Table Of ContentVampire bat, Domodus rotundu.1 (Photo taken by Uwe Schmidt )
Natural History
of
Vampire Bats
Editors
Arthur M. Greenhall
Office of Scientific Authority
U.S. Fish and Wildlife Service (retired)
Washington, D.C.
and
Research Associate
American Museum of Natural History
New York, New York
and
Uwe Schmidt
Zoological Institute
University of Bonn
West Germany
Boca Raton London New York
CRC PressC isR anC im Pprrinets osf, thIenc.
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First published 1988 by CRC Press
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Library of Congress Cataloging-in-Publication Data
Natural history of vampire bats / editors, Arthur M. Greenhall and Uwe Schmidt.
p. cm.
Bibliography: p.
Includes index.
ISBN 0-8493-6750-6
1. Vampire bats. I. Greenhall, Arthur Merwin, 1911-II. Schmidt, Uwe. Dr.
QL737.C52N38 1988
599.4--dc19 87-24622
A Library of Congress record exists under LC control number: 87024622
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PREFACE
Although vampire bats have been known by the inhabitants of the Americas for thousands
of years, it is Jess than 150 years since these bats were scientifically identified. Prior to the
1930s, when the vampire bats became a veterinary public health problem in Panama and
Trinidad, knowledge of these bats was only fragmentary. During the past 50 years however,
when funds became available to study vampires, a large amount of knowledge accumulated
demonstrating that vampire bats are not only unique among bats because of their diet but
also because of their unusual morphological, physiological, and behavioral adaptations di-
rected toward obtaining their food. Most of this information concerns the common vampire
bat, Desmodus rotundus, and much is directed toward their control. The white-winged
vampire bat, Diaemus youngi, and the hairy-legged vampire bat, Diphylla ecaudata, have
existed largely uninvestigated as they have played no major role in the transmission of
disease. When some measure of control and viable vaccines were obtained, funds for vampire
bat study diminished and have now all but disappeared. As this book was being written it
became quite clear that much more information about the three vampire species will unfold
when general interest is stimulated and funding again becomes available.
A major problem with these bats is that whatever information exists is scattered throughout
the literature or is not recorded. In 1982, CRC Press, Inc., approached Arthur Greenhall to
write or edit a book on bats. Since there are some excellent books on the ecology and
biology of bats with very little on vampire bats, it seemed appropriate to publish what is
known about them in one volume. Since we had worked together harmoniously in Mexico
on a United Nations project investigating paralytic rabies and vampire bats, it was natural
to invite Uwe Schmidt to be co-editor. Every biologist we asked to contribute a chapter was
enthusiastic about the value of such a book that would provide an in-depth presentation of
these unique animals. That the chapters were completed expeditiously emphasized the whole-
hearted support of the authors. We are deeply grateful to all the authors for their contributions,
time and ideas.
The community of bat biologists was particularly saddened by the death of William A.
Wimsatt of Cornell University on January 9, 1985. Although extremely ill, he had consented
to write the chapter on reproduction. When he realized he would not be able to complete
the chapter, he requested that Christel Schmidt assume responsibilty for this subject.
Now there is a book which encompasses the different aspects of the natural history of the
vampire bats: their biology, economic importance, management, and folklore, and it also
indicates the gaps in our knowledge.
Arthur M. Greenhall
Uwe Schmidt
EXORDIUM
My interest in vampire bats began in the summer of 1933, when I first saw a live Des modus
rotundus at the Gorgas Memorial Laboratory in Panama. At the time I was assistant to
Raymond L. Ditmars, Curator of Mammals and Reptiles at the New York Zoological Park,
who was with me in Panama collecting animals.
Herbert C. Clark and Lawrence H. Dunn were investigating these bats because they had
found vampire bats were transmitting a protozoan, Trypanosoma hippicum, to equines,
causing the disease "Murrina" that is fatal to equines, but not to bovines or humans.
We collected the first vampire bat ever exhibited alive (Figure 1). Back at the zoo, we
carefully observed our captive bat using its long thumb as a foot for quadrupedal locomotion
and saw that it did not suck blood as had long been supposed, but appeared to lap it. We
searched the literature, but found nothing regarding vampire bats. We recorded the bat's
nimble walking on motion picture film, but felt that it was important to substantiate our
observations with additional live Desmodus.
The following summer (1934) I arrived in Trinidad at the height of a paralytic rabies
outbreak that had been fatal to thousands of livestock and a number of humans. The gov-
ernment team investigating the outbreak consisted of: 1. Lennox Pawan, Pathologist, who
had isolated rabies virus from Desmodus rotundus (Figure 2); Eric de Verteuil, in charge
of public health; H. V. Metivier, in charge of veterinary affairs; F. W. Urich, Zoologist,
and LudolfWehekind, his assistant. None of these scientists had heard that there was Murrina
in Panama.
In Pawan's laboratory I prepared bat brains to be examined for Negri bodies. With de
Verteuil, in the Colonial Hospital, I spoke to patients who had been bitten by vampire bats,
including two patients who later died. With Metivier I saw stricken cattle in the field as
well as experiments with bats and cattle at the Government farm in North Central Trinidad.
I went into the field to collect vampire bats with Urich and Wehekind, who also allowed
me to observe and photograph their captive vampire and "blood donor" goat (Figure 3).
Urich told me about the habits of Desmodus rufus (as it was then called) while Wehekind
regaled me with tales of Trinidad's colorful folklore creatures: "Soucouyant" that sucks
human blood, "Dwens" babies who died without baptism, whose feet tum backwards,
"Papa Bois" who watches over the wild animals in the forest and "Mama d'Leau", his
consort, who watches over the aquatic creatures.
We returned to the Bronx Zoo with more live vampire bats. More bats, in fact, than the
zoo thought it needed for display, so for two years I kept 20 live Desmodus in my New
York City apartment.
In 1935 Ditmars and I published our paper ''The Vampire Bat'' Zoologica 191, 53, 1935).
Almost 20 years later, through the kind offices of Ludolf Wehekind, I was able to return
with my family to live in Trinidad. A large outbreak of paralytic rabies occurred soon after
we arrived. Aurelio Malaga-Alba was sent to Trinidad by the Pan American Health Organ-
ization to advise the Government on the outbreak.
Malaga (a most dynamic, energetic investigator) and I collected the white-winged vampire
bat, Diaemus youngi, which until then was known only from 17 skins reported by Colin C.
Sanborn of Chicago's Field Museum of Natural History.
Although ill at the time, Pawan allowed us to house and observe the Jive Diaemus in his
laboratory. There we discovered the oral glands whose purpose remains a mystery. Later
that year, Victor C. Quesnel, microbiologist, made the extraction tests mentioned elsewhere
in this book.
On Malaga's recommendation, the Trinidad Government appointed me Zoologist in charge
of the Anti-Rabies Bat Collectors. When I notified Sanborn about capturing live Diaemus,
he immediately came to Trinidad to collect bats, including more Diaemus. We decided on
FIGURE 1. Raymond L. D1trnan, and the fm.t vamp1re bat, Desmodus ro-
tundus ever exhlbtted alive (Photo taken by Nma Leen. Wtth perrmsston )
a collaborative study on the bats of Trinidad and Tobago. I worked up some of the material
at the Field Museum later that year.
After Sanborn's untimely death, Harold E. Anthony, American Museum of Natural His-
tory, came to Trinidad and suggested that George G. Goodwin, at the American Museum,
continue the studies started with Sanborn. In 1961 we jointly published "A Review of the
Bats of Trinidad and Tobago" which appeared in the Museum's Bulletin 122.
The Trinidad Regional Virus Laboratory was established by the Rockefeller Foundation
in 1953, directed by Wilbur G. Downs, and the following year I was appointed Consultant
Zoologist with laboratory facilities. The Laboratory was primarily concerned with arthropod-
borne viruses, whereas rabies investigation was the purview of the Trinidad Government.
The entire staff of the laboratory was most helpful, but I am especially grateful to those
who shared their virological expertise: W. G. Downs, T. H. G. Aitken, R. R. Anderson,
D. B. Worth, L. Spence, 0. R. Causey, and E. S. Tikasingh. Later, a Rockefeller grant
made it possible for me to meet rabies experts H. N. Johnson, K. Habel and Denny G.
Constantine. At that time, W. L. Jennings was investigating bats in Florida, since the first
rabid bat in the U.S. had been reported in 1953 from Florida, and both of us investigated
the epizootic of rabies in raccoons there.
The isolation of Histoplasma capsulatum in Trinidad brought me in contact with C. W.
Emmons, National Institutes of Health, and L. Ajello, Centers for Disease Control; the
former came to Trinidad to investigate this mycotic disease.
A number of mammalogists came to Trinidad (while we were living there) to observe
OFFICIAL FIRST DAY COVER
FIOlJRE 2. J. Lennox Pawan who isolated rabies virus from Desmodus rotundus tn Trimdad (PermissiOn granted
by the Government of Trinidad and Tobago to reproduce FlfSt Day Cover.)
Trinidad bats. These included: Karl F. Koopman, R. Van Gelder, S. Anderson, P. Her-
skovitz, and R. Peterson.
There were many Trinidadians (in addition to those mentioned earlier) who contributed
to my understanding of vampire bats and paralytic rabies. These include Oswald Siung, who
followed Pawan as Government Pathologist; H. P. Gillette, Chief Medical Officer; Leonard
Charles, Franklin Thurab, a meticulous worker who had been Pawan's laboratory assistant
and then Siung's (eventually Thurab took charge of my laboratory), and Vernon Mahabir-
singh, an excellent field man.
During our 10 years in Trinidad, my substantive position was as Curator of the Royal
Victoria Institute Museum (now the National Museum). 1. R. Bullbrook, my Assistant
Curator, was a geologist and a distinguished archeologist with an encyclopedic knowledge
of West Indian history. L. Wehekind (mentioned earlier) became Associate Curator. J.
Kenny, Fisheries Officer for the Trinidad Government (now Head, Department of Zoology,
University of the West Indies, St. Augustin) conducted numerous bat studies and observations
with me.
Back in the U.S. in 1963, I accepted the position of Chief of the Mammal Section, U.S.
Fish and Wildlife Service's laboratory in the National Museum of Natural History, Smith-
sonian Institution. John L. Paradiso and I shared the burden of work, and we jointly produced
the service's "Bats and Bat Banding'' in 1968 as the Service's Resource Publication 72.
One memorable week was spent with G. Mann, visiting from Chile, who described his
observations of the feeding mechanism of Desmodus rotundus as well as his experience with
Desmodus feeding upon sea lions off the coast of Chile.
The vampire bat problem continued to escalate in Latin America. Aurelio Malaga-Alba,
on behalf of the Mexican Government, requested assistance from the United Nations De-
velopment Program in 1966, as did a number of other countries in Latin America. The
World Health Organization, Pan American Health Organization, and Food and Agriculture
Organization jointly sent a Preparatory Assistance Mission to investigate the paralytic rabies
situation in each of the countries requesting assistance to determine which one would be
most suitable for a 4-year project. As ecologist with this Mission I met many people, among
FIGURE 3. Desmodu> rotundus feedwg on a goat. (Photo taken by Arthur M
Greenhall; m Annual Report of the Board of Regents of the Snnth,on1an Imt!tutlon,
U.S. Government Pnntmg Off1ce, Wa,hmgton. 1937, 277)
them; J. A. Crespo of Argentina, Edgardo Mondolfi of Venezuela, and Bernardo Villa-R.
of Mexico (an old friend). The Mission recommended that the UNDP/FAO Project be placed
in Mexico.
In 1968, I was seconded from the U.S. Fish and Wildlife Service to become the Ecologist
with tht\ VNDP/FAO Research Project on Paralytic Rabies together with Pierre Sureau of
the Pasteur lfi!ltitute as Project Manager and Epizootiologist, and Gosse Bijlenga as Virologist.
ln. 1969, Vwe Schmidt was appointed my Associate Expert and William Lopez-Forment
was selected as Mexican Associate Expert, since he had conducted much field work with
Aurelio MaJaga-Alba and Bernardo Villa. At the end of one year Schmidt returned to West
Germany, taking with him 52 live Desmodus, some of which are still thriving in his laboratory
with the offspring of the original bats, continuing to reproduce.
I am tnost grateful that Uwe Schmidt accepted my invitation to co-edit "The Natural
History of Vampire Bats".
This volume was cmnpleted on non-official time and with personal resources.
Arthur M. Greenhall
THE EDITORS
Arthur M. Greenhall, B.A., M.S., F.L.S. Zoologist (Retired), Office of Scientific
Authority, U.S. Fish and Wildlife Service, Washington, D.C. was graduated from the
University of Michigan with a B.A. in 1934 and a M.S. in 1935. His studies with vampire
bats preceded his graduate work. In Panama in 1932 he found that Desmodus rotundus
transmitted a disease to equines, as well as attacking people. He collected the first vampire
bat ever to be publicly exhibited alive and observed unrecorded habits of the bat. To confirm
these observations, he went to Trinidad in 1934 and found that Desmodus had recently been
found to be the transmitter of paralytic rabies which had killed thousands of cattle and 89
humans. As a result of previous studies in Panama, the Trinidad Government invited him
to be a member of the Commission studying paralytic rabies. His master's dissertation was
the first major study of the vampire bat and included comments about the supernatural
vampire.
Graduate studies at Columbia University and New York University toward his Ph.D. were
intem1pted by World War II. He became the Director of the Portland Zoo in 1942, and then
in 1947 became the General Curator of the Detroit Zoo and Aquarium. After an interval of
12 years he was invited to return to Trinidad, and in 1953 became the Curator of the Royal
Victoria Institute Museum, Director of the Emperor Valley Zoo, and Consultant to the
Trinidad Regional Virus Laboratory of the Rockefeller Foundation. In 1954 another vampire
bat paralytic rabies outbreak occurred, and the Government appointed Greenhall to become
the Zoologist in charge of the Anti-Rabies Programme. He continued until 1963, when he
returned to the United States to the U.S. Fish and Wildlife Service.
In 1963 he became Chief, Mammal Section, Bird and Mammal Laboratories, National
Museum of Natural History, Washington, D.C. In 1968 he was seconded to the United
Nations to become Animal Health Officer (Bat Ecologist), Food and Agriculture Organization
(FAO) Research on Paralytic Rabies Project in Mexico. In 1972 he returned to Washington
as Zoologist, National Fish and Wildlife Laboratory at the National museum of Natural
History (Smithsonian Institution). In 1977 he became Staff Zoologist, Office of Scientific
Authority for U.S. Convention on International Trade in Endangered Species of Wild Fauna
and Flora (CITES).
Various awards and honors include Research Associate, American Museum of Natural
History; Research Associate, National Museum of Natural History (Smithsonian Institution);
Fellow, Linnean Society of London (F.L.S.); Honorary Member, Phi Sigma, Chi Gamma
Phi, Beta Beta Beta; Honorary Consultant, International Union for the Conservation of
Nature and Natural Resources (IUCN)/Survival Service Commission (SSC); Member of the
SSC Chiroptera Specialist Group; Special Achievement Award, U.S. Fish and Wildlife
Service.
Greenhall has published over 200 papers and articles, both technical and popular. The
most significant have been the monographic "Review of the Bats of Trinidad and Tobago,"
"Bats and Bat Banding," "House Bat Management," "Key to the Bats of Argentina,"
"Effects of Vampire Bats and Paralytic Rabies on Livestock Production."
His current research and consultantships include the management and control of vampire
bats and paralytic rabies, control of house bats, neotropical mammals, conservation of
endangered animals, establishment of national parks and wildlife sanctuaries, and interpre-
tation of the Convention of International Trade in Endangered Species of Wild Fauna and
Flora (CITES).
