of 
which serve as "outgroups." 

If the electron-dense, enzyme-rich secretory 
granules are regarded as plesiomorphic in micro- 
chiropteran bats, then it would be reasonable to 
regard a granule with less electron-dense material 
as apomorphic (derived). The glossophagine gen- 
era examined (Glossophaga and Leptonycteris) are 
representative of a phyllostomid evolutionary trend 
in which dentition, tongues, and associated mus- 
culature were modified for feeding on fruit, pollen, 
and nectar (Park & Hall, 1951; Phillips, 1971; 
Greenbaum & Phillips, 1974; Griffiths, 1982, 1983; 
Smith 8l Hood, 1 984). In these genera the electron- 
dense component has been reduced in comparison 
to the phyllostomines. Carollia perspicillata, which 
is omnivorous, fits into this category in that the 
parotid secretory granules contain relatively little 
electron-dense material. In this bat the secretory 
granules are very distinctive because the electron- 
dense inclusions are often in the form of cagelike 
geodesic structures (Phillips & Tandler, 1987; 
Tandler et al., in press). 



224 



FIELDIANA: ZOOLOGY 



Perhaps the most interesting systematic finding 
in our data lies within the nominal subfamily 
Stenodermatinae. These bats represent an evolu- 
tionary trend toward frugivory that includes ex- 
treme gastric adaptation at the gross, histological, 
histochemical, and cellular levels (Forman, 1972; 
Forman et al., 1979; Phillips & Studholme, 1982; 
Phillips et al., 1984). The pale parotid secretory 
granules in Artibeus and Ariteus are synapomor- 
phous, whereas the parotid granules in Sturnira 
are electron-dense, enzyme-rich, and more nearly 
like the plesiomorphic granules of the phyllosto- 
mines, in particular Trachops cirrhosus (figs. 2-3). 
This example is interesting because Sturnira al- 
ways has been something of an enigma. Although 
de la Torre (1961) allied this genus with Vampy- 
rops-\\\it stenodermatines and Smith (1976) in- 
cluded it with the "long-faced" stenodermatines, 
others (e.g., Walton & Walton, 1968) previously 
had placed the genus in a separate subfamily (Stur- 
nirinae) based on a variety of morphological fea- 
tures that seemed inconsistent with the other, more 
traditional, stenodermatines such as Artibeus. In- 
deed, Slaughter (1970) pointed out that Sturnira 
has some distinctive dental features that possibly 
link the genus to the glossophagines. The differ- 
ences in the parotid secretory granules thus are in 
keeping with a variety of other phenotypic differ- 
ences. While it is reasonable to suggest that secre- 
tory granule differences of this magnitude repre- 
sent a major interspecific difference in the 
operational segment of the genomes of homolo- 
gous parotid acinar cells, the eventual systematic 
value of such data will await availability of data 
about still other phyllostomid genera. 

Although our comparative data clearly docu- 
ment great microscopically detectable plasticity in 
the secretory product of parotid acinar cells, many 
questions about their evolution remain unan- 
swered. For example, what has been the role of 
diet? At first glance, our data suggest that insec- 
tivorous and animalivorous species have enzyme- 
rich, electron-dense granules, whereas frugivores 
have enzyme-poor, pale secretory granules. Yet, 
what about Sturnira lilium, which certainly in- 
cludes large amounts of fruit in its diet (Gardner, 
1977)? 

At least three factors interfere with any effort to 
correlate parotid acinar cell product with diet. First, 
acinar cells are but one cell type among several 
that influence the biochemistry of parotid saliva. 
Second, and possibly more important, the par- 
otid is but one salivary gland among a host of 
glands (the submandibular and sublingual and mi- 



nor glands) that are located throughout the oral 
region and that contribute substances to the saliva. 
Some data suggest that different glands in different 
species might have been more responsive, in an 
evolutionary sense, to changes in diet. For ex- 
ample, it is the accessory submandibular glands 
of Trachops and Megaderma that are unique in 
histology and might correlate with feeding on frogs 
(Phillips &. Tandler, 1985; Phillips et al., 1987); 
the parotid in Trachops is similar to that of other 
phyllostomines. 

A third factor, for which we presently have no 
data, is the possibility that parotid acinar cells can 
respond in some way to diet at the individual level. 
Our sample sizes are large enough to convince us 
of the near uniformity of secretory granule mor- 
phology within a population. However, Schick et 
al. (1984) have recently published the first report 
of a secretory cell responding to dietary intake by 
a shift in enzyme production. This is the first such 
detailed molecular data known to us and, although 
pancreatic acinar cells in laboratory rats were the 
source of the data, the implications for compar- 
ative studies of salivary glands are worth noting. 

A final factor for consideration is the complex 
role that salivary glands play in the lives of mam- 
mals. Salivary glands in Artibeus, Chiroderma, and 
Ametrida might contribute to gastric cytoprotec- 
tion (Studier et al., 1983; Phillips et al., 1984). In 
Tonatia sylvicola an unusual organelle found in 
submandibular seromucous cells of males might 
be related to species isolation or sex recognition, 
or both (Nagato et al., 1984). If these examples 
are typical for bats, then digestion is but one of 
several major functions of salivary glands. 

In conclusion, this first systematic microscopic 
survey of a secretory cell product has demonstrat- 
ed a previously unknown, extraordinary degree of 
variation within a group of related sjjecies. Based 
on our findings, one might conclude that Neo- 
tropical microchiropteran bats will serve as a sig- 
nificant model for study of how secretory cells 
have evolved in mammals; meanwhile, data from 
comparative investigations will contribute to our 
knowledge of genie relationships among these an- 
imals. 



Acknowledgments 

Financial support for field and laboratory re- 
search that led to the data reported here came from 
a variety of sources, which we are pleased to ac- 
knowledge: Research Corporation Grant C-1251 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



225 



(to Phillips); NIH Grant DE 03455-02 (to Phil- 
lips); NSF Grant CDP-80 1-8653 (to Phillips); 
Hofstra University HCLAS Executive Committee 
grants (to Phillips); NIH Grant AM-08305 (to 
Tandler); NIH Grant RO DE 07648-01 Al (to 
Tandler & Phillips); the Alcoa Foundation (through 
Dr. Hugh H. Genoways, Carnegie Museum of Nat- 
ural History); and the Graham Netting Research 
Fund via a grant from the Cordelia S. May Char- 
itable Trust (through Genoways). In addition to 
recognizing our financial support, we also wish to 
thank several colleagues whose field companion- 
ship in such places as Suriname and Jamaica, hard 
work, and willingness to share time and ideas in 
their laboratories played a major part in our study. 
These are: Hugh H. Genoways, Carnegie Museum 
of Natural History; Gary W. Grimes and Dorothy 
E. Pumo, Hofstra University; Robert J. Baker, 
Texas Tech University; and Henry A. Reichart, 
formerly of STINASU, in Suriname. Field assis- 
tance was provided by K. M. Studholme, S. L. 
Williams, N. M. Sposito, J. Groen, R. L. Honey- 
cutt, B. Koop, M. Arnold, B. A. Oxberry, P. Bil- 
leter, J. Bickham, and J. Patton. We also acknowl- 
edge technical assistance of Carol Ayala, Case 
Western Reserve University, and typing by Linda 
Cossen, Special Secretarial Services, Hofstra Uni- 
versity. Artwork and lettering were done by Helen 
Tandler. All specimens were collected and im- 
ported under proper permits issued by the relevant 
authorities in Mexico, Jamaica, Suriname, and the 
United States. 

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228 



FIELDIANA: ZOOLOGY 



Appendix 



Specimens Examined 

Voucher specimens of each species and each 
locality are deposited in the collections of the Car- 
negie Museum of Natural History and The Mu- 
seum, Texas Tech University. All specimens ex- 
cept those of Leptonycteris sanborni (Jalisco, 
Mexico) and Ariteus flavescens and Eptesicus lynni 



(Jamaica) were collected in Brokopondo Province, 
Suriname. 

Pteronotus parnellii, 10 (5 S6, 5 29); Tonatia bi- 
dens, 3 {2 SS, 1 2); T. sylvicola, 2 {66); Trachops 
cirrhosus, 2 (1 3, 1 2); Phyllostomus elongatus, 3 
(\6,2 22); P. latifolius, 1 (2); Glossophaga soricina, 
3 (2 $6, 1 2); Leptonycteris sanborni, 2 (3<5); Sturnira 
lilium, 5 (2 35, 3 22); Artibens jamaicensis, 3 (22); 
Ariteus flavescens, 1 (2); Eptesicus lynni, 2 {$$); E. 
brasiliensis, 3 (22); Tadarida brasiliensis, 1 (S); Mo- 
lossus molossus, 8 (4 66, 4 22). 



PHILLIPS ET AL.: SALIVARY GLANDS IN BATS 



229 



Distribution of the Species and Subspecies of 
Cebids in Venezuela 

Roberta Bodini and Roger Perez-Hernandez 



ABSTRACTS 

Thirteen species of Primates representing nine genera in the family Cebidae are found in 
Venezuela. The geographic distribution of these species exhibits four main patterns. Alouatta 
and Cebus are widely distributed in all parts of the country. Four genera, Saimiri, Callicebus, 
Cacajao, and Chiropotes are restricted to south-central Venezuela; their distribution is centered 
in the Amazonian lowlands. Aotus and Ateles are each represented by one species or subspecies 
in south-central Venezuela and another in northwestern Venezuela. Pithecia is mainly restricted 
to the Guianan highlands in eastern Venezuela, with a single known outlying locality in south- 
central Venezuela. Of the nine cebid genera in Venezuela, all occur in south-central Venezuela, 
four occur in northwestern and north-central Venezuela, and only three occur in eastern Ven- 
ezuela. Distribution maps for the 1 3 species in Venezuela are presented with exact localities of 
specimens. 

Nueve generos de cebidos representados por trece especies se hallan en Venezuela. La re- 
particion geografica de las especies se resuelve en cuatro patrones geograficos principales. De 
este modo, Alouatta y Cebus son extensamente repartido por todo el pais. La distribucion de 
los generos, Saimiri, Callicebus, Cacajao y Chiropotes esta restringida al sur-central de Vene- 
zuela con concentracion en las tierras bajas Amazonicas. Aotus y Ateles son representatados, 
cada cual, por una especie (o subespecie), en el sur-central, y otra especie in el noroeste del 
pais. Pithecia se reparte en las alturas guayanas venezolanas del este, y es conocido, a la vez, 
por un solo dato de captura del sur-central venezolano. En resumen, la totalidad de los nueve 
generos esta presente en el sur-central de Venezuela, cuatro de ellos en el noroeste y norte- 
central, y tres en la Venezuela oriental. La reparticion de las trece especies esta documentado 
por mapas. 

Treze especies de Primatas, representando nove generos da familia Cebidae, sao encontradas 
na Venezuela. A distribuifao geografica destas especies exibem quatro padroes principais. Al- 
ouatta e Cebus espalham-se por todas partes do pais. Quatro generos, Saimiri, Callicebus, 
Cacajao, e Chiropotes, limitam-se ao centro-sul de Venezuela; a Bacia Amazonica sendo seus 
focos de distribuifoes. Aotus e Ateles sao representados por uma especie ou subespecie cada 
no centro-sul da Venezuela, e por outra especie no noroeste do pais. Pithecia limita-se ao 
planalto Guianense, no leste da Venezuela, com apenas uma localidade conhecida no centro- 
sul do Pais. Em suma, todos nove generos de Cebidae ocorrem no centro-sul da Venezuela, 
quatro ocorrem no centro-norte e noroeste do Pais, e apenas tres ocorrem no regiao leste da 
Venezuela. Apresentam-se mapas de distribui96es das 13 especies na Venezuela, com locali- 
dades exatas dos especimes colecionados. 



From the Institute de Zoologia Tropical, Universidad 
Central de Venezuela, Apartado 47058, Los Chaguara- 
mos, Caracas 1041 -A, Venezuela. 



BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 231 



Introduction 

Primates of the family Cebidae are some of the 
most conspicuous mammals in the Neolropics, as 
they are diurnal, often forage in large troops, and 
may be quite vocal. However, we actually know 
very little about the exact distributions of most 
species of cebids. In Venezuela, several authors 
have discussed cebids as part of generic revisions 
(see Elliot, 1912; Hershkovitz, 1949, and later 
works; HiU, 1960, 1 962; Kellogg & Goldman, 1944) 
and in works on the status or collections of specific 
species (i.e., Bodini, 1983; Handley, 1976; Mon- 
dolfi & Eisenberg, 1979; Rudran & Eisenberg, 
1982). 

Herein we summarize the distribution of cebids 
in Venezuela based on recent collections, speci- 
men records from several museums, and the lit- 
erature. Specimens reported on are housed in the 
following collections: Estacion Biologica Rancho 
Grande (EBRG); Museo de Biologia, Universidad 
Central de Venezuela (MBUCV); Museo de Cien- 
cias Naturales (MCN); and Museo de Historia 
Natural La Salle (MHNLS). Our objectives are to 
provide accurate locality records and distribution 
maps that may be utilized in future investigations. 



Generic Distributions 

The nine genera of cebids that inhabit Venezuela 
exhibit four main patterns of geographic distri- 
bution (figs. 1-7). (1) Two geneT^—Alouatta and 
Cebus— arc widely distributed in all paris of the 
coimtry. (2) Four genera— Sa/m/r/, Callicebus, 
Cacajao, and Chiropotes—UTe restricted to south- 
central Venezuela, centering on the Amazonian 
lowlands (see Eisenberg & Redford, 1 979). (3) Two 
genera— .4orw5 and Ateles—are each represented 
by one species or subspecies in south-central Ven- 
ezuela and another in northwestern Venezuela or 
northwestern and north-central Venezuela, which 
is the region of the northeastern spurs of the An- 
dean chain and the enclosed Maracaibo basin. (4) 
One genus— Pithecia— is mainly restricted to the 
Guianan highlands in eastern Venezuela, with a 
single known outlying locality in south-central 
Venezuela. In summary, of the nine cebid genera 
in Venezuela, all occur in south-central Venezuela 
(including the outlier record of Pithecia), four oc- 
cur in northwestern and north-central Venezuela, 
and only three occur in eastern Venezuela. 



Specific and Subspecific Distributions 
Saimiri 

In a recent revision of the squirrel monkeys, 
Hershkovitz (1984) recognized four species: Sai- 
miri boliviensis, S. oerstedi. S. sciureus, and S. 
ustus. All those squirrel monkeys found in north- 
em South America he referred to the single species 
S. sciureus (fig. 1 ). Squirrel monkeys from Ama- 
zonian Venezuela and adjacent Brazil and Colom- 
bia he regarded as 5. sciureus cassiquiarensis. 
Hershkovitz (1984) plotted several localities in 
Territorio Federal Amazonas, but listed only a 
single specific locality: "Casiquiare, Rio (mouth), 
2°01'N, 67"X)7'W." Hill (1960) previously had 
mapped the distribution of 5. sciureus as occurring 
throughout all of Venezuela, although he listed 
only a single locality in the state of Bolivar (Camp 
Canaracuni, 4''36'N, 64''10'W). The name cassi- 
quiarensis Lesson is based on Humboldt's descrip- 
tion of a captive female from the banks of the Rio 
Casiquiare, Amazonas, Venezuela. 

Specimens Examined— Total 29. Bolivar: Ca- 
naracuni (4°36'N, 64«'10'W). Territorio Federal 
Amazonas: Campo Cacuri (4°49'N, 65°26'W); Cano 
Yureba, Rio Ventuari (3''35'N, 66°46'W); Rio Pu- 
runame, 40 km from union with Rio Orinoco 
(3°19'N, 65°15'W); Rio Ventuari (3°59'N, 
67'^2'W); San Fernando de Atabapo (4°02'N, 
67»37'W); San Juan de Manapiare (5°14'N, 
66'^2'W). 

Aotus 

Night monkeys are found from Panama 
throughout much of Amazonian South America 
to Paraguay. Historically, it has generally been as- 
sumed that all night monkeys represented the sin- 
gle species Aotus trivirgatus. However, in a recent 
revision of the genus, Hershkovitz (1983) recog- 
nized nine allopatric species. A tenth species, Aotus 
hershkovitzi from Colombia, has recently been 
proposed by Ramirez-Cerquera (1983). Two 
species of Aotus, A. lemurinus and A. trivirgatus. 
have been reported from Venezuela (fig. 2). 

Aotus lemurinus griseimembra Elliot is known 
in Venezuela only fi-om the extreme northwestern 
region, the states of Cojedes and Merida (Hersh- 
kovitz, 1983), and the states of Tachira, Trujillo, 
and Zulia (Handley, 1976). 

Specimens Examined— Total 10. Zulia: Campo 
a Rosario, EHstrito Catatumbo (l''44'N, 67°03'W); 



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Reserva San Manuel, Los Canaguatos (9*t)4'N, 
71''56'W); Kunana, Rio Negro, Sierra de Perija 
(9*75'N, 72*25'W); Rio Tocuco (72''25'W). 

Aotus trivirgatus trivirgatus Humboldt is known 
from the region south of the Rio Orinoco in Ter- 
ritorio Federal Amazonas and the state of Bolivar. 
The Venezuelan national collections contain 
specimens from Cano Manapiare and Rio Anta- 
vari, and two literature records report specimens 
from the right bank of the Rio Caroni (INP- 
ARQUES, 1982;MARNR-DGS-POA-SFS, 1982). 

Our recent discovery of Aotus trivirgatus in 
Guyana to the east of the Rio Caura, well outside 
of the Rio Orinoco basin, led to an investigation 
of geographic variation in the species. Preliminary 
results suggest that these A. trivirgatus represent a 
new geographic race (Bodini & Ferreira, in prep.). 

Specimens Examined— Total 1 1. Bolivar: Cano 
Manapiare (7»1 1'N, 66°40'W); Rio Antavari 
(S'^O'N, 63°10'W). Territorio Federal Amazonas: 
Alto Manapiare (S'^S'N, 66°02'W); San Juan de 
Manapiare (S^M'N, 66°02'W); near Isla Cigarron, 
Rio Negro (r44'N, 67°03'W); recently collected 
in La Neblina, east of Rio Varia (0°59'N, 66°10'W). 

Callicebus 

The genus Callicebus is the sole representative 
of the subfamily Callicebinae and is found from 
northern South America south to northern Para- 
guay. Hershkovitz (1963) and Kinzey (1982) rec- 
ognized three species, Callicebus moloch, C per- 
sonatus, and C. torquatus, of which only C. 
torquatus is found in Venezuela. 

Callicebus torquatus lugens Humboldt is known 
in Venezuela south of the Rio Orinoco (fig. 3). Hill 
(1960) originally suggested that C. torquatus is 
found as far east as Guyana, although this was 
questioned by both Handley (1976) and Hersh- 
kovitz (1963), who reported Venezuelan speci- 
mens only from Territorio Federal Amazonas. 
Handley (1976) reported 31 specimens from 
southern Territorio Federal Amazonas. Bodini 
(1981) reported specimens from Maripa, Camp 
Canaracuni, and the Rio Antavari in Venezuela. 
Recent reports by Kinzey ( 1 982) and Bodini (1981) 
confirm the presence of C torquatus in Guiana 
Region. 

Specimens Examined— Total 26. Bolivar: Cana- 
racuni (4'36'N, 64°10'W); Maripa, 150 km from 
Ciudad Bolivar (7*22'N, 65'^9'W); Rio Antavari 
(5»20'N, 63M0'W); Alto Paragua (4°30'N, 
63*WW). Territorio Federal Amazonas: Alto Cano 
Caname, Departamento Atabapo (3*'33'N, 



67"06'W); Alto Ventuari (3°50'N, 67°04'W); Boca 
Cano Maica, Rio Ventuari (66°30'W); Cano Ya- 
gua, Cerro Cucurito, Departamento Atabapo 
(3''38'N, 66»25'W); Cacuri, Alto Ventuari (4°49'N, 
65"26'W); La Esmeralda (3"X)8'N, 65°32'W); Rio 
Cunucunuma (3°10'N, 66°0rW); Rio Puruname 
(3°19'N, 65''15'W); La Neblina, east of Rio Varia 
(0°59'N, 66°10'W). 

Alonatta 

Alouatta is the only genus within the subfamily 
Alouattinae and is represented by some six species 
widely distributed in Central and South America. 

A single species, Alouatta seniculus, is abundant 
and widely distributed in Venezuela. Hill (1962) 
recognized three subspecies of A. seniculus as oc- 
curring in Venezuela. Alouatta seniculus seniculus 
Linnaeus is foimd in extreme northwestern Ven- 
ezuela, primarily in the states of Apure, Tachira, 
and Zulia, with a single record from Barinas (Ti- 
coporo Forest, 8'X)6'N, 70*'40'W). Alouatta seni- 
culus arctoidea Cabrera, called the "Caracas howler 
monkey" by Humboldt, inhabits all the coastal 
region from Falcon to the state of Miranda. Ca- 
brera (1 958) proposed Caracas as the type locality. 
A third subspecies, A. seniculus stramineus Hum- 
boldt, inhabits all of Venezuela south of the Rio 
Orinoco in Territorio Federal Amazonas and the 
state of Bolivar (fig. 4). 

In addition to these three subspecies, we believe 
a fourth, previously unrecognized, form exists and 
is widely distributed throughout the Venezuelan 
llanos. This undescribed subspecies is character- 
ized by coloration and size. The southern limit of 
its distribution is clearly defined by the Rio Ori- 
noco, but its northern and western limits in the 
Andean piedmont are as yet undetermined. 

Howler monkeys are extremely adaptable to a 
wide array of environments, and we believe they 
are found throughout Venezuela. Their apparent 
absence in certain regions probably reflects lack of 
collecting rather than true distributional gaps, as 
our records demonstrate for the state of Anzoa- 
tegui. The problem of current and historical dis- 
tribution and systematic relationships of the var- 
ious populations in Venezuela is in need of study. 

Specimens Examined— Total 80. Anzoategui: 
Los Cocos, Rio Caris (8''30'N, 64'^5'W). Apure: 
Caiio San Agustin, Selva de San Camilo (7"'19'N, 
7r57'W); Hato El Frio (7°44'N, 68°54'W); Las 
Raicitas, El Saman (7°55'N, 68°40'W). Aragua: 
Asentamiento Los Castillos, Turagua (10°09'N, 
67»3rW); Cumbre de Guacamaya (10°2rN, 
67'40'W); Los Picachos, Rancho Grande ( 1 0*2 1'N, 



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6T4l'W). Barinas: Reserva Forestal Ticoporo 
(8°09'N, 70°40'W). Bolivar: Guayoba, Rio Caura, 
4 km from Maripa (7°20'N, 65°10'W); Hato Bella 
Vista, El Palmar (8°00'N, 62°00'W); Represa del 
Guri, Operacion Rescate (7°40'N, 63°00'W); San 
Martin de Turumban, Anacoco-Cuyuni (6°42'N, 
61°02'W). Carabobo: Las Quiguas, San Esteban 
(10''25'N, eS^OrW). Cojedes: Hato Barbasco 
(9°04'N, 68°08'W); Hato Itabana, 38 km from Las 
Vegas (9°17'N, 68*'13'W); Las Queseras, El Baul 
(8'*25'N, 68°17'W); Montaria Las Lomas, San Car- 
los (9°38'N, 68°34'W). Distrito Federal: Hacienda 
El Limon (10°28'N, 67''1 7'W). Falcon: Sanare, De- 
partamento Silva (10°23'N, 68°25'W); Sierra San 
Luis (1 1°15'N, 69°10'W). Guarico: Corozo Pando 
(8°14'N, 67°17'W); Hato Mapurite, 40 km N of 
Calabozo (9°17'N, 67°24'W); Manapire, near San 
Antonio (9°1 7'N, 66''1 1 'W); Rio Tiznado (8°1 8'N, 
67°48'W). Lara: Cumbre de Las Trojas, 45 km S 
of Cabudare (9°45'N, 69°07'W). Miranda: Rio Ne- 
gro (10°20'N, ee'lS'W); La Guzmanera, Guatopo 
(10'X)0'N, 66°15'W). Portuguesa: Agua Blanca 
(9°40'N, 69°07'W). Tachira: Cerro El Teteo, Bura- 
gua (7°30'N, 7 r57'W); La Fria (8°1 3'N, 72°14'W); 
Paramo Tama (7°27'N, 72°26'W). Territorio Fed- 
eral Amazonas: Cacuri, Alto Ventuari (4°49'N, 
65°26'W); Caiio Yureba, Ventuari (3°35'N, 
66°46'W); Rio Hacha, Alto Ventuari (3°47'N, 
65''38'W); Rio Puriname, 40 km from confluence 
of Rio Orinoco (3°19'N, 65°15'W); San Juan de 
Manapiare (9°05'N, 66°02'W); Cerro La Neblina, 
E of Rio Varia (0°59'N, 66°10'W). Territorio Fed- 
eral Delta Amacuro: Caiio Caneima (9°05'N, 
60°55'W); Guiniquina (9°10'N, 61°06'W). Zulia: 
Laguna de Manaties, Departamento Catatumbo 
(9°27'N, 72°02'W); La Victoria, Rio Negro (9°36'N, 
72°1 5'W); Rio Guasare ( 1 1°02'N, 72°05'W); Sierra 
de Perija (9°00'N, 72°00'W). 



Chiropotes 

The bearded sakis comprise two species limited 
to northern South America (Cabrera, 1958; Hersh- 
kovitz, 1977;Mittermeier«Sc.Coimbra-Filho, 1981). 
One species is found in southern Venezuela rep- 
resented by a single subspecies, Chiropotes satanas 
chiropotes Humboldt (fig. 5). All of our records 
and those reported by Handley (1976) are from 
Territorio Federal Amazonas. Cruz Lima (1945) 
and Rudran and Eisenberg (1982) proposed the 
occurrence of this species in the state of Bolivar 
on the basis of Humboldt's description; Mondolfi 
(1976) reported observations of Chiropotes at 



Maripa and Caiio Maniapure (Bolivar), but we are 
aware of no specimens from this region. 

Specimens Examined— Total 29. Territorio 
Federal Amazonas: Cacuri, Alto Ventuari (4°49'N, 
65''26'W); Caiio Yureba, Departamento Atabapo 
(3°3rN, 66''44'W); Caiio Yagua, Cerro Cucurito 
(3''3 1 'N, 66°44' W); Laguna de Chiripo, Caiio Blan- 
co (3°27'N, 66°40'W); Rio Ocamo, Alto Orinoco 
(2°44'N, 65°11'W); Puerto Ayacucho (5°36'N, 
67''35'W); Rio Orinoco, S of San Fernando de Ata- 
bapo (4'^0'N, 67°38'W); San Fernando de Ata- 
bapo (4°02'N, 67°37'W); San Juan de Manapiare 
(5°14'N, 66'^2'W). 



Cacajao 

The genus Cacajao, or uakaris, contains two 
species which are found in northern South Amer- 
ica (Cabrera, 1958; Hershkovitz, 1972; Mit'ter- 
meier & Coimbra-Filho, 1981). One species, Ca- 
cajao melanocephalus, is found in Venezuela 
restricted to the upper Orinoco region of southern 
Territorio Federal Amazonas (fig. 5). Although few 
specimens exist in collections, they appear to be 
abundant. 

Specimens Examined— Total 2. Territorio Fed- 
eral Amazonas: Alto Cano Atacavi, Departamento 
Casiquiare (3°05'N, 67'>02'W); La Nebhna, E of 
Rio Varia (0°59'N, 66°10'W). 



Cebus 

The capuchin monkeys are found from Hon- 
duras south through Central America and the 
northern two-thirds of South America. Four species 
currently are recognized (Cabrera, 1958; Hersh- 
kovitz, 1972). Three species of Cebus are repre- 
sented in Venezuela: Cebus albifrons, C apella, 
and C. «/^rm//a/z^s (Hershkovitz, 1949, 1958). 

Three subspecies of Cebus albifrons are found 
in Venezuela (fig. 6). Cebus albifrons adustus 
Hershkovitz was described on the basis of three 
specimens from "near head of Rio Cogollo (Apon) 
at eastern base of Sierra de Perija, about 5 kilo- 
meters northwest of Machiques, Lake Maracaibo 
region, Zulia" (Hershkovitz, 1949, p. 369). We 
report additional specimens from Rio Guasare and 
Kasmera. This subspecies is restricted to the Sierra 
de Perija of extreme northwestern Venezuela and 
adjacent Colombia. Cebus albifrons leucocephalus 
Gray is found in extreme western Venezuela, in 
the region south of the Lago de Maracaibo basin. 



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and the states of Apure, Merida, Tachira, and Zu- 
lia. Cebus albifrons unicolor Spix is found in ex- 
treme southern Venezuela in Territorio Federal 
Amazonas. Hershkovitz (1949) reported speci- 
mens from Marimonda, Rio Orinoco, and from 
Solano, Rio Casiquiare. Handley (1976) reported 
specimens from Rio Mavaca and Tamatama. All 
locality records for this subspecies in Venezuela 
are south of the Rio Ventuari. 

Specimens Examined— Total 15. Apure: Caiio 
San Augustin, Selva de San Camilo (7°19'N, 
7r57'W). Merida: Palmichoso, S of Las Virtudes 
(9'^9'N, 70°57'W). Tachira: La Fria (8°13'N, 
72°14'W). Territorio Federal Amazonas: Cano 
Yagua, Cerro Cucurito, Departamento Atabapo 
(B^SS'N, 66''25'W); near Boca Padamo, left side of 
Rio Orinoco (3°02'N, 65°13'W). Zulia: Kasmera, 
Perija (10°05'N, 72°45'W); Kunana (9°36'N, 
72°15'W); Rio Bravo, Distrito Catatumbo (9°05'N, 
72°22'W); Rio Guasare (1 1°02'N, 72°05'W). 

Cebus apella is represented in Venezuela by two 
subspecies (fig. 6). Cebus apella apella Linnaeus is 
restricted to Amazonian Venezuela, the Territorio 
Federal Amazonas, and is found along both banks 
of the upper Orinoco. Cebus apella margaritae 
HoUister is endemic to and restricted to Margarita 
Island. The 800-km gap between the ranges of 
these two subsp>ecies is striking and unexplained. 

Specimens Examined— Total 10. Nueva Espar- 
ta: Sierra de Copey (1 1°03'N, 63°56'W). Territorio 
Federal Amazonas: Alto Caiio Caname (3''22'N, 
67°08'W); Caiio Yapacana (3°30'N, 66°45'W); San 
Fernando de Atabapo (4°02'N, 67°37'W). 

Cebus nigrivittatus is widely distributed in Ven- 
ezuela and represented by perhaps five subspecies 
(fig. 6) (Hershkovitz, 1949; Cabrera, 1958). Cebus 
nigrivittatus apiculatus was described by Elliot 
(1912) on the basis of specimens from La Union, 
Rio Caura, near its confluence with the Rio Ori- 
noco. It is distributed throughout central Vene- 
zuela south of the Orinoco between the Rio Caroni 
and the Rio Ventuari. 

Cebus nigrivittatus brunneus was described by 
J. A. Allen (1914) from specimens from Aroa, a 
station on the Bolivar Railway, Yaracuy, north- 
western Venezuela. Hershkovitz (1949) reported 
an additional specimen from the Paria Peninsula. 
These records plus our specimens suggest that C. 
n. brunneus is continuously distributed through- 
out the Cordillera de la Costa of extreme northern 
Venezuela. Cebus nigrivittatus nigrivittatus Wag- 
ner is restricted in Venezuela to the Amazonian 
region of Territorio Federal Amazonas. Cebus ni- 
grivittatus olivaceus Schomburg is found in south- 
eastern Venezuela. Hershkovitz (1949, p. 348) re- 



ports the type locality as "Vicinity of 'Our Village,' 
said to be situated at latitude 4°57'N., 61°rW., 
altitude 3,100 feet above sea level, southern foot 
of Mount Roraima." 

We believe that an undescribed subspecies, Ce- 
bus nigrivittatus subsp., is widely distributed 
throughout central and northern Venezuela north 
of the Orinoco. The status of this population is 
currently under study. 

The Orinoco Delta region, which Eisenberg and 
Redford ( 1 979) excluded from their consideration 
of biogeographic regions due to insufficient data, 
is now shown by the distributions of Alouatta, 
Cebus, and Pithecia to be clearly aligned with the 
Guyana highlands. The Llanos region now extends 
up to the western edge of the delta, but does not 
include it, as demonstrated by the distributions of 
Alouatta and Cebus. 

Specimens Examined— Total 75. Anzoategui: 
Los Cocos, Rio Caris (8°30'N, 64°05'W); 10 km 
W of Laguna de Unare (10°02'N, 65°12'W); Mor- 
ichal Largo [between Anzoategi and Monagas] 
(8°18'N, 63°15'W). Aragua: Rancho Grande 
(10°10'N, 67°19'W). Barinas: Reserva ForestalTi- 
coporo, on Barinas-Pedregal road (8°03'N, 
70°18'W). Bolivar: Caiio La Urbina (7°15'N, 
66°25'W); Carretera Caicara-S. Juan de Mana- 
piare, km 1 75 (6°02'N, 66°29'W); Carretera El Do- 
rado-Santa Elena, km 33 (6°12'N, 6ri4'W); Cur- 
aima. El Palmar (8°01'N, 61°26'W); El Dorado- 
Santa Elena, km 121 (5°18'N,6ril'W);Guayopo, 
Rio Caura, 14 km from Maripa (7°09'N, 65°10'W); 
Canaracuni (4°17'N, 64°05'W); Guri, Operacion 
Rescate (7°18'N, 63°00'W); Rio Antavari (5'i09'N, 
63°05'W); Rio Marajano, Meseta de Jaua (4°08'N, 
64°1 1'W); Rio Villacoa, 4 km N of mouth (6°16'N, 
67'^5'W); San Martin de Turumban, Rio Cuyuni 
(6"'19'N, 61'^9'W). Carabobo: Bahia de Patanemo 
(10"'12'N, 67''26'W); Urama (10°12'N, 68°08'W). 
Cojedes: Cerro Azul, La Blanquera (8°26'N, 
68°07'W); Montana Las Loma, San Carlos (9°1 7'N, 
68°16'W); Pica, Las Vegas (9°1 5'N, 68°1 7'W). Dis- 
trito Federal: El Avila, Caracas (10° 14'N, 66°13'W); 
Hacienda El Limon (lO^B'N, 67°08'W). Falcon: 
Sanare, Distrito Silva (8''23'N, 68°12'W). Guarico: 
Hato Flores Moradas, Calabozo (8°23'N, 67"! 3'W); 
Hato Mapurite, 40 km N of Calabozo (9°08'N, 
67°irW); Parmana (7°28'N, 65°18'W); San Jose 
de Tiznados (9°16'N, 67''16'W). Lara: La Pastora, 
1 1 km SSW of Sanare (9°2rN, 70°07'W). Miran- 
da: Rio Negro (10''20'N, 66°17'W); La Guzma- 
nera, Guatopo (10°00'N, 66°15'W). Territorio 
Federal Amazonas: Alto Manapiare (5''13'N, 
66°0rW); Alto Ventuari (4°45'N, 65°20'W); Caiio 
Yureba, Rio Ventuari (3°16'N, 66''21'W); Cano 



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Mayaba, Rio Ventuari (4°07'N, 66°16'W); Cano 
Morrocoy, Alto Ventuari (5°08'N, 66°02'W); La 
Esmeralda (S-^S'N, 65°32'W); Ocamo, Rio Oca- 
mo (2°20'N, 65°15'W); Puruname, 40 km from 
Rio Orinoco (3°19'N, 65°15'W); La Neblina, E of 
Rio Varia (0°59'N, 66°10'W). Territorio Federal 
Delta Amacuro: Cano Araguabisi (9°12'N, 
60*'27'W); Guiniquina (9''10'N, 61'X)3'W); Tobe- 
juba, Guayo (9°09'N, 6r25'W). Yaracuy: Agua 
Negra (10°14'N, 68''14'W); Carretera Boca de 
Aroa, 20 km from Palmasola (lO^Ol'N, 69°27'W). 



Ateles 

Spider monkeys are widely distributed from 
northeastern Mexico throughout tropical South 
America. Four species are recognized, of which 
only Ateles belzebuth occurs in Venezuela (fig. 7). 

Ateles belzebuth belzebuth GeofFroy is found in 
southern Venezuela, south of the Rio Orinoco; 
most records are from Territorio Federal Ama- 
zonas. Kellogg and Goldman ( 1 944) indicate a wide 
distribution in Guyana and report the Venezuelan 
localities of La Union, Rio Mato, and El Llagual 
(on both banks of the Rio Caura). 

Ateles belzebuth hybridus Geoffroy is known 
from northern and western Venezuela (Hershko- 
vitz, 1949; Cabrera, 1958; Kellogg & Goldman, 
1944). Handley (1976) reported specimens from 
Apure and Trujillo in western Venezuela. Our 
specimens are from the states of Barinas, Tachira, 
and Zulia. Mondolfi and Eisenberg (1979) report- 
ed it from Cupira and Guatopo, state of Miranda, 
suggesting a discontinuous distribution on the 
coast. 

Specimens Examined— Total 16. Barinas: Re- 
serva Forestal de Ticoporo, Sabana de Anare 
(8°06'N, 70°40'W). Bolivar: Canaracuni (4°06'N, 
64°10'W). Miranda: Cupira (10°10'N, 65°44'W). 
Tachira: La Fria (8°13'N, 72°14'W). Territorio 
Federal Amazonas: Cacuri, Rio Ventuari (4°49'N, 
65°26'W); Rio Ocamo, Alto Orinoco (2°44'N, 
65°11'W); Salto del Oso, Alto Ventuari (4''55'N, 
65°25'W); San Juan de Manapiare (S^M'N, 
66°02'W). Zulia: Rio Guasare ( 1 1°02'N, 72°05'W). 



Lagothrix 

Woolly monkeys have not been collected in 
Venezuela; however two subspecies of Lagothrix 
lagothricha are to be expected: L. lagothricha la- 
got hricha should be found in Territorio Federal 



Amazonas south of the Rio Ventuari, and L. I. 
lugens, in the Selva de San Camilo, state of Apure 
(Fooden, 1963; Hemandez-Camacho & Cooper, 
1976). 



Pithecia 

Sakis are found only in northern South America; 
Hershkovitz (1979) recognized four monotypic 
species, of which only Pithecia pithecia Linnaeus 
is found in Venezuela. P. pithecia is found in Ven- 
ezuela south of the Rio Orinoco and throughout 
the Guianas and northeastern Brazil. In Venezuela 
specimens have been reported primarily from the 
extreme northeastern region, the state of Bolivar 
and Territorio Federal Delta Amacuro (fig. 5), with 
a single outlier locality record for Belen, Rio Cunu- 
cunuma, Territorio Federal Amazonas (3°39'N, 
65°46'W) (Handley, 1976). If the distribution pro- 
posed by Hershkovitz (1979) and Mittermeier and 
Coimbra-Filho (198 1) is correct, P. pithecia should 
inhabit the entire region between the upper Rio 
Orinoco and the Rio Caroni, an enormous area 
for which no specimens have been recorded. 

Specimens Examined— Total 24. Bolivar: Rio 
Curumo (7°15'N, 61°20'W); Rio Grande (8°16'N, 
6 ri 7'W); Gurisoco, El Palmar (8°62'N, 6 1°26'W); 
La Trinidad, El Palmar (7°12'N, 6r23'W); Guri, 
Operacion Rescate (7°18'N, 63°00'W); Rio Boto- 
namo, near Rio Cuyuni (6°59'N, 6rirW); San 
Martin de Turumban, Rio Cuyuni (6''59'N, 
61°02'W). Territorio Federal Delta Amacuro: Al- 
tiplanicie de Nuria (7°50'N, 61°18'W); Yotacuay, 
SW of Cupiare (8°30'N, 61°00'W). 



Literature Cited 

Allen, J. A. 1914. New South American Monkeys. 
BviUetin of the American Museum of Natural History, 
33: 653. 

BoDiNi, R. 1981. Musculatura locomotora de la viudita 
(Callicebus torquatus lugens). Sus implicaciones fun- 
cionales y iilogeneticas. Memoria de la Sociedad de 
Ciencias Naturales La Salle, 16(XL): 1-165. 

. 1983. Distribuci6n, status de la investigacion 

y conservaci6n de los cebidos en Venezuela, p. 1 39. 
In Symjwsio sobre Primatologia en Latinoamerica. IX 
Congreso Latinoamericano de Zoologia, Arequipa, 
Peru. 

Cabrera, A. 1957(1958]. Calalogo de los mamiferos 
de America del sur. Revista Museo Argentine de Cien- 
cias Naturales, "Bernardino Rivadavia," 4(1): xviii + 
138 pp. 

Cruz Lima, E. 1945. Mammals of Amazonia. I. Gen- 
eral introduction and Primates. Contribution from the 



BODINI & PEREZ-HERNANDEZ: CEBIDS IN VENEZUELA 



243 



Museu Paraense Emilio Goeldi de Historia Natural e 
Etnografia, Belem do Para, Rio de Janeiro, Brasil, 274 
pp., 42 pis. 

EiSENBERG, J. F., AND K. Redford. 1979. A biogeo- 
graphic analysis of the mammalian fauna of Vene- 
zuela, pp. 31-36. In Eisenberg, J. F., ed., Vertebrate 
Ecology in the Northern Neotropics. Smithsonian In- 
stitution Press, Washington, D.C., 271 pp. 

Elliot, D. G. 1912. A Review of the Primates. Vols. 
I and II. American Museum of Natural History, New 
York. 

FooDEN, J. 1963. A revision of the woolly monkey 
(genus Lagothrix). Journal of Mammalogy, 44: 213- 
247. 

Handley, C. O., Jr. 1976. Mammals of the Smith- 
sonian Venezuelan project. Brigham Young Univer- 
sity Sciences Bulletin, Biological Series, 20(5): 1-91. 

Hernandez-Camacho, J., and R. W. Cooper. 1976. 
The nonhuman Primates of Colombia, pp. 35-69. In 
Thorington, R. W., Jr., and P. G. Heltne, eds.. Neo- 
tropical Primates: Field Studies and Conservation. 
National Academy of Sciences, Washington, D.C. 

Hershkovitz, P. 1949. Mammals of northern Colom- 
bia. Preliminary report No. 4: Monkeys (Primates), 
with taxonomic revisions of some forms. Proceedings 
of the United States National Museum, 98(3232): 323- 
427. 

. 1958. Type localities and nomenclature of some 

American primates, with remarks on secondary hom- 
onyms. Proceedings of the Biological Society of Wash- 
ington, 71: 53-56. 

1963. A systematic and zoogeographic account 



of the monkeys of the genus Callicebus (Cebidae) of 
the Amazonas and Orinoco river basins. Mammalia, 
27: 1-79. 
. 1972. The Recent mammals of the Neotropical 



region: A zoogeographic and ecological review, pp. 
31 1-431. In Keast, A., F. C. Erk, and B. Glass, eds.. 
Evolution, Mammals, and Southern Continents. State 
University of New York Press, Albany, N.Y. 543 pp. 
1977. Living New World Monkeys (Platyrrhi- 



ni) with an Introduction to Primates. University of 
Chicago Press, Chicago, 1117 pp. 

1 979. The species of sakis, genus Pithecia (Ce- 



bidae, Primates), with notes on sexual dichromatism. 
Folia Primatologica, 31: 1-22. 

1983. Two new species of night monkeys, ge- 



nus Aotus (Cebidae, Platyrrhini): A preliminary report 
on Aotus taxonomy. American Journal of Primatol- 
ogy, 4: 209-243. 



. 1984. Taxonomy of squirrel monkeys genus 

Saimiri (Cebidae, Platyrrhini): A preliminary report 
with description of a hitherto unnamed form. Amer- 
ican Journal of Primatology, 7: 155-210. 

Hill, W. C. O. 1 960. Primates. Comparative Anatomy 
and Taxonomy. IV. Cebidae, Part A. Edinburgh Uni- 
versity Press, London, 523 pp. 

. 1962. Primates. Comparative Anatomy and 

Taxonomy. V. Cebidae, Part B. Edinburgh University 
Press, London, 537 pp. 

INPARQUES. 1982. Guia de los Parques Nacionales 
y Monumentos Naturales de Venezuela. Ediciones 
Fundacion de Educacion Ambiental, Caracas. 

Kellogg, R., and E. A. Goldman. 1944. Review of 
the spider monkeys. Proceedings of the United States 
National Museum, 96(3186): 1-45. 

Kjnzey, W. G. 1982. Distribution of primates and for- 
est refuges, pp. 455-482. In Prance, G. T., ed.. Bio- 
logical Diversification in the Tropics. Columbia Uni- 
versity Press, New York, 7 1 4 pp. 

MARNR-DGS-POA-SFS. 1982. Informe Nacional de 
Fauna Silvestre. Ministerio del Ambiente y de los Re- 
cursos Naturales Renovable, Caracas. 

Mittermeier, R. a., and A. Coimbra-Filho. 1981. 
Systematic: Species and subspecies, pp. 29-109. In 
Coimbra-Filho, A. F., and R. A. Mittermeier, eds.. 
Ecology and Behavior of Neotropical Primates. Aca- 
demia Brasileira de Ciencias, Rio de Janeiro. 

MoNDOLFi, E. 1976. Fauna silvestre de los bosques 
humedos tropicales de Venezuela, pp. 113-181. In 
Hamilton, L. S., J. Steyermark, J. P. Veillon, and E. 
Mondolfi, eds., Conservacion de los bosques humedos 
de Venezuela. Sierra Club— Consejo de Bienestar Ru- 
ral, Caracas. 

MoNDOLR, E., AND J. F. EiSENBERG. 1 979. Ncw records 
for A teles belzebuth hybridus in northern Venezuela, 
pp. 93-96. In Eisenberg, J. F., Vertebrate Ecology in 
Northern Neotropics. Smithsonian Institution Press, 
Washington, D.C, 271 pp. 

Ramirez-Cerquera, J. 1983. Reporte de una nueva 
especit de Primates del genero Aotus de Colombia, p. 
146. Symposio sobre Primatologia en Latinoamerica. 
IX Congreso Latinoamericano de Zoologia. Arequipa, 
Peru. 

RuDRAN, R., AND J. F. Eisenberg. 1982. Conservation 
and status of wild primates in Venezuela, pp. 52-59. 
In Olney, P. J. S., ed.. International Zoo Yearbook, 
Vol. 22. The Zoological Society of London, London, 
488 pp. 



244 



FIELDIANA: ZOOLOGY 



Host Associations and Coevolutionary Relationships 
of Astigmatid Mite Parasites of New World Primates 
I. Families Psoroptidae and Audycoptidae 



Barry M. OConnor 



ABSTRACTS 

Coevolutionary patterns among mites of the families Psoroptidae and Audycoptidae and 
New World Primates are reviewed. Host records for primate parasites originally described from 
artificial situations are compared with field collections from Peruvian primates, with most host 
associations verified. A new sp)ecies of Audycoptidae, Saimirioptes hershkovitzi, is described 
from Cebus apella. The psoroptid subfamily Cebalginae is hypothesized to be a monophyletic 
group based upon 10 synapomorphies. Phylogenetic relationships within the Cebalginae are 
reviewed, with cospeciational histories supported for the genera Alouattalges and Schizopodalges 
and their hosts, and a more diffuse cospeciational pattern observed for the genera Cebalgoides, 
Cebalges, and Fonsecalges and their hosts. Historical relationships of the genus Procebalges 
remain problematical. 



Patrones coevolutivos entre acaros de las familias Psoroptidae y Audycoptidae y primates 
del Nuevo Mundo son revisadas. Registros de huespedes para parasitos de primates originar- 
iamente descritos de situaciones artificiales son comparados con colecciones de campo de 
primates peruanos, con la mayoria de asociaciones de huesped verificadas. Una nueva especie 
de Audycoptidae, Saimirioptes hershkovitzi, es descrita de Cebus apella. Los psoroptidos de 
la subfamilia Cebalginae son hipotetizados ser un grupo monofiletico basado en 10 sinapo- 
morfias. Relaciones filogeneticas entre los Cebalginae son revisadas, con historias coesp>ecia- 
cionales soportadas F>or el genero Alouattalges y Schizopodalges y sus huesp>edes, y un patron 
coespeciacional mas difuso observado en los generos Cebalgoides, Cebalges y Fonsecalges y sus 
huesjjedes. Relaciones historicas del genero Procebalges p>ermanecen problematicas. 



Padroes coevolucionarios entre os acarinos das familias Psoroptidae e Audycoptidae, e os 
primatas do Novo Mundo, sao revisados. Os registros de hospedes primatas de parasitas, que 
foram descritos em situa^oes artificials (cativeiros), sao comparados com cole96es de campo 
de primatas peruanos, e a maioria das associagoes hospedeiras atualmente registradas foram 
averiguadas. Uma nova esjjecie de Audycoptidae, Saimirioptes hershkovitzi, encontrada em 
Cebus apella e descrita. Baseando-se num estudo de 1 sinapomorfias, prop6e-se ser a subfamilia 
psoroptidea, Cebalginae, um grupo monofiletico. As rela96es filogeneticas entre os Cebalginae 
sao revisadas, e as historias de coesF>ecializa9ao entre os generos Alouattalges e Schizopodalges, 
e seus respectivos hospedes, sao confirmadas. Os padroes de coespecializa9ao entre os generos 
Cebalgoides, Cebalges e Fonsecalges, e seus hospedes, sao mais difusos. Os relacionamentos 
historicos do genero Procebalges permanecem incertos. 



From the Museum of Zoology and Department of Bi- 
ology, The University of Michigan, Ann Arbor, MI 48 109. 

OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 245 



Introduction 

Coevolutionary patterns among hosts and par- 
asites have been the focus of much recent study 
and discussion (Brooks, 1979, 1981, 1985; Brooks 
& Glen, 1982; Futuyma & Slatkin, 1983; Nitecki, 
1983). Phylogenetic analysis of evolutionary re- 
lationships among parasite groups or host groups 
can provide additional data sets (e.g., treating par- 
asite distributions as character states of their hosts), 
or such analyses can be used as tests for hypotheses 
regarding evolutionary relationships of the other 
lineages of associated organisms. These phyloge- 
netic analyses are especially useful when the par- 
asite groups are host specific and indicate little 
history of secondary colonization of new hosts. 
The associations between mites and primates are 
particularly amenable to such analysis because a 
number of acarine groups are specifically associ- 
ated with primates, and these lineages exhibit 
enough within-group diversity to allow the con- 
struction and comparison of phylogenetic hypoth- 
eses. I have previously detailed such hypotheses 
for several groups of astigmatid mites parasitic on 
primates (OConnor, 1984). 

Formulation of hypotheses regarding the his- 
tory of associations between primates and their 
associated mites requires three steps. First, the 
parasite taxa must be described and their natural 
host and geographic ranges discovered. Second, 
taxa above the species level in classically derived 
classifications must be tested for naturalness (i.e., 
monophyly). Finally, phylogenetic relationships 
among all taxa must be elucidated. In the present 
paper, each of these questions will be addressed 
for the associations among certain groups of as- 
tigmatid mites and New World Primates. For the 
reader interested in summary information regard- 
ing known host-parasite relationships for New 
World primates, an exhaustive list of literature 
records may be found in Hershkovitz (1977). 

One major difficulty in the application of this 
methodology to the study of the history of mite- 
primale associations is the scanty knowledge of 
the distribution and identity of the parasite sp)ecies 
and their natural host ranges. Although many aca- 
rine parasites of primates have been described, 
much of the material has originally come from 
zoos and primate research centers where the f>os- 
sibility of unnatural interspecific contact between 
host species makes the transfer of parasites a real 
problem. Many other sjjecies have been described 
from preserved host specimens in museum col- 



lections where contamination in the field or in the 
museum may also have been a problem. Finally, 
the actual field locality from which either parasites 
or hosts were collected is known for extremely few 
of the known primate parasites. The uncertainties 
involved in the host range and geographic distri- 
bution of so many of the known species of primate 
parasites render the phylogenetic analyses pro- 
posed earlier (OConnor, 1984) subject to some 
doubt. 

In 1 98 1 , 1 was invited by Philip Hershkovitz to 
collect parasites from specimens of a number of 
primate species which had been collected or ob- 
tained during field studies in Peru in 1 980. A very 
large number of mites was collected, providing a 
unique survey of the primate parasites in a small 
area in Peru and a test for previously reported 
host-parasite associations. These collections yield- 
ed specimens belonging to three families of astig- 
matid mites: Psoroptidae, Audycoptidae, and 
Atopomelidae. In the present paper, the collec- 
tions of Psoroptidae and Audycoptidae will be 
discussed. The collections of Atopomelidae, con- 
sisting of a number of described and undescribed 
species of the genus Listrocarpus, will be studied 
separately. 



r 



Materials and Methods 



Primate specimens were collected in the field by 
Hershkovitz or obtained from local individuals. 
These specimens were prepared by removing and 
simply drying the skins at the time of collection. 
Upon their arrival in the United States, one or 
more skins of each species collected were made 
available to me for parasite removal before the 
skins were sent for tanning. The following species, 
all identified by Hershkovitz, were examined 
(number examined in parentheses): Cebus apella 
(3); C. albifrons (1); Lagothrix lagothricha (1); L. 
flavicauda ( 1 ); Alouatta seniculus ( 1 ); Pithecia hir- 
suta (3); Callicebus moloch (1); Aotus nancymai 
(4); and Saimiri sciureus (5). 

Parasites were removed in two ways. First, all 
skins examined were vigorously brushed over white 
paper, with the dislodged parasites collected under 
a dissecting microscop>e. Finally, one skin of each 
species was soaked in water and mild soap until 
soft (the single specimen oi L. flavicauda was not 
soaked). These skins were then gently washed, the 
wash water filtered through a 200-mesh sieve (mesh 



246 



HELDIANA: ZOOLOGY 



openings 75 micrometers), and the residue ex- 
amined under a dissecting microscope. Mites col- 
lected were preserved in 70% ethanol for subse- 
quent study. 

In the laboratory, mites were cleared in lacto- 
phenol and mounted in Hoyer's medium, with 
some specimens retained in alcohol in the cases 
of large series. Voucher specimens of nominal 
species are deposited in Field Museum of Natural 
History, Chicago, and, when available, will be 
placed in the following institutions: Museum of 
Zoology, The University of Michigan, Ann Arbor; 
The United States National Museum of Natural 
History, Washington, D.C.; LTnstitut Royale des 
Sciences Naturelles, Brussels, Belgium; and the 
collection of F. S. Lukoschus, Katholieke Uni- 
versiteit, Nijmegen, Netherlands. 



Species Accounts 



tritonymphs. Among 1 1 protonymphs in the pres- 
ent collection, eight exhibit characteristics similar 
to the male tritonymphs (i.e., posterior opistho- 
somal lobes sclerotized; coxal fields III well scler- 
otized) while three exhibit characteristics of female 
tritonymphs (posterior lobes unsclerotized; scler- 
otization of coxal fields III much weaker). I inter- 
pret these differences as evidence for sexual di- 
morphism at the protonymphal instar in this 
species. 

Material Examined— Total 133. Twenty-nine 
females, 43 males (of which 3 1 were in tandem 
with female tritonymphs), 38 female tritonymphs, 
8 male tritonymphs, 3 female protonymphs, 8 male 
protonymphs, 4 larvae from Cebus albifrons. 
PERU, Loreto: Nauta, Rio Tigre, 6 km above Rio 
Tigrillo; 18 December 1980; P. Hershkovitz (9264). 
Host now a tanned skin (fmnh 122795). Mites 
labeled bmcx: 81-081 1-3. No specimens were re- 
covered from Cebus apella. 



Family PSOROPTIDAE 

Seven species of mites in the family Psoropti- 
dae, subfamily Cebalginae, are known to parasitize 
New World primates. These species, with their 
known hosts and distributions, are listed below, 
with new records from the Peruvian collections of 
Hershkovitz indicated under "Material Exam- 
ined." Keys to most of these species may be found 
in Fain (1963c). 



Cebalgoides cebi Fain, 1963 

Cebalgoides cebi Fain, 1 963, Bull. Ann. Soc. Roy. Ent. 
Belg., 99: 331. Fain, 1963, Bull. Inst. Roy. Sci. Nat. 
Belg., 39(32): 91. 

This species was briefly diagnosed by Fain 
(1963a) from specimens collected from Cebus al- 
bifrons which originated in "Amerique du Sud" 
and died in the Antwerp (Belgium) Zoo. It was 
more thoroughly described and illustrated by Fain 
(1963c), who listed specimens from Cebus albi- 
frons from Venezuela (type collection), C apella 
from "Amerique du Sud," and Leontocebus (Oed- 
ipomidas) oedipus (= Saguinus oedipus) from Co- 
lombia. All these hosts had died in the Antwerp 
Zoo. 

Fain (1963c) noted sexual dimorphism in tri- 
tonymphs of C cebi. The two protonymphs he 
examined exhibited the characteristics of the male 



Alouattalges corbeti Fain, 1963 

Alouattalges corbeti Fain, 1963, Bull. Inst. Roy. Sci. 

Nat. Belg., 39(32): 122. Fain, 1966, Acarologia 8: 

103. 
Rosalialges cruciformis Lavoipierre, 1 964, Acarologia, 

6: 348. 

This species was briefly described from the ho- 
lotype female collected from a preserved specimen 
of Alouatta seniculus macconnelli which had been 
collected at Paramaribo, Surinam and preserved 
in the British Museum (Natural History) (Fain, 
1 963c). Although only the holotype was described, 
several specimens were apparently recovered from 
this host. Almost simultaneously, Lavoipierre 
(1964a) described and figured the female based 
upon two specimens collected from an "Aotes'''' 
(sic) sp. which had died in San Francisco, Cali- 
fornia, after its importation from Peru. Fain (1966) 
provided illustrations of the female and illustrated 
but did not describe the male beyond length and 
width measurements. 

Material Examined— Total 15. Five females, 
8 males, 2 female tritonymphs from Alouatta se- 
niculus. PERU, Loreto: Nauta, Rio Samiria; 18 
November 1980; P. Hershkovitz (9050). Host now 
a tanned skin (fmnh 122789). Mites labeled bmoc 
81-0809-5. No specimens were recovered from 
Aotus nancymai, suggesting the possibility that the 
two specimens collected by Lavoipierre (1964a) 
represent contamination. 



OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



247 



Schizopodalges lagothricola Fain, 1 963 

Schizopodalges lagothricola Fain, 1963, Bull. Ann. Soc. 
Roy. Ent. Belg., 99: 469. Fain, 1963, Bull. Inst. Roy. 
Sci. Nat. Belg., 39(32): 100. 

This species was described from numerous spec- 
imens from two juvenile Lagothrix lagothricha 
which died in the Antwerp Zoo. The origin of the 
hosts was stated as "Amerique du Sud" (Fain, 
1 963b). Fain ( 1 963c) provided illustrations of male 
and female and indicated that the hosts originated 
in Colombia. 

Material Examined— Total 30. Eleven fe- 
males, 1 1 males, 3 tritonymphs, 2 protonymphs, 
3 larvae from Lagothrix lagothricha. PERU, Lo- 
reto: Nauta, Rio Samiria; 15 November 1980; P. 
Hershkovitz (9032). Host now a tanned skin (fmnh 
122790). Mites labeled bmcx: 81-0809-10. 



Cebalges gaudi Fain, 1 962 

Cebalges gaudi Fain, 1962, Rev. Zool. Bot. Afr., 66: 
160. Fain, 1963, Bull. Inst. Roy. Sci. Nat. Belg., 
39(32): 81. 

This species was briefly described from numer- 
ous sp)ecimens collected from a preserved speci- 
men of Cebus capucinus with no locality infor- 
mation (Fain, 1962). Full descriptions and figures 
were provided later (Fain, 1963c). 

Material Examined— Four females from Ce- 
bus apella. PERU, Loreto: Nauta, Rio Samiria; 18 
November 1980; P. Hershkovitz (9049). Host now 
a tanned skin (fmnh 122792). Mites labeled bmoc 
81-0811-2. 



Fonsecalges johnjadini Fain, 1962 

Fonsecalges johnjadini Fain, 1 962, Rev. Zool. Bot. 
Afr., 66: 161. Fain, 1963, Bull. Ann. Soc. Roy. Ent. 
Belg., 99: 468. Fain, 1963, Bull. Inst. Roy. Sci. Nat. 
Belg., 39(32): 86. 

This species was briefly described from several 
Caliithrixjacchus which had died in captivity (Fain, 
1 962). Fain ( 1 963b) provided an illustration of the 
male, and Fain (1963c) gave a complete descrip- 
tion with figures of both male and female. The 
hosts, which were listed as eight Hapale jacchus 
(= Callithrix jacchus) imported from the "bassin 
de I'Amazone," died in Antwerp. Additional sjiec- 



imens were recorded from a specimen of Hapale 
jacchus leucocephalus (= Callithrix jacchus geof- 
froyi) collected in Bahia, Brazil, and preserved in 
Brussels. 
Material Examined— None. 



Fonsecalges saimirii Fain, 1963 

Fonsecalges saimirii Fain, 1963, Bull. Ann. Soc. Roy. 

Ent. Belg., 99: 330. Fain, 1963, Bull. Inst. Roy. Sci. 

Nat. Belg., 39(32): 90. Fain, 1966, Acarologia, 8: 

107. 
Dunnalges lambrechti Lavoipierre, 1 964, Acarologia, 

6: 343. 

This species was briefly described from numer- 
ous specimens collected from two Saimiri sciureus 
from "Amerique du Sud" and which died in the 
Antwerp Zoo in 1959 and 1963 (Fain, 1963a). 
Fain (1963c) provided further descriptive infor- 
mation and indicated the original hosts had come 
from "Amazonie" and had died shortly after their 
arrival at the Antwerp Zoo. He also mentioned 
additional specimens from "Tamarins spp. ori- 
ginaires d' Amazonie." The inexact locality infor- 
mation leaves the actual identity of the type host 
in doubt following the revision of the genus Sai- 
miri by Hershkovitz (1984). Lavoipierre (1964a) 
described and illustrated this species as Dunnalges 
lambrechti, from "a long series of sjiecimens com- 
prising all stages" from several Tamarinus nigri- 
collis (= Saguinus nigricollis) from eastern Peru 
which died in San Francisco, California. Fain 
( 1 966) illustrated parts of this species from the 
type specimens. 

Fain (1963c) did not observe sexual dimor- 
phism in the nymphs of this species, as only two 
tritonymphs and some shed tritonymphal cuticles 
were examined. Sexual dimorphism was noted in 
the tritonymphs during the present study. All lar- 
vae and protonymphs examined bear three pairs 
of sclerotized apophyses in the ventrolateral re- 
gion: a rounded apophysis immediately posterior 
to trochanter II, a pointed apophysis lateral to the 
base of leg III, and a pointed apophysis poster- 
iolaterally near the posterior lobes. Tritonymphs 
in which these apophyses are retained are here 
interpreted as males, while those in which all 
apophyses are lost are interpreted as females. 

Material Examined— Total 59. Three females 
from Saimiri sciureus macrodon. PERU, Loreto: 
Nauta, Rio Tigre, 5 km above Rio Tigrillo; 17 
December 1980; P. Hershkovitz (9257). Host now 



248 



FIELDIANA: ZOOLOGY 



a tanned skin (fmnh 122810). Mites labeled bmoc 
81-0809-15. Two females, 1 male from same host 
species and locality; P. Hershkovitz (9258, fmnh 
122811, BMOC 8 1 -0809- 1 6). Twenty-four females, 
8 males, 5 female tritonymphs, 8 male trito- 
nymphs, 5 protonymphs, 1 larva from same host 
species. PERU, Loreto: Nauta, Rio Tigre, 6 km 
above Rio Tigrillo; same date; P. Hershkovitz 
(9268, FMNH 1 228 1 6, bmoc 8 1 -0809- 1 8). Two fe- 
males from same host sjjecies and locality; P. 
Hershkovitz (9267, fmnh 1 228 1 5, bmoc 8 1 -0809- 
19). 



Procebalges pitheciae Fain, 1963 

Procebalges pitheciae Fain, 1963, Bull. Ann. Soc. Roy. 
Ent. Belg., 99: 332. Fain, 1963, Bull. Inst. Roy. Sci. 
Nat. Belg., 39(32): 96. 

This species was briefly described from speci- 
mens collected from a Pithecia monachus im- 
ported from "Amerique du Sud" and which died 
in the Antwerp Zoo (Fain, 1963a). The descrip- 
tions were completed and illustrations provided 
later (Fain, 1963c). 

Material Examined— Total 242. Nineteen fe- 
males, 28 males (7 in tandem with female trito- 
nymphs), 15 female tritonymphs, 1 protonymph 
from Pithecia hirsuta. PERU, Loreto: Nauta, Rio 
Samiria; 30 November 1980; P. Hershkovitz 
(9 1 1 5). Host now preserved as a tanned skin (fmnh 
122797). Mites labeled bmoc 81-0809-7. Forty- 
seven females, 8 1 males ( 1 7 in tandem with female 
tritonymphs), 22 female tritonymphs, 8 male tri- 
tonymphs, 13 protonymphs, 8 larvae from same 
host and locality; P. Hershkovitz (9088, fmnh 
122796, BMOC 81-0809-9). 



Family AUDYCOPTIDAE 

Three species of hair follicle inhabiting mites of 
the family Audycoptidae have been previously de- 
scribed from New World primates, all from squir- 
rel monkeys identified as Saimiri sciureus. With 
the recent recognition of several valid species in 
the genus Saimiri (Hershkovitz, 1984), the exact 
identification of the reported hosts is problemat- 
ical. Specimens representing a fourth sr>ecies were 
recovered from Cebus apella from the present col- 
lections. 



Audycoptes greeri Lavoipierre, 1 964 

Attdycoptes greeri Lavoipierre, 1 964, Ann. Natal Mus., 
16: 194. 

This species was described from females col- 
lected from the sinus-hair follicles of Saimiri sci- 
ureus collected in eastern Peru and kept in captiv- 
ity in California (Lavoipierre, 1964b). In the 
absence of more detailed collection information, 
the true host may have been either S. sciureus or 
S. boliviensis, as both occur in eastern Peru (Hersh- 
kovitz, 1984). 

Material Examined— Five females from "5fl/- 
muri sciurea" (sic), without further collection data, 
from the Lavoipierre collection. University of Cal- 
ifornia, Davis. 



Audycoptes lawrencei Lavoipierre, 1964 

Audycoptes lawrencei Lavoipierre, 1964, Ann. Natal 
Mus., 16: 199. 

This species was described from the same hosts 
and habitat as Audycoptes greeri (Lavoipierre, 
1964b), so the actual specific identity of the host 
remains uncertain, as indicated for greeri. 

Material Examined— Two females from "^a/- 
muri sciurea"" (sic), without further collection data, 
from the Lavoipierre collection, University of Cal- 
ifornia, Davis. 



Saimirioptes paradoxus Fain, 1968 

Saimirioptes paradoxus Fain, 1968, Acarologia, 10: 
286. 

This species was described from a single female, 
containing a larva, collected from a Saimiri sci- 
ureus which had died in the Antwerp Zoo (Fain, 
1968). No information concerning the geographic 
origin of the host was given, making the specific 
identity of the host uncertain. 

Material Examined— None. 



Saimirioptes hershkovitzi, new species 

In the following description, all measurements 
are given in micrometers and are presented as ho- 
lotype (range of three measured specimens). 

Female (figs. 1-2)— Body elongate, cylindrical 



OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



249 




Fig. 1 . Saimirioptes hershkovitzi, female. A, dorsum; b, venter. 



to somewhat flattened dorsoventrally; length in- 
cluding gnathosoma 4 1 5 (4 1 5—42 1 ), width at level 
of transverse coxal apodemes III 146 (135-146); 
entire body with transverse striations which are 
very thick anteriorly, thin posteriorly. 

Dorsum (fig. la)— Prodorsal sclerite narrow an- 
teriorly, much widened medially, narrowed pos- 
teriorly and fused internally with apodemes of cox- 
al fields II in most posterior region. Sclerite 
traversed by anterior transverse striations later- 
ally. Length of sclerite 70 (70-77). Paired dorsal 
protuberances present mesal to setae h and sh. with 



striations encircling protuberances along most of 
their length; lengths 29 (28-29). Idiosomal chae- 
totaxy as follows: scapular setae filiform, in a 
transverse line posterior to prodorsal sclerite, 
lengths sci 4 (4-5), see 39 (29-39); setae J, and /, 
very short and rounded, subdivided into two dis- 
tinct lobes; setae h and sh filiform, lengths 9 (6- 
9); setae di subdivided into three parts, a ventral 
filiform part, length 26 (23-26), and two dorsal 
rounded lobes; setae d^ similar to ^2 but filiform 
part shorter, length 19 (14-19), and middle lobe 
more elongate; remaining setae filiform; /j 29 (28- 



250 



FIELDIANA: ZOOLOGY 




< > 



OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



251 



29); /, ventrally positioned, 18 (18-19); ^^4 24 (19- 
24); ^5 21 (no variation); l^ ventral, 5 (4-5); l^ 
ventral, 30 (27-32); a, ventral, 14 (no variation); 
a, 30 (27-30). Four pairs of idiosomal cupules 
present: ia dorsal between setae h and sh; im dorsal 
between and lateral to setae d2 and d^; ip ventral 
slightly anterior to seta /j; ih ventral, between and 
mesal to setae U and /j. Opisthosomal glands pres- 
ent, small, opening between and lateral to setae d^ 
and /,. Anus terminal. Opening to bursa copulatrix 
terminal, above anal opening. 

Venter (fig. lb)— Coxal apodemes well devel- 
oped. Anterior apodemes of coxal fields I straight, 
length 66 (57-66), fused medially to form a ster- 
num of length 23 (23-24). Anterior apodemes of 
coxal fields II curving mesally, length 75 (75-79); 
posterior apodemes of coxal fields II transverse, 
fused ventromedially with anterior apodemes of 
coxal fields III, extending dorsally, length of ven- 
tral portion 50 (46-50), length of dorsal portion 
31 (28-31). Anterior apodemes of coxal fields III 
extending almost longitudinally from trochanters 
III to F>osterior apodemes of coxal fields II, then 
bending at right angles to fuse with the latter, length 
of longitudinal portion 49 (49-51), transverse 
portion 39 (39-44). Anterior apodemes of coxal 
fields IV very wide, anteriomedially directed, length 
50 (50-53). Ovipore located between coxal fields 
III and IV; a crescentic epigynal apodeme posi- 
tioned anterior to ovipore; genital valve elongate, 
disappearing posteriorly under transverse flap; a 
pair of small thickenings along posterior, internal 
portion of valves; two pairs of vestigial genital 
papillae lateral to genital ojiening. Three pairs of 
filiform coxal setae present: cxI at posterior end 
of coxal fields I between coxal apodemes II, length 
12 (9-12); cxIII on median apex of coxal apo- 
demes IV, length 9 (6-9); cx/P'mesad of the base 
of leg IV, length 7 (6-7). Genital setae absent. 

Gnathosoma (fig. 2g)— Gnathosoma elongate, 
somewhat widened posteriorly, then tapering to a 
point ventrally, length (excluding chelicerae) 62 
(59-62), maximum width 26 (24-26). Chelicerae 
elongate, with small, toothed digits, length 46 (44- 
46). Palps apparently two-segmented; basal seg- 
ment bearing two very short, filiform setae, 1 prox- 
imodorsal, 1 medioventral; distal segment with 
short solenidion and 5 sclerotized, pxjinted pro- 
jections. Subcapitulum with pair of short, filiform 
setae medioventrally; with pair of sclerotized, 
pointed projections positioned slightly anterior to 
subcapitular setae, and unpaired projection ante- 
riomedially; median rutellar lobes with sclerotized 
points medially. 



Legs (figs. 2a-f) — Legs I-II similar in structure, 
with all segments free, lengths 84 (80-90). Femora 
I-II bearing two ventral apophyses, one filiform 
seta vF. lengths: I, 9 (8-9); II, 33 (30-33). Genua 
I-II each with large ventral apophysis; setae cG 
expanded basally, then tapering, lengths 22 (20- 
25), setae mG filiform, lengths 43 (40-45). Tibiae 
I-II each with large ventral apophysis; setae gT 
short, filiform, lengths 6 (5-6); solenidia api- 
codorsal, lengths: I, 20 (18-20); II, 22 (21-23). 
Tarsi I-II each with pointed dorsal apophysis, two 
distal hooked apophyses; chaetotaxy of tarsi I-II 
similar; setae d as strongly hooked, apical claws, 
lengths 5 (no variation); e filiform, lengths 40 (43- 
50); /filiform, lengths 7 (6-7); la filiform, lengths 
6 (5-6); wa very short, not measurable; ra filiform, 
lengths 6 (5-6); solenidia «, blunt apically, lengths: 
I, 14 (no variation); II, 26 (24-26). Pretarsi I-II 
similar, total lengths 40 (37-40); pretarsi divided 
into long ambulacral stalk and rounded ambula- 
cral disc; ambulacral disc with condylophore guide 
and ventral "rays" (fig. 2b); condylophores atten- 
uate, appearing to divide at base of ambulacral 
disc. 

Legs III-IV similar, each with fused trochanter- 
femur bearing a large, ventral apwaphysis. Genua 
III-IV glabrous. Tibiae III-IV each with setae kT 
enlarged, bifurcate, and deeply rooted; tibia III 
with solenidion 0, length 6 (6-7), IV absent. 
Tarsi III-IV each with apical, pointed apophysis, 
three setae: fi^ apical, lengths: III, 100(91-100); IV, 
97 (86-97); r filiform, lengths 7 (6-7); w filiform, 
lengths 3 (no measurable variation). 

Types— Holotype and two paratype females from 
Cebus apella. PERU, Loreto: Nauta, Rio Samiria; 
18 November 1980; P. Hershkovitz (9049). Host 
now a tanned skin (fmnh 122792). Mites labeled 
BMOC 81-0811-2. Holotype and 1 paratype de- 
posited in Field Museum of Natural History, Chi- 
cago; 1 paratype in Museum of Zoology, Univer- 
sity of Michigan, Ann Arbor. 

Systematic Position— Saimirioptes hershko- 
vitzi shares with 5". paradoxus, the tyF>e-sp)ecies and 
only other member of the genus, the presence of 
dorsal lobes between setae ^1 and t/j^ the similar 
form of setae t/, and /, (rounded, bifurcate, and 
without filiform part), and the expansion of the 
bases of setae cG of genua I-II. The new sF>ecies 
differs from the type-species in the relative lengths 
of the scapular setae (sce:sci 5-6:1 in S. hershko- 
vitzi, 1.5:1 in S. paradoxus); the shorter filiform 
fKJrtion of setae f/, (not reaching dy in S. hersh- 
kovitzi, extending past d^ in S. paradoxus); the 
shorter lengths of setae 4 (29 in S. hershkovitzi. 



252 



HELDIANA: ZOOLOGY 



67 in S. paradoxus); the presence of setae U (absent 
in S. paradoxus); the longer pretarsi I-II (37-40 
in S. hershkovitzi, 17 in S. paradoxus); and the 
greater lengths of setae d on tarsi III-IV (at least 
twice as long as the entire leg in S. hershkovitzi, 
less than half the leg length in S. paradoxus). 



Discussion 

The Peruvian collections reported upon here 
support the hypothesis that most of the records 
of primate-Cebalginae associations previously re- 
ported reflect natural host-parasite associations. 
The association between Alouattalges corbeti and 
Aotus species was not verified and remains ques- 
tionable. As there were no field collected repre- 
sentatives of the Callitrichidae examined during 
this study, the occurrence of cebalgine mites on 
these hosts remains generally untested in natural 
situations, with only Fonsecalges johnjadini re- 
corded from noncapti ve callitrichids (Fain, 1 963c). 

Having established that most records of pri- 
mate-cebalgine associations reflect natural asso- 
ciations, two additional hypotheses must be pro- 
posed and tested before any coevolutionary 
hypotheses may be tested. These preliminary hy- 
potheses concern the monophyly of the Cebalginae 
and the phylogenetic relationships of the taxa 
within the group. I have previously discussed these 
questions (OConnor, 1 984) but, due to space con- 
straints, was unable to detail the reasoning behind 
my conclusions. 



Monophyly of the Cebalginae 

Before any hypothesis of historical associations 
may be tested, at least one of the lineages must be 
hypothesized to be monophyletic (Brooks, 1981). 
In this study, the psoroptid subfamily Cebalginae, 
which comprises all psoroptid mites parasitizing 
New World Primates, must be tested for mono- 
phyly. In last defining this subfamily. Fain ( 1 963c) 
listed 21 character states for the group. Because 
the comprehensive morphological studies of Fain 
(1963c) were carried out before the methods of 
phylogenetic systematics became widely discussed 
and utilized, no distinction between ancestral and 
derived states was made in the diagnosis of the 
Cebalginae. In order to test whether the Cebalgi- 
nae represents a monophyletic group, I have ex- 
amined the characters listed by Fain (1963c) using 



outgroup comparison to polarize the states. I have 
previously hypothesized that the taxa comprising 
the four subfamilies of Psoroptidae which para- 
sitize the Primates form a monophyletic group 
(OConnor, 1984). These taxa include the Makial- 
ginae (sensu OConnor, 1984; i.e., including the 
Cheirogalalginae and Galagalgidae of Fain), par- 
asites of the Strepsirrhini; the Paracoroptinae, par- 
asites of African Cercopithecidae and Hominidae; 
the Nasalialginae, parasites of Asian Cercopithe- 
cidae; and the Cebalginae, from New World pri- 
mates. This grouping of taxa is regarded as the 
ingroup in the following analysis. Outgroups used 
in defining the character state polarities were the 
other subfamilies of Psoroptidae and earlier de- 
rivative groups in the Astigmata. 



Character Analysis 

1 . Presence of retrograde apophyses on coxa! 
fields III. This state is unique to the cebalgine 
genera, with such apophyses not occurring in other 
taxa in the ingroup or outgroup. 

2. Male body size much smaller than female. 
In the outgroups and other ingroups, males are 
similar in size to females or somewhat larger or 
smaller. The substantial reduction in body size of 
male Cebalginae is unique. 

3. Female opisthosoma more or less squared 
posteriorly and laterally, reinforced with sclero- 
tized areas. This condition is exhibited by all Ce- 
balginae, but not in any outgroup. Among the in- 
groups, the squared body is unique to the 
Cebalginae, but sclerotized reinforcement is also 
found in the Nasalialginae and some Paracorop- 
tinae (Pangorillalges). Among outgroup taxa, 
sclerotized reinforcement is present in some Pso- 
ralginae (Edentalges), but again without the squared 
body form. 

4. Male with legs III very modified, with 3-4 
terminal segments fused and bearing medially di- 
rected projections. The modification of the third 
pair of legs in male Cebalginae is unique and pres- 
ent in all taxa. No similar modification occurs 
anywhere in ingroup or outgroup. 

5. Reduction or loss of paranal suckers in males. 
Paranal suckers are present in males in the out- 
groups and ingroups. In the Cebalginae, the suck- 
ers are very reduced or absent. 

6. Tarsus I with 2 apical solenidia. I regard the 
apical displacement of solenidion oj, in the Ce- 
balginae as derived. In the other ingroups and most 
outgroups, this solenidion is median or basal on 



OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



253 



the tarsus. Apical displacement of this solenidion 
also occurs in the subfamily Psoroptinae and the 
monobasic Marsupialginae, conditions I regard as 
convei^ent. 

7. Loss of dorsal seta ^,. Seta d^ is absent in all 
Cebalginae and retained in the other ingroups and 
most outgroups. This seta is also lost in the Lis- 
tropsoralginae and the psoroptid parasites of ro- 
dents (Echimyalges. Myoproctalges, and Coen- 
dalges), conditions I regard as convergent. 

8. Reduction or loss of apophyses from the pos- 
terior tarsi of the female. Apophyses are present 
on the posterior tarsi of some of the outgroups 
such as the Listropsoralginae and are retained in 
some ingroups, the Makialginae and Paracorop- 
tinae. I consider the presence of these apophyses 
to be plesiomorphic for the primate-associated 
psoroptid lineage. In the Cebalginae, taxa in which 
females have well-developed posterior legs retain 
vestiges of these apophyses, while taxa in which 
the legs are reduced retain no traces. I consider 
this reduction to be a derived state for the Cebal- 
ginae and regard the loss of these structures in the 
Cebalginae and Nasalialginae as convergent. 

9. Posterior edge of female opisthosoma with 
2 pairs of long, strong setae. In most ingroups and 
psoroptid outgroups, seta /, is long and strong while 
seta di is shorter and thinner. Of the ingroups, 
setae l^ and d^ are equally well developed in the 
Cebalginae and Nasalialginae. However, in the Ce- 
balginae, setae /, and d^ are closely associated, usu- 
ally on a single projection, while in the Nasalial- 
ginae these setae are separate, on distinctly different 
projections. I regard these two conditions as con- 
vergent. I also regard as convergence the elonga- 
tion of seta di in some Psoralginae. 

10. Dorsal seta l^ sometimes absent. Seta I4 is 
absent in all Cebalginae except Cebalgoides cebi. 
The seta is retained in that species, the other in- 
groups, and most outgroups. The loss of seta I4 
could be regarded as a synapomorphy defining a 
group containing all Cebalginae except Cebal- 
goides. However, this hypothesis conflicts with 
groupings suggested by all other characters (see 
page 256). At this point, it is more parsimonious 
to regard the loss of seta U as a synapomorphy for 
the Cebalginae, with a reversal in Cebalgoides. 

1 1 . Loss of retrograde apophyses on the ante- 
rior legs. Retrograde apophyses are present on the 
anterior legs in some of the outgroups, notably the 
Listropsoralginae and Marsupialginae in the Pso- 
roptidae and in the related families Audycoptidae, 
Rhyncoptidae, and Myocoptidae. Among the in- 
groups, these apophyses are retained in the Ma- 



kialginae but lost in the Cebalginae, Paracorop- 
tinae, and Nasalialginae. I regard this loss as 
characterizing these latter three groups as a natural 
unit and thus plesiomorphic for the Cebalginae. 

12. Female bursa copulatrix subterminal or 
ventral. In the outgroups, the female bursa cop- 
ulatrix is terminal, as it is in the Cebalginae, Ma- 
kialginae, and Nasalialginae among ingroups. I re- 
gard the dorsal position of the bursa in the 
Paracoroptinae as derived. Thus, this character 
retains the plesiomorphic condition in the Cebal- 
ginae. 

13. Base of the gnathosoma with retrograde 
apophyses. In some outgroups as well as in the 
Cebalginae, Makialginae and some Paracordpti- 
nae (i.e., Pangorillalges), the base of the gnatho- 
soma bears retrograde apophyses. These are ab- 
sent in certain outgroups (e.g., the Psoroptinae) 
and in Paracoroptes and Nasalialges among in- 
group taxa. I regard the retention of these apoph- 
yses in the Cebalginae as plesiomorphic. 

14. Absence of retrograde apophyses on coxae 
I-II. Retrograde apophyses occur on coxae I-II 
only in the Makialginae. They do not occur on 
both coxal fields in any other ingroup or outgroup 
taxa, although the Listropsoralginae and Myocop- 
tidae have apophyses on coxae II. I regard, these 
structures as synapomorphies characterizing the 
Makialginae, and their absence in the Cebalginae 
as plesiomorphic. 

15. Well-developed "claws" on tarsi I-II. 
Clawlike apophyses on the apices of tarsi I-II are 
present in many outgroup taxa, including both 
mammal and bird parasites. I regard the presence 
of this character state as ancestrally characterizing 
all Psoroptidia, and thus plesiomorphic for the 
Cebalginae. 

16. Tarsus II with solenidion apically dis- 
placed. Solenidion oj is apically displaced in the 
Cebalginae, Paracoroptinae, and Nasalialginae. In 
most outgroups and in the Makialginae, the so- 
lenidion retains its ancestral, basal position. I re- 
gard this displacement as diagnosing a natural 
group consisting of the above three primate-as- 
sociated subfamilies and thus plesiomorphic for 
the Cebalginae. 

17. Female tarsal chaetotaxy 7-7-6—4 or -5. 
Possession of 7 setae on tarsi I-II involves the loss 
of seta ba in the Paracoroptinae and Cebalginae 
and probably the Nasalialginae. Retention of 8 
setae on these segments (including ba) is charac- 
teristic for most outgroup taxa and the Makial- 
ginae. I regard this state as diagnosing a natural 
group comprising the Cebalginae, Paracoroptinae, 



254 



HELDIANA: ZOOLOGY 



and Nasalialginae and thus plesiomorphic for the 
Cebalginae. 

1 8. Angles of female opisthosoma generally with 
strong hooks. Such hooks are found in the cebal- 
gine genera Cebalges, Cebalgoides. and Fonse- 
calges, but not in other taxa in the outgroup or 
ingroup. I regard the presence of these structures 
as characterizing a monophyletic unit within the 
Cebalginae but not the group as a whole. 

19. Posterior legs of female normal or atro- 
phied. Legs III-IV of the female are reduced in 
size and may exhibit fusion of segments in the 
genera Cebalges and Fonsecalges but not in other 
Cebalginae nor other members of the ingroup. 
Well-developed legs are characteristic of most out- 
group taxa, although similar reduction of the pos- 
terior legs occurs in some Psoroptinae and Pso- 
ralginae. I regard the atrophied legs of some 
Cebalginae as characterizing a smaller monophy- 
letic unit within the group and convergent with 
the outgroup taxa noted above. 

20. Posterior legs of nymphal stages normal or 
short and atrophied and bearing a long seta. The 
reduction of the legs in the nymphs is found in all 
cebalgine genera except Procebalges. It is also found 
in a few outgroup taxa (e.g., the Psoroptinae and 
Psoralginae), but not in most outgroups or in other 
ingroup taxa. I regard this state as diagnosing a 
group within the Cebalginae and not characteriz- 
ing the group as a whole. 

2 1 . Dorsal seta d^ sometimes absent. Seta d^ is 
absent in Schizopodalges lagothricola and Fon- 
secalges johnjadini (but not F. saimirii), and pres- 
ent in other Cebalginae, other ingroups, and most 
outgroups. The distribution of this derived state 
within the Cebalginae strongly suggests the inde- 
pendent loss of this seta in the two species sharing 
the state. This character at best diagnoses a group 
within the Cebalginae, but is more likely a case of 
convergence. In no way does this character diag- 
nose the group as a whole. 



Conclusions 

On the basis of the above analysis, I conclude 
that, of the 21 character-states listed by Fain 
(1963c) as diagnosing the Cebalginae, five repre- 
sent unique synapomorphies which diagnose the 
Cebalginae and occur in no other group (character- 
states 1-5), five are synapomorphies which diag- 
nose the Cebalginae but which also occur as con- 
vergent states in certain other taxa (character-states 
6-10), seven are symplesiomorphies diagnosing 



larger groups which include the Cebalginae (char- 
acter-states 11-17), and four represent within- 
group apomorphies diagnosing smaller groups 
within the Cebalginae (character-states 1 8-2 1 ). The 
ten synapomorphies diagnosing the Cebalginae 
leave no doubt that the group is a natural one. 



Phylogenetic Relationships 
Within the Cebalginae 

I have previously presented a hypothesis of phy- 
logenetic relationships among genera in the Ce- 
balginae based upon 17 characters (OConnor, 
1 984). This cladogram is reproduced here (fig. 3), 
with numbers on the cladogram referring to the 
derived states of the characters listed below. In the 
earlier study, space limitations prevented discus- 
sion of the different states of these characters and 
the reasons for interpreting their polarity. Addi- 
tionally, an error in character 1 2 appeared in the 
list of character-states. As I have hypothesized that 
the Cebalginae form a monophyletic group within 
a larger lineage comprising the Makialginae, Par- 
acoroptinae, and Nasalialginae, taxa in these three 
lineages were used as outgroups to polarize the 
character-states within the Cebalginae. 

1. Female with coxal apodemes III-IV fused. 
In all taxa in the outgroups, coxal apodemes III- 
IV end freely. Within the Cebalginae, these apo- 
demes are fused together on either side in the gen- 
era Procebalges, Schizopodalges, and Fonsecalges. 
Although I regard the fused condition to be de- 
rived, conflicts with many other characters suggest 
that these apodemes have fused independently in 
the three genera. 

2. Female epigynum fused with coxal apo- 
demes I. In most outgroup taxa, the female ovi- 
pore is located between coxal fields II and III. 
There has been a trend toward anterior displace- 
ment of the ovip)ore in a number of psoroptid 
lineages (e.g., Psoralginae, Listropsoralginae, Pso- 
roptinae), and the more anterior p>osition occurs 
in certain taxa among the primate associated lin- 
eage as well (e.g., Lemuralges in the Makialginae). 
Within the Cebalginae, this derived state occurs 
in the genera Procebalges and Schizopodalges. Be- 
cause this character-state distribution conflicts with 
many other characters, I regard the presence of 
this state in the two genera as convergence. 

3. Male with paranal suckers lost. In all out- 
groups, the male retains paranal suckers. Within 
the Cebalginae, the suckers are retained only in 



OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



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V, D 





• 11 




















• 2 










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• 1 


■ 12 








• 












■ 14 










• 8 






■ 13 




■ 15 












■ 7 








* 






■ 6 








■ 10 






• 5 








■ 9 


• 2 






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Fig. 3. Phylogenetic relationships among genera in the subfamily Cebalginae. Numbers refer to derived states of 
characters discussed in the text. Hosts of each mite taxon given in parentheses. 



Procebalges. I regard the loss of these structures 
as derived and characterizing the sister group of 
Procebalges. 

4. Immature stages with pretarsi III-IV lost. 
Described immature stages of outgroup taxa all 
retain the pretarsus on legs III-IV. Within the Ce- 
balginae, only Procebalges retains this ancestral 
state. In all other Cebalginae, pretarsi III-IV are 
lost in the immature stages, a condition I regard 
as derived. 

5. Gnathosomal apophyses lost in both sexes. 
As discussed in the previous section, I regard the 
presence of gnathosomal apophyses as ancestral 
in the Cebalginae. These structures are lost in both 
sexes in the genera Schizopodalges and Alouat- 
talges within the Cebalginae, a condition I regard 



as derived. I regard this state as a synapomorphy 
for these two genera and convergent with the sim- 
ilar state occurring in the genera Lemuralges 
(Makialginae), Paracoroptes (Paracoroptinae), and 
Nasalialges (Nasalialginae). I have treated loss of 
these apophyses in the female only as a separate 
character (16, below). 

6. Female with seta 5 of tarsi I-II in the form 
of a strongly hooked claw. In the outgroup taxa, 
seta s is simple and filiform. Within the Cebalgi- 
nae, this state is retained in all taxa except Schi- 
zopodalges and Alouattalges where the seta is en- 
larged and hooked. I regard the latter state as 
derived and a synapomorphy for the two genera. 

7. Male with a large spur on tarsus III. This 
condition is unique to the cebalgine genera Schi- 



256 



FIELDIANA: ZOOLOGY 



zopodalges and Alouattalges. In the outgroups and 
other ingroup taxa, such a spur does not exist. I 
regard the presence of this character-state as in- 
dicative of common ancestry of these two genera. 

8. Male with ridges on leg III restricted to tibial 
element. Ridges on the distal portion of leg III of 
the male characterize the Cebalginae (see above). 
Such ridges do not occur in any outgroup. I regard 
the restriction of the ridges to a spur extending 
from the tibial portion of the leg as a derived state 
defining the group Schizopodalges + Alouattalges 
because this condition forms a functional complex 
with the spur on tarsus III (character 7) and a 
dorsal spur on femur IV. This latter character was 
not considered in the previous analysis (OConnor, 
1984). The conjunction of the three spurs serves 
to lock legs III and IV together into a unit. 

9. Female with recurved hooks on posterior 
border of opisthosoma. Strong hooks are present 
in this position in females of Cebalgoides, Ce- 
balges. and Fonsecalges. Weaker development in 
this area is characteristic of other cebalgine genera, 
while in the outgroups, no projections exist. I re- 
gard the possession of strong hooks as a derived 
condition defining a lineage comprising the three 
genera noted above. 

10. Male with pretarsus III lost. Males retain a 
pretarsus on leg III in most outgroup taxa. This 
pretarsus is lost in the genera Galagalges and Chei- 
rogalalges (Makialginae), and also in the cebalgine 
genera Cebalgoides, Cebalges, and Fonsecalges. I 
regard the loss of pretarsus III as a synapomorphy 
for the latter three genera within the Cebalginae, 
with independent loss in the makialgine lineage 
comprising the former two genera. 

1 1 . Female without opisthosomal sclerite. A 
median sclerite is present in females of most out- 
group taxa and is retained in all Cebalginae except 
Schizopodalges. I regard this loss as derived in the 
latter genus. Convergence in this character occurs 
with some outgroup taxa. This sclerotization is 
also lost in Gaudalges caparti, and Lemuralges 
(Makialginae), and Nasalialges (Nasalialginae). 

12. Female with seta s of tarsi III-IV enlarged 
and clawlike. In the previous study (OConnor, 
1984), a lapsus occurred in that the state "seta 5 
of tarsi III-IV reduced" was listed as a derived 
condition for the genus Alouattalges. In fact, pos- 
session of a small, filiform seta s on these tarsi 
must be regarded as the ancestral condition, as it 
occurs in all outgroup taxa as well as in most Ce- 
balginae. In the genus Alouattalges, seta 5 is en- 
larged and recurved on tarsi III-IV, a condition I 
regard as the true derived state. 



13. Male with opisthosomal lobes widely 
spaced. In most outgroup taxa, the opisthosomal 
lobes of the male are close together, a condition 
also found in most Cebalginae. These lobes are 
widely spaced in the genus Cebalgoides within the 
Cebalginae, a condition I regard as derived. Con- 
vergence occurs, with this state also occurring in 
Nasalialges (Nasalialginae). 

1 4. Male with apodeme between genital and anal 
region strongly reduced. In the outgroup taxa, males 
do not possess a transverse apodeme between the 
genital and anal regions. A large apodeme is pres- 
ent in this position in most male Cebalginae. In 
the prior analysis (OConnor, 1 984), I stated that 
this apodeme was absent in Cebalgoides as indi- 
cated by Fain (1963c) and regarded this as a re- 
versal. Closer examination of a number of speci- 
mens indicates that a very small apodeme is present 
in some males of this taxon. I regard the presence 
of the apodeme as a derived condition for the 
Cebalginae, and its reduction or loss in Cebal- 
goides as a further derived state. 

1 5. Female with tibiae-tarsi III-IV fused. In the 
outgroups, the tibia and tarsus of legs III-IV are 
freely articulated. This condition is present in most 
Cebalginae as well. In the genera Cebalges and 
Fonsecalges, these segments are fused in the fe- 
male, a condition I regard as derived within the 
Cebalginae. Similar fusions occur in some other 
psoroptid subfamilies but not within the primate- 
associated lineage. 

16. Female with gnathosomal apophyses lost. 
Gnathosomal apophyses are retained in females 
of most outgroup taxa and are retained in some 
Cebalginae. These apophyses are lost in the female 
but retained in the male in the genus Fonsecalges. 
I regard this loss as independent of the loss of 
apophyses in the Schizopodalges-Alouattalges lin- 
eage, where the apophyses are lost in both sexes 
(character 5). 

17. Female with pretarsi III-IV lost. Pretarsi 
III-IV are retained in the female in all outgroup 
taxa and in all Cebalginae except Fonsecalges. I 
regard this loss as derived and convergent with the 
similar loss in females in other psoroptid groups 
(e.g., some Psoroptinae, Psoralginae). 



History of Primate-Cebalgine Evolution 

The present knowledge of the diversity and host 
associations of the family Audycoptidae is not suf- 
ficient for formulating hypotheses concerning the 



OCO^fNOR: MITE PARASITES OF NEW WORLD PRIMATES 



257 



history of their host associations. However, given 
the phylogenetic relationships among cebalgine 
taxa presented in the cladogram (fig. 3), two meth- 
ods exist for using these relationships to make hy- 
potheses regarding the history of the associations 
between the New World primates and the Cebal- 
ginae. The first method would involve comparing 
the phylogenetic hypothesis for the parasite group 
with a similar hypothesis for the hosts. The hy- 
pothesis tested by this comparison is whether the 
current pattern of host-parasite associations di- 
rectly results from strict cospeciation between the 
hosts and their parasites. Given a phylogenetic 
hypothesis for each group, such a hypothesis is 
easily tested. The second type of conclusion which 
might be drawn from the parasite cladogram is a 
phylogenetic hypothesis concerning host relation- 
ships. This method is dependent upon the as- 
sumption of cospeciation, or at least noncoloni- 
zation, between the two lineages. 

The current state of knowledge concerning the 
phylogenetic relationships among New World pri- 
mates is somewhat confused. Many early hypoth- 
eses were based upon classical methodology in 
which ancestral and derived character-states were 
not differentiated in proposing hypotheses of re- 
lationships. Schwartz et al. (1978) summarized the 
state of knowledge only a few years ago by saying 
that "Platyrrhini apF>ears in and of itself to be a 
natural group, although both its wider relation- 
ships and the relationships among its members 
remain unclear" (p. 128). Much new information 
relating to this question was presented in symp>osia 
in 1978-1979 and published in a volume edited 
by Ciochon and Chiarelli (1980). In the following 
discussion, I refer to phylogenetic hypotheses for 
New World primates suggested by Rosenberger 
(1977) and contributors to the Ciochon and Chi- 
arelli volume as representing the most modem 
phylogenetic thinking regarding Platyrrhine rela- 
tionships. 

Returning to the question of cospeciation be- 
tween primates and cebalgine mites, the first dif- 
ficulty encountered is the lack of a consensus con- 
cerning phylogenetic relationships among all New 
World primates. Phylogenetic hypotheses based 
on the following types of data sets are in strong 
disagreement: dentition (Rosenberger, 1977; 3 
separate hypotheses); integumentary characters 
(Perkins & Meyer, 1980); karyology (Chiarelli, 
1980); immunological methods (Sarich & Cronin, 
1980); and other immunological data and protein 
sequence data (Baba et al., 1 980). Despite the strong 
disagreement among these data sets, some patterns 



are common to several or all of these. These may 
be compared with the cladogram of mite relation- 
ships to test the cospeciational hypothesis. 

One host relationship which is supported by al- 
most all data sets is the relationship between Al- 
ouatta and the Atelinae (including Lagothrix). Of 
the hypotheses cited above, only the karyological 
evidence did not support this relationship, and 
then only because the karyotype of Lagothrix is 
so derived that Chiarelli ( 1 980) made no hypoth- 
esis as to its relationships. This relationship be- 
tween Alouatta and Lagothrix is mirrored by the 
sister group relationship between their parasites, 
Alouattalges and Schizopodalges, which is among 
the most strongly supported relationships among 
the Cebalginae. 

A second host relationship which is partially 
mirrored in the mile phylogeny is the relationship 
between the genera Cebus and Saimiri, which was 
supported by all data sets except the integumen- 
tary data (Perkins & Meyer, 1980). In this case, 
the mite phylogeny suggests a cospeciational pro- 
cess, but some additional hypotheses are required 
to explain the distributions of the genera Cebal- 
goides, Cebalges, and Fonsecalges. The relation- 
ships among these three parasite genera are com- 
patible with a cospeciational scenario if an early 
sjjeciation event between the Cebalgoides and the 
Cebalges-Fonsecalges lineages occurred in con- 
junction with a common ancestor of Cebus and 
Saimiri. Extinction of the Cebalgoides lineage on 
the Saimiri line, with cospeciation of the Cebalges- 
Fonsecalges lineage in both host lines, leads to the 
present distribution on the Cebidae. In all cases, 
colonization events are required to explain the 
presence of both of these lineages on the Callitrich- 
idae, given the probable monophyly of that taxon. 

Certain other relationships among the parasite 
taxa are not mirrored by host phylogenies. The 
sister group relationship between Procebalges and 
all other cebalgines is not reflected in any hyp)oth- 
esis of host relationships. The relationship of Pi- 
thecia to the Alouatta-Lagothrix lineage was sug- 
gested by both dental data (Rosenberger, 1977) 
and karyology (Chiarelli, 1 980). Interestingly, the 
integumentary data (Perkins &. Meyer, 1980) sug- 
gest that Pithecia retains the most plesiomorphic 
skin characteristics of any mite-bearing New World 
primate (Aotus was regarded as even more plesio- 
morphic). As cebalgine mites are skin inhabitants, 
it might be suggested that plesiomorphic skin re- 
tains plesiomorphic mites, a hypothesis which 
would require the ancestral possession of a Pro- 
cebalges lineage on other cebids with subsequent 



258 



HELDIANA: ZOOLOGY 



extinction on all but Pithecia in order to retain a 
basically cospeciational history. Discovery of ce- 
balgine mites on hosts related to Pithecia (Chi- 
ropotes, Cacajao) will provide a test of these hy- 
potheses. 

The utility of using the proposed mite phylogeny 
to infer a host phylogeny at higher levels than 
mentioned above depends upon the degree to which 
cospeciational patterns outweigh colonizations or 
extinctions in the hypothesized historical relation- 
ships. Among the entire psoroptid lineage para- 
sitizing all Primates, cospeciational patterns ap- 
pear to be supported in a large majority of cases 
(OConnor, 1984). However, noncosj)eciational 
patterns such as the distribution of the genus Le- 
muralges in the Makialginae, or the cebalgine par- 
asites of the Callitrichidae, make the use of these 
mites as consistent indicators of host phylogeny 
at least somewhat suspect. Given the large dis- 
parity in phylogenetic hypotheses generated from 
subsets of the overall character matrix for the New 
World Primates, this parasite data deserves to be 
at least considered by future workers in this area. 



Acknowledgments 

I would like to thank Philip Hershkovitz, with- 
out whose cooperation and interest this study could 
not have been attempted. I also thank Robert 
Timm and Bruce Patterson, Field Museum of Nat- 
ural History, for their hospitality and cooperation 
during the processing of host specimens and for 
their critical review of the manuscript. I thank J. 
H. S. Klompen, University of Michigan, for his 
comments on the manuscript. I thank Margaret 
van Bolt, Museum of Zoology, University of 
Michigan, for assistance with the illustrations. 



Literature Cited 

Baba, M., L. Darga, and M. Goodman. 1980. Bio- 
chemical evidence on the phylogeny of Anthropoidea, 
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Brooks, D. R. 1979. Testing the context and extent of 
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studying the evolution of ecological associations. An- 
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Brooks, D. R., and D. R. Glen. 1982. Pinworms and 
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Chiarelli, A. B. 1980. The karyology of South Amer- 
ican Primates and their relationship to African and 
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Fain, A. 1 962. Diagnoses d'acariens nouveaux. Revue 
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. 1 963a. Nouveaux acariens psoriques parasites 

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. 1 963b. Un nouvel acarien producteur de gale 

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1 963c. Les acariens producteurs de gale chez 



les lemuriens et les singes avec une etude des Psorop- 
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1 966. Les acariens producteurs de gale chez les 



lemuriens et les singes II. Nouvelles observations avec 
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. 1968. Notes sur trois acariens remarquables 



. 1981. Hennig's parasitological method: A pro- 
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. 1985. Historical ecology: A new approach to 



(Sarcoptiformes). Acarologia, 10: 276-291. 

FuTUYMA, D. J., AND M. Slatkin, EDS. 1983. Cocvo- 
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Hershkovitz, P. 1977. Living New World Monkeys 
(Platyrrhini), vol. 1 . University of Chicago Press, Chi- 
cago, 1117 pp. 

. 1984. Taxonomy of squirrel monkeys genus 

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Lavoipierre, M. M. J. 1964a. A note on the family 
Psoralgidae (Acari: Sarcoptiformes) together with a 
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parasitic on primates. Acarologia, 6: 342-352. 

. 1964b. A new family of acarines belonging to 

the Suborder Sarcoptiformes parasitic in the hair fol- 
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191-208. 

NiTECKi, M. H., ED. 1983. Coevolution. University of 
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OConnor, B. M. 1984. Co-evolutionary patterns be- 
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Griffiths, D. E., and C. E. Bowman, eds., Acarology 
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Perkins, E. M., and W. C. Meyer. 1980. The phy- 
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OCONNOR: MITE PARASITES OF NEW WORLD PRIMATES 



259 



331-346. In Ciochon, R. L., and A. B. Chiarelli, eds., R. L., and A. B. Chiarelli, eds., Evolutionary Biology 

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260 HELDIANA: ZOOLOGY 



Notes on Bolivian Mammals 

2. Taxonomy and Distribution of Rice Rats 

of the Subgenus Oligoryzomys 



Nancy Olds and Sydney Anderson 



ABSTRACTS 

There are at least three kinds of small, long-tailed rice rats of the subgenus Oligoryzomys 
(genus Oryzomys) in Bolivia. We use the names Oryzomys microtis (including O. fornesi), O. 
chacoensis, and O. longicaudatus for these taxa. The correctness of these names is less certain 
and resolution of the nomenclatorial questions awaits study of specimens from outside Bolivia. 
The ranges of O. chacoensis and O. microtis are at low elevations and probably overlap to some 
degree geographically. The range of O. longicaudatus is in the highlands. No sample from one 
locality includes specimens of more than one species; therefore, ecological and microgeographic 
differences at places where any two of the three species meet are unknown. Further study may 
reveal other and more cryptic species within Bolivia. No one measurement or other characteristic 
that we have studied will unequivocally distinguish all adult specimens of any one of the three 
kinds from Bolivia. Geographic variation probably occurs within as well as beyond Bolivia in 
at least two of the three species (the most uniform seems to be O. chacoensis), but more material 
is needed to describe such patterns. We refrain from using subspecies names in consideration 
of an ignorance of both geographic patterns of variation and the status of available names. The 
only name in the subgenus with a Bolivian type locality, O. chaparensis, is tentatively considered 
a synonym of O. microtis. 



Existen en Bolivia por lo menos tres clases de pequefias ratas arroceras de cola larga del 
subgenero Oligoryzomys (genero Oryzomys). Nosotros usamos los nombres de Oryzomys mi- 
crotis (incluyendo O. fornesi), O. chacoensis, y O. longicaudatus para estos taxa. La exactitud 
de estos nombres es menos cierta y la resolucion de problemas de nomenclature necesita estudio 
de esp>ecimenes de fuera de Bolivia. Los rangos de distribucion de O. chacoensis y O. microtis 
estan en bajas elevaciones y probablemente se superponen geograficamente en algun grado. 
Ninguna muestra de una localidad contiene especimenes de mas de una especie. Diferencias 
ecologicas y microgeograficas son desconocidas en lugares donde cualquiera de las dos o tres 
especies se encuentran. Ulterior estudio puede revelar otras y mas cripticas especies en el interior 
de Bolivia. Ninguna medida u otra caracteristica que nosotros hemos estudiado separara cla- 
ramente todos los especimenes adultos de Bolivia. Variacion geografica ocurre probablemente 
dentro asi como fuera de Bolivia en al menos dos de las tres especies (la mas uniforme parece 
ser O. chacoensis), pero mas material es necesario para describir tales patrones de variacion. 
Nos abstenemos de usar nombres subespecificos en consideracion de la ignorancia de los 
patrones de variacion y el estado de los nombres disponibles. El unico nombre en el subgenero 
con una localidad tipica en Bolivia, O. chaparensis, es tentativamente considerado como O. 
microtis. 



From the Department of Mammalogy, American Mu- 
seum of Natural History, New York, NY 10024. 

OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 261 



Existem no minimo tres tipos de pequenos ratos-de-arroz, de cauda longa, pertencentes ao 
subgenero Oligoryzomys (genero Oryzomys) na Bolivia. A estes taxa damos os nomes de Ory- 
zomys microtis (incluindo O. fornesi), O. chacoensis, e O. longicaudatus. A precisao destes 
nomes e incerta, e resolu^oes da nomenclatura aguardam estudos de especimes nao bolivianos. 
O. chacoensis e O. microtis ocorrem em eleva?6es baixas, e as duas esjiecies provavelmenle 
coocorrem em algumas areas. Oryzomys longicaudatus ocorre em areas montanhosas. Amostras 
de um so local nao incluem mais do que uma esF)ecie, e nao se conhecem diferen9as ecologicas 
ou microgeograficas em areas onde duas ou mais especies possam coocorrer. Com futuros 
estudos, novas especies, mais ocultas, poderao ser encontradas na Bolivia. Nenhum unico 
carater que estudamos pode inequivocamente separar especimes adultos dos tres tipos de 
Oligoryzomys na Bolivia. Varia96es geograficas provavelmente ocorrem na Bolivia, bem como 
em outras areas, mas mais material e necessario para poder documentar quaisquer padroes 
geograficos (O. chacoensis parece ser a esp>ecie mais uniforme em aparencia). Dada a falta de 
conhecimento, ambos dos padroes geograficos, e da disponibilidade dos nomes, nao usamos 
nomes de subespecies. O unico subgenero com localidade de tipo na Bolivia, O. chaparensis, 
e aqui lentativamente considerado como sinonimo de O. microtis. 



Introduction 

Bolivian landscapes range from less than 300 m 
to more than 4000 m, and habitats range from the 
humid lowland Amazonian tropical forests and 
subtropical savannahs to the high barren plains 
and snow-capped peaks of the Andean altiplano 
(fig. 1 ). Habitats may change abruptly, often within 
only a few kilometers. The mammals of Bolivia 
are also diverse, and provide excellent opportu- 
nities for ecological and taxonomic studies of broad 
scope. However, the animals are poorly known 
(Mares & Genoways, 1982); before satisfactory 
general conclusions can be reached, the Bolivian 
species need to be clearly delimited, both mor- 
phologically and geographically. 

Mice of the genus Oryzomys occur throughout 
South America and are important members of 
small mammal communities (Myers & Carleton, 
1981; Mares et al., 1981; Alho, 1982; O'Connell, 
1982;Streilein, 1982a-c; Viega-Borgeaud, 1982). 
We examined critically one subgenus of Oryzo- 
mys, Oligoryzomys, in Bolivia to determine how 
many species are present and where they occur. 
For a general description of the subgenus, see Myers 
and Carleton (1981, pp. 9-12). 

The subgenus Oligoryzomys needs revision. 
There are few published studies of the more than 
45 named forms (Tate, 1932; Ellerman, 1941; Ca- 
brera, 1961; Myers & Carleton, 1981; Honacki et 
al., 1982). Myers and Carleton (1981) studied 
Oligoryzomys from Paraguay, where they recog- 
nized three species: Oryzomys nigripes. O. cha- 



coensis, and O. fornesi. They also clarified no- 
menclatorial questions relating to the 'name 
Oryzomys nigripes. Since Paraguay borders Boliv- 
ia, this recent study was used as our starting point. 
We used the same measurements and comparable 
analyses. We assumed that O. chacoensis and O. 
fornesi occurred also in southeastern Bolivia, near 
the Paraguayan border. If O. nigripes occurs in 
Bolivia, the most probable place for it is in eastern 
Santa Cruz, from which no specimens are now 
available. 

At least six names have been used in the liter- 
ature or in museum collections for Bolivian sjdcc- 
imens of the subgenus Oligoryzomys: Oryzomys 
longicaudatus and O. stolzmanni (of the high- 
lands); O. chaparensis (from the lowlands of Co- 
chabamba); O. nigripes (used for all forms); O. 
delicatus (used for a few specimens at middle el- 
evations); and 0./7ave5ce/ts (lowland). These names 
have been used with considerable uncertainty in 
the past (see summaries in Tate, 1932, and espe- 
cially Myers & Carleton, 1981). One cause of this 
problem is that the original descriptions are vague 
or apply equally well to more than one species of 
Oligoryzomys. 

The taxonomic confusion can be resolved by 
detailed study of adequate numbers of museum 
specimens, and the nomenclatorial confusion re- 
solved by comparisons with type specimens. Re- 
newed interest in South American mammals has 
resulted in more specimens, which will help in 
these tasks. 



262 



FIELDIANA: ZCX)LOGY 



Brasi 




Chile 



Argentina 



V _ 



- II 



Fig. 1 . Map of Bolivia showing the departments and the 500- and 3000-m contour lines. Be, Beni; Ch, Chuquisaca; 
Co, Cochabamba; LP, La Paz; Or, Onaro; Pa, Pando; Po, Potosi; SC, Santa Cniz; Ta, Tarija. 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



263 



Methods 

We examined specimens in the collections of 
the American Museum of Natural History 
(AMNH); Academy of Natural Sciences in Phila- 
delphia (ANSP); California Academy of Sciences 
(CAL): Field Museum of Natural History (FMNH); 
Museum of Vertebrate Zoology, University of Cal- 
ifornia (MVZ); Museum of Zoology, University of 
Michigan (UMMZ); and United States National 
Museum of Natural History (USNM). 

Measurements were obtained as follows: exter- 
nal measurements are from the original labels or 
field notes, when available, or remeasured (fluid- 
preserved specimens only). The following cranial 
measurements were taken to the nearest 0.0 1 mm 
on a craniometer (see Anderson, 1 968) following 
Myers and Carleton (1981) and Musser (1979): 

ZN = depth of zygomatic notch 
LR = length of rostrum from tip of nasals to 
posterior edge of zygomatic notch 
GLS = greatest length of skull 
ZB = zygomatic breadth 
BB = breadth of braincase 
BIC = breadth of interorbital constriction 
LIF = length of incisive foramen 
LPB = length of palatal bridge 
LMl, LM2, LM3 = crown lengths of upF>er mo- 
lars 
WMl, WM2, WM3 = crown widths of upjjer 
molars 
MM = greatest breadth across molars (labial 
edges) 
LMX = crown length of upp>er toothrow 
LNP = length of nasal projection 
LD = length of diastema 
LB = length of bulla 
HB = height of bulla 
TL = total length, including tail 

T = tail length 
HP = length of hind foot, including claw 
E = length of ear, from notch 

Specimens were sorted by age, using dental cri- 
teria as outlined here (following Myers & Carleton, 
1981): 

Age Class I: M' not erupted or newly erupted, 

M- unworn. 
Age Class II: M^ slightly to moderately worn, 

but not flat; M- slightly worn; enamel island 

formed by the isolation of the internal part of 

the mesoflexus of M-. 



Age Class III: M ' flat or slightly concave; enam- 
el island of above well isolated; M' and M- 
substantially worn. 

Age Class IV: M^ concave; enamel island oblit- 
erated; teeth well worn, but main cusps still 
discernible. 

Age Class V: M ' and M- flat or concave; folding 
pattern obliterated. 

Individuals in age class I were examined, mea- 
sured, and included in mapping geographic ranges, 
but were excluded from statistical treatment of 
character variation. 

Sp)ecimens examined are listed by locality and 
museum catalogue number in the Appendix. Lo- 
calities are plotted in Figure 2. Statistical analyses 
were done using the computer facilities of the City 
University of New York (CUNY) and programs 
from SAS (Statistical Analysis System) Institute, 
Inc. (1982). 



Taxonomy 

Our knowledge of geographic variation and the 
status of some of the names is sketchy at best, so 
we do not use subspecific names. The name Ory- 
zomys (Oligoryzomys) longicaudatus stolzmanni 
was first used by Hershkovitz (1940, p. 81), by 
inference for Bolivian populations, but we are not 
certain that O. stolzmanni (Thomas, 1894; type 
locality Huambo, 3700 ft, department of Ama- 
zonas, Peru) and O. longicaudatus (type locality 
restricted to Valparaiso, Chile) are conspecific or 
that the Bolivian specimens are conspecific with 
either. The resolution of these problems awaits 
further study beyond Bolivia. 

We noticed no diflerence between Bolivian spec- 
imens from Beni and Brazilian SF)ecimens from 
the vicinity of the typ>e locality of O. microtis (Low- 
er Rio Solimoes, 50 mi above mouth), and thus 
the subspecific name Oryzomys microtis microtis 
might be applied to Bolivian specimens. However, 
we need to know more about geographic variation 
in the species (see Remarks under O. microtis). 
Some other names that may refer to consjsecific 
populations and thus be relevant as possible sub- 
specific epithets are as follows: 

1. Oryzomys destructor from lowland eastern 
Peru has been assigned to longicaudatus but may 
prove to be consp)ecific with microtis; if so, de- 
structor is the senior synonym. If destructor and 
microtis are conspecific, the northern Bohvian mice 



264 



HELDIANA: ZOOLOGY 




Fig. 2. Distribution of Bolivian Oligoryzomys: • = Oryzomys microtis; O = O. longicaudatus; X = O. chacoensis. 

might be known as Oryzomys destructor destruc- Bolivia on geographic and ecological grounds, al- 

tor, or perhaps O. destructor microtis. though no specimens are presently available. Myers 

2. Oryzomys fornesi from northern Argentina and Carleton (1981) referred specimens from San 

might also be expected to occur in southeastern Joaquin, Beni, to O. fornesi. 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



265 



3. Oryzomys mattogrossae from Brazil just to 
the east of Bolivia is conspecific with O. microtis 
and has its type locality nearest the department of 
Beni, where most of the Bolivian specimens have 
been taken; thus, for geographic reasons, it must 
be considered in any future subspecific determi- 
nations. The uncertain specific status of utiariten- 
sis from the same type locality as mattogrossae is 
noted below. If it is conspecific with mattogrossae, 
utiaritensis should be regarded as a synonym 
thereof at the subspecies level also. 

We concur with Myers and Carleton (1981) in 
the recognition of two species groups of Oligory- 
zomys. These are the smaller-bodied, small-toothed 
Oryzomys microtis. O.Jlavescens, and O. delicatus, 
and the larger-bodied, larger-toothed O. chacoen- 
sis, O. longicaudatus, and O. nigripes. 

In the Species Accounts section, synonymies 
cover only Bolivian records. Other relevant names 
are discussed elsewhere in the text. 



Results of Statistical Analyses 
Sex and Age Variation 

We examined sex and age variation in Oryzomys 
microtis. O. longicaudatus. O. chacoensis. and O. 
flavescens (from Uruguay). Males are slightly larg- 
er in general; the average size differences, consid- 
ering all characters for each species, are 2.7%, 1 .6%, 
2.5%, and 2. 1%, respectively. We examined sexual 
dimorphism in all measurements for each species 
by one-way and two-way analyses of variance of 
sex and of sex and age for each species. Results of 
two-way analyses of variance of sex and age (using 
all specimens of tooth wear class II or greater) on 
specimens of O. microtis from the department of 
Beni, Bolivia, are presented in Table 1 . Our results 
are roughly comparable to those of Myers and 
Carleton (1981). We did not separate sexes in fur- 
ther statistical analyses, although we watched for 
unbalanced sex ratios in samples when interpret- 
ing results. 

Age variation is more difficult to assess, as no 
detailed study has been published. Mice of this 
subgenus probably continue to grow for most of 
their lives (see Myers &. Carleton, 1981), although 
the rate slows with age. We performed least-squares 
regression analyses (General Linear Models pro- 
cedures of SAS) on five variables: greatest length 
of skull (GLS), total length (TL), zygomatic breadth 



(ZB), length of the first upper molar (LMl), and 
length of the hind foot (HF), by species, to plot i 
graphically the relationship between relative age I 
and size. Based on study of these graphs, we de- 
cided to include age classes II-V in further anal- 
yses, as did Myers and Carleton (1981). Most spec- I 
imens were juveniles (age class I) or young adults 
(age classes II-III). Few specimens of age classes 
IV-V were present among the species we exam- 
ined. See Tables 2 and 3 for the mean adult ages 
of the specimens studied. 



Principal Components 

We analyzed principal components (with the 
Princomp procedure of SAS on a correlation ma- 
trix) using measurements taken on individuals of 
all sp)ecies (the approximate numbers of specimens 
are in tables 2 and 3, some specimens were ex- 
cluded because of missing measurements). When 
plotted, the first principal component tends to sep- 
arate the smaller species {O. microtis and O. fla- 
vescens) from the larger species {O. chacoensis and 
O. longicaudatus). The percentage of the totJil vari- 
ance accounted for by the first three components 
is 68.7%. 



Discriminant Analysis 

We performed several different discriminant 
analyses (using the SAS programs Discrim, Step- 
disc, and Candisc). A stepwise discriminant anal- 
ysis chose the following characters (in order of 
selection): tail length, breadth of interorbital con- 
striction, diastema length, length of incisive fo- 
ramina, rostral length, bullar length, length of 
hind foot, molar breadth, length of nasal projec- 
tion, length of zygomatic notch, total length, zy- 
gomatic breadth, length of palatal bridge, and length 
of maxillary toothrow. 

Using the SAS program Discrim, we were able 
to test the posterior probability of group mem- 
bership. In all analyses, we used as "known" groups 
samples of the species Oryzomys microtis from 
Beni, Bolivia; O. flavescens from Uruguay; O. lon- 
gicaudatus from the department of La Paz. Boliv- 
ia; and O. chacoensis from Paraguay. Plots of the 
individual mice on the first two canonical corre- 
lates showed little overlap among the four species. 
We assumed that the following "unknowns" be- 
longed to one of the four "known" groups. We 
submitted as "unknowns" the holotypes of O. mi- 



266 



HELDIANA: ZOOLOGY 



crotis, O. mattogrossae, and O. chaparensis; the 
paratype of O. chaparensis; and two specimens 
from Beni assigned by Myers and Carleton (1981) 
to O. chacoensis (see Remarks under O. microtis). 
The holotypes of O. microtis and O. mattogrossae 
were assigned to the O. microtis species sample, 
with a posterior probability of more than 95%. 
The holotype of O. chaparensis was also assigned 
to O. microtis (P = 0.987). The paratype of O. 
chaparensis was assigned to O. microtis (P = 0.985). 
Several other specimens that we wanted to test, 
including the type of O. delicatus, had missing 
values and could not be used in the analysis. We 
recognize that the validity of the taxonomic con- 
clusions based on these analyses depends on the 
correctness of the initial assumption, and that it 
needs further testing. Not all measurements were 
available for all skulls, but in each comparison 
the largest possible subset was used. 



Species Accounts 

Subgenus Oligoryzomys Bangs, 1 900 

Diagnosis— Within the genus Oryzomys, the 
subgenus Oligoryzomys is distinguished by small 
size and delicate structure throughout, tail rela- 
tively long, hind foot long and slender; skull small, 
delicate, interorbital region narrow, outer edges of 
frontals squarish but unbeaded, braincase smooth 
and unridged, zygomatic plate narrow and with 
slight forward projection, molar teeth and incisors 
small and delicate but with cusp pattern like other 
Oryzomys (description adapted from Bangs, 1 900). 
Oryzomys of the subgenus Microryzomys are also 
small and delicate, but they differ from Oligory- 
zomys in having a more slender rostrum, shallow- 
er zygomatic notch, shorter and more rounded 
braincase, sphenofrontal foramen and squamo- 
soalisphenoid groove present, and karyotype with 
a low FN/2n ratio (Myers & Carleton, 1981, p. 
12). 



Oryzomys chacoensis Myers and Carleton, 1981 

Oryzomys chacoensis Myers and Carleton, 1 98 1 , p. 19 
(typ)e locality "419 km by road NW Villa Hayes 
[alongside the Trans Chaco Highway], Dept. Bo- 
queron, Paraguay"). 

Diagnosis— C>ryzc>my5 chacoensis Myers and 
Carleton (1981, p. 20) was diagnosed as "A me- 



Table 1 . Nongeographic variation in Oryzomys mi- 
crotis. Results of analysis of variance of Bolivian O. mi- 
crotis. 



Char- 


Error 






F (inter- 


acter 


d.f. 


F(sex) 


F (age) 


action) 


ZN 


98 


3.75 


0.44 


0.52 


LR 


89 


26.31*** 


1.87 


1.35 


GLS 


81 


22.69*** 


1.77 


0.75 


ZB 


86 


13.65*** 


1.19 


1.33 


BB 


92 


6.57* 


0.41 


2.55* 


BIC 


96 


0.74 


1.79 


1.22 


LIF 


95 


4.64* 


1.71 


0.51 


LPB 


93 


4.32* 


0.81 


1.01 


LMl 


98 


0.08 


2.38* 


0.76 


LM2 


99 


1.38 


5.37*** 


5.57*** 


LM3 


95 


4.23* 


1.93* 


1.82 


WMl 


98 


0.51 


1.38 


1.68 


WM2 


99 


0.82 


1.43 


1.06 


WM3 


95 


0.17 


1.90* 


1.24 


MM 


95 


3.07 


3.08** 


1.81 


LMX 


94 


1.53 


2.38* 


1.36 


LNP 


87 


15.05*** 


1.26 


1.73 


LD 


96 


10.87** 


1.23 


1.58 


LB 


92 


5.45* 


0.48 


0.71 


HB 


91 


1.77 


0.41 


0.83 


TL 


91 


5.36* 


1.38 


1.21 


T 


91 


0.02 


1.65 


2.34* 


HF 


92 


6.41* 


0.95 


0.67 


E 


48 


0.02 


1.53 


0.84 



* P < 0.05; ** P < 0.01; *** P < 0.001. 

dium-sized species of the subgenus Oligoryzomys 
unique in its whitish underside with hair white to 
the base on the chin and throat, relatively long 
ears having hairs on inner surface with unusually 
short or absent dark basal bands, small but dis- 
tinctive tufts of orangish hairs anterior to the ears, 
and karyotype with 2n = 58, FN = 74." 

Distribution in Bolivia— We examined spec- 
imens (see Appendix) from the departments of 
Cochabamba, Santa Cruz, and Tarija (fig. 2). Myers 
and Carleton (1981) reported two specimens from 
Beni. These specimens have been restudied and 
are here reassigned to Oryzomys microtis (see Re- 
marks under that species). Habitat of O. chacoen- 
sis is grassland and thomscrub. The highest known 
elevation is 640 m (Rio Lipeo). 

General Description and Comparisons— 
Cranial and external measurements are listed in 
Table 2. The dorsal pelage is rufous and heavily 
lined with black hairs. The venter is white and 
sharply distinguished from sides, which are clearer 
than dorsum. Frequently a thin orange line sep- 
arates the sides from the belly. The cheeks are 
slightly paler than the rest of the face. The tail is 
dark gray, weakly bicolored, and long relative to 
the body. The hind feet are whitish above. Juve- 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



161 



Table 2. Mean, standard deviation, and range (in mm) for Oryzomys chacoensis and O. longicaudatus. 







O. chacoensis 




O. longicaudatus 




Bolivia 


Paraguay 


Argentina 


Bolivia 


Character 


(N = 15) 


(N = 16) 


(N = 5) 


(N = 34) 


ZN 


1.15 ± 0.28 


1.30 ± 0.20 


1.15 ± 0.12 


1.02 ± 0.20 




(0.88-1.59) 


(0.90-1.61) 


(0.94-1.24) 


(0.57-1.36) 


LR 


7.27 ± 0.45 


7.39 ± 0.57 


7.67 ± 0.34 


7.14 ± 0.72 




(6.62-7.94) 


(6.57-8.86) 


(7.25-8.07) 


(4.49-8.15) 


GLS 


24.90 ± 0.82 


24.52 ± 1.09 


26.12 ± 0.32 


25.01 ± 1.39 




(23.80-26.69) 


(22.52-27.12) 


(25.79-26.43) 


(21.06-27.29) 


ZB 


13.10 ± 0.59 


12.76 ± 0.62 


13.64 ± 0.09 


13.17 ± 0.65 




(11.94-13.98) 


(11.91-14.27) 


(13.54-13.71) 


(11.55-13.98) 


BB 


11.33 ± 0.41 


11.16-0.34 


11.39 ± 0.06 


11.25 ± 0.37 . 




(10.42-11.87) 


(10.59-11.59) 


(11.35-11.45) 


(10.58-11.96) 


BIC 


3.89 ± 0.20 


3.76 ± 0.10 


3.86 ±0.13 


3.63 ± 0.22 




(3.52-4.27) 


(3.64-3.95) 


(3.76-4.04) 


(3.35-4.17) 


LIF 


4.62 ± 0.26 


4.71 ± 0.39 


5.07 ± 0.29 


4.68 ± 0.27 




(4.25-5.20) 


(4.17-5.56) 


(4.64-5.31) 


(4.23-5.26) 


LPS 


4.14 ± 0.22 


3.98 ± 0.26 


4.22 ± 0.31 


3.97 ± 0.21 




(3.65-4.54) 


(3.50-4.65) 


(3.75-4.52) 


(3.49-4.41) 


LMl 


1.68 ± 0.10 


1.63 ± 0.07 


1.74 ± 0.5 


1.59 ± 0.09 




(1.47-1.84) 


(1.54-1.79) 


(1.70-1.83) 


(1.42-1.81) 


LM2 


1.09 ± 0.06 


1.10 ± 0.07 


1.14 ± 0.07 


1.05 ± 0.04 




(0.96-1.18) 


(1.00-1.28) 


(1.08-1.26) 


(0.97-1.15) 


LM3 


0.84 ± 0.05 


0.80 ± 0.04 


0.84 ± 0.04 


0.81 ± 0.05 




(0.74-0.92) 


(0.73-0.88) 


(0.78-0.88) 


(0.67-0.92) 


WMl 


1.07 ± 0.07 


1.03 ± 0.03 


1.11 ± 0.05 


1.04 ± 0.06 




(0.93-1.24) 


(0.96-1.08) 


(1.05-1.17) 


(0.91-1.15) 


WM2 


1.02 ± 0.07 


1.00 ± 0.04 


1.04 ± 0.06 


0.99 ± 0.06 




(0.86-1.14) 


(0.93-1.06) 


(0.97-1.11) 


(0.87-1.09) 


WM3 


0.87 ± 0.05 


0.85 ± 0.03 


0.90 ± 0.05 


0.83 ± 0.06 




(0.78-0.97) 


(0.78-0.92) 


(0.84-0.94) 


(0.67-0.95) 


MM 


4.66 ± 0.20 


4.57 ± 0.17 


4.70 ± 0.13 


4.61 ± 0.27 




(4.32-4.99) 


(4.34-5.09) 


(4.58-4.87) 


(4.03-5.08) 


LMX 


3.60 ±0.12 


3.50 ± 0.13 


3.71 ± 0.15 


3.46 ± 0.14 




(3.29-3.78) 


(3.23-3.68) 


(3.61-3.98) 


(3.20-3.77) 


LNP 


1.43 ± 0.24 


1.59 ± 0.17 


1.49 ± 0.17 


1.40 ± 0.25 




(1.04-1.77) 


(1.35-2.07) 


~ (1.29-1.64) 


(0.99-2.04) 


LD 


5.66 ± 0.34 


5.63 ± 0.44 


5.89 ± 0.24 


6.04 ± 0.47 




(5.17-6.33) 


(5.02-6.44) 


(5.63-6.18) 


(4.96-7.07) 


LB 


3.54 ± 0.14 


3.60 ± 0.14 


3.75 ± 0.03 


3.47 ± 0.20 




(3.28-3.81) 


(3.28-3.82) 


(3.72-3.78) 


(3.07-3.84) 


HB 


2.75 ± 0.18 


2.73 ± 0.15 


2.72 ± 0.06 


2.76 ± 0.32 




(2.43-3.04) 


(2.40-2.96) 


(2.66-2.76) 


(2.23-3.32) 


TL 


227.93 ± 12.45 


219.50 ± 12.29 


229.00 ± 8.34 


217.68 ± 16.88 




(204-252) 


(194-241) 


(215-237) 


(172-252) 


T 


134.27 ± 10.26 


126.06 ± 8.46 


138.00 ± 5.70 


124.15 ± 12.18 




(112-150) 


(112-139) 


(130-145) 


(87-145) 


HF 


25.33 ± 1.33 


24.38 ± 1.54 


23.40 ± 2.30 


25.13 ± 1.20 




(23-28) 


(23-29) 


(21-26) 


(22-28) 


E 


16.70 ± 1.33 


15.07 ± 1.69 


18.40 ± 1.14 


15.03 ± 1.72 




(14-18) 


(12-17) 


(17-20) 


(12-18) 


AGE 


2.65 ± 0.82 


2.31 ± 0.68 


2.55 ± 0.87 


3.04 ± 0.88 




(2-4) 


(2-4) 


(2-4) 


(2-5) 



268 



HELDIANA: ZOOLOGY 



niles are grayer, as are all young Oligoryzomys in 
Bolivia, but this is especially noticeable on the 
venter, which has mixed gray and white hairs. In- 
cisive foramina extend posteriorly to the anterior 
edge of M' or slightly beyond. Alisphenoid strut 
(a strut of the alisphenoid bone that covers the 
lateral part of the alisphenoid canal, see Musser, 
1 982, p. 29) is generally absent (table 4). The sides 
of the interorbital constriction are divergent pos- 
teriorly (less parallel-sided than in O. longicau- 
datus) (fig. 3). The following quotes are from Myers 
and Carleton (1981, first and second quotes, p. 2 1 , 
third quote, p. 24): 

Oryzomys chacoensis differs from O. cha- 
parensis Osgood (1916) primarily in color 
pattern: the type of chaparensis is much 
darker and less hispid dorsally, yellowish 
ventrally, lacks the orange tufts anterior to 
the ears, and has a grayish throat. The distal 
portions of the nasals of the holotype flare 
laterally to an extent not seen in chacoensis. 



amined the two darker ones. When skulls are com- 
pared, the two darker mice are among the older 
and larger individuals in the series. They seem to 
have relatively broader braincases than most, but 
there is no character shared by these two that is 
not also seen in one or more of the others. In a 
discriminant analysis (SAS program Discrim) the 
posterior probability of membership (in the four 
reference species) allies these two mice with O. 
chacoensis (ansp 18187, P = 0.997, and ansp 
18188, P = 0.954 with chacoensis). We have as- 
signed these mice to O. chacoensis. Since Rio Li- 
pto is in the area where these two species meet, 
further study there should reveal whether sym- 
patry exists and if species differences remain dis- 
tinct. 

Specimens from Argentina (listed in the Ap- 
pendix) are similar in coloration to both Para- 
guayan and Bolivian specimens, but are woolier. 
This sample extends the known range of O. cha- 
coensis into the department of Jujuy, northwestern 
Argentina. 



Oryzomys chacoensis can be distinguished 
from fornesi, with which it occurs sympat- 
rically, by its larger size (maxillary toothrow 
usually > 3.3mm, ears usually > 15mm), 
characteristic karyotype, lack of preputial 
glands, and in most specimens by its lack of 
buff" on the belly. 

. . . the hind feet of chacoensis are relatively 
short compared to those of the more terres- 
trial fornesi. 

In reference to the last point, however, our cal- 
culations of length of hind foot relative to length 
of head and body (using data from tables 2 and 3) 
are about 26% for both O. chacoensis and O. mi- 
crotis (including fornesi). Comparison of speci- 
mens also reveals no noticeable difference. 

Remarks— This species is clearly distinct from 
O. microtis (here including O. fornesi). In most 
morphometric characters, O. chacoensis grossly 
resembles O. longicaudatus from Bolivia and O. 
nigripes from Brazil. 

Our specimens from Santa Cruz show some 
variation in coloration: two specimens (amnh 
247772-247773) are pale. The specimens from 
Tarija resemble the Paraguayan samples more 
closely than do the specimens from Santa Cruz. 

A series of 13 sp)ecimens from Rio Lipeo in- 
cludes two with darker pelage that resemble O. 
longicaudatus. We measured these mice and ex- 



Oryzomys longicaudatus (Bennett, 1832) 

Mus longicaudatus Bennett, 1832, p. 2 (type locality 
"In trees in Chile," restricted to Valparaiso by Ca- 
brera, 1961, p. 391). 

Oryzomys longicaudatus: Thomas, 1898, p. 3 (Aguai- 
renda Mission, San Francisco, perhaps not O. lon- 
gicaudatus); Thomas, 1926, p. 194 (Tupiza). 

Oryzomys Stolzmanni: Thomas, 1902, p. 130 (Cha- 
ruplaya, Choro); Neveu-Lemaire and Grandidier, 
1 9 1 1 , p. 9 (Charuplaya, Choro). 

Oryzomys ^X>;flavescens groMTp: Thomas, 1925, p. 578 
(Carapari, perhaps not O. longicaudatus). 

Oryzomys stolzmanni stolzmanni: Sanborn, 1950, p. 
2 (Rio Aceramarca, Cocapunco, Nequejahuira, 
Okara, Pitiguaya, Pongo). 

Diagnosis— Bolivian specimens referred to this 
species differ from other Bolivian Oligoryzomys 
in that they are larger (especially in the size of the 
teeth) than O. microtis and lack the buffy wash on 
the venter; are grayer-bellied, darker, and less griz- 
zled dorsally than O. chacoensis; and occur at gen- 
erally higher elevations (at least in northern Bo- 
livia). 

Distribution in Bolivia— O. longicaudatus is 
found in the valleys and mountains of the Andes 
from at least middle elevations (1200 m, Entre 
Rios) up to 3720 m (Poopo). Specimens from near 
Camiri and Cuyambuyo in southern Bolivia are 
from elevations of 780 to 1000 m. Specimens (see 
Appendix) have been reported or examined from 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



269 




Fig. 3. Dorsal and ventral views of skull of (left), Oryzomys longicaudatus stolzmanni; (middle), O. chacoensis, 
and (right), O. microtis microtis. Specimens in the American Museum of Natural History. Scale at lower right represents 
10 mm. 



the departments of Chuquisaca, Cochabamba, La 
Paz, Oruro, Polosi, Santa Cruz, and Tarija (see 
map, fig. 2). 

General Description and Comparisons— 
Cranial and external measurements are presented 
in Table 2. The belly is gray, rarely washed with 
buff; upper parts are a dark brown lined with black 
hairs, and often the sides are more rufous. A thin 
orange lateral line may be present. Nose and face 



are dark with paler cheeks. Often there are pale 
spots just behind the ears. Hind feet are pale above. 
The tail is brown above, weakly to strongly bi- 
colored, and relatively long compared to that of 
any other Bolivian Oligoryzomys. Incisive foram- 
ina generally extend to the anterior edge of M' or 
slightly behind. Alisphenoid strut is generally ab- 
sent (table 4). The interorbital area is slightly more 
constricted on the average than that of O. cha- 



270 



HELDIANA: ZOOLOGY 



coensis, and has relatively parallel sides (see fig. 
3). 

Remarks— Oryzomys longicaudatus may be a 
composite of more than one species. Oryzomys 
destructor and O. stolzmanni were considered sub- 
species of 6>. longicaudatus by Cabrera ( 1 96 1 ) and 
full species by Soukup (1961). Gardner and Patton 
(1976) found four karyotypic variants in speci- 
mens they assigned to O. longicaudatus, and com- 
mented that these may represent four separate 
species. One of their karyotypic variants, no. 2, is 
represented by 1 7 specimens from Balta, depart- 
ment of Loreto (now Acayali), Peru, at an eleva- 
tion of about 300 m. Myers and Carleton (1981, 
p. 26) mentioned that the karyotype of these spec- 
imens as reported by Gardner and Patton (1976) 
agreed better with results they obtained for O.for- 
nesi (= O. microtis) than for either O. chacoensis 
or O. longicaudatus. We have examined these 
specimens and judge that they are clearly different 
from O. longicaudatus and belong with either O. 
microtis or O. destructor (Balta is near the type 
locality of O. destructor). The inclusion of karyo- 
type no. 3 under O. longicaudatus, represented by 
one female from Venezuela, also needs verifica- 
tion. 

We have five specimens from Caracato, de- 
partment of La Paz (amnh 248977-248981), that 
are quite different from other Bolivian Oryzomys 
longicaudatus. The pelage closely resembles that 
of O. nigripes from Paraguay in color and texture; 
it is paler and more obviously lined with black 
hairs dorsally than the pelage of typical O. lon- 
gicaudatus. They are also slightly larger bodied, 
but four of the five are old individuals (age classes 
IV-V) and this may account for the larger size. 
Because of the small sample size and the single 
locality, we chose not to distinguish this popula- 
tion taxonomically. 

It is possible that Andean populations may oc- 
cupy areas that are rather widely scattered and 
physically isolated in different valleys or side can- 
yons. If so, there may be considerable geographic 
variation. More specimens are needed to detect 
and describe such patterns. 

The sjjecimens from near Cuyambuyo are at 
comparatively low elevations (980 to 1000 m) for 
O. longicaudatus, although the species occurs at 
progressively lower elevations farther south (Os- 
good, 1916). The two localities near Cuyambuyo 
are only about 60 km from Rio Lipeo, where O. 
chacoensis occurs. The SAS Discrim program 
analysis allied the largest adult specimen from near 
Cuyambuyo (ummz 155891) with O. longicauda- 



tus (F = 0.956). There are problems in distinguish- 
ing O. longicaudatus and O. chacoensis solely by 
cranial morphology in all areas. 



Oryzomys microtis Allen, 1916 

Oryzomys (Oligoryzomys) microtis Allen, 1 9 1 6, p. 525 
(type locality Lower Rio Solimoes, 50 mi above 
mouth [80 km from its confluence with Rio Negro, 
Amazonas, Brazil]). 

Oryzomys chaparensis Osgood, 1 9 1 6, p. 205 (holotype 
and paratype, type locality Todos Santos, on Rio 
Chapare, Department of Cochabamba, Bolivia); 
Gyldenstolpe, 1932, p. 25 (Todos Santos); Myers 
and Carleton, 1981, p. 38 (Todos Santos); all in 
reference to the same specimens. 

Oryzomys fornesi Massoia, 1973, p. 22 (type locality 
Naineck, Dept. Rio Pilcomayo, province of For- 
mosa, Argentina); as used by Myers and Carleton, 
1981, p. 25 for five specimens fVom San Joaquin, 
Beni, Bolivia. 

Diagnosis— Allen described Oryzomys microtis 
as being readily distinguished from other Olig- 
oryzomys by its pale coloration, relatively small 
ears, and tail less than half total length. However, 
the tail of the holotype is more than half of the 
total length, a discrepancy noted by Goodwin 
(1953). Massoia (1973) diagnosed O. fornesi, here 
considered a junior synonym of O. microtis, as the 
smallest species of the subgenus in Argentina, hav- 
ing the ears covered with short ochraceous hairs, 
the pterygoids short (shorter than the molar se- 
ries), the interorbital constriction narrow, and the 
incisive foramina not extending to the molar se- 
ries. In Bolivia, it can be distinguished by its small 
body size, toothrow generally less than 3.3, gray- 
buffy color of the belly, and relatively short tail in 
proportion to body. 

Distribution in Bolivia— Specimens (see Ap- 
pendix) from the lowlands of Bolivia, in the de- 
partments of Beni, Cochabamba, Pando, La Paz, 
and Santa Cruz have been examined. Habitat in- 
cludes marshes and wet forests up to an elevation 
of 1 800 m (Guanay, see fig. 2). 

General Description and Comparisons — 
Smallest of the Bolivian Oligoryzomys, with small 
teeth, toothrow generally less than 3.3 (averaging 
3.10). Massoia ( 1 973) gave general measurements 
useful in distinguishing this species from other Ar- 
gentine species of Oryzomys (Oligoryzomys) as fol- 
lows: length of hind foot generally less than 24, 
length of ear generally less than 13, breadth of 
braincase less than 10.8, and length of incisive 
foramina generally less than 4.5. All of his mea- 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



271 



Table 3. Mean, standard deviation, and range (in mm) for Oryzomys microtis and O. flavescens. 







O. microtis 




O. flavescens 




Bolivia 


Brazil 


Peru 


Uruguay 


Character 


(N = 67) 


(N = 20) 


(N = 26) 


(N = 20) 


ZN 


1.01 ± 0.17 


0.98 ±0.18 


0.84 ±0.13 


1.07 ± 0.15 




(0.53-1.46) 


(0.78-1.48) 


(0.61-1.18) 


(0.79-1.32) 


LR 


6.82 ± 0.51 


6.82 ± 0.42 


6.60 ± 0.41 


6.75 ± 0.43 




(5.17-8.16) 


(6.16-7.46) 


(5.96-7.36) 


(5.89-7.39) 


GLS 


23.54 ± 1.06 


23.79 ± 0.88 


23.27 ± 0.82 


23.41 ± 0.94 




(21.24-25.78) 


(22.47-25.04) 


(21.75-24.53) 


(21.66-24.81) 


ZB 


12.28 ± 0.51 


12.40 ± 0.44 


11.86 ± 0.53 


12.05 ± 0.49 




(11.42-13.73) 


(11.59-13.32) 


(11.01-12.93) 


(11.25-12.76) 


BB 


10.78 ± 0.36 


10.71 ± 0.23 


10.47 ± 0.32 


10.70 ± 0.26 • 




(9.38-11.78) 


(10.35-11.06) 


(9.49-10.87) 


(10.29-11.22) 


BIC 


3.76 ± 0.17 


3.72 ± 0.14 


3.63 ±0.11 


3.35 ±0.17 




(3.38-4.19) 


(3.44-3.93) 


(3.45-3.94) 


(2.95-3.77) 


LIF 


3.96 ± 0.27 


4.01 ± 0.22 


4.00 ±0.18 


4.59 ± 0.35 




(3.38-4.53) 


(3.65-4.37) 


(3.73-4.33) 


(3.54-5.20) 


LPB 


4.04 ± 0.32 


4.06 ± 0.22 


3.98 ± 0.26 


3.77 ± 0.19 




(3.14-4.76) 


(3.64-4.38) 


(3.42-4.51) 


(3.51-4.06) 


LMl 


1.46 ± 0.10 


1.43 ± 0.07 


1.51 ± 0.06 


1.47 ± 0.06 




(1.22-1.64) 


(1.32-1.56) 


(1.39-1.66) 


(1.37-1.60) 


LM2 


0.95 ± 0.06 


0.95 ± 0.05 


0.93 ± 0.08 


1.01 ± 0.06 




(0.81-1.09) 


(0.84-1.06) 


(0.67-1.04) 


(0.83-1.09) 


LM3 


0.70 ± 0.07 


0.68 ± 0.05 


0.66 ± 0.05 


0.73 ± 0.04 




(0.54-0.85) 


(0.59-0.78) 


(0.55-0.77) 


(0.66-0.80) 


WMl 


0.94 ± 0.06 


0.93 ± 0.06 


0.99 ± 0.04 


0.97 ± 0.04 




(0.78-1.08) 


(0.80-1.04) 


(0.91-1.10) 


(0.93-1.07) 


WM2 


0.88 ± 0.06 


0.89 ± 0.06 


0.93 ± 0.04 


0.93 ± 0.05 




(0.70-1.01) 


(0.78-0.99) 


(0.82-1.00) 


(0.86-1.03) 


WM3 


0.73 ± 0.06 


0.74 ± 0.06 


0.80 ± 0.05 


0.78 ± 0.04 




(0.55-0.85) 


(0.58-0.85) 


(0.71-0.93) 


(0.70-0.85) 


MM 


4.33 ± 0.21 


4.39 ± 0.23 


4.33 ± 0.18 


4.26 ±0.15 




(3.83-4.93) 


(3.99-4.81) 


(4.07-4.70) 


(3.98-4.50) 


LMX 


3.10 ± 0.17 


3.07 ± 0.16 


3.14 ± 0.13 


3.22 ±0.12 




(2.64-3.43) 


(2.75-3.34) 


(2.93-3.39) 


(2.96-3.47) 


LNP 


1.32 ± 0.22 


1.25 ± 0.32 


1.20 ± 0.20 


1.33 ± 0.14 




(0.91-1.74) 


(0.75-1.70) 


■ (0.81-1.62) 


(1.16-1.68) 


LD 


5.75 ± 0.41 


5.78 ± 0.36 


5.32 ± 0.40 


5.35 ± 0.35 




(4.82-6.62) 


(5.13-6.51) 


(4.49-6.01) 


(4.67-5.90) 


LB 


3.29 ± 0.14 


3.18 ± 0.19 


3.01 ± 0.12 


3.30 ±0.15 




(2.94-3.57) 


(2.90-3.63) 


(2.75-3.27) 


(2.91-3.53) 


HB 


2.46 ± 0.20 


2.10 ± 0.23 


2.22 ±0.12 


2.64 ± 0.23 




(1.96-2.87) 


(1.72-2.45) 


(1.96-2.38) 


(2.02-2.94) 


TL 


185.88 ± 11.12 


185.00 ± 10.60 


179.77 ± 9.16 


198.56 ± 17.66 




(165-214) 


(164-202) 


(160-196) 


(160-223) 


T 


101.00 ± 6.67 


95.44 ± 7.88 


101.69 ± 8.13 


110.37 ± 11.05 




(87-116) 


(75-105) 


(77-115) 


(85-127) 


HF 


22.81 ± 1.21 


21.68 ± 1.34 


21.37 ± 1.60 


25.21 ± 1.85 




(19-25) 


(20-25) 


(18-25) 


(20-28) 


E 


14.49 ± 1.04 
(12-16) 






13.97 ± 1.05 
(12-16) 


AGE 


2.57 ± 0.73 


2.35 ± 0.59 


2.97 ± 0.88 


2.18 ± 0.35 




(2-5) 


(2-4) 


(2-5) 


(2-3) 



272 



FIELDIANA: ZOOLOGY 



surements fall within the extremes of our values 
for O. microtis, except for measurements of the 
toothrow (LMX), in which his are on the upper 
end of our range. Values of cranial and external 
measurements are listed in Table 3. The venter is 
white mixed with gray to gray-bufFy and clear huf- 
fy. The dorsum is rufous brown, lined with black 
hairs, and has paler sides. Often there is no clear 
demarcation between sides and belly. There are 
no white spots behind ears. Juveniles are similar 
to adults, but show a grayer belly. The tail is dark 
above and weakly bicolored. Occasionally orange- 
tipped hairs lie anterior to the ear. Hind feet are 
white above. Incisive foramina generally extend 
posteriorly to the anterior edge of M' and not 
beyond. In comparison with adults of O. cha- 
coensis and O. longicaudatus, O. microtis has a 
shorter skull, longer braincase, shorter incisive fo- 
ramina, a relatively broader interorbital constric- 
tion with divergent sides, and smaller, narrower 
teeth; also, an alisphenoid strut is more often pres- 
ent (table 4). 

Remarks— This species is similar to Oryzomys 
flavescens of Uruguay and Argentina (see Myers 
& Carleton, 1981). Specimens of both species were 
reported from the same locality, Capitan Solari, 
in the province of Chaco, northern Argentina, by 
Contreras and Berry (1983). Measurements of 6>. 
flavescens from Uruguay are listed in Table 3 (see 
also Langguth, 1 963). In comparison with O. mi- 
crotis, specimens of O. flavescens have slightly 
larger teeth, larger bullae, longer incisive forami- 
na, and are generally larger. Oryzomys flavescens 
needs to be more clearly defined. Massoia and 
Fomes ( 1 967) once synonymized O. flavescens with 
O. nigripes, but according to the later analyses of 
Massoia (1973) and Myers and Carleton (1981) 
these taxa are not consjiecific. Uruguayan speci- 
mens oi O. flavescens are clearly different from O. 
nigripes from eastern Paraguay. Future studies may 
reveal additional relationships, including the pos- 
sibility that O. microtis and O.fiilvescens of Cen- 
tral America and northern South America are con- 
specific (Handley, 1976, referred Venezuelan 
specimens to O. fiilvescens). 

The name Oryzomys microtis flyrnesi might be 
used to reflect the conspecific status oiflyrnesi and 
microtis. This subspecific name would apply to at 
least the five specimens identified by Massoia 
(1973), pending more detailed studies that would 
test whether geographic differences warrant con- 
tinued subspecific recognition. 

Until the relationships of the Andean ""longi- 
caudatus -group" are clarified, it is also possible 



Table 4. Presence or absence of alisphenoid strut in 
five species of Oryzomys (Oligoryzomys) expressed as a 
percentage; number of specimens in parentheses. 

Alisphenoid strut 



Species 


Present 


Absent 


O. chacoensis 


2.4(1) 


97.6(41) 


O. longicaudatus 


5.6 (2) 


94.4 (34) 


O. microtis 


35.8 (54) 


64.2 (97) 


O. flavescens 


3.2(1) 


96.8 (30) • 


O. nigripes 


8.3 (3) 


91.7(33) 



that one or more of the earlier names currently 
assigned to this group actually belong with Ory- 
zomys microtis, and have priority. For example: 
O. stolzmanni (named in 1894, type locality listed 
under synonymy) and O. destructor (named in 
1 844, type locality: "the house mouse of the 'Plan- 
tagen at the border of the forest," eastern Peru, 
above 6000 ft, according to Tate, 1932, p. 9; re- 
stricted by Cabrera, 1961, to the haciendas along 
the Rio Chinchao in the department of Huanuco, 
between 900 and 1000 m) have been associated 
with O. longicaudatus. These type localities are at 
low elevations, which suggests the possible affinity 
of these forms with microtis rather than with lon- 
gicaudatus. We have examined a series from low 
elevation in the department of Pasco, Peru, that 
is clearly O. microtis. This locality is near the 
type locality of destructor (Cole, 1984). If future 
study of the holotype of destructor proves it to be 
conspecific with O. microtis, the name of the species 
should be O. destructor instead of O. microtis. 

We have examined the type specimens of Ory- 
zomys microtis (amnh 37091, type locality Lower 
Rio Solimoes, 50 mi above mouth, Brazil), O. 
mattogrossae (amnh 37542, type locality Utiarity, 
Mato Grosso, Brazil), O. utiaritensis (amnh 37541, 
type locality Utiarity, Rio Papagaia, Mato Grosso, 
Brazil), and O. delicatus (amnh 7317/5925, type 
locality Trinidad, West Indies). We judge that the 
small lowland Bolivian specimens are O. microtis 
and also represent the same species as specimens 
referred by Myers and Carleton to O. fornesi (O. 
microtis has priority). The Bolivian specimens dif- 
fer slightly from Massoia's description of O. for- 
nesi in having a shorter average toothrow, other- 
wise in being slightly larger. If O. fornesi and O. 
microtis are conspecific, it is possible that this dif- 
ference in measurements reflects his small sample 
size (N = 5), or that there may be significant geo- 
graphic variation between Argentine and Bolivian 
populations. We further treat O. mattogrossae as 
a synonym of O. microtis (O. mattogrossae is not 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



111 



included in the synonymy above because the name 
has never been used in print for a Bolivian spec- 
imen). As mentioned by Myers and Carleton 
( 1 98 1 ), O. utiaritensis may belong with O. nigripes. 
More specimens from Mato Grosso are needed to 
establish the characteristics of the species there. 
Oryzomys microtis was considered a synonym of 
O. delicatus by Alho ( 1 982) and Pine ( 1 973); how- 
ever, because of the slightly damaged condition of 
the skin and skull of the holotype of O. delicatus 
and the remoteness of its type locality (on the is- 
land of Trinidad, Allen & Chapman, 1 897), we 
defer any decision on this question. 

We have reexamined the two specimens from 
Beni that Myers and Carleton (1981) referred to 
Oryzomys chacoensis (usnm 39 1 297, 46074 1 ) and 
have assigned them to O. microtis. These speci- 
mens are young adults (age class II), have external 
measurements that fit well with O. microtis and 
that are rather small for O. chacoensis (TL 200, 
200; T 111, 110; HF 19, 24, respectively), and 
have toothrows measuring 3.34 and 3.40, respec- 
tively. The toothrows fall on the upper end of the 
range for O. microtis and the lower end of the range 
for O. chacoensis (tables 2-3). We then compared 
their measurements through a discriminant anal- 
ysis with the samples ofO.flavescens, O. microtis, 
O. chacoensis. and O. longicaudatus. The posterior 
probability of membership for both was greatest 
for O. microtis (usnm 391297, P = 0.937; usnm 
460741,/' = 0.953). 

We refer the holotype (fmnh 21330) and the 
one paratype (amnh 40787) o^ Oryzomys chapa- 
rensis to Oryzomys microtis. The holotype is the 
largest and one of the oldest individuals we have 
identified as O. microtis. Its greatest skull length 
is 26.8, zygomatic breadth 13.4, breadth of brain- 
case 1 1.2, crown length of maxillary toothrow 3.5, 
tail length 111, and length of hind foot 26. The 
skull length, the toothrow, and hind foot length 
are larger than other Bolivian O. microtis (see table 
3). The pelage coloration of this specimen falls 
within the range of variation in O. microtis. For 
a comparison of pelage coloration between O. cha- 
parensis and O. chacoensis, see the account of O. 
chacoensis under General Description and Com- 
parisons. 

Geographic Variation— We were unable to 
detect significant differences in p>elage coloration 
between our samples from Brazil, Bolivia, and 
Peru. The Brazilian specimens are much like the 
Bolivian specimens. A / test showed only three 
variables that differ significantly: bullar height (/ = 
5.45, P < 0.001), tail length (/ = 2.61, P < 0.05), 



274 



and length of hind foot (/ = 3.24, P < 0.01); all 
have greater average values in Bolivian specimens, | 
but the ranges of values overlap. One possible con- 
founding factor is that the Brazilian sample is com- 
posed of mostly males (ca. 95%) and has a lower 
average adult age than the Bolivian or Peruvian ' 
samples (see tables 2-3). If the degree of sexual 
dimorphism in this species is considered (2.72%, 
see p. 266), the difference may actually be less than i 
noted in our tests. For these reasons the subspecies ! 
name Oryzomys microtis microtis might be ap- I 
propriate for specimens from northern Bolivia. 

More differences exist between Peruvian and 
Bolivian samples. The Bolivian specimens are 
generally larger: the zygomatic notch is deeper, 
diastema longer, braincase broader, zygomatic 
breadth greater, and interorbital constriction 
broader. However, Bolivian specimens have sig- 
nificantly narrower teeth. Brazilian specimens, as 
noted above, are similar to Bolivian specimens 
and differ from the Peruvian material about as 
much as Bolivian specimens do. The Peruvian mice 
used in this comparison come from a lowland site 
in Pasco, the department just south of Huanuco, 
which is near the type locality of Oryzomys de- 
structor. 

One skin (amnh 247776) from near Villa Tunari 
in Cochabamba is dorsally paler and ventrally 
whiter (not bufly) than any skin from Pando or 
Beni. A skin (amnh 246809) from near Buena Vis- 
ta in Santa Cruz is comparable to the darker spec- 
imens from Beni, but is ventrally less bufl[> . 



Concluding Remarks 

Given the considerable pelage differences noted 
in reference to the few available specimens from 
Cochabamba and Santa Cruz in the accounts of 
Oryzomys chacoensis and O. microtis, the prob- 
lems noted in reference to sF>ecimens from south- 
em Tarija in the accounts of O. chacoensis and O. 
longicaudatus, the difficulty of identification that 
results from overlapping ranges of measurements, 
the sketchy knowledge of geographic variation, and 
the small samples from most critical areas, the 
taxonomy of Bolivian Oligoryzomys will probably 
prove to be more complicated than that presented 
here. We hojie our summary will provide infor- 
mation useful in further taxonomic work on the 
subgenus. All of our data and detailed analyses are 
deposited in the Department of Mammalogy, 



FIELDIANA: ZOOLOGY 



American Museum of Natural History, and are 
available for the use of other interested workers. 
More than 400 additional specimens, many in- 
cluding karyological and biochemical prepara- 
tions, have been collected in 1984, 1985, and 1986 
since this paper was written. Study of these will 
certainly help to clarify remaining taxonomic 
problems. 



Acknowledgments 

This study was partially funded by a grant from 
the Undergraduate-Graduate Research Fund Pro- 
gram, which is supported by the Susan Greenwall 
Foundation and administered by the American 
Museum of Natural History. We thank Dr. Leslie 
F. Marcus for his statistical advice. Masaaki Yone- 
da kindly provided us with a list of SF>ecimens in 
the Museo Nacional de Historia Natural, La Paz, 
Bolivia (MNHN). We are grateful to the curators 
of the collections referred to here, especially Dr. 
Philip Myers, Dr. Guy G. Musser, and Dr. Michael 
D. Carleton, for their generous advice and assis- 
tance in various ways. We also thank Dr. Robert 
S. Voss, Dr. James L. Patton, and Dr. Robert M. 
Timm for reviewing the manuscript. 



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OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



275 



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malia, 46: 335-359. 



276 



HELDIANA: ZOOLOGY 



Appendix ^ 

Specimens Reported or Examined 

The latitude and longitude are abbreviated; 1657/6523 means 16*'57'S, 65''23'W, for example. Where 
references are given (to Thomas), the specimens have not been examined by us. As noted, there are a 
few other specimens that we have not seen. 



O. chacoensis 

Specimens: 88. 



Boquer6n 

4 1 9 km by road NW Villa Hayes, 1 amnh (248397). 



BOLIVIA 
Santa Cruz 

1623/6059, San Ignacio, province of Velasco, 10 
USNM (390120-390121, 390664-390666, 
391522-391526). 

1808/6312, 7 km E and 3 km N Ingeniero Mora, 
13 AMNH (247758-247761, 247765-247773). 



Tarua 

2056/6321, 2 km S and 10 km E of Tiquipa, 4 

amnh (246799-246801, 246822). 
21 19/6325, 8 km S and 10 km E of Villa Monies, 

8 AMNH (246802-246806, 246810, 146912, 

246914). 
2128/6317, 35 km by road SE Villa Montes, Ta- 

ringuiti, 5 ummz (155937-155938, 156332- 

156334). 
2241/6426, Rio Lipeo, 13 ansp (18176-18188). 



PARAGUAY 

Presidente Hayes 

Rio Pilcomayo, 15 mi W Rio Paraguay, 3 amnh 
(143892-143894). - - 

NuEVA AsuNa6N 

19 km by road WSW, km 588 Trans Chaco, 8 

amnh (249255-249262). 



Chaco 

50 km WNW Fortin Madrejon, 9 amnh (248398- 
248406). 



ARGENTINA 

JUJUY 

Yuto, 10 AMNH (167855, 179976, 179980, 182570- 
182571, 182738-182739, 183312, 185226- 
185227, 185269). 

Sta. Barbara, 4 amnh (185224-185225, 185228, 
186954). 



O. longicaudatus 

Specimens: 134. 

BOLIVIA 

Chuquisaca 

1927/6407, Tola Orko, Tomina Province, 6 usnm 
(271588-271590, 545226-545227, 545229). 

1929/6433, Horcus, 80 km SE Sucal, 3 mvz 
(134654-134656). 

1931/6409, Monte Canto, 1 usnm (271591). 

?, Chuyayacu, 1 fmnh (72889). 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



zm. 



CCXTHABAMBA 

1620/6645, Yungas, 1 cm (5276). 

1654/6642, El Choro, 1 amnh (1 19405), 12 fmnh 

(74902-74913), 1 bm (Thomas, 1902). 
1714/6541, Incachaca, 19 amnh (38525-38542, 

38550, not seen), 2 cm (5081-5082). 
1734/6621, Parotani, 2 amnh (38543-38544), 1 

fmnh (21668). 
1742/6509, 20 mi E Totora, 1 mvz (1 19916). 
1807/6509, Aiquile, 2 fmnh (50970-50971). 



22 1 2/6436, 8 km by road N of Cuyambuyo, 2 
UMMZ (156326-156327). 

2213/6436, 4 km by road N of Cuyambuyo, 10 
UMMZ (155889-155891, 156312-156313, 
156315-156317, 156319-156320). 



Department Uncertain 
?, Zapial, 1 usNM (270911). 



La Paz 

1530/6824, Cocapunco, 1 amnh (72644). 
1535/6843, Tacacoma, 1 amnh (91540). 
1535/6843, Tacacoma-Sorata, 2 amnh (91541- 

91542). 
1539/6824, Okara, 2 amnh (72704-72705). 
1543/6840, 10 km by road N of Sorata, 3 ummz 

(156301-156303). 
1547/6840, Sorala, at base of Ml. Sorata, 4 amnh 

(91536-91539). 
1600/6516, Charuplaya, 16 bm (Thomas, 1902). 
1618/6753, Rio Aceramarca, 2 amnh (72693- 

72694). 
1619/6752, Nequejahuira, 3 amnh (72722, 72724- 

72725). 
1620/6808, Mt. Chacaltaya, 1 ummz (1 1571 1). 
1620/6756, Pongo, 15 amnh (72702-72703, 

72706-72715, 72726, 81283, 241612). 
1621/6747, Pitiguaya, 4 amnh (72716, 72729- 

72731). 
1659/6749, Caracato, 5 amnh (248977-248981). 
1823/6659, Poopo, 1 mnlp (not seen). 

POTOSi 

1 844/6609, 3 km SE of Pocoata, 1 amnh (255946). 
2127/6543, Tupiza, 1 bm (Thomas, 1926). 



Santa Cruz 

1754/6429, 5 mi W Comarapa, 1 mvz (1 19917). 
2005/6334, nr. Camiri, 1 cal (13803). 



Tarua 

2131/6445, 10 mi NW Tarija, 1 mvz (1 19918). 
2 1 32/64 1 2, Entre Rios, 3 USNM (27 1 4 1 1-27 1 4 1 2, 
271432). 



O. microtis 

Specimens: 230. 

BOLIVIA 
Beni 

1049/6525, Guayaramerin, 2 amnh (210050- 

210051). 
1110/6522, 4 km below Santa Cruz, 1 "amnh 

(211727). 
1 142/65 1 6, 4 km S Santa Rosa, 4 amnh (2 1 1 729- 

211730, 211791-211792). 
1 200/6502, Rio Itenez, 20 km from mouth, 2 amnh 

(211756-211757). 
1200/6506, Puerto More, Rio Itenez, 3 amnh 

(211758-211760). 
1 2 1 3/65 1 3, Mamore River on bank opposite Cas- 

cajal, 16 amnh (211724-211725, 211754, not 

seen, 211762-211774). 
1226/6511, Mamore River, 2 amnh (211721, 

211761). 
1229/6417, Rio Itenez, opposite Costa Marques, 

Brazil, 11 amnh (210122, 210038-210045, 

210365-210366). 
1229/6418, Rio Itenez, 1.5 km below Costa Mar- 
ques, Brazil, 1 amnh (210364). 
1230/6415, Pampa de Meio, 2 amnh (210046- 

210047). 
1 230/64 1 8, Baures River mouth, 6 amnh (2 1 0028- 

210031, 210382, 210383, not seen). 
1240/6330, Curiche River mouth, 6 amnh 

(210032-210037). 
1241/6432, 15 km above Horquilla on Rio Ma- 

chupo, 1 amnh (210053). 
1244/6318, Versalles, 1 amnh (210052). 
1244/6428, Las Perias, 3 fmnh (1 17079-1 17081), 

1 USNM (460741). 



278 



FIELDL\NA: ZOOLOGY 



1300/6515, Mamore River, 1 amnh (21 1722). 
1304/6449, San Joaquin, 1 fmnh (1 17075), 6 usnm 

(364735, 364738, 391299, 460273, 460742- 

460743). 
1312/6410, Cachuelita, 1 usnm (460739). 
1 3 1 2/65 1 5, 8 km N Exaltacion, 6 amnh (2 1 1 775- 

211780). 
1313/6221, 20 km W Larangeira, Bahia de los 

Casara, 2 amnh (210048-210049). 
1313/6409, Boroica, 1 usnm (460740). 
1322/6520, Palacios, province of Yacuma, 1 usnm 

(461082). 
1324/6442, Chaco Lejo, 20 km SE San Ramon, 2 

usnm (391295-391296). 
1325/6435, Tacuaral, 1 usnm (391297). 
1343/6521, Puerto Caballo, 2 amnh (211785, 

214597). 
1420/6455, 10 km W San Pedro, on Mamore Riv- 
er, 5 AMNH (21 1786-21 1790). 
1428/6734, Rurrenabaque, 2 amnh (247774- 

247775). 
1434/6455, 23 km W San Javier, 1 amnh (214760). 
1437/6457, Ibare River mouth, 2 amnh (21 1783- 

211784). 
1446/6451, Ibare River, 24 km from mouth, 1 

amnh (21 1755, not seen). 
1454/6422, 6 km W of Casarabe, 7 amnh (255947- 

255953). 
1503/6658, 1 km E of La Embocada, 2 ummz 

(155940, 156292). 
1515/6415, El Triunfo, 1 usnm (391298). 
1550/6441, 5 km NW Grande River mouth, 2 

AMNH (21 1781-21 1782). 
?, Centenela, 1 fmnh (1 17074). 
?, "Beni" only, 1 amnh (232699). 



Pando 

1117/6855, Rio Nareuda, 11 amnh (248982- 
248990. 248993-248994). 



Santa Cruz 

1 703/6335, 7 km N Santa Rosa, 1 amnh (25460 1 ). 
1724/6344, 7 km N and 17 km W of Buena Vista, 
3 AMNH (246809, 246820, 246935). 



BRAZIL 

Amazonas 

Solimoes, 10 amnh (37088-37097, 37091 = ho- 

lotype of O. microtis). 
Guatsue, 1 amnh (37100). 
Lower Solimoes, 1 amnh (37157). 

ParA 

Capim, 150 mi SE Belem, 1 amnh (188964). 
Capim, Est. B.R. 14, km 97, 1 amnh (203400). 
Rio Xingu; Porto de Moz, 1 amnh (95983). 
Rio Xingu; Villarinho do Monte, 4 amnh (95984- 
95986, 95997). 



GoiAs: Rio Madeira 



COCHABAMBA 

1648/6508, Todos Santos, 1 amnh (paratype of 
O. chaparensis, 40787), 1 fmnh (holotype, 
21330). 

1657/6523, 2 km E of Villa Tunari, 4 amnh 
(247662-247664, 247776). 



Auara Igarape, 5 amnh (91874, 91876-91878, 

94245). 
Sta. Antonio du Uayara, 4 amnh (92258-92261). 
Rosarinho, Lago Miguel, 1 1 amnh (92705-92715). 
Rosarinho, Lago Sampaio, 2 amnh (9271 6-927 1 7). 



Mato Grosso 



La Paz 

1 5 1 5/68 1 0, Mapiri, 5 amnh (7272 1 , 730 1 1 , 72732, 

72697-72698). 
1528/6752, Guanay, 1 amnh (72701). 
1528/6818, Ticunhuaya, 1 amnh (72700). 
1540/6742, 4 km NW Alcoche, 2 ummz (126777, 

127167). 



Utiarity, Papagaio River, 3 amnh (37540-37542; 
37541 = type of O. utiaritensis, 37542 = type 
of 6). mattogrossae). 



? Amazonas 

S bank R. Amazon: Villa Bella Imperatriz, 2 amnh 
(91899-91900). 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 



279 



PARAGUAY 

Caaguazu 

24 km NNW Carayao, 1 ummz (133821). 

Presidente Hayes 

24 km NW Villa Hayes, 3 UMMZ (133830-1 33831, 
133833). 

PERU 

Ayacucho 

Hda. Luisiana on Rio Apurimac, 1 amnh (242484). 

Pasco 

Ca. 1 km N Puerto Bermudez, 38 amnh (24555 1- 
245588). 



Maldonado 

Barra del Arrojo Maldonado, 9 km ENE Punta 
del Este, 1 amnh (205994). 



Montevideo 

Rio Santa Lucia, 1 km SE Santiago Vasquez, 2 
AMNH (205995-205996). 



ROCHA 

22 km SE Lascano, 18 amnh (205997-206009, 
206011-206015). 



San Jose 

Dept. only, 3 amnh (232216-232218). 

Soriano 

3 km E Cardona, 1 amnh (206016). 



O. flavescens 

Specimens: 33. 



Treinta y Tres 

16 km SSW Boca del Rio Tacuari, 1 amnh 
(206017). 



URUGUAY 

Artigas 

6 km NNW Belen, 2 amnh (205986-205987). 

Cerro Largo 

Sierra de Vaz, Rio Tacuari, 20 km SE Melo, 1 
amnh (205988). 

Lavalleja 

12 km WSW Zapican, 4 amnh (205989-205905). 



O. nigripes 

Specimens: 31. 

BRAZIL 

GOIAS 

Anapolis, 7 amnh (134528-134530, 134532, 
134534, 134538, 134540). 



280 



FIELDIANA: ZOOLOGY 



MiNAS Gerais PARAGUAY 

(Serra do Caparao), Rio Caparao, 8 amnh (80369- Ainambay 

80370, 80372, 80375-80379). 

(Serra do Caparao), Boa Espera, 1 amnh (80391). 4 km by road SW Cerro Cora, 1 ummz (125523). 



Mato Grosso CaaguazO 

Maracaju, 9 amnh (134541-134542, 134544- N of Coronel Oviedo, 1 ummz (124212). 

134547, 134551, 134838, 134902). 24 km NNWCarayao, 3 ummz (133835-133837). 



SAo Paulo 

Piquete, 1 amnh (36496). 



OLDS & ANDERSON: RICE RATS OF SUBGENUS OLIGORYZOMYS 281 



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

Jose L. Yaftez, Juan C. Torres-M iira, Jaime R. Rau, and Luis C. Contreras 



ABSTRACTS 

New records of Euneomys from central and southern Chile are given. Specific assignment of 
this material is difficult, owing to the lack of good diagnostic characters, small number of 
specimens, and spotty distribution. Absence of clear-cut differences between Euneomys localities 
suggests that there is only a single species, E. chinchilloides. Subspecific assignments are deferred, 
because at present they can only be made geographically, and the gaps between localities seem 
to be more apparent than real. 

Se citan nuevos registros de Euneomys para Chile Central y Sur. La determinacion es- 
pecifica de este material es dificil debido a la carencia de buenos caracteres diagnosticos, al 
bajo niimero de especimenes y a la distribucion localizada con grandes hiatos intermedios. La 
ausencia de claras diferencias entre Euneomys de distintas localidades sugiere la existencia de 
tan solo una especie, E. chinchilloides. Las nominaciones subespecificas se posponen, ya que 
al moment© estas solo se pueden realizar basadas en las localidades geograficas y los hiatos 
parecen ser mas aparentes que reales. 

Sao citados novos registros de Euneomys para o Sul e Centro do Chile. Dada a falta de bons 
carateres diagnosticos, o baixo numero de especimes, e a distribui^ao localizada com hiatos 
intermediarios, e dificil a determina9ao esfiecifica deste material. A ausencia de caracteristicas 
distintas entre Euneomys de localidades diferentes sugere a existencia de uma especie apenas: 
E. chinchilloides. Nao foram designadas nomina96es subespecificas ja que, no momento, estas 
so poderiam ser realizadas na base das localidades geograficas, e os hiatos podem ser mais 
aparentes do que reais. 



Introduction 

The genus Euneomys is one of the least studied 
cricetid rodents in southern South America. These 
herbivorous, volelike rodents have a relatively 
large, heavy body and a short tail and ears (Os- 
good, 1943). Euneomys has close affinities with 
the phyllotine and sigmodontine groups (Hersh- 



From Museo Nacional de Historia Natural, Casilla 
787, Santiago, Chile (Yanez); Dei>artamento Biologia y 
Quimica, Facultad de Ciencias, Universidad de Talca, 
Casilla 747, Talca, Chile (Torres-Mura and Contreras); 
and Estacion Biologica de 'Doiiana' (Sevilla), Apartado 
1056, Sevilla 41013, Espana (Rau). 



kovitz, 1 962; Mann, 1978). The genus has a known 
distribution from 33°00'S to Cape Horn (fig. 1), 
mainly along and to the east of the Andes (Hersh- 
kovitz, 1962; and subsequent reports). 

In Chile Euneomys is known from Lago Fag- 
nano and Hermite Island (Tierra del Fuego), Punta 
Arenas (Magallanes), Torres del Paine and Laguna 
Lazo (Ultima Esperanza), Puerto Ibaiiez (General 
Carrera), Campo Bandera (Aysen), Pino Hachado 
(Malleco), and Lo Valdes and Farellones (Santia- 
go) (Osgood, 1943; Mann, 1944, 1978; Markham, 
1971; Greer, 1965; Reise & Venegas, 1974; Pine 
et al., 1979). In Argentina Euneomys is certainly 
known from Bahia del Buen Suceso (Tierra del 
Fuego), upper Rio Chico (Santa Cruz), Paso de 



YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS 



283 




Fig. 1. Collecting localities of Euneomys. 1, Farellones; 2, Lo Valdes; 3, Bartos del Flaco; 4, Pino Hachado; 5, 
Lastarria; 6, Puerto Ibariez; 7, Torres del Paine; 8, Rio Baguales and Cueva del Milodon; 9, Estrecho de Magallanes 
and Palli Aike; 10, Punta Arenas; 11, Lago Fagnano; 12, Hermite Island; 13, Grevy Island; 14, San Rafael; 15, Cueva 
Traful; 16, Cerro Leones; 17, Paso de los MoUes, Pilcaniyeu; 18, Upper Rio Chico, Santa Cruz; 19, Bahia del Buen 
Suceso. Open circles represent new localities. 



284 



FIELDIANA: ZOOLOGY 



Table 1. Mandibular measurements of Euneomys found in bam owl pellets from Lastarria and Baiios del Flaco 
and from animals trapped in Farellones near Santiago. 



Character 



Farellones* 



BaAos del Flaco 



Lastarria 



MH-1 


4.07 ± 0.12(3.8-4.1) 


4.1, 3.9 


2.6,t4.0, 3.1t 


MHC 


8.8 ± 0.44(8.1-9.1) 


9.2, 9.1 


7.9,6.1 


MAL 


6.07 ± 0.08 (5.9-6.2) 


6.1, 6.1 


5.9,6.0,6.1 


MDL 


4.03 ± 0.24 (3.6-4.4) 


3.6, 4.1 


3.6,3.9,3.1 


SL 


6.24 ± 0.16(6.1-6.5) 


6.2, 6.3 


5.8, 6.0, 5.5 



MH- 1 = Mandibular height at the first molar; MHC = mandibular height from condyle to angular process; MAL = 
mandibular alveolar length; MDL = mandibular diastema length; SL = symphysis length. 

Values from Farellones near Santiago are X ± 2 SE (and range; N = 7). Individual measurements are given for 
Baiios del Flaco and Lastarria. 

* LCM collection, f Juvenile, the last cheektooth is not fully erupted. 



Los MoUes, Pilcaniyeu, and Cerro Leones (Rio 
Negro), Cueva Traful (Neuquen), and San Rafael 
(Mendoza) (Hershkovitz, 1962; Pine et al., 1978; 
Pearson & Pearson, 1982; Massoia, 1982). In this 
paper we report new records of Euneomys in Chile 
and review the scant literature on the taxonomy 
and geographic distribution of this poorly known 
genus. 



New Records and Current Status 

Some new records are from bam owl (Tyto alba) 
pellets from Lastarria (near Temuco), 39°14'S, 
70°40'W, and from Bafios del Flaco (near San Fer- 
nando), 34°57'S, 70°26'W. In addition, six other 
specimens of Euneomys were recently captured on 
Grevy Island (Tierra del Fuego; 55°32'S, 67°37'W) 
and at Palli Aike (Magallanes; 52°25' S, 69''42'W) 
and Rio Baguales (Ultima Esperanza; 51°02'S, 
72°30'W); all are deposited in the Museo del Ins- 
tituto de la Patagonia (MIP), Punta Arenas. Data 
obtained from these new specimens have been in- 
corporated by locality into Table 2. 

Determination of this material to genus was not 
difficult, using Reise's (1973) key and studies by 
Hershkovitz (1962) and Mann (1978). However, 
determination to species is difficult, owing to a 
lack of good diagnostic characters, the small num- 
ber of specimens, and their spotty distribution. 

Osgood recognized two species of Euneomys in 
Chile: E. chinchilloides (with two subspecies) in 
Tierra del Fuego and Magallanes and E. petersoni 
in Ultima Esperanza and Aysen. Subsequently, 
Mann ( 1 944) described E. noei from Lo Valdes in 
the Andes near Santiago. Hershkovitz (1962) con- 
sidered E. noei as doubtfully separable from E. 
mordax, with type locality at San Rafael (Men- 



doza Province, Argentina), and thought that mor- 
daxwas probably a subspecies of £". chinchilloides. 
Greer (1965) referred his four Malleco specimens 
to E. chinchilloides petersoni, despite their greater 
similarity to noei or mordax in measured char- 
acters. Miller and Rottmann (1976) identified all 
their Euneomys from near Santiago as E. mordax. 
Mann (1978) thought all Chilean specimens rep- 
resented a single polytypic species: E. chinchil- 
loides chinchilloides from Grande de Tierra del 
Fuego Island and the adjacent mainland, E. c. 
petersoni from the eastern Andes of Ultima Es- 
peranza and Aysen, and E. c. noei from the Andes 
outside Santiago. Subsequently, Pine et al. (1979) 
suggested that two sympatric species of Euneomys 
occur in Farellones near Santiago, but declined to 
identify the other species until comparisons could 
be made with specimens representing the entire 
range of the genus. Karyotypes of nine specimens 
from Farellones show insignificant differences 
(Spotomo, pers. comm.). 

Thus, Hershkovitz ( 1 962) and Mann (1978) rec- 
ognized only one polytypic species, whereas Pine 
et al. (1979) and Tamayo and Frassinetti (1980) 
regarded specimens from different areas as differ- 
ent species. Tamayo and Frassinetti ( 1 980) thought 
Greer's Malleco specimens also represented Eu- 
neomys sp. Unfortunately, this taxonomic uncer- 
tainty is difficult to dispel, as only seven intact 
skulls of Euneomys from the Andes outside of 
Santiago are deposited in Chilean collections (La- 
boratorio Citogenetica de Mamiferos [LCM], Fac. 
Medicina, Universidad de Chile), and none is de- 
posited in collections in Mendoza or San Rafael, 
Argentina (R. A. Ojeda, pers. comm.; H. A. La- 
giglia, pers. comm.). 

Euneomys material recently obtained from owl 
pellets and deposited in the Museo Nacional de 
Historia Natural de Chile is incomplete, consisting 



YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS 



285 



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286 



HELDIANA: ZOOLOGY 



mainly of mandibles and maxillae. The limited 
samples and their disintegration make a statistical 
analysis impossible. Comparisons of the owl pellet 
material with the seven specimens collected near 
Santiago (table 1) show no differences. The same 
results are obtained when comparisons are ex- 
tended to include all known Chilean specimens 
(table 2). Using these criteria it is impossible for 
us to assign specimens from Lastarria and Banos 
del Flaco to either E. c. mordax or E. c. petersoni. 



Conclusion 

The absence of clear-cut differences between lo- 
calities in Table 2 suggests that there is only a 
single species represented, E. chinchilloides. Be- 
cause subspecific assignments at present can only 
be made geographically, treatment of subspecies 
is deferred until an adequate sample of specimens 
is available. The fact that Euneomys are not taken 
in traps at the same localities where they are found 
in owl pellets (e.g., Reise & Venegas, 1976; Pear- 
son &. Pearson, 1982; present study) suggests that 
current gaps in the distribution o^ Euneomys might 
be attributable to sampling error rather than to 
truly disjunct distributions. The new records re- 
ported here fill both morphological and geograph- 
ical gaps. 



Acknowledgments 

We thank Angel Spotomo and Claudio Venegas 
for making specimens of Euneomys available to 
us. Bruce D. Patterson, Ronald H. Pine, and an 
anonymous reviewer made valuable comments. 
Mrs. Veronica Aguirre typed the manuscript, and 
Luz Uribe assisted with the figure. This report is 
dedicated to P. Hershkovitz in recognition of his 
pioneering contributions to our understanding of 
Neotropical mammals. 



Literature Cited 

Greer, J. K. 1965. Mammals of Malleco Province, 
Chile. Publications of The Museum, Michigan State 
University, Biological Series, 3: 51-151. 

Hershkovitz, P. 1962. Evolution of Neotropical cric- 
etine rodents (Muridae). Fieldiana: 2kx)logy, 46: 1- 
524. 

Mann, G. 1 944. Dos nuevas especies de roedores. Bio- 
logica (Santiago), 1: 95-126. 

. 1978. Los pequenos mamiferos de Chile. Gay- 
ana: Zoologia, 40: 1-342. 

Markham, B. 1971. Catalogo de los anfibios, reptiles, 
aves y mamiferos de la Provincia de Magallanes, Chile. 
Publicaciones Instituto de la Patagonia, Serie Mono- 
grafias N2. 3, 64 pp. 

Massoia, E. 1982. Restos de mamiferos recolectados 
en el paraje Paso de Los Molles, Pilcaniyeu, Rio Negro. 
Revista Investigaciones Agropecuarias INTA, Buenos 
Aires, 17: 39-53. 

Miller, S. D., and J. Rottmann. 1976. Guia para el 
reconocimiento de mamiferos chilenos. Ed. Gabriela 
Mistral, Santiago, 200 pp. 

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

Pearson, O. P., and A. K. Pearson. 1982. Ecology 
and biogeography of the southern rainforests of Ar- 
gentina, pp. 129-142. In Mares, M. A., and H. H. 
Genoways, eds.. Mammalian Biology in South Amer- 
ica. A Symposium Held at the Pymatuning Laboratory 
of Ecology, May 10-14, 1981. Sf>ecial Publication Se- 
ries, Pymatuning Laboratory of Ecology, University 
of Pittsburgh, 6: 1-539. 

Pine, R. H., J. P. Angle, AND D. Bridge. 1978. Mam- 
mals from the sea, mainland, and islands at the south- 
em tip of South America. Mammalia, 42: 105-1 14. 

Pine, R. H., S. D. Miller, and M. L. S. Schamberger. 
1979. Contributions to the mammalogy of Chile. 
Mammalia, 43: 339-376. 

Reise, D. 1973. Clave para la determinacion de los 
craneos de marsupiales y roedores chilenos. Gayana: 
Zoologia, 27: 1-20. 

Reise, D. L., and W. Venegas. 1974. Observaciones 
sobre el comportamiento de la fauna de microma- 
miferos en la Region de Puerto Ibaiiez (Lago General 
Carrera), Aysen, Chile. Boletin Sociedad de Biologia, 
Concepcion, Chile, 47: 71-85. 

Tamavo, M., and D. Frassinetti. 1980. Catalogo de 
los mamiferos fosiles y vivientes de Chile. Boletin 
Museo Nacional de Historia Natural, Chile, 37: 323- 
399. 



YANEZ ET AL.: NEW RECORDS AND STATUS OF EUNEOMYS 



287 



Morphological Variation, Karyology, 

and Systematic Relationships 

of Heteromys gaumeri (Rodentia: Heteromyidae) 

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



ABSTRACTS 

Morphological variation was assessed within and among populations of Heteromys gaumeri 
using univariate and multivariate statistical analyses of external and cranial measurements. 
Although patterns and amount of nongeographic variation in H. gaumeri were similar to other 
heteromyines, geographic variation was relatively conservative. Mean values of most characters 
were statistically homogeneous among localities and spatially unpattemed. Consequently, no 
association was found between levels of within- and among-sample variation for individual 
characters (the "Kluge-Kerfoot phenomenon"). Populations of//, gaumeri were chromosomally 
monomorphic. The lack of morphological and chromosomal variation in //. gaumeri contrasts 
sharply with patterns in other heteromyines. Heteromys gaumeri is morphologically and chro- 
mosomally distinct from the //. desmarestianus species group (to which it is currently assigned) 
and appears to share some primitive characters with Liomys (the sister group of Heteromys). 
We recommend that H. gaumeri be removed from the H. desmarestianus group. 

La variacion morfologica intra e interpoblacional de Heteromys gaumeri fue evaluada usando 
analisis estadisticos univariados y multivariados de medidas extemas y craneales. A pesar de 
que los patrones y cantidad de variacion intrapoblacional en //. gaumeri fue similar a la de 
otros heterominos, la variacion geografica fue relativamente conservadora. Los valores pro- 
medio de la mayoria de los caracteres fueron estadisticamente homogeneos entre las localidades, 
sin mostrar ningun patron de variacion espacial. En conservencia, no se encontro asociacion 
alguna entre los niveles de variacion intra e interpoblacional para caracteres individuals ("fen- 
omeno Kluge-Kerfoot"). Las poblaciones de //. gaumeri fueron monomorficas cromosomica- 
mente. La falta de variacion tanto morfologica como cromosomica en H. gaumeri contrasta 
marcadamente con los patrones encontrados anteriormente para otros heterominos. Heteromys 
gaumeri es morfologica y cromosomicamente distinguible del grupo //. desmarestianus (al cual 
se asigna actualmente) y aparentemente comparte algunos caracteres primitivos con Liomys (el 
grupo hermano de Heteromys). Nosotros recomendamos que se remueva a H. gaumeri del 
grupo //. desmarestianus. 

Avalia-se a variafao morfologica intra- e interpopulacional de Heteromys gaumeri, atraves 
de analises estatisticas uni- e multivariadas de medidas extemas e craniais. Apesar dos padroes, 
e da quantidade de varia^ao intrapopulacional em H. gaumeri serem similares aos de outros 
heteromideos, a varia9ao geografica e relativamente conservadora. Os valores medios da maior 
parte dos carateres examinados sao estatisticamente homogeneos entre as localidades, e nao 



From Department of Biology, Angelo State Univer- 
sity, San Angelo, TX 76909 (Engstrom); Section of 
Mammals, Carnegie Museum of Natural History, 4400 
Forbes Avenue, Pittsburgh, PA 1 5206 (Genoways; pres- 
ent address. University of Nebraska State Museum, 212 
Morrill Hall, Lincoln, NE 68588); and Department of 
Wildlife and Fisheries Sciences, Texas A&M University, 
College Station, TX 77843 (Tucker, present address, The 
Jackson Laboratory, Bar Harbor, ME 04609). 

ENGSTROM ET AL.: HETEROMYS GA UMERI 289 



surgiu nenhum padrao de variances locals. Consequentemente, nao foram encontradas asso- 
cia96es entre os niveis de varaiafoes intra- e interpopulacionais para carateres individuals (o 
"fenomeno KJuge-Kerfoot"). Popula^oes de H. gaumeh mostraram-se cromossomicamente 
monomorficas. A falta de varia9ao morfologica ou cromossomica em H. gaumeri e altamente 
contrastante aos padroes encontrados em outros heteromideos. Heteromys gaumeri dlstingue- 
se tanto morfologica quanto cromossomicamente do grupo H. desmarestianus. ao qual esta 
atualmente deslgnado, e aparentemente possue carateres primitivos em comum com Liomys— 
grupo irmao de Heteromys. Recomendamos que H. gaumeri seja removido do grupo H. des- 
marestianus. 



Introduction 

Spiny mice of the genus Heteromys (Hetero- 
myidae: Heteromyinae) are Neotropical rodents 
that typically occur in rainforest and cloudforest 
habitats from east-central Mexico south to north- 
em South America. Currently, the genus is par- 
titioned into two subgenera {Xylomys and Hetero- 
mys), and two species groups {desmarestianus and 
anomalus groups) are recognized in the nominate 
subgenus (Goldman, 1911; Hall, 1981; Rogers & 
Schmldly, 1982). Hall and Kelson (1959) and Hall 
(1981) noted that taxonomlc relationships within 
Heteromys and in particular the H. desmaresti- 
anus species group were problematical and in need 
of revision. Recently, Rogers and Schmldly ( 1 982) 
reviewed morphological variation among repre- 
sentatives of the H. desmarestianus species group 
from northern Middle America, exclusive of H. 
gaumeri, and concluded that only two species (//. 
desmarestianus and H. goldmani) were represent- 
ed in the material they examined. 

Heteromys gaumeri, the third species currently 
recognized in the desmarestianus group, is endem- 
ic to the Yucatan Peninsula. Ecologically this 
species characteristically occurs in relatively dry 
deciduous and subdeciduous-subperennlal forest. 
Systematic relationships of H. gaumeri are enig- 
matic. In his review of the subfamily Heteromyi- 
nae, Goldman (1911) placed H. gaumeri in his 
desmarestianus group for convenience, but noted 
(p. 29) that it was "not closely related to any known 
species." In this paper, we review morphological 
and chromosomal variation within H. gaumeri and 
comment on the systematic relationships of this 
species to the H. desmarestianus species group and 
the subgenus Heteromys. 



Materials and Methods 

A total of 322 specimens o{ Heteromys gaumeri 
was examined in the morphological analyses. 



Specimens examined are listed in the Appendix, 
and institutions housing those specimens are as 
follows (abbreviations for North American col- 
lections follow Choate &. Genoways, 1 975): Amer- 
ican Museum of Natural History (amnh); British 
Museum (Natural History), London (bmnh); Field 
Museum of Natural History (fmnh); Museum of 
Natural History, University of Kansas (ku); James 
Ford Bell Museum of Natural History, University 
of Minnesota (mmnh); Royal Ontario Museum 
(ROM); Texas Cooperative Wildlife Collections, 
Texas A«&M University (tcwc); The Museum, 
Texas Tech University (ttu); Museum of Zoology, 
University of Michigan (ummz); Unlversidad Na- 
cional Autonoma de Mexico (unam); and National 
Museum of Natural History (usnm). 

Each specimen was assigned to one of five pre- 
sumptive age classes (I-V, from youngest to oldest) 
based on pelage characteristics and state of erup- 
tion and relative wear of the maxillary toothrow, 
following Genoways (1973). For each adult and 
selected subadult specimens, the following four 
external (recorded from specimen labels) and 10 
cranial measurements (measured to the nearest 0. 1 
mm using dial calipers) were taken, as defined by 
Genoways (1973): total length (TL); length of tail 
(TV); length of hind foot (HF); length of ear (LE); 
greatest length of skull (GLS); zygomatic breadth 
(ZB); interorbital constriction (IOC); mastoid 
breadth (MB); length of nasals (LN); length of ros- 
trum (LR); length of maxillary toothrow (MTR); 
depth of braincase (DBC); Interparietal width 
(IW); and interparietal length (IL). Each measure- 
ment was chosen because it was examined in pre- 
vious studies of variation in other heteromyine 
rodents and was geographically variable in some 
taxa (Goldman, 1911; Genoways, 1973; Rogers & 
Schmldly, 1982). 

Initially, four qualitative cranial characters and 
dorsal coloration also were examined, as described 
by Genoways (1973) for Liomys (the sister group 
of Heteromys). Unlike patterns found in species 
of Liomys, each of these characters was equally or 



290 



FIELDIANA: ZOOLOGY 



more variable within populations of H. guumeri 
as among populations, and the characters varied 
in no apparent geographic pattern. Consequently, 
qualitative cranial characters and dorsal color- 
ation were not analyzed further. 

Nongeographic variation in the 14 mensural 
characters was examined in one sample of 94 spec- 
imens collected near Campo Experimental Fores- 
tal "El Tormento," 7.5 km W Escarcega, Cam- 
peche. These specimens were collected within a 
two square kilometer area of transitional tropical 
evergreen-tropical deciduous forest and were here 
considered to represent a single population. All 
calculations were performed using subprograms of 
the Statistical Analysis System (SAS; SAS Insti- 
tute, Inc., 1982). Standard statistics (mean, range, 
standard deviation, standard error, coefficient of 
variation, skewness, and kurtosis) were calculated 
for each variable within each subgroup (MEANS 
and UNIVARIATE procedures). Student's / test 
(or an approximation of variances were unequal) 
was used to test for significant differences between 
sexes within each age class (TTEST procedure). A 
model I, one-way analysis of variance (ANOVA) 
was used to test for significant differences among 
age classes with sexes pooled (GLM procedure). 
Subsequently, a Duncan's multiple range test 
(DUNCAN) was used to determine maximally 
nonsignificant subsets of age classes. 

Straney (1978) criticized the use of F tests in 
unbalanced ANOVAs to determine patterns of 
nongeographic variation. To augment hypothesis 
testing, variance partitioning of a model II, two- 
way ANOVA (VARCOMP procedure) was used 
to estimate the relative contributions of sex, age, 
sex by age interaction, and error to within sample 
variation. Age and sex were considered random 
factors (see discussion in Leamy, 1983) and the 
percent contribution of each factor was estimated 
from variance components. The main effects (sex 
and age) in the two-way ANOVAs were not in- 
dependent because the data were unbalanced 
(Searle, 1971). Consequently, ANOVAs were run 
with sex entered into the model first, then again 
with age entered first. 

For analysis of geographic variation of mensural 
data, adult specimens were assigned to one of 1 1 
grouped localities to increase sample size (fig. 1). 
In no instance did a grouped locality cross a major 
physiographic or previously recognized taxonomic 
boundary. The specific geographic composition of 
samples is as follows: Group 1— Chuntuqui and 
Laguna de Sotz (= Zotz), Peten, Guatemala; 103 
km SE Escarcega (= Francisco Escarcega), Cam- 



peche, Mexico. Group 2 — 7.5 km W Escarcega, 
Campeche, Mexico. Group 3— Apazote, 7 km N, 
5 1 km E Escarcega, and La Tuxpeiia, Campeche, 
Mexico. Group 4— Dzibalchen and San Jose Car- 
pizo, Campeche, Mexico. Group 5— Esmeralda and 
Santa Rosa, Yucatan, Mexico. Group 6— Chichen 
Itza and Piste, Yucatan, Mexico. Group 7— Ti- 
zimin and Tunkas, Yucatan, Mexico. Group 8 — 
La Vega, Pueblo Nuevo Xcan, and Puerto Mo- 
relos, Quintana Roo, Mexico. Group 9— Felipe 
Carrillo Puerto, Quintana Roo, Mexico. Group 
10— Bacalar, Quintana Roo, Mexico. Group 1 1 — 
Kate's Lagoon and Rockstone Pond, Belize, Be- 
lize. 

For each mensural character, we analyzed two 
aspects of geographic variation: (1) statistical het- 
erogeneity of mean values among geographic sam- 
ples, and (2) significant departure of means from 
spatial randomness (see Sokal & Oden, 1978). The 
null hypothesis of homogeneity of means among 
grouped localities for each character was tested 
using a model I, one-way ANOVA (SAS:GLM). 
By ANOVA, the variance of each character was 
partitioned into among and within (error) locality 
effects, and the percentage of variation attributable 
to each effect was estimated from variance com- 
ponents (SAS: VARCOMP). Homogeneity among 
grouped localities across all characters was ex- 
amined using a multivariate analysis of variance 
(SAS:GLM, MANOVA). 

The null hypothesis of no geographic pattern 
among grouped locality means was examined by 
testing for significant association between geo- 
graphic and phenetic distance matrices. For each 
character, a phenetic distance matrix was con- 
structed in which the elements were calculated as 
the absolute differences between means for all pairs 
of localities (Sokal, 1979). Multivariate phenetic 
matrices of taxonomic distance (Sneath & Sokal, 
1973) were calculated for all 14 mensural char- 
acters and a set restricted to those showing sig- 
nificant heterogeneity, using the NT-SYS library 
of computer programs (Rohlf et al., 1982). Ele- 
ments of the geographic distance matrix were the 
actual map distances (in km) between all pairs of 
grouped localities (taken from the center of each 
grouped locality); all connections between pairs of 
localities were maintained because there are no 
obvious physiographic or ecological barriers to gene 
flow among populations of//, gaumeri on the Yu- 
catan Peninsula. Three test statistics (Mantel's Z; 
Spearman's rho, R; and a component of Kendall's 
tau, K^) were used to test for significant association 
between each phenetic distance matrix and the 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



291 



90 



21 



18 



87 

"T" 




KILOMETERS 
50 100 

I H r-" 

25 50 
MILES 



± 



21 



18 



90 



87 



Fig. 1 . Geographic distribution of Heteromys gaumeri and approximate geographic areas included in the 1 1 
grouped locahties used in analyses of morphological variation. Closed circles denote specimens used in analyses of 
geographic variation and open circles denote specimens examined but not included in statistical analyses. 



geographic distance matrix (Dietz, 1983). Values 
of P associated with each statistic were estimated 
from 2,000 random permutations using a FOR- 
TRAN program supplied by E. J. Dietz. 



In the chromosomal analysis, standard karyo- 
types were examined for 26 specimens oi Hetero- 
mys gaumeri. Additionally, karyotypes of one 
specimen of H. desmarestianus and 20 specimens 



292 



HELDIANA: ZCX)LOGY 



of H. anomalus were examined for comparison. 
Standard karyotypes were prepared from speci- 
mens sampled from natural populations using the 
in vivo bone marrow technique of Patton (1967), 
as modified by Lee (1969). Terminology regarding 
relative chromosome arm ratios is that of Patton 
(1967). In the calculation of Fundamental Num- 
bers (FN), relative chromosome arm ratios were 
scored conservatively. Several chromosomes re- 
corded as acrocentric here had telomeric knobs of 
chromatin in elongated preparations. Only chro- 
mosomes which consistently displayed second 
arms of chromatin, regardless of state of contrac- 
tion of the preparation, were scored as biarmed. 
The following voucher specimens are deposited in 
the Texas Cooperative Wildlife Collection, Texas 
A&M University, and the Carnegie Museum of 
Natural History (sample size in parentheses): Het- 
eromys gaumeri (total 26)— MEXICO. Campeche: 
7.5 km W Escarcega (8). Quintana Roo: 2 km N, 
8 km W Bacalar (8); 8 km NNE Felipe Carrillo 
Puerto (3); 2.5 km NNE Felipe Carrillo Puerto (1). 
Yucatan: Chichen Itza (1); Cenote Seco, 2 km E 
Chichen Itza (5). Heteromys desmarestianus (total 
l)-MEXICO. Chiapas: 9.4 km S Palenque (1). 
Heteromys anomalus (total 20)— VENEZUELA. 
Miranda: 25 km N Altagracia de Oricuto (6); 24 
km N Altagracia de Oricuto (1). Monagas: Cari- 
pito (4). Sucre: 40 km SW Caripito (9). 



Results 

Morphological Variation 

NoNGEOGRAPHic VARIATION— The Sample of 
Heteromys gaumeri (N = 94) from 7.5 km W Es- 
carcega, Campeche, was used to estimate within- 
sample variation of the 1 4 mensural characters. 
Two approaches were taken, hypothesis testing and 
estimation of variance components. 

Initially, / tests were used to test for significant 
differences between sexes in each of age classes I- 
IV. Males average larger than females for most 
characters in most age classes; however, significant 
differences between sexes were found for only one 
measurement in age class I (length of rostrum) and 
seven measurements in age class III (table 1). 

Results of ANOVA among age classes with sexes 
pooled are given in Table 1 . Separate analyses of 
the age effect within each sex gave results similar 
to those in Table 1 and are not presented. Signif- 
icant variation with age was found in each of the 



14 measurements. Each measurement tended to 
increase with age, although this pattern was not 
pronounced for length of maxillary toothrow. In 
the a posteriori DUNCAN analysis, age class V 
averaged significantly larger than the other age 
classes for two characters (interorbital constriction 
and interparietal length), whereas age classes IV- 
V formed a homogeneous subset for the remaining 
12 characters. Patterns of variation among age 
classes I-III were less consistent, although age 
classes I-III differed significantly from each other 
and age classes IV-V in six of 10 cranial mea- 
surements. 

To complement hypothesis testing, the relative 
contributions of sex, age, sex by age interaction, 
and error (residual variation) to a two-way AN- 
OVA were estimated from variance components 
(table 2). Separate analyses with either sex or age 
entered into the model first generally yielded re- 
sults that differed by 1% to 4%. Because of the 
similarity of these estimates (and because sex most 
often is analyzed before age in studies of nongeo- 
graphic variation), only the results in which sex 
was entered into the model first are presented. 

Most of the variation (average 97%) in the AN- 
OVA was attributable to the effects of age and 
error. Age contributed the largest proportion of 
variance for most characters (average 53%); error 
was nearly as important, contributing an average 
of 44%. For length of hind foot, length of maxillary 
toothrow, interparietal width, and interparietal 
length, age contributed a relatively small propor- 
tion of the variance, and variation was mostly 
attributable to error. Homogeneity of estimates 
across characters was examined using z-transfor- 
mations and a subsequent c/z/-square test (Sokal 
&, Rohlf, 1981). Estimates of age and error were 
significantly heterogeneous across characters. 

Sex and interaction were relatively unimportant 
factors in the ANOVA for most characters, con- 
tributing an average of 1% and 2% of the variance, 
respectively. The only noteworthy exception to this 
pattern was interparietal length, for which inter- 
action accounted for 23% of the variance. Despite 
this exception, estimates of the effects of sex and 
interaction were statistically homogeneous across 
characters. Given that only a small proportion of 
the variance of each character was attributable to 
the effect of sex, the significant sexual dimorphism 
found for seven of 1 4 characters in age class III in 
preliminary / tests (table 1) probably was due to 
trivial differences accentuated by large sample size 
(N = 47). For most characters, the largest pro- 
portion of variance was attributable to age and 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



293 



Table 1. Age variation in 14 external and cranial measurements of //er^romy^^aM/wer/ from 7.5 km WEscarcega, 
Campeche, Mexico. 



Age 












class N 


Mean (range) ± 


2SE 


CV 


Total Length 












V 


2 


265.0 (237.0-272.0) 


+ 


10.00 


2.7 


IV 




14 


261.2(235.0-288.0) 


± 


9.82 


7.0 


III* 


42 


242.6(186.0-275.0) 


+ 


5.87 


7.8 


II 1 17 


220.6 (200.0-252.0) 


+ 


6.70 


6.3 


I 1 4 


190.3(179.0-210.0) 


± 


13.60 


7.1 


F= 19.45*** 












Length of Tail 












IV 


14 


145.5(137.0-159.0) 


± 


4.58 


5.9 


V 


2 


139.5(139.0-150.0) 


± 


1.00 


0.5 


HI* 




42 


132.5 


(87.0-152.0) 


± 


3.65 


8.9 


U 


17 


120.1(101.0-136.0) 


± 


4.62 


7.9 


I 1 4 


99.8 


(96.0-117.0) 


± 


6.85 


6.9 


F = 20.27*** 












Length of Hind Foot 












V 


3 


35.0 










IV 




16 


34.2 


(32.0-36.0) 


± 


0.66 


3.9 


in* 


47 


33.2 


(30.0-35.0) 


± 


0.40 


4.1 


n 


19 


33.0 


(30.0-35.0) 


± 


0.70 


4.6 " 


I 1 6 


31.0 


(29.0-34.0) 


± 


1.71 


6.8 


F = 6.76*** 












Length of Ear 










* 


V 


3 


17.7 


(17.0-18.0) 


± 


0.67 


3.3 


IV 


16 


17.4 


(17.0-19.0) 


± 


0.31 


3.6 


in 


47 


16.1 


(14.0-18.0) 


± 


0.26 


5.6 


u 


19 


15.6 


(14.0-18.0) 


± 


0.44 


6.1 


I 1 6 


14.8 


(14.0-16.0) 


± 


0.61 


5.1 


F= 15.77*** 












Greatest Length of Skull 










V 2 


35.8 


(35.5-36.0) 


± 


0.50 


1.0 


IV 15 


35.2 


(33.9-37.5) 


± 


0.49 


2.7 


HI* 1 47 


32.9 


(30.2-34.5) 


± 


0.26 


2.7 


n 1 19 , 


31.2 


(29.6-34.2) 


± 


0.50 


3.5 


I 1 6 


28.8 


(27.8-29.8) 


± 


0.65 


2.7 


F= 71.20*** 












Zygomatic Breadth 












V 


3 


16.3 


(16.1-16.5) 


± 


0.23 


1.2 


IV 


16 


16.1 


(15.3-16.7) 


± 


0.19 


2.4 


ni* 1 44 


15.1 


(14.0-16.3) 


± 


0.14 


3.1 


n 1 .19 


14.4 


(13.7-15.1) 


± 


0.20 


3.0 


I 1 6 


13.5 


(12.9-13.8) 


± 


0.33 


3.0 


F = 59.28*** 












Interorbital Constriction 










V 1 3 


9.3 


(9.2-9.5) 


± 


0.20 


1.9 


IV 1 16 


8.7 


(8.3-9.0) 


± 


0.10 


2.4 


ni* 1 47 


8.2 


(7.5-9.3) 


± 


0.12 


4.8 


n 1 19 


7.8 


(7.2-8.6) 


± 


0.16 


4.3 


I 1 6 


7.5 


(7.0-7.7) 


± 


0.20 


3.3 


F= 29.71*** 












Mastoid Breadth 












V 


3 


15.5 


(15.3-15.8) 


± 


0.30 


1.7 


IV 


15 


15.3 


(14.8-15.9) 


± 


0.18 


2.3 


III 1 47 


14.7 


(14.0-15.4) 


± 


0.10 


2.3 



294 



FIELDIANA: ZOOLOGY 



Table 1. Continued. 



Age 
class 



N 



Mean (range) ± 2 SE 



CV 



19 
6 

F = 27.94*** 



14.4 (13.4-15.2) ± 0.21 3.2 

13.7 (13.1-14.1) ± 0.29 2.6 



V 

IV 

III 

II 

I 



Length of Nasals 

2 14.5 (14.0-15.0) ± 1.00 4.9 

16 14.3 (13.2-15.6) ± 0.37 5.1 

47 13.0 (11.8-14.2) ± 0.18 4.7 

19 11.9 (10.8-13.4) ± 0.29 5.3 

6 10.5 (10.0-10.9) ± 0.26 3.0 

F = 59.06*** 



V 

IV 

III* 

II 

I* 



Length of Rostrum 

2 15.6 (15.1-16.0) ± 0.90 4.1 

16 15.2 (14.2-16.3) ± 0.27 3.6 

47 14.0 (12.7-15.3) ± 0.18 4.3 

19 12.9 (11.9-13.9) ± 0.23 3.9 

6 11.6 (11.0-12.1) ± 0.37 3.9 

F = 65.77*** 



IV 

II 

V 

III 

I 



Length of Maxillary Toothrow 

16 4.9 (4.7-5.2) ± 0.07 3.0 

17 4.8 (4.5-5.1) ± 0.08 3.3 
3 4.8 (4.7-4.8) ± 0.07 1.2 

46 4.7 (4.3-5.2) ± 0.05 3.4 

5 4.7 (4.5-4.8) ± 0.17 2.8 
F = 3.72** 



V 

IV 

III 

II 

I 



Depth of Braincase 

3 9.1 (8.9-9.4) ± 0.29 2.8 

16 9.0 (8.5-9.5) ± 0.13 3.0 

46 8.8 (8.3-9.7) ± 0.08 3.3 

19 8.5 (7.9-8.8) ± 0.13 3.3 

6 8.5 (8.3-8.8) ± 0.16 2.3 

F= 12.68*** 



V 

IV 

II 

III 

I 



Interparietal Width 

3 9.3 (8.9-9.8) ± 0.54 5.1 

15 8.8 (8.0-10.0) ± 0.31 6.8 

19 8.6 (7.6-9.4) ± 0.21 5.3 

45 8.5 (7.3-9.7) ± 0.17 6.7 

6 8.0 (7.3-8.6) ± 0.36 5.6 

F=4.11** 



V 

IV 

III 

II 

I 



Interparietal Length 

3 5.5 (5.1-6.1) ± 0.61 9.6 

15 5.1 (4.2-5.8) ± 0.21 8.0 

45 5.0 (4.2-5.7) ± 0.10 6.8 

19 4.9 (4.2-5.3) ± 0.14 6.3 

6 4.7 (3.8-5.3) ± 0.45 11.9 

F= 3.23* 



Vertical lines alongside age classes denote nonsignificant subsets. Asterisks after F statistics indicate levels of 
significance (* P < 0.05; ** P < 0.01; *** P < 0.001). Males and females in age classes I-IV were used to test for 
significant mean differences due to sex (/ test). Asterisks following age classes indicate significant sexual dimorphism 
{P < 0.05). 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



295 



Table 2. Percentage of total variation attributable to 
sex (S), age (A), sex by age interaction (S x A), and error 
(E) for 14 external and cranial measurements of Hetero- 
mys gaumeri from 7.5 km W Escarcega, Campeche, 
Mexico. 



Char- 




Variance components 




acters* 


S 


A 


S X A 


E 


TL 





61.3 


1.5 


37.2 


TV 





63.6 





36.4 


HF 


4.8 


29.9 





65.3 


LE 





50.3 





49.7 


GLS 





84.1 





15.9 


ZB 





77.5 


4.6 


17.9 


IOC 





66.7 


0.5 


32.8 


MB 





64.1 


2.0 


33.9 


LN 





79.7 


1.4 


18.9 


LR 





82.1 


0.8 


17.1 


MTR 


2.2 


21.1 





76.7 


DBC 


1.2 


42.4 


0.6 


55.8 


IW 


3.9 


19.1 





77.0 


IL 





1.1 


22.7 


76.2 


Mean 


0.9 


53.1** 


2.4 


43.6** 



* Abbreviations of characters are defined in text. Es- 
timates were calculated from variance components (con- 
sidering sex as a random factor). 

** Significant heterogeneity {P < 0.01) of estimates 
among characters. 



(TL) of the total variance. For those characters 
that were significantly heterogeneous, an average 
of 1 7% of the variance was attributable to locality. 
In a MANOVA across all mensural characters, 
locaHties were marginally heterogeneous {P < 0.05) 
by two test statistics (Hotelling-Lawley Trace and 
Wilk's Criterion), but homogeneous by a third test 
statistic (Pillai's Trace), suggesting an overall lack 
of mensural differentiation among localities. 

Because significant spatial patterning is theo- 
retically possible (although unlikely) even with sta- 
tistically homogeneous means (Sokal & Oden, 
1 978), each character was tested for significant de- 
partures from spatial randomness. We found no 
significant association between matrices of phe- 
netic and geographic distance for any character by 
any of three test statistics. Multivariate taxonomic 
distance matrices calculated by using all characters 
and using only those characters that were signifi- 
cantly heterogeneous among localities were also 
incongruent with geographic distance. According- 
ly, mean values of most mensural characters were 
homogeneous among grouped localities and not 
spatially patterned as a simple function of geo* 
graphic distance. 



error, and the main effects in the model (sex and 
age) were independent. 

Variance components also were estimated from 
a two-way ANOVA restricted to age classes IV 
and V (results available on request). In this anal- 
ysis, the pattern of variance partitioning changed 
considerably. For each character, error contrib- 
uted the largest proportion of variance (average 
64%), whereas sex, age, and interaction had small- 
er average contributions ( 1 0%, 1 2%, and 1 4%, re- 
spectively). These estimates, however, should only 
be regarded as approximations because of the small 
sample size of age class V (N = 3). Estimates de- 
rived from larger subsets of age classes included a 
large average contribution (> 40%) attributable to 
the effect of age. Based on these results, geographic 
analyses were restricted to age classes IV-V with 
sexes pooled. 

Geographic Variation— Standard statistics 
were calculated for each mensural character in each 
grouped locality. By ANOVA, the variance of each 
character was partitioned into among and within 
(error) locality effects. Only six of 14 characters 
were significantly heterogeneous among grouped 
localities (table 3). For seven characters (HF, ZB, 
IOC, MB, DBC, IW, IL), the among-locality vari- 
ance component was zero and for the remaining 
characters, locality accounted for 6% (LE) to 30% 



Karyology 

Heteromys gaumeri (2n = 56, FN = 76; fig. 
2a)— The autosomal complement comprises grad- 
ed series of 1 1 pairs of large- to small-sized meta- 
centric and submetacentric chromosomes and 1 6 
pairs of large- to small-sized acrocentric elements. 
The X chromosome is large and submetacentric 
and the Y is medium-sized and subtelocentric. No 
variation was noted among individuals. 

Heteromys desmarestianus (2n = 60, FN = 
66; fig. 2b)— The autosomal complement com- 
prises four pairs of submetacentric and metacen- 
tric chromosomes, two large, one medium-sized, 
and one small, and 25 acrocentric pairs graded 
from large to small. The X chromosomes are pre- 
sumed to be large and submetacentric (no males 
were examined). 

Heteromys anomalus (2n = 60, FN = 68; fig. 
2c)— The autosomal complement comprises two 
large, one medium-sized, and two small pairs of 
submetacentric and metacentric chromosomes and 
24 acrocentric pairs graded from large to small. 
The X chromosome is large and submetacentric 
and the Y is medium-sized and subtelocentric. No 
variation was noted among individuals. 



296 



FIELDL\NA: ZOOLOGY 



Discussion 



r Patterns of nongeographic variation in mensural 
characters of Heteromys gaumeri are generally 
concordant with those observed in other hetero- 
myine rodents (Genoways, 1973; Rogers & 
Schmidly, 1982; see also Straney, 1978). In all 
species examined, age contributes a large propor- 
tion of within-sample variance for most charac- 
ters; only a minor component of total variance is 
attributable to sex. Male heteromyines generally 
average larger than females, however, and with 
large sample size secondary sexual dimorphism 
often appears significant for some characters (Gen- 
oways, 1973; age class III, this study). 

I Relative levels of variability of individual char- 
acters among species of heteromyines were com- 
pared using coefficients of variation of adults for 
the 12 external and cranial measurements com- 
mon to each study of intralocality variation (total 
length, length of tail, and the 10 cranial measure- 
ments included in this study; see Genoways, 1 973; 
Rogers & Schmidly, 1982). Coefficients of varia- 

f tion (CVs) appeared congruent among species for 
individual characters, although CVs appeared het- 
erogeneous among characters within species (ex- 
ternal measurements and interparietal width and 
length consistently were more variable than other 
characters). There was no indication of a reduction 
of within-sample variation for any character in H. 
gaumeri relative to other heteromyines. 

Although patterns and level of intralocality vari- 
ation in Heteromys gaumeri appear similar to oth- 
er heteromyines, geographic variation in H. gau- 
meri is relatively conservative. In H. gaumeri, 
mean values for most characters were homoge- 
neous among localities and geographically unpat- 
temed; only a small proportion of variance (av- 
erage 8%) was attributable to interlocality variation. 
We found no positive statistical correlation be- 
tween levels of within- and among-sample vari- 
ance (the "Kluge-Kerfoot phenomenon," KJuge & 
Kerfoot, 1 973) for mensural characters in H. gau- 
meri (tested using Kendall's rank correlation be- 
tween the W, and A, statistics suggested by Sokal, 
1976; but see Rohlf et al., 1983). These data con- 
trast with studies of geographic variation within 
species of Liomys (Genoways, 1973) and other 
members of the H. desmarestianus species group 
(Rogers & Schmidly, 1982), in which population 
samples were statistically heterogeneous and ap- 
peared spatially patterned. 
The relative lack of interlocality variance in 



Heteromys gaumeri might be attributable to a re- 
stricted geographic distribution, to relative envi- 
ronmental homogeneity on the Yucatan Penin- 
sula, and/or to a lack of genetic divergence among 
populations. Compared to wide-ranging species of 
Liomys and Heteromys, the geographic area oc- 
cupied by H. gaumeri is relatively small, with little 
topographic or climatic relief Plant communities 
grade gradually from lowland tropical evergreen 
forest in the south and east, to deciduous forest in 
the northwestern and north-central portions of the 
peninsula, to a scrub zone bordering the northern 
coast (Paynter, 1955). For statistically heteroge- 
neous but spatially unpattemed characters, char- 
acter distributions might be determined mainly by 
stochastic factors (e.g., genetic drift) in the absence 
of strong migration or selective gradients (Sokal 
&. Oden, 1978). Peromyscus yucatanicus, which 
has a similar geographic range and occupies sim- 
ilar habitats to H. gaumeri, is geographically vari- 
able in color and cranial size (Lawlor, 1 965; Huck- 
aby, 1980), and at least color closely tracks 
vegetational changes on the peninsula. Conse- 
quently, lack of interlocality differentiation in col- 
or, qualitative, and mensural characters in H. gau- 
meri might not result solely from selective 
responses or stochastic processes in a homoge- 
neous environment. 

The reduced level of geographic variation in 
Heteromys gaumeri also is consistent with a hy- 
pothesis of little genetic divergence among pop- 
ulations. If populations of H. gaumeri are genet- 
ically similar, similarity probably is not a product 
of panmixia, but more likely of a reduction of 
genetic variation through a genetically depauper- 
ate founding population (see Johnston, 1976; 
Johnston &. KJitz, 1977; Baker, 1980) or genetic 
bottleneck. Although we have little direct evidence 
of genetic variation among populations of//, gau- 
meri, individuals sampled appear chromosomally 
monomorphic. In //. desmarestianus, interlocality 
polymorphism in fundamental number is pro- 
nounced (D. S. Rogers, pers. comm.). More sen- 
sitive estimation of the level of interpopulational 
genetic divergence of //. gaumeri awaits study of 
genie variation. 



Systematic Relationships of H. gaumeri 

MorpholcxjV— Allen and Chapman (1897) de- 
scribed Heteromys gaumeri from seven individ- 
uals from Chichen Itza, Yucatan. In his review of 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



297 



Table 3. Geographic variation in six external and cranial measurements of Heteromys gaumeri. 



Grouped 
locality 



Percentage of variation 



N 



Mean (range) ± 2 SE 



Locality 



Error 



1 
2 
3 

4 
S 

6 

7 
8 
9 

10 
11 



3 
16 
10 

S 

9 
17 

4 
3 
3 

2 
4 



288.7 
261.7 
276.9 
272.0 
280.0 
286.8 
294.5 
281.7 
263.3 
256.5 
270.5 



Total Length 

276.0-298.0) ± 13.13 

235.0-288.0) ± 8.63 

245.0-302.0) ± 10.98 

262.0-286.0) ± 8.83 

263.0-295.0) ± 7.51 

265.0-324.0) ± 8.29 

288.0-300.0) ± 4.93 

275.0-290.0) ± 8.82 

250.0-280.0) ± 17.64 

253.0-260.0) ± 7.00 

250.0-290.0) ± 18.65 



23.7 



76.3 



3.75* 



Length of Tail 



1 


3 


154.7 ( 


148.0-166.0) 


+ 


11.39 13.6 


86.4 


2.02* 


2 


16 


144.8 ( 


129.0-159.0) 


+ 


4.12 






3 


10 


150.4( 


124.0-173.0) 


+ 


9.73 






4 


5 


154.4( 


141.0-166.0) 


+ 


9.48 




• 


5 


9 


161.0( 


144.0-193.0) 


+ 


9.22 






6 


17 


154.4( 


135.0-183.0) 


± 


5.53 






7 


4 


162.8 ( 


160.0-166.0) 


± 


2.75 




- 


8 


3 


152.7 ( 


151.0-155.0) 


± 


2.40 




V- 


9 


3 


145.7 ( 


140.0-150.0) 


+ 


5.93 






10 


2 


147.5 ( 


147.0-148.0) 


+ 


1.00 






11 


4 


146.5 ( 


129.0-160.0) 


+ 


12.92 












Greatest Length of Skull 






1 


3 


35.8 


(35.2-36.2) 


+ 


0.61 15.6 


84.4 


2.30* 


2 


17 


35.3 


(33.9-37.5) 


± 


0.44 








9 


35.0 


(33.5-36.3) 


+ 


0.57 








S 


34.9 


(33.6-36.2) 


± 


0.88 








9 


35.7 


(34.3-36.7) 


+ 


0.52 








23 


35.9 


(34.1-38.2) 


+ 


0.41 








6 


36.4 


(35.8-37.0) 


± 


0.41 






8 


3 


35.2 


(33.8-36.6) 


± 


1.62 






9 


4 


35.5 


(34.2-36.6) 


± 


1.10 






10 


3 


34.4 


(34.0-34.7) 


+ 


0.40 






11 


2 


34.6 


(34.3-34.8) 


+ 


0.50 












Length of Nasals 






1 


3 


15.2 


(14.7-15.7) 


+ 


0.58 13.8 


86.2 


2.19* 


2 


18 


14.3 


(13.2-15.6) 


+ 


0.34 








10 


13.9 


(12.8-14.5) 


+ 


0.40 








5 


14.2 


(13.6-15.0) 


+ 


0.61 








9 


14.6 


(13.7-15.5) 


+ 


0.39 








25 


14.8 


(13.6-15.8) 


+ 


0.27 








6 


14.8 


(14.0-15.5) 


+ 


0.47 








4 


14.6 


(13.7-15.8) 


+ 


0.87 








4 


14.6 


(13.9-15.2) 


± 


0.54 






10 


3 


13.8 


(13.2-14.5) 


± 


0.75 






11 


2 


13.9 


(13.3-14.5) 


± 


1.20 












Length 


of Rostrum 








3 


15.4 


(14.8-16.3) 


± 


0.92 10.5 


89.5 


2.16* 




18 


15.4 


(14.2-16.3) 


± 


0.25 








10 


15.1 


(14.1-15.8) 


± 


0.38 








5 


15.2 


(14.2-16.0) 


± 


0.61 








8 


15.5 


(14.8-16.0) 


± 


0.33 








25 


15.6 


(14.6-16.6) 


± 


0.23 








6 


16.0 


(15.1-16.7) 


± 


0.62 






8 


4 


15.4 


(14.4-16.4) 


± 


0.84 







298 



FIELDIANA: ZOOLOGY 



Table 3. Continued. 



Grouped 


N 


Mean (range) + 2 SE 


Percentage of variation 




locality 


Locality Error 


F 


9 
10 
11 


4 
3 
2 


15.6 (15.2-16.0) ± 0.37 
14.8 (14.2-15.5) ± 0.75 
14.6 (14.4-14.8) ± 0.40 







Length of Maxillary Toothrow 

18.9 



1 


3 


4.9 


(4.7-5.1) 


± 0.23 


2 


19 


4.9 


(4.7-5.2) 


± 0.07 


3 


10 


4.9 


(4.6-5.2) 


± 0.13 


4 


5 


5.0 


(4.9-5.1) 


± 0.06 


5 


8 


4.8 


(4.6-5.0) 


± 0.08 


6 


26 


5.0 


(4.4-5.3) 


± 0.08 


7 


6 


5.2 


(4.9-5.6) 


± 0.21 


8 


4 


4.9 


(4.7-5.1) 


± 0.17 


9 


4 


4.8 


(4.6-5.0) 


± 0.17 


10 


4 


4.9 


(4.8-5.1) 


± 0.13 


11 


2 


5.2 


(5.0-5.4) 


± 0.40 



81.1 



2.76** 



Asterisks following F statistics indicate levels of significance (see footnote to Table 1). Percentages of variation are 
based on variance components estimated from the entire data set. Grouped localities are defined in text and outlined 
in Figure 1. 



the Heteromyinae, Goldman (1911) placed H. 
gaumeri in his H. desmarestianus group, but noted 
that it was aberrant and not closely related to any 
known species. Although H. gaumeri possesses 
characters diagnostic for Heteromys, including up- 
per and lower molars with three lophs and lower 
permanent premolars with three lophs, it also 
shares some characters with the sister group Lio- 
mys not shared with other Heteromys, including 
posterior sole of hind foot haired as in Liomys 
rather than naked as in other Heteromys and early 
disappearance of the enamel island between cin- 
gulum and metaloph(id) of upper and lower mo- 
lars, a condition somewhat intermediate between 
the genera (Goldman, 191 l;Genoways, 1973). As- 
suming Heteromys is monophyletic, the characters 
shared between H. gaumeri and Liomys probably 
are primitive (based on the "ojierational rule" out- 
group procedure outlined by Watrous & Wheeler, 
1981). Dorsal coloration of H. gaumeri also is 
unusual in that the dorsum has a definite ochra- 
ceous cast with a bright ochraceous lateral line 
resembling Liomys pictus and L. spectabilis more 
than other, darker Heteromys. However, dorsal 
coloration in H. gaumeri might be convergent on 
L/omys through their common occupation of xeric 
forest and thorn scrub habitats rather than more 
mesic forest characteristic of most other species 
of Heteromys. 

Karyology — Compared to the nominate 
species in the Heteromys desmarestianus and H. 



anomalus species groups, the karyotype of//, gau- 
meri also is divergent (see fig. 2). Superficially, the 
karyotypes of //. desmarestianus (2n = 60, FN = 
66) and //. anomalus (2n = 60, FN = 68) appear 
similar and might differ by a single rearrangement 
(heterochromatic addition/deletion or pericentric 
inversion), whereas the karyotype of //. gaumeri 
(2n = 56, FN = 76) differs by a minimum of seven 
rearrangements from //. desmarestianus and six 
from //. anomalus. The karyotype reported herein 
for //. desmarestianus differs from that reported 
by Genoways (1973; 2n = 60, FN = 82) in having 
a larger number of acrocentric elements. Popula- 
tions of //. desmarestianus are variable in fun- 
damental number and the karyotype reported 
herein is among the lowest FN karyotypes known 
for the species (D. S. Rogers, pers. comm.). 

Conclusions— We agree with Goldman (1911, 
p. 30) that Heteromys gaumeri "is a somewhat 
aberrant species, presenting characters which set 
it off from all the others [species of the H. des- 
marestianus group]." The distinctive morpholog- 
ical, ecological, and karyotypic features of//, gau- 
meri distinguish it from other members of the //. 
desmarestianus group (including //. goldmani; D. 
S. Rogers, pers. comm.). Shared (probably prim- 
itive) characters with Liomys suggest that H. gau- 
meri might represent an early branch of the lineage 
leading to other Heteromys (which share probable 
derived states for these characters). At present, we 
believe the distinctness and unresolved phyloge- 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



299 



XI &{ii SI M >(» 99 MM 



AiS Mr A« 



A^it/s. 



OA Af> A* OH Awn. /in Ml Aft 
r%^ AA AA 4 



X Y 



iri B Kn .. 11 a it 

9A tl DC «» ai t* It im 



DA lid t\ »t tt) U M AA 



OA nn 60 Oft Aft »» 



X X 



10 n QO AA A9 00 to A« 



i^a i<( (in ft A #0 #11 ak aq 

HA (fS llW 411 HA #14 jf^ 



X X 




Fig. 2. Representative karyotypes of Heteromys: A, karyotype of a male H. gaumeri (2n = 56, FN = 76) from 
7.5 km W Escarcega, Campeche, Mexico; B, karyotype of a female H. desmarestianus (2n = 60, FN = 66) from 9.4 
km S Palenque, Chiapas, Mexico; C, karyotype of a female H. anomalus {2n = 60, FN = 68) from Caripito, Monagas, 
Venezuela. Insert shows sex chromosomes of a male. 



300 



FIELDIANA: ZOOLOGY 



netic position of H. gaumeri would best be em- 
phasized by removing it from the H. desmaresti- 
anus group, and recognizing it as a divergent species 
at the same cladogenic level as the species groups 
in the subgenus Heteromys. Further investigation 
of heteromyines might indicate that H. gaumeri 
warrants subgeneric recognition. 



Species Account 

I Heteromys gaumeri Allen and Chapman, 1897 

Heteromys gaumeri Allen and Chapman, 1897, Febru- 
ary. Bull. Amer. Mus. Nat. Hist., 9: 9. 

HoLOTYPE— Adult male, skin and skull, amnh 
12028/10461; from Chichen Itza, Yucatan, Mex- 
ico. Type examined. 

Distribution— Northern Belize; Peten, Gua- 
temala; and Campeche, Quintana Roo, and Yu- 
catan, Mexico (see fig. 1); subdeciduous-subpe- 
rennial tropical rain forest, tropical deciduous 
forest, and thorn scrub forest from sea level to 
100 m. 

Description— Dorsal coloration of adults ranges 
from dark to medium gray, with heavy admixture 
of orange buff hairs lending an overall ochraceous 
cast to the otherwise gray dorsum (adults in worn 
pelage appear more ochraceous and molting in- 
dividuals often have a 'salt and pepper' appear- 
ance); rich orange buff lateral line, usually broad 
and conspicuous, extending from cheeks to base 
of tail, ochraceous hairs often extending onto dor- 
sal and ventral surfaces of ankle; margins of fore- 
arm orange buff, interrupted on dorsal surface by 
white line; venter and feet white; tail well haired, 
grayish brown above, dull white below, with con- 
spicuous terminal tuft of hairs; ears dusky, lightly 
edged with dull white; sole of hind foot haired, 
posteriorly. Subadults medium to dark gray above, 
dorsum without ochraceous hairs; ochraceous lat- 
eral line faint and narrow. Juvenile pelage similar 
to that of subadults, but spiny hairs on dorsum 
absent. Tail longer than head and body; soles of 
hind feet with six tubercules; body size medium 
for the genus. Skull size medium, with relatively 
large auditory bullae; lower permanent premolar 
with three lophs; upp>er and lower molars with 
three lophs, enamel island formed between meta- 
loph(id) and cingulum disappearing quickly with 
wear. 2n = 56, FN = 76. 

Couv\v.\so^s— Heteromys gaumeri is geo- 
graphically isolated from all other heteromyines 



except H. desmarestianus, which it might contact 
at the southern edge of the Yucatan Peninsula (see 
Jones et al., 1 974). Heteromys gaumeri differs from 
H. desmarestianus in averaging smaller in most 
external and cranial measurements; having a broad, 
bright ochraceous lateral line, extending onto 
cheeks and ankles (a narrow, pale ochraceous lat- 
eral line often is present in H. desmarestianus, but 
seldom extends onto cheeks or ankles); having soles 
of hind feet haired posteriorly (this area is naked 
in H. desmarestianus and all other Heteromys); 
having a relatively well-haired tail with terminal 
tuft (the tail is sparsely haired in H. desmaresti- 
anus, without a conspicuous terminal tuft); having 
relatively large auditory bullae; and in having a 
diploid number of 56 chromosomes (compared to 
60 in H. desmarestianus). For additional com- 
parisons with other heteromyines, see Goldman 
(191 1) and Genoways (1973). 

Remarks— Laurie (1957, p. 387) assigned eight 
specimens from three localities in the state of Yu- 
catan, Mexico (Tekom, 2; X-Cala-Koop, 1; Chi- 
chen Itza, 5) to Heteromys desmarestianus. One 
of us (HHG) reexamined her material, which is 
stored in alcohol; based on size, coloration, and a 
haired posterior sole of the hind foot, all are as- 
signable to H. gaumeri. Consequently, the north- 
ernmost locality for H. desmarestianus on the Yu- 
catan Peninsula is 85 km W Chetumal, Quintana 
Roo, Mexico (Jones et al., 1974). Two other spec- 
imens, from Kate's Lagoon, Belize, identified by 
Laurie (1957, p. 387) as H. desmarestianus, are 
referable to H. gaumeri. These specimens, along 
with additional material from Honey Camp La- 
goon (reported by Izor &. McCarthy, 1984) and 
Rockstone Pond reported here, suggest that H. 
gaumeri is distributed throughout northern Belize. 



Acknowledgments 

Permits to collect specimens in Mexico were 
kindly provided by the Director General of the 
Departmento de Conservacion de la Fauna Sil- 
vestre. Especial thanks are extended to Biol. Fran- 
cisco Rodriguez Gallegos, Ing. Antonio Sanchez, 
and personnel of Campo Experimental Forestal 
"El Tormento" for their hospitality during field- 
work near Escarcega, Campeche. We also thank 
the many curators who permitted examination of 
specimens in their care. E. Jaquelin Dietz provided 
the computer program for testing the association 
between distance matrices. Kenneth Schoenly pre- 
pared the final copy of Figure 1. D. S. Rogers 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



301 



provided information and helpful discussion on 
character-states within the Heteromyinae. Por- 
tions of this study were funded by the Texas Ag- 
ricultural Experiment Station (project H-1977 to 
D. J. Schmidly) and a Faculty Organized Research 
Grant, Angelo State University. Fieldwork in Ven- 
ezuela was supported by NSF Grant DEB 79-2 1519 
to J. W. Bickham and the Carnegie Museum of 
Natural History through the M. Graham Netting 
Research Fund, established by a grant from the 
Cordelia S. May Charitable Trust. 



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HELDIANA: ZOOLOGY 



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Appendix 
Specimens Examined 

MEXICO: Campeche: Apazote, near Yohaltun, 

6 (usnm); Campeche, 1 (usnm); 5 km S Cham- 
poton, 10 m, 3 (ku); Dzibalchen, 1 (ku); 7 km N, 
5 1 km E Escarcega (= Francisco Escarcega), 4 (ku); 
7.5 km W Escarcega, 65 m, 94 (15 ku, 79 tcwc); 

7 km E Escarcega, 4 (ku); 103 km SE Escarcega, 
2 (ku); La Tuxpena, Champoton, 1 2 (usnm); San 
Jose Carpizo, 3 (fmnh); San Jose Carpizo, 45 km 
S Campeche, 19 (ummz); San Juan, 4 (fmnh). 
Quintana Roo: 2 km N, 8 km W Bacalar, 1 1 (tcwc); 

8 mi NNE Felipe Carrillo Puerto, 4 (tcwc); 4 km 
NNE Felipe Carrillo Puerto, 30 m, 9 (ku); 2.5 mi 
NNE Felipe Carrillo Puerto, 1 (tcwc); La Vega, 5 
(usnm); Pueblo Nuevo Xcan, 10 m, 1 (ku); 1.5 
km S, 1 km E Pueblo Nuevo Xcan, 1 (ku); Puerto 
Morelos, 4 (usnm). Yucatan: Calcehtok, 55 km SW 
Merida, 2 (1 ku, 1 ummz); Chichen Itza, 10 m, 36 
(15 amnh, 11 bmnh, 2 fmnh, 1 ku, 7 usnm); 
Chichen Itza, Cenote Xtoloc, 1 (tcwc); 2 km E 



Chichen Itza, Cenote Seco, 5 (tcwc); 4 km E Dzi- 
tya, 1 (mmnh); Esmeralda, Laguna de Chichan- 
canab, 2 (1 ku, 1 ummz); Esmeralda, 45 km S Peto, 
4 (ummz); 1 km SW Casa de la Esmeralda, Laguna 
de Chichancanab, 2 (1 ku, 1 unam); 2 km SSW 
Casa de la Esmeralda, Laguna de Chichancanab, 
3 (1 AMNH, 2 unam); La Ceiba, 2 km SE Laguna 
de Chichancanab, 1 (unam); 66 km NE Merida, 

1 (ku); 14 km SW Muna, 1 (rru); Oxkutzcab, 
Ebiztic (cave deposit), 6 (usnm); Peto, 3 (ku); 3 
km N Piste, 16 (7 ku, 6 mmnh, 3 ttu); 2 km N 
Piste, 2 (ku); Piste, 10 m, 2 (ku); Progreso, 1 (usnm); 
Santa Rosa, 1 (ummz); Santa Rosa, 20 km S Peto, 

2 (ummz); Santa Rosa, 25 km S Peto, 1 (ummz); 
1 km SSW Santa Rosa, 3 (1 ku, 1 ummz, 1 unam); 
Tekom, 2 (bmnh); 6 km N Tizimin, 1 (ku); Tun- 
kas, 7 (usnm). Buena Vista, 1 (fmnh) and X-Cala- 
Koop, 1 (bmnh) were not located exactly. GUA- 
TEMALA: Peten: Chuntuqui, 2 (usnm); Laguna 
de Sotz [= Zotz], 2 (usnm). BELIZE: Belize: Kate's 
Lagoon, 2 (bmnh); Rockstone Pond, 1 1 (rom). Or- 
ange Walk: Honey Camp Lagoon, 2 (fmnh). 

Additional Records (Hatt et al., 1953, p. 64, 
unless noted otherwise)— MEXICO: Yucatan: Ac- 
tun Chacaljas, 3 km SSW Calcehtok (cave de- 
posit); Actun Coyok [= Coyoc], 3.5 km SSE 
Oxkutzcab (cave deposit); Actun Has, 3.5 km WSW 
Yokat (cave deposit); Actun Lara, 3 km SW Yokat 
(cave deposit); Actun Oxkintok, 3 km SW Santa 
Cruz (cave deposit); Actun Spukil, 4.5 km SSW 
Calcehtok (cave deposit); Loltun, 5 km SW 
Oxkutzcab (cave deposit); Xbac (Gaumer, 1917, 
p. 13); Yaxcach (Gaumer, 1917, p. 13). GUA- 
TEMALA: Peten: 1 1 km NE JHores, 3.2 km inland 
SE shore Laguna Peten Itza, ca. 100 m (Ryan, 
1960, p. 11). 



ENGSTROM ET AL.: HETEROMYS GAUMERI 



303 



Species Groups of Spiny Rats, 

Genus Proechimys (Rodentia: Echimyidae) 



James L. Patton 



ABSTRACTS 

Nine species groups of the spiny rat subgenus Proechimys, family Echimyidae, are defined 
on the basis of bacular characters and qualitative features of the cranium. The latter include 
the structure of the incisive and mesopterygoid foramina, temporal ridge and infraorbital canal 
development, and counterfold pattern of the cheekteeth. 

Three groups are apparently monotypic, including the decumanus -group of southwestern 
Ecuador and adjacent Peru, the canicollis-group of northeastern Colombia and adjacent Ven- 
ezuela, and the s i mo nsi -group of the western Amazon Basin from Colombia to northern Bolivia. 
The remaining six groups are polytypic, but the number of species in each remains unclear. 
The semispinosus-group ranges from Central America south to southwestern Ecuador in the 
Pacific lowlands; its only Amazonian representative is P. oconnelli from east-central Colombia. 
The longicaudatus-group ranges from southeastern Colombia through the western Amazon 
Basin into the northern Parana Basin of Brazil and northern Paraguay. The goeldii-group ranges 
throughout the Amazon Basin from eastern Peru to eastern Brazil. The guyannensis-group 
occurs from the coastal Guianan region through the Rio Negro and eastern half of the Amazon 
Basin in Brazil, with an isolate in Goias and Minas Gerais states. The cuvieri-group has a 
similar distribution, but extends further up the Amazon into northern Peru, with one isolate 
in east-central Peru. Finally, the trinitatus-gcoup is found from north-central Colombia eastward 
across northern Venezuela to Trinidad. 



Nueve grupos de especies de la rata espinosa subgenero Proechimys, familia Echimyidae, son 
definidos primariamente en las bases de caracteres baculares que son soportados por razgos 
cualitativos del craneo. El ultimo incluye la estructura del foramen incisivo y mesopterigoideo, 
arista temporal y desarroUo del canal infraorbital y patron de contraplegamiento de los dientes 
postcaninos. 

Tres grupos son aparentemente monotipicos, incluyendo el grupo decumanus del suroccidente 
del Ecuador y adyacente Peru, el grupo canicollis del nororiente de Colombia y adyacente 
Venezuela, y el grupo simonsi del occidente de la cuenca amazonica desde Colombia hasta el 
norte de Bolivia. Los seis grupos remanentes son politipicos, pero el numero de especies en 
cada uno p>ermanece obscuro. El grupo semispinosus se extiende desde el sur de Centroamerica 
hasta el suroccidente ecuatoriano en las tierras bajas del Pacifico; su solo representante de la 
cuenca amazonica el P. oconnelli del centroriente colombiano. El grupo longicaudatus se dis- 
tribuye desde el suroriente de Colombia a traves del occidente de la cuenca amazonica hasta 
el norte de la cuenca del Parana en Brazil y norte de Paraguay. El grupo goeldii se distribuye 
a traves de la cuenca amazonica desde el Peru oriental hasta el Brazil oriental. El grupo 
guyannensis ocurre desde la region costera guayanesa a traves de rio Negro y la mitad oriental 
de la cuenca amazonica en Brazil, con un aislamiento de poblaciones en los estados de Goias 



From Museum of Vertebrate 2^ology, University of 
California, Berkeley, CA 94720. 



PATTON: SPECIES GROUPS OF PROECHIMYS 305 



y Minas Gerais. El grupo cuvieri tiene una distribucion similar, pero se extiende mas arriba del 
Amazonas en el interior del norte del Peru, con un aislamienlo de poblaciones en el centroriente 
peruano. Finalmente, el grupo trinitatus es encontrado desde el centronorte de Colombia hacia 
el oriente a traves del norte de Venezuela hasta Trinidad. 



Nove grupos de especies de ratos-de-espinho, do subgenero Proechimys, familia Echimyidae, 
sao definidos principalmente na base de carateres baculares que concordam tambem com 
caracteres qualitativos do cranio. Estes incluem: a estrutura dos incisivos e do forame mesop- 
terigoideo, o desenvolvimento das temporas e do canal infraorbital, e o padrao dos molares. 

Tres grupos sao aparentemente monotipicos, incluindo o grupo decumanus do sudoeste do 
Equador, o grupo canicollis do nordeste da Colombia e das areas adjacentes na Venezuela, e o 
grupo simonsi da Bacia Amazonica ocidental, desde a Colombia ate o norte da Bolivia. Os seis 
grupos restantes sao politipicos, mas o numero de especies em cada continua incerto. O grupo 
semispinosus estende-se da America Central ao sudoeste do Equador, nas planicies do Pacifico. 
Seu unico representante na Bacia Amazonica e P. oconnelli, do Centro-leste da Colombia. O 
grupo longicaudatus estende-se do sudeste da Colombia, atraves da Amazonia ocidental, ate o 
norte da Bacia do Parana no Brasil e no norte do Paraguai. O grupo goeldii ocorre na Bacia 
Amazonica, do leste do Peru ao leste do Brasil. O grupo guyannensis ocorre da costa guianense, 
ate o Rio Negro e a regiao oriental da Bacia Amazonica no Brasil, com uma especie isolada 
nos estados de Goias e de Minas Gerais. A distribuifao do grupo cuvieri e parecida, mas este 
ocorre tambem ate o norte do Peru, com uma especie isolada no centro-leste do Pais. Por final, 
o grupo trinitatus e encontrado do centro-norte da Colombia, atraves da Venezuela, ate Trin- 
idade. 



Introduction 

Spiny rats of the genus Proechimys represent one 
of the most diverse groups of Neotropical rodents; 
with the possible exception of tuco-tucos, Cteno- 
mys, the number of taxa of spiny rats is unmatched 
by any other caviomorph (Woods, 1984). The ge- 
nus extends throughout lowland forests from Nic- 
aragua to northern Paraguay and the coastal re- 
gions of Brazil. Despite this diversity and large 
geographic range, however, the group is taxonom- 
ically one of the most poorly understood among 
all of the Neotropical mammals. Only a few stud- 
ies have succeeded in recognizing the number of 
taxa sympatric at any single locality (e.g., Moojen, 
1948; Patton & Gardner, 1972), and no study has 
been able to follow geographic character trends 
within a clearly defined taxon over any but the 
shortest distances. Diagnosis of species and hence 
definition of natural units in Proechimys have been 
severely hampered by the often extreme level of 
variability within and between populations for 
most morphological characters that have been ex- 
amined (Moojen, 1948; Hershkovitz, 1948; Pat- 
ton &. Gardner, 1972). Even karyotypes, which 
have proven useful in differentiating sympatric taxa 
of spiny rats (Patton & Gardner, 1972), are often 



highly variable geographically (Reig & Useche, 
1976;Reigetal., 1 980; Gardner & Emmons, W84). 

In this report I will challen