U.S. Department of the Interior

U.S. Geological Survey

Circular 1329

Prepared by the USGS National Wildlife Health Center

Bat Rabies and 

Other Lyssavirus Infections
Front cover photo (D.G. Constantine)

A Townsend’s big-eared bat.
Bat Rabies and

Other Lyssavirus Infections

By Denny G. Constantine

Edited by David S. Blehert

Circular 1329

U.S. Department of the Interior

U.S. Geological Survey
U.S. Department of the Interior

KEN SALAZAR, Secretary

U.S. Geological Survey

Suzette M. Kimball, Acting Director

U.S. Geological Survey, Reston, Virginia: 2009

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Suggested citation:

Constantine, D.G., 2009, Bat rabies and other lyssavirus infections: Reston, Va., U.S. Geological Survey Circular 1329, 

68 p.

Library of Congress Cataloging-in-Publication Data

Constantine, Denny G., 1925–

Bat rabies and other lyssavirus infections / by Denny G. Constantine.

 p. cm. - - (Geological circular ; 1329)

ISBN 978–1–4113–2259–2

1. Rabies. 2. Rabies in animals. 3. Bats as carriers of disease. 4. Bats -- Diseases.

I. Title.

RA644.R3C665 2009

616.9’53 - - dc22

 2008038349

Photo on previous page:

A pallid bat in flight. (D.G. Constantine)
Prepared by the USGS National Wildlife Health Center 

in cooperation with the U.S. Fish and Wildlife Service

Disease Emergence and Resurgence: 

The Wildlife-Human Connection

U.S. Department of the Interior

U.S. Geological Survey

Circular 1285

Disease Emergence and Resurgence:

The Wildlife-Human Connection

Prepared by the USGS National Wildlife Health Center

in cooperation with the U.S. Fish and Wildlife Service

Circular 1285

U.S. Department of the Interior

U.S. Geological Survey

Field Manual of

Wildlife Diseases

General Field Procedures and Diseases of Birds

U.S. Department of the Interior

U.S. Geological Survey

Prepared by the USGS National Wildlife Health Center 

in cooperation with the U.S. Fish and Wildlife Service

Tularemia

U.S. Department of the Interior

U.S. Geological Survey

Circular 1297

Prepared by the USGS National Wildlife Health Center 

in cooperation with the U.S. Fish and Wildlife Service

Circular 1297

U.S. Department of the Interior

U.S. Geological Survey

Tularemia

Related USGS Publications

Disease Emergence and Resurgence—

The Wildlife-Human Connection,

by Milton Friend, USGS Circular 1285

(also available on CD-ROM)

Tularemia, by Milton Friend, 

USGS Circular 1297

(also available on CD-ROM)

Field Manual of Wildlife Diseases—

General Field Procedures and Diseases

of Birds, edited by Milton Friend and 

J. Christian Franson, USGS Information 

and Technology Report 1999–001

v

Foreword

In many ways, bats are the nocturnal equivalent of birds (Kunz, 1982). Like birds, they consume 

large numbers of insects that cause harm to agriculture and to humans; researchers estimate 

that the value of bat control of cotton crop pests in eight counties in south-central Texas ranges 

from $121,000 to $1,725,000 per year (Cleveland and others, 2006). Bats also play an important role as pollinators and in spreading plant seeds by eating the fruits of some plants. The 

estimated 1,100 species of bats worldwide comprise about 20 percent of all mammal species 

(Tudge, 2000), are second only to rodents in number of species, and are likely the most numerous in total individuals of any type of mammal (Kunz 1982). Despite their important ecological 

roles and benefits to humans, bats are among the most maligned and mysterious animal group 

in existence, largely due to folklore and unfamiliarity with the true nature of these primarily 

nonhematophagous, or nonblood feeding, species (King, 1993).

The notoriety associated with rabies in vampire bats of the tropical and subtropical regions of 

the New World is unfortunately transposed by many to also reflect the demeanor and consequences of the presence of other types of bats. Clearly, vampire bat rabies was a prominent 

challenge for early Spanish colonists and their domestic animals during their settlement within 

Latin America (Baer, 1975a; Lopez and others, 1992). Because of vampire bat rabies, the presence of bats in urban environments of the Northern Hemisphere commonly elicits fear among 

many adults, but wonderment among the young and the young-at-heart. The former tend to 

associate the presence of bats with rabies, while the latter see only the uniqueness of these 

highly specialized flying mammals and often place themselves at risk by attempting to capture 

and handle these animals for closer examination.

Rabies in a nonhematophagous North American bat was not documented until 1953, when a 

yellow bat (Dasypterus floridanus, now Lasiurus intermedius, the Northern yellow bat) was 

examined following its aggressive biting of a 7-year-old boy in Florida. Fortunately, despite no 

previous documentation of rabies in insectivorous bats in the United States, the child received 

prompt postexposure prophylactic treatment for rabies and did not become clinically ill (Venters 

and others, 1954, Scatterday, 1954). Subsequent investigations resulted in retrospective 

diagnoses of bat rabies in the USA in 1951 and, possibly, even 30 years earlier, suggesting 

the presence of rabies in insectivorous bats long before its recognition (Baer, 1975b). During 

recent decades, an average of 700 to 800 cases of rabid insectivorous bats has been diagnosed 

annually in the USA (Rupprecht and others, 1995). During 2006, 1,692 cases of rabies were 

reported in bats, a 14.5 percent increase over 2005 and 24.4 percent of the total animal rabies 

cases. Only raccoons accounted for a greater number of rabies cases in the USA during 2006 

(Blanton and others, 2007).

Although the annual number of human rabies deaths in the USA remains very small, most of 

those deaths are attributed to unrecognized exposures to rabies associated with bats, primarily the silver-haired bat and the tricolored bat, two infrequently encountered bat species. 

The ecological relationships driving such outcomes extend beyond evolutionary increases 

in viral infectivity reported for these bat species (Messenger and others, 2003). Equally, if 

not more important, are factors that result in human contact with these species. Humaninduced landscape changes influence the distribution and abundance of animal populations, and these types of changes have had a major role in the ascension of wildlife rabies 
vi

(Rupprecht and others, 1995) and likely will continue to do so. Bat-associated human disease 

emergence during recent years includes Nipah, Marburg, “Melaka,” and Hendra viruses, and 

human Nipah virus infections clearly result from landscape changes that cause the bats to 

relocate their roosts to sites where interspecies disease transmission is facilitated (Daniels 

and others, 2007). That outcome emphasizes the importance of understanding bat ecology as a 

component of disease ecology.

Knowledge of bat ecology is especially important when humans and bats share environments. 

This information is needed to properly educate people about bats and for the application of preemptive actions, including education, for minimizing disease risks that may be present. Simple, 

inexpensive means for bat exclusion noted in the text can be installed to permit bats to depart 

from but prevent re-entry into buildings. This text also stresses the importance of house cats in 

the transmission of bat rabies, a situation that can largely be controlled by preemptive actions 

identified by the author and that many readers may apply to their daily lives with their pet cats.

Of special importance is the potential for contact between children and bats. Young people are 

often at higher risk for exposure to rabid bats because their uninhibited curiosity may result in 

their handling of moribund and injured bats, including those that may have been captured by the 

family cat. This writer had several personal experiences evaluating bats that were picked up by 

students, brought to grade school biology classes, and that proved to be rabid. Fortunately, no 

human rabies cases resulted from those events.

Education about bats provides valuable balance so that disease risks are not overstated, 

fostering fear of these species, or understated to the extent that people are placed at undue 

risk. Personal experiences have indicated that private sector presentations by those interested 

in bat conservation often understate disease considerations and demonstrate risky behaviors, 

such as handling bats without protective gloves, in efforts to overcome negative perspectives of 

bats advanced by others. These same types of indiscretions also appear in various publications 

(King, 1993).

The best defense against potential disease risks posed by bats, or any animal species, is a 

sound understanding of disease ecology and the wise application of that knowledge. Through 

this publication, Dr. Constantine strives to provide the reader with these attributes by sharing 

over a half-century of personal experiences and scientific investigations of bat rabies ecology. 

During his career, Dr. Constantine has significantly contributed to the scientific knowledge of 

this complex disease and challenged existing dogma in ways that have enhanced our current 

understanding of rabies transmission (see literature citations associated with the text). This 

unique and highly informative presentation incorporates anecdotes as well as hard science to 

provide both nonscientists and specialists with numerous points to ponder and a better understanding of bats as part of our biological world. The presentation is enhanced by the many 

illustrations that portray diversity among bat species, some of their adaptive features, and other 

aspects of bat biology. Bats have successfully colonized most regions of earth except for Antarctica and some other treeless northern areas of extreme cold (Kunz, 1982). These species need 

to be conserved for their ecological importance and for the benefits that they provide humans, 

rather than feared because of disease.

Milt Friend

Emeritus Scientist

USGS–National Wildlife Health Center
vii

Acknowledgments

This publication is due in large part to the help and cooperation provided the author by a great 

variety of workers during a lifetime of his investigations. Those individuals worked in fields 

such as public health, wildlife health, wildlife ecology and conservation, animal control, and 

also included landowners and land managers. Space does not allow detailing their individual 

contributions or listing all of them. Instead, I thank the multitude of those who have assisted my 

efforts and note those field companions who have made relevant contributions recalled during 

the development of this report. I have greatly enjoyed probing the secrets of nature with all of 

these individuals, some of whom are listed below by geographic area of shared exploration.

Africa: Dr. Lars Karstad.

Alaska: Dr. Robert L. Rausch, Dr. Raymond J. Hock.

California: Dr. Thomas J. Cade, Dr. William R. Dawson, Harry J. Fletcher, Christopher Henne, 

Gerry P. Mulcahy, Dr. Kenneth E. Stager, Dr. Joseph M. Szewczak, Raymond Williams, and 

Dr. Sherwin F. Wood.

California and Australia: Dr. Phillip Leitner, Dr. Elizabeth D. Pierson, and Dr. William E. Rainey.

Southwestern US, Mexico, and/or Central America: Devil B. Adams, Michael Balizan, 

Dr. Raúl Flores Crespo, Donald M. Goede, Dr. Louis N. Locke, Dr. G. Clay Mitchell, 

Richard W. Russell, Dr. Karl M. Schmidt, Dr. Bernardo Villa-R., Elliott J. Wescott, Jr., 

Loran M. Whitelock, and John R. Williams.

Trinidad, West Indies: Dr. Leonard V. Butcher, Dr. Vincent G. Moe, and Dr. Farouk Muradali.

Venezuela: Dr. Rexford D. Lord.

Brazil: Elisa Band, Dr. Nistan Fidalgo Peres, Dr. Takashi Fujii, and Dr. Otávio Diniz.

Thailand: Dr. David E. Davidson, Jr., Dr. Richard Luce, Dr. Boonlert Lumbertdacha, 

Dr. Charles E. Rupprecht, and Dr. Chakrapol Sriaroon.

Japan: Dr. Tatsuo Udagawa. 

Worldwide: Kenneth G. Constantine

I also thank Drs. David Blehert and Milton Friend for their great assistance in the organization 

and development of this publication.

ix

Contents

Foreword ....................................................................................................................................................... v

Acknowledgments..................................................................................................................................... vii

Overview........................................................................................................................................................ 1

Background................................................................................................................................................... 1

Causative Agent........................................................................................................................................... 2

Geographic Distribution.............................................................................................................................. 5

Patterns and Trends .................................................................................................................................... 5

Species Susceptibility................................................................................................................................. 9

Human Infections.............................................................................................................................. 10

Bat Infections .................................................................................................................................... 14

Infections in Other Species............................................................................................................. 18

Obtaining a Diagnosis ............................................................................................................................... 19

Disease Ecology......................................................................................................................................... 19

Bat Activity and Disease Patterns ................................................................................................. 19

Temperature and Rabies Virus Virulence ..................................................................................... 22

How Colonial Bats Avoid Extinction by Lyssaviruses ................................................................. 22

Paralytic and Furious Disease Courses ....................................................................................... 22

Viral and Host Species Diversity .................................................................................................... 25

Intraspecies and Interspecies Transmission of Bat Lyssaviral Variants................................. 25

Latin America............................................................................................................................ 27

North America.......................................................................................................................... 27

Europe ....................................................................................................................................... 28

Africa ....................................................................................................................................... 29

Asia ....................................................................................................................................... 29

Australia .................................................................................................................................... 29

Environmental Persistence ............................................................................................................. 30

Points to Ponder: Interspecies Interactions in Potential Bat Rabies Transmission Settings ....... 32

Bat Colonies in Manmade Structures ........................................................................................... 32

Bat Colonies in Caves and Mine Tunnels...................................................................................... 37

Flying Bats that Enter Buildings at Night ...................................................................................... 37

Unprovoked Attacks by Flying Rabid Bats.................................................................................... 38

Translocation of Bats ....................................................................................................................... 38

Vampire Bats and Rabies ................................................................................................................ 38

Attacks on People.................................................................................................................... 42

Attacks on Livestock ............................................................................................................... 42

Attacks on Domestic Dogs and Cats .................................................................................... 42

Attacks on Other Bats............................................................................................................. 43

Disease Prevention and Control.............................................................................................................. 43

Humans............................................................................................................................................... 43

Wildlife Rehabilitators............................................................................................................. 43

Cave Explorers.......................................................................................................................... 44

Animals............................................................................................................................................... 45

References Cited........................................................................................................................................ 45

Glossary....................................................................................................................................................... 53
x

Topic Highlight Boxes

1. Vampires, Werewolves, and Settlement of the Americas ................................................... 3

2. The Course of Bat Rabies in Humans.................................................................................... 11

3. Human Survival of Clinical Rabies—An Exception to the Rule......................................... 12

4. Seasonal Bat Behavior and Disease Transmission ............................................................ 16

5. Bats—Diverse Mammals with Diverse Diets ...................................................................... 20

6. Immunity to Lyssaviruses in Colonial Bats ........................................................................... 23

7. Bat Versus Bat—Interspecies Transmission of Bat Lyssaviruses................................... 28

8. Lyssaviruses—Deadly Hitchhikers........................................................................................ 31

9. Domestic Cats and Lyssavirus Transmission....................................................................... 34

10. The Tripartite Vampire Bat Clan ............................................................................................. 40

Illustrations

1. Unrooted phylogenetic tree based on complete lyssavirus N gene 

nucleotide sequences..................................................................................................................4

2–3. Maps showing:

2. Number of bat species reported with rabies infections within the Americas ..........6

3. Old World distribution of lyssavirus infections (excluding rabies virus)

reported in bat species and humans in Africa, Asia, Australia,

Europe, the Indian subcontinent, and the Middle East .................................................7

4–6. Graphs showing:

4. Number of rabid bats identified in the United States from 

1955 to 2006...........................................................................................................................8

5. Percentage of rabies-suspect bats identified as rabid in the United States 

from 1964 to 2006..................................................................................................................9

6. Number of rabies- and other lyssavirus-infected bat species by continent...........10

7–17. Photographs showing:

7. Circular pattern of earth excavation resulting from confined 

body movement associated with posterior paralysis in 

a rabies-infected Zebu......................................................................................................18

8. Townsend’s big-eared bats awakening from hibernation...........................................19

9. Brazilian free-tailed bats attempting to find roosting space 

among a dense colony of cohorts on a cave ceiling...................................................23

10. Canyon bat in flight............................................................................................................24

11. Western small-footed myotis bat in flight......................................................................24

12. California myotis bat in flight ...........................................................................................24

13. Big brown bat in flight.......................................................................................................24

14. A mixed group of pallid and Brazilian free-tailed bats in an attic .............................26

15. A Brazilian free-tailed bat and a Yuma myotis bat cooperating 

to conserve body heat ......................................................................................................26

16. A dead Brazilian free-tailed bat being consumed by dermestid 

beetles commonly found in association with bat colonies.........................................26

17. Brazilian free-tailed bat. ...................................................................................................26

18. Diagram showing factors involved in cyclic fluctuations and 

environmental persistence of bat populations and lyssaviral infections.................30
xi

19–26. Photographs showing:

19. An evening foraging flight of Brazilian free-tailed bats .............................................32

20. Brazilian free-tailed bats roosting under a railroad trestle........................................33

21. A dead Brazilian free-tailed bat lying on the guano-covered floor 

of a bat cave .......................................................................................................................33

22. Exclusion devices permit bats to depart from but prevent 

re-entering the interior of a building ..............................................................................36

23. Researcher among a Brazilian free-tailed bat colony.................................................37

24. Common vampire bat ........................................................................................................39

25. Application of anticoagulant jelly to the back of a vampire bat ................................39

26. Vampire bat-bitten burro ..................................................................................................39

Tables

1. Lyssavirus infections in mammals: agents, hosts, identification, 

and effectiveness of antirabies prophylaxis measures for humans ....................................2

2. General geographic area of rabies-related lyssavirus infections 

reported in bats.............................................................................................................................5

3. Bat families in the USA reported with bat rabies....................................................................9

4. Insectivorous bat origins of human rabies infections recognized 

in the USA and Canada from 1951 to 2006 ..............................................................................11

5. Human nonrabies lyssaviral infections reported in Africa, Asia, 

Australia, and Europe.................................................................................................................14

6. Examples of incubation periods and times to death relating to 

human mortalities from bat-associated lyssavirus exposure.............................................15

7. Examples of nonbat species infected by rabid bats .............................................................18

8. Examples of vampire bat rabies variants found in other species 

in Latin America ..........................................................................................................................27

9. Diseases other than rabies that may be transmitted by vampire bats ..............................42

10. Preemptive actions for minimizing human exposures to lyssaviruses..............................44

Appendixes

A. Bat Species Reported Infected with Rabies in the USA from 

1953 to 2007..................................................................................................................................57

B. Bat Species Reported Infected with Rabies in the Americas 

through 2006. ..............................................................................................................................58

C. Common and Scientific Names for Species Cited ................................................................61

D. Lyssavirus Infections in Bats Reported in Africa, Asia, Australia, 

and Europe...................................................................................................................................64

E. Course of Bat Lyssavirus Infections in Humans....................................................................66

Words in bold type in the text, the topic highlight boxes, and the tables are defined in the 

Glossary.
Top: A big brown bat drinking on the wing is reflected in the water below. (D.G. Constantine)

“What shall I do? What can I do? 

How can I escape from this 

dreadful thing of night and gloom 

and fear?”

(Stoker, 1897)

“He who shall eat the tongue

or the heart of a bat

shall flee from water and die.”

(Rhazes, 10th century Persian physician, quoted by Aldrovandi, 1599)
Bat Rabies and

Other Lyssavirus Infections

Synonyms

Rabia en murciélagos (Spanish)

Raiva em morcegos (Portuguese)

Rage de chauves-souris (French)

Tollwut bei fledermäusen (German)

Overview

Rabies constitutes a nearly global viral disease of 

mammals that affects the central nervous system, is usually transmitted by bite, and typically results in the death of 

the host. The causative agent is a member of the lyssavirus

group (Niezgoda and others, 2002). Each Lyssavirus species is divisible into strains called variants, each of which 

usually is found only in an individual reservoir host species 

of carnivore (Order Carnivora) or bat (Order Chiroptera), 

within which the virus multiplies (Table 1; Rupprecht and 

others, 2001). Humans and other species are infected through 

contact with a host animal. Bats are the focus for this synopsis (Box 1), and they are known to be infected throughout 

the Americas and in Europe, Asia, Africa, and Australia 

(Niezgoda and others, 2002). Both suborders of bats, the 

Megachiroptera or large fruit-eating bats of the Old World 

tropics, and the Microchiroptera, consisting of all other bats 

distributed globally to the limits of tree growth, are known 

to become infected with variants of lyssavirus. Bat species 

known to be rabies infected in the USA and the Americas are 

listed in Appendixes A and B, respectively. 

By Denny G. Constantine

Background

The origin of rabies, as of many other infectious diseases, is a matter of speculation. It has been postulated that 

rabies viruses may have originated from an insect virus 

that infected insect-eating bats some 7,000 to 12,000 years 

ago (Badrane and Tordo, 2001). Three such insect viruses, 

proposed for inclusion in the Lyssavirus genus, are known 

today. Moreover, data and calculations have been presented 

to support the view that after rabies viruses evolved in bats 

between 900 to 1,500 years ago, the infection subsequently 

spilled over into carnivorous mammals (Badrane and Tordo, 

2001). Bats and carnivores are essentially the only natural or 

persistent reservoir hosts of rabies known today, and individuals of other mammal species usually die without spreading 

the infection.

North American bats north of Mexico, nearly all of them 

insectivorous, were ignored as potential rabies hosts until 

1953 when the father of a bitten child, aware of the infection 

spread by vampire bats, insisted that the biting bat be tested. 

After the bat was determined to be rabid (Venters and others, 

1954), surveys were conducted until the virus was found 

throughout the continent in all bat species that were extensively tested. Whereas only 0.1 percent of seemingly healthy 

bats were infected, 10 percent of bats that were rabies-suspect, because they were found ill or were dead, were positive 

for rabies infection (Constantine, 1988a).
2 Bat Rabies and Other Lyssavirus Infections

Findings in North America soon stimulated surveys 

throughout the world. These surveys focused on sampling 

colonies of healthy bats, and they generally yielded negative 

results, such as those from North America. However, the testing of symptomatic bats eventually produced rabies viruses 

that differ from typical carnivore rabies isolates (Hentschke 

and Hellman, 1975; Mohr, 1957; Pitzschke, 1965). Interest 

in bat rabies has varied with time. For example, European 

bat rabies was identified in the laboratory only 14 times over 

30 years. Then, in 1985, three European human rabies deaths 

following bat bites were reported (Lumio and others, 1986; 

Selimov and others, 1986), causing such increased effort that 

by the end of 1989, 4,705 European bats had been tested, 

379 (8 percent) of which were found to be infected (Kappeler, 1989). Additional cases in bats and other mammals 

were discovered in Africa and Asia. In Australia, where rabies 

was believed to be absent, rabies infection was reported in 

adequately sampled bat species ranging from small insecteaters to large fruit-eating bats known as flying foxes.

Causative Agent

Classical rabies has been known for centuries and it is 

caused by the most widespread of seven similar virus species 

(also called genotypes, as each species is defined by phylogenetic analysis of viral genes) within the genus Lyssavirus 

(McColl and others, 2000). Four additional putative Lyssavirus species were discovered recently and their taxonomic

positions are under study (Kuzmin and others, 2005) (Fig. 1). 

Discoveries of additional lyssaviruses are likely to follow. 

Whereas classical rabies is well known, other lyssavirus species were not identified prior to 1956; thus, far less is known 

about them. For practical purposes, all lyssavirus species 

potentially produce rabies-like infections, so all are included 

in this publication. Lyssavirus is an appropriate name for 

these viruses as “lyssa” means rage, fury, or canine madness 

in Greek.

Table 1. Lyssavirus infections in mammals: agents, hosts, identification, and effectiveness of antirabies prophylaxis measures 

for humans.

Lyssavirus type Known hosts

Diagnostic 

capabilities 3

Effectiveness 

of antirabies 

prophylaxis 

for humans Name Genotype Reservoirs 4 1 Victims

Rabies 1 Carnivores, American 

bats

Humans, other 

mammals

Most government 

laboratories

Satisfactory.

Australian bat 7 Australian bats Humans Most government 

laboratories

Satisfactory.

European bat-2 6 European bats Humans Major regional Reduced.

Irkut bat Unclassified Asian bats Ferrets 2 Major regional Reduced.

Aravan bat Unclassified Asian bats Hamsters 2 Major regional Reduced.

Khujand bat Unclassified Asian bats Ferrets 2 Major regional Reduced.

European bat-1 5 European bats Humans, cats, 

stone marten, sheep

Major regional Poor.

Duvenhage bat 4 African bats Humans Major regional Poor.

Mokola 2 African shrews and 

insectivorous 

rodents

Humans, dogs, cats Major regional None.

Lagos bat 3 African bats Dogs, cats, mongooses Major regional None.

West Caucasian bat Unclassified Eastern European bats Hamsters 2 Major regional None.

1

 A species that maintains the virus in nature. Reservoir species may also succumb to infection.

2

 Experimental infections.

3

 Disease diagnostic laboratories capable of isolating the virus and identifying the genotype.

4

 Degree of protection estimates based primarily on laboratory animal responses (Hanlon and others, 2005). 
Vampires, Werewolves, and Settlement of the Americas 3

Vampires, Werewolves, and Settlement of the Americas Box 1

Bat-borne rabies, perhaps especially involving 

common vampire bats, may have inspired or 

influenced early tales of human vampires in Europe. 

The patterns of serial transmission by bite and 

the dreadful transformation of victims are strikingly 

similar. Correspondingly, rabies may have contributed 

to tales of werewolves and the transformation of 

their worldwide counterparts into various carnivorous 

mammals, perhaps influenced by cases of “barking” 

human rabies victims or of attacks by some infected 

human victims on other people.

Vampire bats also were likely causes of rabies prior to the arrival of European 

explorers in the Americas, and the rabies-like deaths of explorers in tropical America 

caused by “venomous bites” of vampire bats were reported in the 16th century. 

At that time, native peoples attempted to prevent rabies infections by cleansing the 

sites of vampire bat bites and cauterizing the wounds with wood embers (d’Anghiera, 

1555; De Oviedo y Valdes, 1526). Subsequent outbreaks of rabies in humans and 

their domestic animals plagued Latin America and prevented the development of 

extensive areas, problems that continue today. The infection eventually was found in 

various other tropical American bat species that specialize in diets consisting of fruit, 

pollen, nectar, insects, fish, or other vertebrates.

Common vampire bat in a Venezuelan 

cave. (D.G. Constantine)
4 Bat Rabies and Other Lyssavirus Infections

Prudence suggests that all lyssaviruses should be 

regarded as dangerous as classical rabies and, thus, likely to 

be lethal for humans and other mammals. The hazards can 

be even greater because not all of these viruses are readily 

identifiable by standard rabies diagnostic methods (Table 1). 

Furthermore, except for Australian bat lyssavirus, other lyssavirus infections are not dependably preventable by available 

rabies vaccines and rabies hyperimmune sera. For example, 

no biologics are available to prevent infection from the Lagos 

bat, Mokola, or West Caucasian bat viruses, and current 

rabies biologics have variably reduced effectiveness against 

the other currently known lyssaviruses (Hanlon and others, 

2005).

Based on knowledge of rabies and most other lyssaviral 

species, each lyssaviral species may prove to be divisible into 

variants. Each variant typically is found in a single species of 

reservoir host, and it apparently develops after its precursor 

is transmitted to a new host species. For example, a different 

bat lyssaviruss may evolve after it is transmitted from one bat 

species to another bat species, within which it subsequently 

reproduces in isolation, becomes differentiated from the original lyssavirus, and as time progresses, it becomes increasingly 

efficient in establishing itself in the second bat species. After 

a variant is transmitted from its reservoir host to an atypical 

or aberrant host species, a dead-end infection usually results 

because the maladapted atypical host quickly dies before the 

virus can be transmitted further. Identification of variants 

that infect atypical hosts, such as humans, usually points to a 

particular natural reservoir host species, such as a species of 

carnivore or bat, as the source for the human infection.

50 changes

Genotype 7: 

Australian bat

lyssavirus

Australia

Bats

Genotype 6: 

European bat 

lyssavirus 2

Europe

Bats

Genotype 1: 

Classical rabies

World

All mammals

Genotype 5: 

European bat lyssavirus 1

Europe

Bats

Aravan lyssavirus 

Irkut lyssavirus 

Khujand lyssavirus 

Genotype 4: 

Duvenhage virus

Africa

Bats

Genotype 3: 

Lagos bat virus

Africa

Bats

Genotype 2: 

Mokola virus

Africa

West Caucasian Bat Virus

Shrews

100

100 82

99 100

100

Figure 1. Unrooted phylogenetic tree based on complete 

lyssavirus N gene nucleotide sequences. Previously recognized 

genotypes are circled, and geographic area and primary host 

species are given; recently discovered lyssaviruses (Khujand, 

Aravan, Irkut, and West Caucasian Bat Virus) are not circled. 

Bootstrap values are presented for key nodes, and branch 

lengths are drawn to scale (Modified from Kuzmin and others, 

2005; see also Kissi and others, 1995).
Patterns and Trends 5

Geographic Distribution

Although bat lyssavirus infections have a broad geographic distribution (Figs. 2 and 3), our knowledge of lyssavirus distribution is incomplete for various reasons, including 

underreporting (Table 2). In some instances, reporting is 

inhibited by apprehension over the potentially suppressive 

effects on development or tourism. At best, reporting requires 

the presence of people to detect infected bats. For example, 

many cases in the USA are reported from cities, but unpopulated or sparsely populated areas often are devoid of reports. 

Nevertheless, lyssavirus infections have received considerable attention in the Americas and they are now known wherever bats are found. Eventually, lyssavirus infections may be 

found globally except for treeless areas and some isolated 

islands.

Patterns and Trends

Changes in human perception and increased research 

efforts have improved understanding of bat lyssavirus infections in bats, humans, and other mammals. Although vampire 

bat-borne rabies infected humans prior to European explorations in tropical America, the viral cause was not discovered 

until over 400 years later (Carini, 1911). Knowledge of 

the existence of vampire bat rabies stimulated research to 

enhance understanding of the extent and the gravity of infection and led to the development of methods for combating 

it. However, resource allocations for these efforts have been 

inconsistent and have fluctuated greatly. Consequently, the 

history of bat lyssavirus infections in general and their suppressive effects on geographic and economic development in 

the Americas are incompletely known. By 1969, the Food and 

Agricultural Organization of the United Nations concluded 

that cattle deaths and indirect losses caused by vampire bats 

cost Latin America over one-third of a billion dollars annually (Steele, 1969). Since then, relief has been provided 

through development of vampire bat control techniques and 

rabies vaccines. These advances have largely been supported 

and coordinated by the Pan American Health Organization 

(Belotto, 2005).

The 1953 discovery in the USA of bat rabies transmitted 

by bats other than the vampire bat (Appendix A) stimulated a 

major research effort that resulted in the disclosure of increasing numbers of cases (Fig. 4). These findings supported the 

popular conclusion that bat rabies was a new and growing 

problem. However, case increases proved to be proportional 

to investigative effort, with growing numbers of bats tested as 

time progressed (Fig. 5) (Constantine, 1967a). News coverage 

is another factor leading to local increases in the submission 

of bats to be tested and to corresponding increases in reported 

cases. In addition, technological advances have increased the 

application of rabies virus variant identification techniques 

and are yielding greater numbers of human rabies cases attributed to bats. During 2006, 1,692 cases of rabies in bats were 

reported in the USA.

Table 2. General geographic area of rabies-related lyssavirus infections reported in bats.

[Unknown indicates that the lyssavirus species was known only to be rabies-related]

Lyssavirus 

species

Number of bat species reported infected

Africa Australia Asia Europe Indian 

subcontinent

Middle 

East

Aravan 1

Australian 5

Duvenhage 1

European-1 9

European-2 7

Irkut 1

Khujand 1

Lagos 4 1

West Caucasion 1

Unknown 1 3 7 2 1

Total: 6 5 6 24 2 2
United States of America

33*

Canada

13

Argentina

8

Chile

4 Uruguay

Paraguay

2

Bolivia

3

Brazil

33

Peru

19

Ecuador 1

Columbia

4

Venezuela

2 Guyana Surinam

Panama 8 French Guiana

Costa Rica 1

Nicaragua 1

Honduras 1

El Salvador 1

Belize 8

Grenada 1 Trinidad 9

Cuba 2

Mexico

31

Guatemala 5

EXPLANATION

Color reflects the number of bat species reported 

rabies-infected within a country

The specific number of bat species infected in 

each country is shown in bold type

 No data

 1−5

 6−10

 11−15

 16−20

 21−25

 26−30

 31−35

* Total includes Alaska; naturally occurring rabid bats 

have not been found in Hawaii

6 Bat Rabies and Other Lyssavirus Infections

Figure 2. Number of bat species reported with rabies infections 

within the Americas (D.G. Constantine, unpub. data).
EXPLANATION

Lyssavirus species. Bolded numbers after virus species indicate the numberof bat species infected with the virus (black) or the number of human victims (orange) in each country.A Australian D Duvenhage E-1 European bat virus-1 L Lagos E-2 European bat virus-2 I IrkutAR Aravan M MokolaK Khujand WCBV West Caucasian bat virusU Unknown, virus not characterized  beyond rabies-related

AR, 1

A, 5A, 2

D, 1

U, 1

D, 3 D, 2

D, 1 D, 1

L, 2

L , 1

L , 1

L , 1 M, 2

I. 1

K, 1

U, 1WCBV, 1

U, 1

U, 2

U, 2U, 1

U, 1

U, 1

U, 2 U, 1–2

E-1, 2

E-1, 1

E-1, 1

E-1, 1

E-1, 1 E-1, 1

E-2, 2

E-1, 1 E-2, 1

E-2, 1

E-2, 1

E-2, 1

E-2, 1

E-2, 1

E-2, 1

E-2, 1

E-1, 2

U, 1

U, 1

U, 1 U, 1

U, 1

U, 2

See inset

L , 1

Figure 3. Old World distribution of lyssavirus infections (excluding rabies virus) reported in bat species (black) and humans (red) in Africa (BoulCrick and others; 1982; Foggin, 1988; Familusi and others, 1972; Foggin, 1988; Markotter and others, 2006a; Meredith and others, 1971; Paweska anand others, 1980; van Thiel and others, 2008), Asia (Arai and others, 2003; Kuzmin and others, 2003; Tang and others, 2005), Australia (Allworth and others, 2002; Warrilow and others, 2003), Europe (Amengaul and others, 1997; Botvinkin and others, 2003; Bruyère and Janot, 2000; Bruyère-Massand others, 2002; Hentschke and Hellman, 1975; Johnson and others, 2006; Kappeler, 1989; Kerekes, 1999; King and Crick, 1988; Khozinski and othe2005; Lumio and others, 1986; Matouch, 1999; Nikolíc and Jelesíc, 1956; Pérez-Jordá and others, 1995; Serro-Cobo and others, 2002; Tunçman, 195Van der Poel and others, 2003; WHO Collaborating Centre for Rabies Surveillance and Research, 1999), the Indian subcontinent (Pal and others, 1and the Middle East (Picard-Meyer and others, 2004).
1,200

1,300

1,400

1,500

1,600

1,700

100

200

300

400

500

600

700

800

900

1,000

1,100

 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005

0

Number of rabid bat indentifications

Year

8 Bat Rabies and Other Lyssavirus Infections

Figure 4. Number of rabid bats identified 

in the United States from 1955 to 2006. 

Compiled from U.S. Centers for Disease 

Control and Prevention data.
1964–70 1971–80 1981–90 1991–2000 2001–06

0

2

4

6

8

10

12

Rabid bats, in percent

Time period

4,072 12,364 14,816 13,161 10,453

Species Sucsceptibility 9

Species Susceptibility

It has long been assumed that all mammals are susceptible to rabies infection, a view that may apply in varying 

degrees to the other lyssaviruses as more is learned about 

them. No gross lesions aid in the identification of lyssavirus 

infections, so diagnosis of living or dead untested subjects is 

tentative and based on history, which usually includes reports 

of behavior that is not specific to the disease. Rabies virus 

is known to be maintained within two reservoir host mammal groups, carnivores and bats, and carnivores host virus 

variants that differ considerably from those hosted by bats 

(Constantine, 1967a). Rabies and other lyssavirus infections 

have been reported from 114 bat species (Appendixes B and 

D), including 33 in the USA (Appendix A; Table 3). Australia 

has the fewest number of bat species (five documented) with 

lyssavirus infections; the numbers of infected species for 

Asia, Africa, and Europe also are low (Appendix D; Fig. 6). 

The lyssaviruses identified from infected bats represent 10 

(including rabies) of the 11 currently named or putative 

genotypes for this virus group, and several of the lyssaviruses from bats remain unclassified (Table 1). Thus, there is 

adequate reason to regard all lyssaviruses to be potentially as 

deadly as classical rabies.

The results of exploratory rabies transmission experiments provided the first indication of the existence of viral 

variants. These experiments demonstrated that the spectrum 

of carnivores susceptible to infection differed according to the 

bat species origin of rabies virus, as did incubation periods, 

duration of illness, and morbid behavior (Constantine, 1967a). 

Maladaptations, such as the death of the host before the virus 

can reach the saliva for transmission, reveal why infections of 

new, unadapted host species may, in effect, be dead end.

Table 3. Bat families in the USA reported with 

bat rabies.

Family Food habits Number of

infected species

Phyllostomidae Omnivorous 1 1

Mormoopidae Insectivorous 1

Molossidae Insectivorous 4

Vespertilionidae Insectivorous 27

1

 Members of this diverse family of neotropical bats feed upon 

insects, small vertebrates, blood, fruit, nectar, and pollen.

Figure 5. Percentage of rabiessuspect bats identified as rabid in 

the United States from 1964 to 2006. 

The first documented bat rabies 

case in humans was identified in 

1953; bat rabies surveillance records 

are unsatisfactory for 1954–63. 

(Bold numbers indicate the number 

of bats tested during the indicated 

time period. Compiled from U.S. 

Centers for Disease Control and 

Prevention data.) 
0

10

20

30

40

50

60

North

America

Central

and 

South

Australia Asia Africa Europe America

Number of infected bat species

10 Bat Rabies and Other Lyssavirus Infections

Human Infections

Each year as many as 100,000 people globally may 

die of rabies (Rupprecht and others, 1995), and ten million 

people take antirabies treatment (Goswami and others, 2005). 

Carnivores, principally dogs, are the vectors for the great 

majority of rabies transmissions to people in developing 

areas. In Latin America, thousands of people must have died, 

and hundreds continue to perish due to vampire bat-borne 

rabies, but case reports are lacking from some countries, particularly from their remote regions. Due to competition for 

limited resources for research of the causes of human mortality, vampire bat rabies infection in humans, especially those 

living in inaccessible areas, did not receive adequate attention until recently (Navarro Vela, 2001; Takoaka and Omoto, 

2001; Vargas Pino, 2001). Reports of human rabies due to 

bats other than vampires are difficult to discern because of 

local inability to differentiate vampire bats from any of the 

great variety of bat species in Latin America.

The recognition of human lyssavirus infections of bat 

origin exists to varying degrees in developed countries. The 

existence of bat rabies and associated human mortality in 

North America north of Mexico was unknown prior to the 

1950s. Between 1951 and 2006, 51 recognized human rabies 

cases in the USA and Canada were determined to be of 

insectivorous bat origin (Table 4), and there is some evidence that bat rabies existed before 1951 in these countries 

(Christensen, 1946; Constantine, 1967b). This revelation of 

bat-associated human rabies mortality as a result of public 

health investigations has been mirrored elsewhere for other 

lyssaviruses (Table 5).

The onset of clinical disease in humans infected by 

rabies virus nearly always results in death (Box 2). Exceptions are so noteworthy as to be the subject for scientific 

reports (Hattwick and others, 1972; Porras and others, 1976; 

Willoughby and others, 2005; Box 3). Examples of the 

course of disease in humans infected by bat lyssaviruses 

(Appendix E) illustrate the consequences of those infections 

and the variability in incubation time between exposure and 

the onset of clinical disease (Table 6).

Figure 6. Number of rabies- and 

other lyssavirus-infected bat species 

by continent.
Species Susceptibility 11

The Course of Bat Rabies in Humans Box 2

Rabies in humans is characterized by a variable 

incubation period, usually from 1 to 3 months, but 

ranging from 10 days to a year or more. The disease 

then generally develops in three phases: prodromal, 

excitatory, and paralytic (Fishbein, 1991). Prodromal 

signs last from 2 to 10 days, are nonspecific, and 

include slight fever, malaise, headache, nausea, 

dilation of the pupils, sore throat, anxiety, irritability, 

and perhaps sensitivity to wind, bright light, and 

noise. More significant are abnormal sensations such 

as pain, burning, cold, and itching or tingling near the 

site of virus entry, such as a bite.

The excitatory phase, present in 80 percent of 

cases, may be absent, rapidly transitory, last 1 

week, or continue until death. During this phase, 

victims experience increasing anxiety, apprehension, 

and melancholia. Muscles around the virus entry 

site may weaken. Eyes may undergo unusual or 

uncoordinated movements, pupils may undergo 

asymmetric dilation or constriction, and eye corneas

may be insensitive to touch. Facial muscles may 

weaken and hoarseness develops. Other signs can 

be alterations in heart rate and intensity, respiratory 

rate and depth, urinary retention followed by 

overflow incontinence, and constipation. The classical 

manifestation of rabies in humans is hydrophobia, 

when upon attempts to swallow liquid, the victim 

forcefully expels it due to the involuntary contraction 

of the muscles involved in swallowing and respiration. 

This reaction is subsequently repeated if the victim 

is provoked by the sight or sound of liquids. The 

convulsive attack may cause death through choking 

and failure to breathe. The victim may suddenly die 

from respiratory arrest or death may be preceded by 

coma.

Paralysis, which predominates in 20 percent of cases, 

may precede death by 1–4 weeks. Swallowing is 

possible but becomes difficult. A general flaccid 

paralysis develops. Apathy progresses to stupor, 

followed by coma, peripheral vascular collapse, 

and death.

Table 4. Insectivorous bat origins of human rabies infections recognized in the USA and Canada 

from 1951 to 2006.

Bat species Family

Number of bat 

carcasses

identified

taxonomically

Number of 

lyssaviruses 

identified to the 

variant level

Total number of 

bats involved in 

human rabies 

infections

Brazilian free-tailed bat Molossidae 7 10 10

Big brown bat Vespertilionidae 2 1 3 3

California myotis Vespertilionidae 1 1 1

Myotis sp. Vespertilionidae 0 1 1

Silver-haired bat Vespertilionidae 1 11 11

Tricolored bat Vespertilionidae 0 17 17

Totals 11 43 2 51 

1

 Variants were an Ef (Big brown bat)-associated, a Ps (Tricolored bat)-associated, and an undetermined variant.

2

 Total includes eight bats for which neither the bat taxonomic identity nor the virus variant were determined.
12 Bat Rabies and Other Lyssavirus Infections

Box Human Survival of Clinical Rabies—An Exception to the Rule 3

Cases of patients surviving rabies infection after the appearance of clinical signs, 

without preexposure vaccination or postexposure prophylaxis or both, are rare. 

However, two cases of patients surviving clinical rabies are well documented. The 

first case was a 6-year-old boy in Ohio (Hattwick and others, 1972), and the second 

was a 15-year-old girl in Wisconsin (Willoughby, Jr., and others, 2005). Both patients 

were exposed to rabies through bat bites. In each case, aggressive supportive 

treatment was given to avoid or decrease the known or anticipated ravages of the 

disease. The reasons for the success of the administered treatments are not fully 

understood, but the positive outcomes offer incentives for continuing research 

efforts.

Case 1

A 6-year-old boy was bitten on his left thumb by a 

rabid big brown bat on October 10, 1970, while he 

was asleep in his family’s farmhouse near Wilshire, 

Ohio. Four days later, a 14-day course of rabies 

vaccine was begun, but no antirabies serum was 

given (Table 1). Recovery was complete 6 months 

after the onset of symptoms.

The physicians (Hattwick and others, 1972) suggested 

that the rabies strain in this case may have been less 

virulent for humans than other strains, however that 

hypothesis was not pursued. Rabies variants from big 

brown bats have been shown to possess extremely 

different abilities to cause infection in animals (Bell 

and others, 1962; Constantine and others, 1968b), 

and geographically distinct rabies variants from this 

species are known in the United States (De Mattos, 

C.C. and others, 2001). For example, a bat identified 

as a big brown bat transmitted lethal rabies by bite 

to a woman in Maryland (Barnhart and others, 1976), 

but the rabies variant proved to be associated with 

the tricolored bat (Messenger and others, 2002). 

Alternately, an unidentified bat transmitted lethal 

rabies by bite to a man in Washington in 1997, and 

the rabies variant was identified as originating from 

a big brown bat (Geyer and others, 1997).

Case history 1.

Day of 

symptoms Symptoms/treatment

1 Neck pain developed 2 days following the vaccine series (60 days after the victim was bitten 

by the bat).

5 High fever, appetite loss, vomiting, and dizziness, but alert and cooperative.

11 Pain in head and limbs, became lethargic, writing and walking were difficult.

13 Speech problems, uncooperative and bizarre behavior, bit and chewed on his saliva collection 

tube, weakness, partial paralysis, and seizures of the left side of the face and left limbs.

15 Coma, heart rhythm irregularities, and seizures. Aggressive treatment continued to prevent 

oxygen deprivation and intracranial hypertension. Excess cerebrospinal fluid removed from 

within the brain.

22 Coma ended and recovery began.
Human Survival of Clinical Rabies—An Exception to the Rule 13

Case history 2.

Day of 

symptoms Symptoms/treatment

1 Fatigue, tingling, and numbness of the left hand about 1 month after the victim was bitten by 

a bat.

3 Unsteadiness, double vision, nausea, and vomiting, but no fever.

5 Fever of 101.8 oF (38.8oC), slurred speech, involuntary eye movements, and tremors of the 

left arm.

6 Bat bite revealed, patient transferred to the Medical College of Wisconsin with a fever of 

100.8oF (38.2oC), muscular twitching, incoordination, difficulty speaking, tremors in the 

left arm, dulled alertness, and hypersalivation. A tracheal breathing tube was installed. 

Because rabies signs had already developed, rabies vaccine or antirabies serum were not 

administered.

7 to 34 Partial paralysis and numbness due to sensory nerve dysfunctions. Extensive supportive 

treatment administered, including mechanical breathing, heart monitoring and treatment, 

intracranial hypertension control, antiviral drugs, and a drug-induced coma permitting the 

patient’s rabies antibodies to increase and destroy the virus. After 7 days of coma, sedation 

medications were discontinued, and the patient became increasingly alert.

35 Patient removed from isolation and started on a rehabilitation program.

80 Patient discharged to her home.

Case 2

A more recent case of survival also involved a bat, 

but the species is unknown. In October 2004, in 

Fond du Lac, Wisconsin, a 15-year-old girl was bitten 

on her left index finger by a bat that she picked up 

and released outside after it had fallen from flight 

while it was inside a church. Nobody considered a 

potential rabies danger, thus rabies prophylaxis was 

not administered. The first symptoms appeared about 

1 month later, and the patient was discharged from 

the hospital 80 days following the onset of symptoms 

(Table 2). Rehabilitation following release from the 

hospital contributed to this success story. A tutor 

helped the victim finish her sophomore year of high 

school so that she could rejoin her classmates in the 

fall. Physical therapy helped her to overcome speech 

problems, weakness in her left hand and foot, and 

abnormal movements in her arms and hands. The 

victim entered college as an 18-year-old freshman in 

the fall of 2007. She regrets that none of the 10 other 

rabies patients subsequently administered the same 

or similar treatment survived, but she continues her 

life with energy and enthusiasm. She is an aspiring 

wildlife conservationist, and she still loves animals, 

including bats.

Coma and Intervention

Although both human rabies survivors experienced coma, a natural and often 

terminal development in clinical rabies, the medical personnel of the first case 

merely acknowledged its existence whereas those in the second case induced coma 

with drugs. The latter authors (Willoughby, Jr., and others, 2005) said it was not clear 

if the induced coma played a role in the victim’s recovery, but they recommended 

that their updated protocol and future updates (Medical College of Wisconsin, 2007) 

be followed in the treatment of subsequent patients. A focus for consideration is 

control of the patient’s body temperature.
14 Bat Rabies and Other Lyssavirus Infections

Bat Infections

Studies of rabies in North American insect-eating bats 

reveal that incubation periods can be from 21 days to at least 

209 days, and the duration of the disease can last from less 

than 1 day to 20 days. Most important, however, is the knowledge that individual bats have transmitted the virus by bite as 

early as 12 days before the appearance of clinical signs and 

24 days before the death of the bat (Constantine, 1994). In 

either situation, a bitten person could develop clinical disease 

or be dead of rabies before clinical signs appear in the biting 

bat, a sobering prospect for persons who prefer to spare the 

life of a biting bat. Worse yet, if the bat becomes chilled, and 

thus lethargic or torpid, all of these periods become extended. 

For example, some bats are known to awaken from hibernation in spring, whereupon they resume development of rabies 

infection that has been placed on hold all winter (Constantine, 

1967a; Box 4).

Paralytic rather than furious rabies appears to be 

characteristic of rabies in most infected bats, but there are 

disturbing exceptions. Infected insectivorous bats display 

neither external nor internal gross lesions nor clinical signs 

specific enough to permit a diagnosis prior to laboratory tests 

performed on brain tissue. They may display a variety of 

nonspecific clinical signs that change as the disease progresses or in response to environmental influences. Basically 

the bat may be expected to suffer nervous system disorders 

that make it overly responsive to sensory stimuli, such as 

touch, and that elicit retaliatory bites, tremors, and perhaps 

wing flapping. However, these responses are not diagnostic, 

because the approach of persons or animals can elicit similar 

behavior in healthy bats. Progressive paralysis of infected 

bats can increase irritability and biting of nearby objects. 

Such biting may also be used as an aid in crawling or climbing as paralysis disrupts coordination. Developing paralysis 

can oblige flying bats to alight on any elevated object, even 

a standing person, rather than land on the ground. Thus, bats 

found in unusual places may be infected. Flight problems 

decrease the bat’s ability or efforts to secure food and water, 

leading to attendant weight loss and dehydration; captive bats 

simply cease to eat. If an infected bat does not fall to earth 

during flight, it may finally die in its shelter. In either situTable 5. Human nonrabies lyssaviral infections reported in Africa, Asia, Australia, and Europe.

[ABL, Australian bat lyssavirus; EBL, European bat lyssavirus]

Location Year Source of

infection 1

Number of

human cases

Lyssavirus

species Reference

Australia 1996 Bat 1 ABL Allworth and others, 1996.

Australia 1997 Bat 1 ABL Hanna and others, 2000.

China 2002 Bat 1 Rabies-like 2 Tang and others, 2005.

Finland 1985 Bat 1 EBL-2 Lumio and others, 1986.

India 1954 Bat 1 Rabies-like 2 Veeraraghavan, 1955.

Kenya 2007 Bat 1 Duvenhage van Thiel and others, 2008.

Nigeria 1968 Unknown 1 Mokola Familusi and others, 1972.

Nigeria 1971 Unknown 1 Mokola Familusi and Moore, 1972.

Russia 1985 Bat 1 EBL-1 Botvinkin and others, 2005.

Scotland 2002 Bat 1 EBL-2 Fooks and others, 2002.

South Africa 1970 Bat 1 Duvenhage Meredith and others, 1971.

South Africa 2006 Bat 1 Duvenhage Paweska and others, 2006.

Ukraine 1977 Bat 1 Rabies-like 2 Botvinkin and others, 2005.

Ukraine 2002 Bat 1 Rabies-like 2 Botvinkin and others, 2005.

1

 Bat species not reported.

2

 Untyped lyssaviruses causing rabies-like clinical signs.
Species Sucsceptibility 15

ation, lack of activity can result in lethargy and consequent 

life extension before death (Constantine, 1967a; Sadler and 

Enright, 1959).

Rabies in experimentally infected vampire bats resembles the previously described infections in insect-eating bats 

in reference to the clinical disease, incubation period, illness 

period, and periods when virus can be detected in saliva 

(Moreno and Baer, 1980). This contrasts with early reports 

that vampire bats could be healthy carriers and excretors of 

rabies virus in saliva (Torres and de Queiroz Lima, 1935; 

Pawan, 1936). However, those early investigators may have 

mistaken as rabies one of a number of other viruses that can 

chronically infect bat salivary glands (Constantine, 1970; 

Constantine, 1988b). Another important characteristic of 

vampire bats is that, unlike most temperate zone bats, the 

tropical vampire bats are not physiologically able to lower 

their body temperatures to become lethargic or dormant, 

which would defer viral disease development (Lyman and 

Wimsatt, 1966).

Most infective bites from insectivorous bats are inflicted 

in a manner similar to bites inflicted by healthy bats for 

self-defense, such as when a bat is caught by a person or an 

animal. Rarely, however, some rabid bats, including insect 

eaters, are known to attack people, most often during daytime. Furious daytime attacks on people and animals by rabid 

common vampire bats have long been known during outbreaks in Latin America (Haupt and Rehaag, 1921; Torres and 

de Queiroz Lima, 1935; Pawan, 1936). Noise produced by or 

associated with the victim evidently can precipitate attacks 

(Constantine, unpub. data). Such attacks, like one experienced 

by the author, are known to be direct and quick, unlike the 

erratic flight of bats in pursuit of insects. Similarly, lyssavirus-infected black flying foxes of Australia are known to 

attack other bats repeatedly (Mackenzie, 1999). Unprovoked 

attacks by bats are clearly suggestive of rabies, but they are 

difficult to differentiate from events described by frightened 

people who think they are under attack by healthy bats that, in 

reality, are in pursuit of nearby flying insects. 

Human perceptions of abnormal behavior by bats are not 

always correct and often result in healthy bats being mistaken 

as ill. Rather than being infected, the bats may be lethargic or 

dormant when at rest in a cool place. In addition, young bats 

may not yet be able to fly, or bats injured from cat attacks 

or other causes may appear partially paralyzed. Species that 

take shelter in the soft foliage of trees, such as hoary bats, 

sometimes are attacked by birds and fall to earth, where their 

tender wings suffer abrasions. These abrasions may cause the 

bats to react with a hissing sound and to extend their wings 

and thrash about in an effort to become airborne. This behavior occurs whether or not the bat is infected with rabies.

Table 6. Examples of incubation periods and times to death relating to human mortalities from bat-associated lyssavirus exposure.

[See appendix E for more detailed descriptions of the nine cases below]

Case1 Exposure 

location

Day of onset of 

symptoms after 

exposure

Day of death 

following onset 

of symptons

Comments

1 South America 27 days 9 Victim bitten on toe while sleeping; bat seen flying from foot when victim 

awoke. 

2 North America 16 days 9 Victim bitten on forearm when bat picked up from ground. 

3 North America About 6 weeks 4 Airborne exposure in cave occupied by millions of bats. 

4 North America 26 days 12 Victim bitten on finger when bat was rescued from attack by dogs; dogs 

potentially exposed. 

5 North America 21 days 7 Victim bitten on ear while sleeping; bite awoke victim, cat ate bat; cat 

potentially exposed. 

6 Europe 51 days 11 Victim bitten by abnormally acting bat that was then released.

7 Africa About 1 month 4 Victim awakened by bat bite on lip. 

8 India About 11 weeks 2 5 Victim bitten on forearm when bat was picked up from ground.

9 Australia Unknown 16 Victim cared for fruit bats.

1

 Case 1, Hurst and Pawan, 1932; Case 2, Sulkin and Greve, 1954; Case 3, Irons and others., 1957; Case 4, Humphrey and others, 1960; Case 5, Humphrey and

others, 1960; Case 6, Lumio and others, 1986; Case 7, Meredith and others, 1971; Case 8, Veeraraghavan, 1955; Case 9, Allworth and others, 1996. 

2

 May be longer, because the day of the onset of symptoms was not recorded.
0

50

100

150

200

250

300

350

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.

Number of rabies-suspect bats

Month

Long-legged myotis. (D.G. Constantine)

16 Bat Rabies and Other Lyssavirus Infections

Box Seasonal Bat Behavior and Disease Transmission 4

Most North American bat species live in temperate 

zones and are heterothermous, in that when a 

bat is inactive or sleeping its body temperature 

corresponds to that of the environment, similar to 

what is observed in cold-blooded (poikilothermous) 

reptiles and amphibians. When a heterothermous 

bat is active, however, its body temperature is 

regulated to a uniform warm level like that of most 

warm-blooded (homoiothermous) 

mammals and birds. Thus, when the 

environmental temperature decreases, 

heterothermous bats become cold and 

eventually lethargic or dormant when 

at rest, similar to a hibernating state 

that results in conservation of energy. 

Their flying insect food responds 

similarly to cold weather and does 

not fly. Many of these bats, the longlegged myotis for example, hibernate 

in winter, whereas others migrate 

to warm areas. Because most North 

American bat species either migrate 

to warmer climates or are inactive in 

cooler areas during cooler months, 

bat rabies similarly disappears during 

the winter. Accordingly, cases of bat rabies in the 

USA routinely start appearing in spring, increase until 

early fall, and decrease thereafter. North America’s 

seasonal cycle of bat rabies has been repeated in 

part and with similar results in Latin America, Europe, 

Asia, Africa, and Australia, but seasonal patterns are 

less pronounced toward the equator.

Submissions of bats suspected of having rabies, by month (reproduced from Blanton 

and others, 2007).
Seasonal Bat Behavior and Disease Transmission 17

Brazilian free-tailed bat. (D.G. Constantine)

A smaller number of temperate North American bat 

species are full-time homoiothermous mammals. 

These bats cannot hibernate and must remain in a 

warm environment at all times. The California leafnosed bat, whose warm geographic range extends 

from Mexico into southern California and Arizona, 

does not migrate southward in winter and often 

inhabits geothermally warmed mine tunnels or caves. 

It selects warmer sites for shelter during winter, often 

within the same underground lair, and it flies outside 

to capture its insect prey as the weather allows.

The Brazilian free-tailed bat, found in warm parts of 

North and South America, is intermediate between 

the homoiothermous and heterothermous bats. 

Although its body temperature decreases if it is 

inactive during short cold periods, it cannot undergo 

true prolonged hibernation. Instead, it migrates to 

relatively warm areas for winter, where the bats and 

their flying insect food both remain active.

California leaf-nosed bat. (D.G. Constantine)
18 Bat Rabies and Other Lyssavirus Infections

Infections in Other Species

In addition to direct exposures of humans 

to rabid bats, other animals, principally 

carnivores, may become infected or contaminated by rabid bats and consequently become 

additional sources for infection (Table 7). 

Experiments have demonstrated that domestic 

and wild carnivores are among the species 

susceptible to rabies variants of North American insectivorous bat origin, although variants 

differ in their degrees of infectivity for various carnivore species (Constantine, 1966a; 

Constantine, 1966b; Constantine, 1966c; 

Constantine and Woodall, 1966; Constantine, 

1967a; Constantine, 1967b; Constantine and 

others, 1988a; Constantine and others, 1988b). 

Laboratory mice have been experimentally 

infected by bites of North American insectivorous bats (Bell and others, 1962). In addition, 

in 2006 following a public exhibition attended 

by approximately 150,000 people, a horse was 

diagnosed postmortem to have been infected with a rabies 

variant associated with big brown bats (Blanton and others, 

2007). As a result, a health alert was issued by the state health 

department and 53 people who had attended the exhibition 

were consulted about potential rabies exposure. Some carnivores and other species are also known to be susceptible to 

other lyssaviruses (Table 1).

Vampire bat-transmitted rabies can kill and decimate 

herds of cattle as well as destroy horses, mules, donkeys, 

burros, goats, sheep, pigs, dogs, and poultry. Poultry, however, often soon die from exsanguination rather than rabies 

(Greenhall, 1988). After an incubation period of about 1 to 2 

months, the disease kills cattle within several days to a week. 

Initially the animal is restless, perhaps excitable, and loses its 

appetite. The next day, its rear legs tremble and are uncoordinated due to developing paralysis, and it salivates profusely 

because of difficulty in swallowing. Some animals, especially 

Zebu cattle, may charge people, but the disease is primarily paralytic rather than furious. By the third day the animal 

is constipated, eventually falls onto its side, and cannot rise. 

On about the fourth day, posterior paralysis is complete, but 

attempted running with the forelegs can result in circular 

movement and consequent circular earth excavation (Fig. 7). 

Death follows in the next day or two (Baer, 1991).

Table 7. Examples of nonbat species infected by rabid bats. 

Species infected Relative

occurrence Natural or laboratory infection Comments

Cattle Common Natural and laboratory infection Major vampire bat problem in Latin America.

Cats Occasional Natural and laboratory infection Increasing problem, presumably by bat bite or bat ingestion.

Dogs Rare Natural and laboratory infection Presumably by bat bite or bat ingestion.

Foxes Rare Natural and laboratory infection By bat bite, bat ingestion, or aerosols in congested free-tailed 

bat caves.

Skunks Rare Natural and laboratory infection Presumably by bat bite or bat ingestion.

Coyotes Rare Laboratory infection and 

presumed in nature

By bat bite, bat ingestion, or aerosols in congested free-tailed 

bat caves.

Laboratory mice Only in laboratory Laboratory infection By bat bite.

Figure 7. Circular pattern of earth excavation (dark area of ground behind calf 

(indicated by an arrow) resulting from confined body movement associated with 

posterior paralysis in a rabies-infected Zebu. (G.C. Mitchell)
Disease Ecology 19

Obtaining a Diagnosis

A bat that is suspected of being infected with a lyssavirus must be euthanized and its brain tested in a laboratory to 

obtain a diagnosis. Standard tests for rabies virus suffice for 

lyssaviruses known to exist in the Americas or Australia, but 

special tests at regional disease diagnostic laboratories may 

be necessary to identify lyssaviruses found in Europe, Asia, 

Africa (Table 1), and perhaps elsewhere.

Because of the consequences for a human infection, the 

handling of bats merits precautionary measures. Local health 

departments generally provide directions to the public regarding submission of bats for laboratory diagnosis. Heavy leather 

work gloves prevent bites from most live insectivorous bats, 

and rubber gloves prevent skin contamination from the saliva 

or tissue fluids of dead bats. The bats often will be retrieved 

by animal control personnel. Dead bats should be double 

bagged and kept cold or frozen, even if they are decomposed 

or partially eaten. It is important to keep the dead bats cold 

or frozen to prevent further decomposition that may render 

test results unreliable, which would force medical personnel 

to decide that the bat was infected and to initiate appropriate 

lifesaving precautions for individuals known to have been 

exposed to the bat.

Complete histories that accompany submitted bats facilitate the rapid notification of persons-at-risk, the initiation of 

timely lifesaving medical management, and disease management involving relevant pets. Pertinent information often is 

not sought until after the bat has been determined to have 

been infected, thus causing the delay of lifesaving measures 

while information is obtained. Histories are most informative 

when they reconstruct the trail of events since the bat was first 

observed, including the kinds of known and potential exposures (bites, scratches, saliva, tissue fluids, etc); dates; places; 

the names, addresses, and telephone numbers of associated 

persons and animal owners; and known as well as potential 

pet or other animal contacts.

Disease Ecology

The ecology of bat rabies and other lyssavirus infections 

is closely associated with bat ecology, including bat feeding 

strategies (Box 5) and the dynamics of infection within various bat species. Thus, those factors provide the focus for this 

section on Disease Ecology, along with differences in global 

virus transmission in relation to vampire bats or to other bats. 

Consideration is also given to the environmental persistence 

of lyssaviruses and their hosts. 

Bat Activity and Disease Patterns

Bat lyssavirus is evident where and when bats are present 

and physiologically active (rather than in hibernation), which 

coincides with the availability of bat food. Most information on this subject is derived from studies in the temperate 

and tropical areas of America, with evidence of comparable 

patterns in these zones globally. Bats are present and active 

in temperate areas during warm seasons, when their flying 

insect food is available, and correspondingly, bat infection is 

also evident at these times. As the cold season approaches, 

evidence of both insects and bats decreases and eventually 

disappears, as do cases of bat lyssavirus. During this period, 

some bat species migrate to warmer areas, where insects are 

available. Other species retire to cold sites like caves to hibernate instead of migrating (Fig. 8). In contrast, bat food, such 

as insects and fruit, is available during all seasons in tropical 

areas, and bats and bat infections may be evident continuously 

in those areas.

Figure 8. Townsend’s big-eared bats 

awakening from hibernation. The ears 

of some bats are still folded beneath 

their folded wings, in typical hibernation 

position. (D.G. Constantine)
20 Bat Rabies and Other Lyssavirus Infections

Box Bats—Diverse Mammals with Diverse Diets 5

Bats are distributed globally to the limits of tree growth and divided into 1,116 species 

(Simmons, 2005). Different groups or species of bats exhibit specialized dietary 

preferences. Bats in temperate zones are less numerous and diverse than those in 

tropical zones. Thus, while insectivorous bats are most common in temperate regions, 

the greater availability and variety of dietary items near the equator supports a diversity 

of food choices and food specializations among tropical bat species, a few of which are 

discussed below.

The diadem leaf-nosed bat of northern Australia 

hangs in ambush from a tree branch, ready to 

fly out and capture passing flying insects. Some 

insectivorous bats specialize in eating aquatic 

insects, which they detect and gaff with elongated 

claws on large feet as the insects break the water 

surface. Similarly, other bats, such as the greater 

bulldog bat, gaff small fish to either supplement 

or replace their insect diet. Some bats snatch 

earthbound insects, sometimes capturing lizards as 

well. Other bats take insects that feed on ripe fruit, 

which the bats may also consume.

The Queensland tube-nosed fruit bat of 

northern Australia feeds on the fruits and 

blossoms of native trees. This bat’s tubular 

nostrils may function as snorkels when 

its muzzle is immersed in mushy fruit 

pulp. Because they do not have to chase 

and kill prey to survive, fruit bats exhibit 

a placid demeanor and seem to expect 

courteous treatment from people in return. 

For example, the pictured tube-nosed bat 

immediately accepted and consumed an 

offered banana, forgoing the opportunity 

to flee. Similar behavior has also been 

observed among other neotropical 

fruit bats.

The fringe-lipped bat, the pictured example observed in a Venezuelan 

cave, eats insects, lizards, and frogs, grabbing its prey between its 

jaws while in flight. It is attracted to frogs by their calls, recognizing 

frog species by their sounds and thus bypassing poisonous frogs.

Fringe-lipped bat. (D.G. Constantine)

Queensland tube-nosed fruit bat. 

(D.G. Constantine)

Diadem leaf-nosed bat (D.G. Constantine)
Bats—Diverse Mammals with Diverse Diets 21

The greater spear-nosed bat is omnivorous, 

eating insects, smaller bats, mice, birds, and 

fruit. The pictured bat, netted in Trinidad, shared 

the author’s dinner the day it was captured, 

eating filet mignon and fruit salad.

The common vampire bat, the pictured example 

captured in Trinidad, is sanguivorous and consumes 

only blood. Literally bloodthirsty, it will die if it does 

not feed for 3 nights. Fearless when under attack, 

vampire bats deftly escape from potential predators, 

a reaction that can be unnerving to a human trying 

to outwit a single bat, to say nothing of an encounter 

with a dozen or more. Regardless of the size 

disparity, vampire bats view humans as a source 

of nourishment. The first vampire that the author 

observed was on all fours, with its wings folded to 

serve as forelegs, staring at him from a ledge near 

the ceiling of a Mexican cave and trying to determine 

its distance to the potential human prey by doing 

“push-ups” and moving its head from side-to-side, 

similar to an owl’s head movements during hunting. 

Several nights later another vampire bat, caught in the 

author’s harp trap (Constantine, 1958) as it tried to 

enter a calf pen in search of a meal, likewise stared at 

the author, calculating its distance to him from within 

its cage. The vampire bat’s predatory nature shows 

courage, determination, and rapid responses when 

it must avoid retaliatory attacks, such as repeated 

kicks when it tries to obtain blood from the hind feet 

of cattle. Vampires are adept at avoiding kicks by 

jumping out of the way, only to swiftly return.

A vampire bat caged with a rat will relentlessly pursue 

the rat as prey and almost invariably triumph, even 

“boxing” the rat with its folded wings to ward off 

retaliatory charges. Similarly, when caged with a large 

snake intent on eating the bat, the vampire is capable 

of repeatedly avoiding strikes and persistently holding 

its course until able to obtain its blood meal from the 

nose of the seemingly hypnotized snake (Greenhall, 

1988). Thus, the impression is fostered that the bat 

is endowed with the supernatural powers of the 

fictitious Dracula (Stoker, 1897).

Greater spear-nosed bat. (D.G. Constantine)

Common vampire bat. (D.G. Constantine)
22 Bat Rabies and Other Lyssavirus Infections

The most effective way to determine if infected bats 

exist in an area is to test clinically ill and dead bats. This type 

of surveillance for lyssavirus infections is often required for 

protection of the public health. Although the prevalence of 

infection among clinically ill bats varies from species to species in the USA, it averages 10 percent, which is similar to the 

8 percent prevalence reported in Europe. To find at least one 

infected bat, one must test a minimum of 22, 29, or 44 clinically ill bats for statistically reliable results at confidence levels of 90, 95, or 99 percent, respectively (Constantine, 1994).

Studies of some bats indicate that they undergo population fluctuations in response to mortality caused by lyssavirus infection, similar to the occurrence of the infection 

in carnivores such as foxes and skunks (Constantine, unpub. 

data). Mortality decreases the population, and accordingly, 

evidence of subsequent disease until the population increases 

to a density that again supports lyssavirus transmission 

through increased frequency of contact. Increased rainfall can 

hasten population growth by providing greater plant growth 

to support the development of bat food, such as insects, and 

thus the greater survival of bats. Some migratory bats will 

forsake areas of drought for more suitable places. At the other 

extreme, however, atypically prolonged rainfall retards the 

flight of both insects and bats and is known to cause mass 

starvation of some bat species (Constantine, 1982a).

Temperature and Rabies Virus Virulence

Temperature can be critical in the manifestation or control of diseases, including rabies. The hypothermia of hibernation increases the resistance of an animal host to various 

viral, bacterial, and metazoan parasites. In many instances, 

disease is prevented or diminished. In other instances, disease 

development is deferred during hypothermia and resumes 

when the host awakens. In addition, viruses usually cannot 

replicate at higher temperatures, and fever in the infected host 

serves to depress viral development. However, certain highly 

pathogenic virus strains have adapted to this condition and are 

able to replicate at higher temperatures. Lower body temperatures may favor less virulent rabies virus strains. The seasonal 

differences in the body temperatures of heterothermic bats 

may account for some virulence differences observed among 

the rabies virus variants from different bat species (Constantine, 1967a; Constantine,1970).

How Colonial Bats Avoid Extinction by 

Lyssaviruses

It would seem that highly colonial animals, such as many 

bat species, would be eliminated quickly after a lyssavirus 

enters the population, but multimillion-bat colonies persist in 

the presence of the disease. Studies of rabies in huge populations of Brazilian free-tailed bats in caves of the southwestern USA and Mexico provide some insight into this question (Constantine and others, 1988a). In essence, maternal 

antibody provides a protective barrier or passive immunity

against disease upon virus exposure within the colonies. Subsequent virus exposure then stimulates the immune system 

to provide protective antibody or acquired immunity for the 

long-term protection of exposed bats (Box 6).

Paralytic and Furious Disease Courses 

The author’s early experiences with bats and bat rabies 

in North America indicated that rabid bats experienced only 

the paralytic form of the disease without obvious aggressiveness or furiousness. For example, he and coworkers spent 

the equivalent of many years in and about caves occupied by 

millions of Brazilian free-tailed bats and were never attacked, 

although one of every 200 bats was infected (Fig. 9). In 

addition, none of the thousands of bats of other colonial bat 

species attacked under similar circumstances. Logic suggested that sufficient biting or other forms of transmission 

had occurred normally and frequently enough among these 

crowded animals to have rendered the furious form of rabies 

relatively superfluous and perhaps obsolete as a viral survival 

mechanism.

However, lyssavirus-infected bats do attack. Initial 

reports of attacks by flying rabid bats were limited to certain species that live as solitary individuals. These included 

the more diminutive of North American bats, primarily the 

canyon bat (Fig. 10), and, less often, the western small-footed 

myotis (Fig. 11) and the California myotis (Fig. 12) (Constantine, 1967a). It was suspected that deliberate attacks by these 

small solitary bats on conspecific bats had been retained as 

a necessary viral survival mechanism. Eventually, however, 

attacks on people by individual rabid bats of various species, 

including Brazilian free-tailed bats and other colonial bats, 

were documented. Fortunately, such attacks are rare events 

that are committed by individual rabid bats separated from 

colonies that otherwise would seem to dominate their interest. 

Thus, unwary investigators were likely spared attacks because 

they could not compete with the noisy bat colonies for the 

attention of any rabid bats primed for attack.
Immunity to Lyssaviruses in Colonial Bats 23

Immunity to Lyssaviruses in Colonial Bats Box 6

Studies of rabies in huge populations of Brazilian 

free-tailed bats in caves of the southwestern USA 

and Mexico revealed that rabies infection is present 

in about 0.5 percent or 1 of every 200 of the mature 

bats (Constantine and others, 1968a). Disease 

transmission by infective saliva was possible from 

bites, licking of saliva into wounds or mucosal 

surfaces, or contamination by or inhalation of cave 

air charged with droplets of saliva expelled by the 

chattering bats (Constantine, 1967a; Constantine, 

1967b). As many as 80 percent of the bats examined 

possessed recognizable blood serum antibodies 

against the virus. Colonial bats develop life-long 

immunity to rabies virus by two mechanisms. Unborn 

and suckling bats receive a temporarily effective 

dose of antibodies by way of the placenta and milk 

(passive immunity), but this immunity is typically 

lost 6 to 10 weeks following birth (Constantine and 

others, 1968a). Many young bats, while they are still 

temporarily immune, are also likely exposed to virus, 

which they resist by passive immunity, but against 

which they consequently developed long-lasting 

antibodies (active immunity). Such bats would be 

expected to respond to future viral exposures as if 

they were receiving booster doses of rabies vaccine. 

Young bats killed by rabies within the colonies were 

presumably produced by non-immune mothers. This 

scenario provides a mechanism for these populations 

to survive in the presence of rabies virus. The virus 

“immunizes” most of the population, but it still claims 

sufficient victims to maintain disease within the 

colony so that the virus persists.

Figure 9. Brazilian free-tailed bats 

attempting to find roosting space among a 

dense colony of cohorts on a cave ceiling 

(bat density equals approximately 300 bats 

per square foot). (D.G. Constantine)

The nasal mucosa may be a site of rabies entry and exit 

from victims in some highly congested bat caves. This 

2-micron section of nasal mucosa from a symptomatic, 

immature Brazilian free-tailed bat (Tadarida brasiliensis) 

from the Frio Cave, Uvalde County, Texas, was stained 

with fluorescent dye that indicates the presence of rabies 

antigen, shown by the apple green areas. 
24 Bat Rabies and Other Lyssavirus Infections

Figure 11. Western small-footed myotis bat in flight. (D.G. Constantine)

Figure 10. Canyon bat in flight. (D.G. Constantine)

Figure 12. California myotis bat in flight. (D.G. Constantine)

Figure 13. Big brown bat in flight. (D.G. Constantine)
Disease Ecology 25

Viral and Host Species Diversity

Current information suggests that each species of lyssavirus may be divisible into several or more strains called variants. Each variant is maintained within populations of a particular reservoir host species to which it is adapted. Typically, 

these hosts are isolated by ecologic, anatomic, and social 

factors from other host species. Further isolation is afforded 

to viral variants harbored within sedentary, nonmigratory 

host species. Recent studies indicate that some sedentary 

reservoir host species populations are divisible into different 

DNA-identifiable host subpopulations that occupy different 

extensive, although often contiguous, geographic areas. Thus, 

these subpopulations provide conditions of isolation that 

favor the development of unique viral variants. For example, 

such a geographic correlation between nonmigratory big 

brown bat (Fig. 13) populations and the distributions of the 

rabies virus variants they maintain has been observed in the 

USA (De Mattos C.C. and others, 2001). The nonmigratory 

common vampire bat rabies virus is similarly divided into at 

least three geographically distinct variants (De Mattos, C.A. 

and others, 2001). In contrast, the Brazilian free-tailed bat, 

whose migratory habits conflict with population isolation and 

differentiation (McCracken and others, 1994), harbors fewer 

and less contrasting variants (Smith, 1996).

Although the association of a given virus variant with 

a particular animal host species can be useful for tracing the 

origin of infection in atypical victims, such as humans, it 

cannot rule out intermediate hosts. For example, cats that 

chew infected bats may subsequently transmit the virus to 

people, either promptly by mechanical transmission or perhaps later if the cat develops rabies. However, the relatively 

rabies-resistant cat may not become infected, eliminating it 

as a suspect intermediate host (Constantine, 1967a; Bell and 

Moore, 1971). Thus, these early studies indicated that rabies 

viruses recovered from various bat species differed in their 

abilities to infect or produce (or both) particular pathological 

responses in a given spectrum of wild carnivore or laboratory 

mammal species. These studies foreshadowed later serological methods followed by protein- and RNA-based analyses 

that now are used to delineate virus variants. The original 

discoveries of virus variants multiplied the challenge of 

defining the public health significance of bat rabies, but at the 

same time, they helped clarify it. Nonetheless, the challenge 

continues to grow by disclosures of rare-to-frequent transmission of variants from one host species to another, sometimes 

with serial transmission thereafter, as described below in 

the section on intraspecies and interspecies transmission. 

Such host interspecies transfer of virus raises the specter of 

potentially unwelcome consequences, such as a change from 

a predominantly paralytic disease to unabated furious disease 

and consequent attacks.

Intraspecies and Interspecies Transmission 

of Bat Lyssaviral Variants

Lyssaviral variants can be transmitted between bats 

of the same species (intraspecies) or among bats of different species (interspecies). Intraspecies transmission seems 

evident when infection with like variants is present in both 

mother and suckling bats in the absence of contact with other 

sources of virus. Such conditions usually prevail in multimillion aggregations of Brazilian free-tailed bats. Similarly, 

the northern yellow bat, a solitary species that takes shelter 

in tree foliage, has been found rabies infected with its litter 

of three rabies-infected suckling offspring (Schneider and 

others, 1957). Also a lactating eastern red bat and her litter 

of four were all rabies infected (Birney and Rising, 1967). 

Newborn bats might be infected by their mother as she severs 

the umbilical cord, by licking infective saliva onto mucosal 

surfaces, by bites, or by aerosol in some environments. 

Conspecific mature bats probably are infected by various 

similar methods as they crowd together in shelters or interact 

at feeding or drinking sites. However, more than one colonial 

species sometimes live in the same shelter, where single or 

multiple interlopers may crowd into another species’ cluster 

(Figs. 14 and 15), possibly to maintain body heat, whereupon 

bites might be exchanged. Thus, interspecies transmission of 

virus seems a logical source of virus variant differentiation 

within the recipient species over time (Box 7).

Although lyssaviruses usually are transmitted by bite, 

alternate exposure routes include aerosols, ingestion, and by 

contamination of wounds or mucosal surfaces with infective 

saliva or tissues (Johnson, 1965; Constantine, 1988a). These 

viruses may survive through additional possible and probable transmission routes, including insects. Although Mokola 

lyssavirus has been shown to replicate in inoculated mosquitoes, biological transmission by arthropods of that or other 

lyssaviruses has not yet been demonstrated.

Mechanical transmission by arthropods however, seems 

likely in certain situations. Dermestid beetles and their larvae, 

for example, consume the tissues of live or dead rabid bats in 

huge cave colonies of Brazilian free-tailed bats (Fig. 16), and 

the author has demonstrated rabies virus on the exteriors of 

both life stages of beetles recovered from the skull cavities of 

bats (Constantine, unpub. data). Adult beetles and larvae seeking flesh to eat will attack living or dead mammals, including 

humans. In so doing, the contaminated insects may be able to 

mechanically transmit the virus. The abdominal contents of 

beetles that feed on bats are known to be consumed by assassin bugs, which routinely attack mammals, including humans, 

for blood meals. Such rabies-contaminated bugs may also be 

able to mechanically transmit virus in this manner. Various 

mites, hordes of which parasitize bats, include species that 

consume fluids available at the edges of mouths, eyes, or 

within the nostrils or ears of bats as well as of human visitors 

to bat caves. Thus, these feeding habits may provide opportunities for interspecies mechanical transmission of virus.
26 Bat Rabies and Other Lyssavirus Infections

Figure 15. A Brazilian free-tailed bat (blue-grey) 

and a Yuma myotis bat (brown) cooperating to 

conserve body heat. (D.G. Constantine)

Figure 16. A dead Brazilian free-tailed bat being consumed 

by dermestid beetles commonly found in association with 

bat colonies. (D.G. Constantine)

Figure 17. Brazilian free-tailed bat. (D.G. Constantine)

Figure 14. A mixed group of pallid (light-colored) 

and Brazilian free-tailed (blue-grey) bats in an attic. 

(D.G. Constantine)
Disease Ecology 27

Table 8. Examples of vampire bat rabies variants found in other species in Latin America.

Country Date Findings

Brazil 1998–2001 22 of 23 dog and cat rabies cases tested (Kotait and others, 2001).

2000 32 of 53 non-vampire bat rabies cases tested, representing 16 different bat species (Silva, 2001).

2001 A cat previously in contact with a bat developed rabies and infected its owner (Kotait and others, 2001).

Costa Rica 2001 An aggressive household cat bit two people who died of rabies of vampire bat origin (Badilla and others, 2003).

Mexico 1999–2001 Fruit bat and dog (Vargas Pino, 2001; Velasco Villa and others, 2001).

2004 Child bitten by cat (Tesora Cruz and others, 2006).

The development of methods to recognize and classify 

viral variants by their host species of origin has helped to 

retrospectively identify the infection source, such as a particular species of bat or carnivore, putatively responsible for 

infecting a human victim (Rupprecht and others, 1991; Smith, 

1989). Recent reports based on these techniques also indicate 

that a large amount of interspecies lyssavirus transmission has 

occurred between mammal species, especially from vampire 

bats to other bats and carnivores.

Latin America

Vampire bat rabies virus has been isolated from other 

bats and from carnivores in Latin America (Table 8). These 

findings indicate direct transmission of rabies by vampire bats 

or by other bats that might be serially transmitting the vampire rabies variant. Moreover, where Brazilian rabies vaccination campaigns had brought urban canine and feline rabies 

under control, the finding of a vampire rabies variant instead 

of a canine rabies variant as the cause of recent rabies cases in 

dogs and cats (Table 8) indicates that the vampire bat variant 

is transmitted serially among pets.

Reports of interspecies transmission of rabies by bats 

other than vampires refer nearly exclusively to insectivorous species. In Chile, where five rabies virus variants were 

studied, the Brazilian free-tailed bat (Fig. 17) was considered 

to be the source of a variant found in one human, dogs, cats, 

cows, and a pig. A variant typical of Western red bats or other 

members of the genus Lasiurus was recovered from a red bat, 

three Brazilian free-tailed bats, and a dog. It was suspected 

that livestock were infected by dogs or cats that had acquired 

the infection by attacking grounded bats, but none of the 

identified variants closely resembled variants associated with 

vampire bats (De Mattos, C.A. and others, 2001; Favi and 

others, 2002).

North America

There have been 51 known or presumed bat-transmitted 

human rabies infections in the USA (Messenger and others, 

2002; Krebs and others, 2003; Krebs and others, 2004; 

Blanton and others, 2006) and Canada (Dempster and others, 

1972; Varughese, 1983; Webster and others, 1987; Krebs and 

others, 2001; Krebs and others, 2004) from 1951 to 2006. 

Human cases were attributed to bat species based on bat 

carcass identification or prior association of rabies variants 

with a given bat species (Table 4). Bat carcasses were often 

unavailable for species identification and for comparison 

with identified virus variants. In cases where bat carcasses 

were available, tricolored bat virus variants were identified 

from a big brown bat, indicating interspecies transmission, 

and from a bat retrospectively identified by an untrained 

observer as an eastern red bat (Hardman and others, 1995). 

However, red bats and tricolored bats have similar pigment 

distribution in their wing membranes and could be confused 

with one another. In studies of rabies in Alaskan bats (Blanton 

and others, 2007), a little brown myotis was infected with a 

rabies variant associated with silver-haired bats, and a Keen’s 

myotis was infected with a variant associated with eastern 

red bats. The frequent absence of a bat contact in the human 

victim’s history also leaves open the possibility that an intermediate host, such as a cat that had chewed a rabid bat, might 

be the source of some human infections.

Transmission of rabies from bats to individual wild or 

domestic carnivores and livestock in the USA was implied in 

various earlier reports based on virus variant characteristics 

(Rupprecht and others, 1991). Notwithstanding the possibility of viral transmission by intermediate hosts, 28 animals 

(11 foxes, 1 skunk, 7 cats, 2 cows, 6 horses, and 1 sheep) 

were suspected of having been infected by 12 big brown bats, 

8 eastern red bats, 2 Brazilian free-tailed bats, and 6 bats 

of undetermined species (Baer and Smith, 1991). A survey 

in Colorado designated a rabies variant identified from big 

brown bats as the infective agent in three skunks and one 

fox (De Mattos, C.C. and others, 2001). A study of rabies 

virus variants obtained from 19 skunks during a skunk rabies 

outbreak in Arizona indicated that those variants were nearly 

identical to a virus variant commonly found in big brown 

bats, the suspected source of the outbreak (Leslie and others, 

2006). These results provide possible evidence of serial trans-
Brazilian free-tailed bat bitten and bled to death in captivity by a 

common vampire bat cage mate. (D.G. Constantine)

28 Bat Rabies and Other Lyssavirus Infections

Box Bat Versus Bat—Interspecies Transmission of Bat Lyssaviruses 7

mission of rabies among North American carnivores following acquisition of virus from bats. In addition, these outbreaks 

contrast with earlier single isolated occurrences of evident bat 

rabies variant transmission to carnivores in areas otherwise 

free of rabies in carnivores (Smith and others, 2001).

Europe

Few interspecies transmissions of lyssaviruses from bats 

have been reported in Europe, however fewer investigations 

have been done in Europe than in other regions. The five 

European human deaths of European bat lyssavirus (EBL) 

origin (Table 5) were mentioned earlier: one from EBL-2 in 

Scotland (Fooks and others, 2002; Fooks and others, 2003), 

one from EBL-1 in Russia (Selimov and others, 1989), one 

from EBL-2 in Finland (Lumio and others, 1986), and two 

from uncharacterized lyssaviruses in the Ukraine (Selimov 

and others, 1989; Botvinkin and others, 2005). In addition, 

three domestic sheep were reported infected with EBL-1 in 

Denmark (Stougaard and Ammendrup, 1998a; Stougaard 

and Ammendrup, 1998b), and a stone marten was infected 

with the same virus in Germany (Müller and others, 2001). 

It seems improbable that insectivorous bats would attack 

sheep, and it is possible that the virus was transmitted by a 

carnivore, such as a fox or cat that was infected as it attacked 

a grounded bat. The EBL-1 variant also was isolated from 

nonnative Egyptian fruit bats in zoos that had previously 

exchanged animals with Denmark, the Netherlands, and Belgium (Ronsholt and others, 1998). Presumably the infection 

originated from a native bat. The EBL-1 virus is most often 

associated with the serotine bat, although it is known from at 

least three other bat species in Europe (Appendix D).

The EBL-2 virus is known from two bat species, the 

pond bat in Denmark and the Netherlands and Daubenton’s 

bat in Germany, Denmark, Switzerland, and England and 

Scotland in the United Kingdom. Transmissions of lyssaviruses from bats to other mammals seem to be rare in Europe, 

where most infected bats have been found in areas apparently 

free of infection in other species, comparable to many areas 

of North America.

In 1957, before rabies virus variants were 

known to exist, the author sought bat 

shelters in Mexico that were occupied 

by common vampire bats and migratory 

Brazilian free-tailed bats from the USA. 

The purpose of that inquiry was to explore 

the possibility that vampire bats might 

occasionally attack and drink the blood of 

insectivorous bats, as had been observed 

among captive bats. Such behavior would 

facilitate virus transmission from infected 

vampire bats. Alternatively, vampires might 

be infected by saliva-contaminated air in 

some free-tailed bat cave shelters or by 

retaliatory bites, and may be infected by 

free-tailed bat lyssavirus variants.

Several caves were found to be occupied 

by the two bat species. About 200,000 of 

the migratory free-tailed bats were in a 

large but shallow cave that also harbored 15 common 

vampire bats in a deep side tunnel. The free-tailed 

bats had little protection from weather, which if cold 

would render them lethargic and helpless, and if wet 

and stormy would discourage their evening departure 

and the departure of the vampires. Either condition 

would make the free-tailed bats readily available to 

the vampires, which die if they do not feed for 3 

nights (McNab, 1973). Lyssavirus variants from the 

two species were later determined to be closely 

related (Franka and others, 2006). Other workers 

also found that captive vampire bats readily fed on 

insectivorous as well as frugivorous bat cagemates, 

and a frugivorous little yellow-shouldered bat in 

South America had a similar bite wound when it was 

captured in a cave shared with vampire bats (Lord and 

others, 1973).
Disease Ecology 29

Africa

Four lyssavirus species are currently known from mammals in Africa: classical rabies, Duvenhage bat, Lagos bat, 

and Mokola. Each of the four is divisible into variants, and 

the existence of multiple hosts for each virus species suggests previous interspecies transmission. Classical rabies is 

present in humans and in domestic and wild mammals, but it 

has not been identified in bats outside of the Americas. Only 

Duvenhage and Lagos bat lyssaviruses are currently known 

in African bats. In 1970, 2006, and 2007 Duvenhage bat 

virus caused the rabies-like deaths of two men in South Africa 

(Meredith and others, 1971; Paweska and others, 2006), and 

of a woman in Kenya (van Thiel and others, 2008). Each of 

these people were bitten or scratched by unidentified bats, 

although the bat in one case was said to have resembled the 

insectivorous long-fingered bat. Duvenhage bat virus has 

since then been recovered in South Africa from another bat 

resembling that species (Paweska and others, 2006) and in 

Zimbabwe from the insectivorous Egyptian slit-faced bat 

(Foggin, 1988). Additionally, an incompletely characterized lyssavirus was found in an unidentified bat in Zambia 

(Ahmadu and Zulu, 1998).

Lagos bat virus is known in straw-colored fruit bats in 

Nigeria and Senegal, Peters’ lesser epauletted fruit bat in the 

Central African Republic, Wahlberg’s epauletted fruit bat in 

South Africa, and in the insectivorous Gambian slit-faced bat 

in Senegal. In addition, it was reported in a domestic dog in 

Ethiopia, two domestic cats in South Africa and Zimbabwe, 

and a water mongoose in South Africa (Markotter and others, 

2006a; Markotter and others, 2006b). Experimental infections of Lagos bat virus were demonstrated in dogs and rhesus 

monkeys (Percy and others, 1973). No human infections are 

known, but during a Lagos bat virus outbreak among fruit 

bats in South Africa, a motorcyclist was pursued and bitten 

on his face by a bat, which escaped. The bite victim received 

antirabies prophylaxis and did not develop clinical disease.

Mokola lyssavirus has not been confirmed in bats. Two 

human cases were identified in Nigeria; one of the victims 

survived (Familusi and others, 1972). The virus has also been 

isolated from an insectivorous rusty-bellied brush-furred rat 

in the Central African Republic (Saluzzo and others, 1984), 

a domestic cat in Ethiopia (Mebatsion and others, 1992), and 

domestic dogs and cats in Zimbabwe and South Africa (Foggin, 1982; Foggin, 1985; Sabeta and others, 2007). Mokola 

was also found in insectivorous Crocidura shrews in Nigeria 

and Cameroon. An association between shrews and bats is 

evident from observation of a Crocidura shrew feeding on an 

Angolian free-tailed bat within an attic colony (Osborne and 

Osborne, 1981). Experimental infections were demonstrated 

in genets, a mongoose, a jackal, dogs, and rhesus monkeys 

(Foggin, 1985; Percy and others, 1973).

Asia

Human lyssavirus infections from uncharacterized lyssavirus species were reported in two people previously bitten 

by bats; one infection was in a person in India who handled 

a disabled bat (Veeraraghavan, 1955) and the other was in a 

person in China, where a survey later disclosed infections in 

3 of 300 unidentified bats (Tang and others, 2005). Similar 

infections were reported in the greater horseshoe bat in Turkey, the northern bat and Daubenton’s bat in the Asiatic part 

of the Russian Federation, and the Indian flying fox in India. 

Aravan lyssavirus was discovered in the lesser mouse-eared 

bat in Kyrghystan (Arai and others, 2003; Botvinkin and others, 2003), and Khujand bat lyssavirus was recovered from 

the whiskered bat in eastern Siberia. However, a 1967 report 

of rabies infection in two bats in Thailand (Smith and others, 

1967) was later reported to the author by the laboratory of 

origin to have likely resulted from laboratory contamination 

rather than originating from the bats in question.

A survey of healthy bats in the Philippine Islands failed 

to yield lyssaviruses from 821 insectivorous or frugivorous

bats, which is similar to the results from earlier surveys of 

clinically normal bats globally, but 9.5 percent of 231 bats 

had antibodies against Australian bat lyssavirus (Arguin and 

others, 2002). Similarly, no lyssaviruses were recovered from 

212 healthy bats in Bangladesh, but 3 of 127 Indian fruit bats 

had antibodies against the bat lyssaviruses Aravan and Khujand (Kuzmin and others, 2006). Sampling of bats in Thailand 

indicated that 15 of 335 Lyle’s flying fox fruit bats and 1 

of 45 lesser dawn fruit bats had antibodies against Aravan, 

Khujand, Irkut, or Australian bat lyssaviruses (Lumbertdacha 

and others, 2005). The survey results were interpreted as evidence of the presence of lyssaviruses but of a low prevalence 

of infection.

Australia

Australia was considered free of lyssaviruses until 1995, 

when a new lyssavirus species (genotype 7), now known as 

Australian bat lyssavirus, was first observed to produce signs 

of aggression and, ultimately paralysis and death in black 

flying foxes (Fraser and others, 1996). Some infected bats 

repeatedly and severely attacked and bit others of the species, including the offspring of one attacker. By 1999, surveys 

found the virus in 42 flying foxes, including members of 3 

additional species: the little red flying fox, the gray-headed 

flying fox, and the spectacled flying fox. In addition, an Australian bat lyssavirus variant was found in four yellow-bellied 

sheathtail bats (Warrilow and others, 2003). Two female bat 

caregivers are now known to have died of the disease, one 

from the sheathtail bat variant and the other from a flying fox 

variant (Mackenzie, 1999). As mentioned above, antibodies 

against Australian bat lyssavirus were detected in bats in the 

Philippine Islands and in Thailand, indicating distribution 

of the virus well beyond Australia (Arguin and others, 2002; 

Lumbertdacha and others, 2005). The existence of multiple 
Spreading infection and mortality

Population recovered:

high frequency of contact

Infected bat: Immigrant 

or maturing incubation

Population increase: 

favorable environmental conditions

Population decline:

low frequency of contact

30 Bat Rabies and Other Lyssavirus Infections

variants of this virus in various host species suggests past, if 

not continuing interspecies transmission, the latter demonstrated by the human cases.

Environmental Persistence

Rabies mortality in bats usually goes unnoticed because 

many bats die in their shelters, while others fall with folded 

wings into dense vegetation. Even multiple mortalities may 

hardly be noticeable for similar reasons. In some ways, bat 

mortality from rabies is comparable to a grass fire. The virus 

is represented by a burning match and the host mammal 

population by a field of dry grass. If a lit match drops where 

the grass is dense, a substantial and noticeable fire results, 

destroying nearly all of the grass. In time the grass regrows 

and the process may be repeated. However, if the grass 

blades are widely separated, only one or a few blades may 

burn unnoticed – the density of the grass being the key factor. Similarly, if a bat population is dense, it supports rabies 

transmission between individuals more readily, and thus the 

population is reduced (Constantine, unpub. data). Rabies 

infections may not be obvious in less dense populations.

Cyclic fluctuations and environmental persistence of 

bat populations and lyssaviral infections are influenced by 

several main factors (Fig. 18). A favorable climate is required 

to produce the plant base of the bat food supply: insects for 

insectivorous bats, fruit for frugivorous bats, and cattle as the 

blood source for vampire bats. Additional factors, differing 

by bat species, include such things as antibody systems and 

the amount of antibodies in the body, solitary or colonial 

habits, hibernation or migration habits, the numbers of young 

or litters produced per year, and alternate virus transmission 

routes. In addition, prolonged incubation periods provide a 

mechanism for virus persistence during the winter (Box 8). 

The overall effect of natural lyssavirus-caused mortality in 

bats may contribute to keeping host populations and food 

resources in balance without doing irreversible harm to either. 

This “balancing act” works despite some ghastly aspects of 

the scenario.

Figure 18. Factors involved in cyclic 

fluctuations and environmental persistence of 

bat populations and lyssaviral infections.
Lyssaviruses—Deadly Hitchhikers 31

Lyssaviruses—Deadly Hitchhikers Box 8

Long incubation periods, in some cases extended 

by hibernation, enable rabies virus and other 

lyssaviruses to survive within seemingly 

healthy bats for more than 7 months before the 

appearance of clinical signs of disease. During 

this period of apparent dormancy, a migratory 

bat may transport a virus along both legs of 

its journey of as many as 4,000 kilometers. For 

example, a big brown bat that was captured in 

North America before winter hibernation, and kept 

active and isolated all winter while fed in captivity, 

developed rabies 7 months later or 1 month 

after it would have awakened from hibernation 

(Moore and Raymond, 1970). Two Townsend’s 

big-eared bats of North America, awakened from 

hibernation in mine tunnels in New Mexico and 

adapted to captivity in a warm environment, 

developed the disease nearly 3 months later, or 

about 7 months after they entered hibernation 

(Constantine, 1967a). Brazilian free-tailed bats 

that were caught in New Mexico and similarly 

held captive after they otherwise would have 

migrated southward died of rabies after periods 

of 7 days to 3 months (Constantine, 1967a). These 

examples illustrate methods by which bat rabies 

virus persists overwinter in the apparent or actual 

absence of bats.

Big brown bat. (D.G. Constantine)

Townsend’s big-eared bat hovering over water 

with tongue extended to drink.

(D.G. Constantine)
32 Bat Rabies and Other Lyssavirus Infections

or quarantined after biting bats were discarded or escaped. 

A cycle of near-hysteria developed as poisons in typically 

unclosed bat harborages continued to act, sickening replacement bats for many years, leading to further poison applications and more frightening news coverage. Finally, efforts by 

those who understood these events prevailed over panic and 

politics, and toxicant use fell into disrepute (Tuttle, 1988).

Bats usually take shelter in elevated sites from which 

they can easily take flight (Figs. 19 and 20). They are noticeable due to their squeaking vocalizations, departures and 

arrivals, accumulations of droppings under their entry sites, or 

odors. Bats ill with lyssavirus infections may occasionally fall 

from these colonies (Fig. 21), whereupon predators such as 

cats, or curious people, are attracted to them resulting in bites 

and disease exposures. Whereas uninformed or fearful people 

often assume that destruction of bat colonies is the evident 

solution, that act is counterproductive, and a compromise 

such as excluding rather than killing the bats may benefit both 

bats and humans.

The exclusion of bat colonies and the sealing of bat 

shelters to prevent reentry are sometimes appropriate, 

especially when large colonies are located in school buildings where children may pick up fallen bats and be bitten. In 

another example, hundreds of bats were lodged in slots under 

an urban concrete bridge, from which boys pried dozens 

of young, flightless bats and threw them through the open 

windows of passing automobiles (Constantine, unpub. data). 

Ordinarily, exclusion of bats should not be undertaken until 

all of the young bats can fly and join mature bats on nightly 

foraging flights, which usually occurs by late summer or 

early fall. Otherwise, the young bats will be sealed inside the 

shelter to die and decompose, or they may crawl into building 

interiors where human and pet contacts may result. Exclusion is achieved by sealing all but the main exit and entry 

passages and arranging simple one-way valve-like systems 

over the exits, permitting bats to leave but preventing re-entry 

(Fig. 22) (Constantine, 1982b; Frantz, 1986).

Points to Ponder: Interspecies 

Interactions in Potential Bat Rabies 

Transmission Settings

Intertwined with lyssavirus disease ecology are the interspecies interactions that develop when humans, their domestic 

animals, or other animals invade or otherwise interact in a setting where the potential for disease transmission exists. Cats 

play an especially important role in bat lyssavirus exposures 

(see Box 9 and Disease Prevention and Control).

Bat Colonies in Manmade Structures

Insectivorous bats have long provided services to mankind by controlling insects that attack and spread diseases to 

humans, livestock, crops, and forests (Constantine, 1970). 

Furthermore, vegetarian bats, as the sole pollinators and seed 

distributors of many major tropical food plants (Constantine, 

1970), have and continue to make possible human and other 

life in tropical areas. Fear of bat rabies, however, has often 

resulted in the persecution of bats associated with manmade 

structures, and this fear has generally been fed by a lack of 

understanding of the ecology of bats.

Upon the discovery of bat rabies in the USA in 1953, 

frightening news stories repeatedly appeared, alarming 

homeowners and others whose buildings sheltered bats. Not 

knowing what to do but agreeing with public demands that 

killing the bats seemed appropriate, government agencies 

readily obliged by turning their mosquito control DDT foggers against bats and encouraged others to do likewise. Other 

toxicants were used after DDT was banned (Tuttle, 1988). 

This reaction resulted in widespread grounding of poisoned 

bats, which resembled rabid bats. An approximately tenfold 

increase in bitten people and pets followed, and most people 

received antirabies treatment and pets either were destroyed 

Figure 19. An evening foraging 

flight of Brazilian free-tailed bats. 

(D.G. Constantine)
Points to Ponder: Interspecies Interactions in Potential Bat Rabies Transmission Settings 33

Figure 20. Brazilian free-tailed 

bats roosting under a railroad 

trestle (inset). (D.G. Constantine)

Figure 21. A dead Brazilian free-tailed bat lying on the guano-covered floor of a bat cave. (D.G. Constantine)
34 Bat Rabies and Other Lyssavirus Infections

Box Domestic Cats and Lyssavirus Transmission 9

It was easy to comprehend the extreme concern of a panicked 

mother who found her baby mouthing a bat delivered by the family cat 

(Constantine, unpub. data). In areas of the world where domestic cats 

are popular as pets or for rodent control, they are the primary predators 

of bats. The irrepressible hunting habits of domestic cats---and their 

practice of bringing home their victims---accounts for the great majority of 

contacts between people and bats. In the USA, approximately 10 percent 

of bats that are home-delivered by domestic cats prove to be infected 

with lyssavirus.

which it can launch itself into flight. Alternatively, 

especially if the bat is ill, it may remain hidden during 

daytime and initiate its upward climb after dark. The 

bat may climb blankets onto a bed or be chilled by 

the night air and seek a warm place, possibly next 

to the body of a sleeping human. As the person 

turns during the night and makes contact with the 

bat, it may inflict a retaliatory bite that might not 

awaken the sleeper. This scenario has occurred many 

times, involving infected as well as uninfected bats, 

and human rabies mortality likely has resulted. Six 

such rabies exposures are recorded in California 

(Constantine, unpub. data).

Cats generally bite and chew to stun, kill, or eat a 

bat, and if the bat is infected then the interior of the 

cat’s mouth can become contaminated with virus. 

Two relevant hazards can result: (1) the contaminated 

mouth of the cat may temporarily have the ability 

to mechanically transmit the virus by bite, by licking 

the virus into even slight wounds or onto mucosal 

surfaces, or by scratches with claws (Tabel and 

others, 1974) that were licked previously during 

cleaning or preening; (2) the cat may ultimately 

develop the disease, especially if it was not 

previously vaccinated, often vanishing to die in hiding 

as overt signs appear. Twenty-six likely human rabies 

exposures in California resulted from people being 

bitten by cats while they were trying to extract rabid 

bats held between the cat’s teeth (Constantine, 

unpub. data). The possibility also exists for virus 

transmission through aerosolized oral or nasal fluids 

expelled as a cat sneezes, which is a demonstrated 

route of transmission for plague (Gage and others, 

2000). Variations of such disease exposures could 

follow unobserved encounters between cats and 

infected bats and may account for many of the 

unexplained human lyssavirus deaths involving virus 

variants characteristic of bats but lacking histories 

involving bats.

Cats readily jump fences to forage widely, whereas 

dogs are restricted to yards or tethered where they 

may be left with a bat courtesy of the family cat. 

Reports from 1 California county over a 14-year period 

revealed that 1 or more resident cats were found with 

428 (66 percent) of the 649 seemingly ill, injured, 

or dead bats that appeared at residences. Dogs and 

cats were present together with 26 of the bats (4 

percent), but only dogs were found with 52 of the 

bats (8 percent), an unknown number of which may 

have been delivered by resident cats (Constantine, 

unpub. data).

As mentioned elsewhere, most individual temperate 

zone bats become inactive when they are resting, 

are ill or injured, or as a consequence of cool ambient 

temperatures. These bats may appear to be dead 

and are easy prey for cats. A cat sometimes enters 

a residence with an unnoticed bat in its mouth, 

and having delivered its catch, the cat usually loses 

interest in the seemingly lifeless bat and leaves it on 

the floor. Later, as the bat warms, it seeks a vertical 

surface where it can climb to safety or a site from 

(Photo by M. Friend)

A Lyle’s flying fox 

in Thailand climbs 

from the ground 

and up a tree 

trunk to reach the 

colonial roosting 

site, demonstrating 

the climbing 

behavior of bats. 

(D.G. Constantine)
(Photos by M. Friend)

Domestic Cats and Lyssavirus Transmission 35

Through variations of the scenarios described above, 

domestic cats are likely intermediate rabies hosts or 

vectors between bats and humans. Cats are relatively 

resistant to rabies; a cat exposed to rabies may never 

develop the disease and thus be above suspicion. 

In laboratory experiments, striped skunks could be 

infected by eating the carcass of a single mouse 

infected with rabies virus of silver-haired bat origin, 

but cats fed 25 such carcasses were not infected 

(Bell and Moore, 1971). Moreover, a cat may eat an 

infected bat and return home with a contaminated 

mouth and claws. Alternately, a cat may eat a rabid 

bat after it has bitten a person, thus precluding the 

bat from being tested and consequently discouraging 

the human bite victim from seeking treatment. This 

happened in Wisconsin in 1959, and it resulted in a 

documented human rabies mortality (Humphrey and 

others, 1960).

Rabies in cats is a “spillover” or incidental infection 

transmitted to a cat by a 

reservoir host: a rabid dog, wild 

carnivore, or bat. In the USA, 

annual reports of rabid cats 

ranged from 200 to 500 cases 

per year between 1938 and 

1960 (Rupprecht and Childs, 

1996). During this same time 

period, however, dog rabies 

cases typically outnumbered 

cat cases by a factor of 10 or 

more (Rupprecht and Childs, 1996). Canine rabies 

vaccination campaigns initiated in the 1950s led to 

a drastic reduction in dog cases, but at the time, far 

less emphasis was placed on cat rabies vaccination. 

Thus by 1981, as dog rabies came under control, cat 

rabies cases increased so that they exceeded dog 

cases and they have maintained their lead since. 

This trend may be explained by the popularity of 

cats as pets, by their relative risk of encountering 

rabid animals when allowed to roam freely, and 

by difficulties in the enforcement of mandated cat 

rabies vaccination (Rupprecht and Childs, 1996). In 

addition, since the discovery of bat rabies in the USA 

in the early 1950s and the consequent alarm when 

cats bring home rabid bats, increasing numbers of 

cats have been sacrificed and tested for rabies. The 

relative rabies resistance of cats, coupled with the 

lower rabies dose received from a typical bat bite, 

probably results in fewer cat rabies infections from 

bats than from bites that roaming cats may receive 

from rabid skunks, foxes, or raccoons. However, 

this does not diminish the danger to people from 

mechanical or biological transmission of rabies by 

pet cats.

The obvious solution to this problem is to keep 

domestic cats confined indoors and to ensure that 

they are up-to-date on vaccinations. Hunting cats 

bring home an amazing variety of diseases and 

disease-transmitting parasites that they either carry 

themselves or have acquired from wildlife. Some 

diseases acquired this way by humans, such as 

plague (Gage and others, 2000; Baldwin and others, 

1991), are treatable if they are recognized early 

enough, but only three or four people are known to 

have survived lyssavirus infections. Stray cats and 

feral cats may also 

transmit diseases 

and parasites---

including some 

feline diseases 

that neutralize the 

protective effects 

of vaccinations-

--to owned cats 

(Pedersen and 

others, 1985; 

Yamamoto and 

others, 1989). The 

prompt testing 

of bats that are 

found under 

circumstances 

resembling any of the foregoing scenarios may reveal 

lyssavirus infection, and thus initiate prophylaxis 

specific to the found variant to potentially infected 

humans. Management of pets as directed under 

Disease Prevention and Control would decrease 

disease transmission to other pets, wildlife, and 

people.
36 Bat Rabies and Other Lyssavirus Infections

Figure 22. Exclusion devices permit bats 

to depart from but prevent re-entering 

the interior of a building. Bats enter 

and exit buildings through very narrow 

openings that may be revealed by soiled 

areas such as A, attic vent louvers, or B, 

fascia. Buildings can be bat-proofed by 

installation of simple valve-like devices 

such as C and D that allow bats to exit but 

prevent entry (Constantine, 1982b).

B

C D

A
Points to Ponder: Interspecies Interactions in Potential Bat Rabies Transmission Settings 37

Bat Colonies in Caves and Mine Tunnels

Most caves or mine tunnels in North America are devoid 

of bats. Some, however, may be inhabited only in summer, 

others only in winter, and still others year-round. If bats are 

present, their numbers usually vary from 1 to 30, but sometimes to several hundred or more. Some bat species hide in 

crevices, but others are more exposed and obvious. Summer 

shelters are warm to hot, and bats may be active. Winter 

shelters of most species are cold, facilitating the hibernation 

of bats, individually or in compact groups.

Some bats do not survive human intrusions due to 

injuries or predation outside or inside the shelter, and human 

intrusion may cause bats to abandon shelters. Hibernating bats 

often slowly awaken and move to safer places after someone 

tries to noiselessly sneak past them, which can sometimes 

result in the loss of sufficient body fat for the bat to survive 

until spring. When a person enters in summer, pandemonium 

can result as bats panic to avoid intruders. Some bats may 

fly outside while others retreat to deeper sites to escape the 

intruder. Flightless suckling bats often become dislodged 

and fall or are panicked into clumsy attempts at flight. Such 

young bats or distressed adults may fall or land on human 

intruders, who may be scratched or even bitten, especially if 

they slap or grab the bat. Thus, the intruder has experienced 

a potential lyssavirus exposure. Human avoidance of bat 

shelters benefits bats by removing the potential for disrupted 

hibernation, and benefits people by removing the potential for 

disease transmission from bats.

The danger of rabies transmission by inhalation of 

airborne virus is known to exist in caves occupied by many 

thousands to millions of Brazilian free-tailed bats (Fig. 23) 

(Constantine, 1967b; see also “Cave Explorations” below). 

This danger becomes greater as bat numbers increase or 

as cave space decreases proportionately, and similar caves 

crowded with other bat species in tropical areas pose similar 

danger. Prohibition of the entry of humans into such shelters 

during seasons of bat occupancy, except for persons willing 

to risk exposure after they have received proved effective 

pre-exposure antirabies immunizations, decreases the risk of 

infection by aerosol or other means of transmission. Depending on location, such visitors may also be exposed to excessive heat, humidity, noxious cave gasses (such as ammonia), 

carnivorous beetles, blood-sucking and other arthropods, 

snakes, mammalian predators ranging from skunks to jaguars, and other hazards, including histoplasmosis (Constantine, 1988a). The latter tuberculosis-like fungal disease is 

acquired by inhaling infective spores in guano dust. Relevant 

warning signs have been erected at some caves in the USA.

Special gates that permit bat passage but that block 

human entry have been installed at the entrances of some bat 

caves and mine tunnels to protect the much smaller groups of 

bats that shelter therein. However, similar gates would probably kill many bats as they rapidly enter or depart en masse at 

multimillion-bat colonies.

Flying Bats that Enter Buildings at Night

A flying bat occasionally enters a residence through 

an open window or door at night, because it recognizes the 

light inside as an attractant for its flying insect food. In this 

situation, the bat is likely to depart the same way it entered if 

doors to other rooms are closed to prevent it from getting lost 

in adjacent rooms. If the bat clings near the ceiling, it can be 

encouraged to leave by intimidation with a broom, or it may 

be captured carefully in a butterfly net or by placing an open 

coffee can over it and sliding a cardboard lid behind the can 

opening to keep the bat inside. Capture technique may have 

to be varied or improvised if the bat falls to the floor or does 

not cooperate. Such a bat is unlikely to be infected, but there 

are no guarantees, and bites or scratches should be regarded 

as potential rabies exposures unless the bat is quickly tested 

and proved uninfected. In the absence of skin contact, the bat 

may be released outside. Window and door screens discourage such bat visits.

Figure 23. Researcher among a Brazilian free-tailed bat colony. 

Note the personal protective clothing and equipment worn, such 

as coveralls, gloves, respirator, and hat; eye protection is also 

recommended. (D.G. Constantine)
38 Bat Rabies and Other Lyssavirus Infections

Alternatively, a bat may enter through an unscreened 

chimney, sometimes attracted by exiting warm air on cool 

evenings, or it may gain entry by unknown means. It may 

enter a darkened building at night seeking temporary rest, 

shelter from wind or rain, warmth, or perhaps a new home or 

stopover site while migrating, and it may be discovered the 

following morning. As with any mysterious appearance of a 

bat, one cannot know if the bat was delivered by a resident—

and thus potentially exposed—pet cat. One would prefer 

to be assured that no humans or pets have been exposed to 

rabies, but that is impossible, especially if the bat has been 

in a bedroom with a sleeping person. The teeth of many bats 

are sharp but short and may be capable of transmitting virus 

without awakening bitten persons or leaving detectable bite 

wounds. Healthy adults evidently have not awakened to bites, 

and others, such as infants, elderly, or medicated persons may 

be unable to perceive or communicate events. At least 30 percent of 51 North American human rabies victims were either 

awakened at night during a rabid bat attack or the bat was 

present in their bedrooms when they awoke in the morning 

(Constantine, unpub. data). Also, sound can precipitate rabid 

bat attacks, and any source of sound in an otherwise quiet 

bedroom may be a target. Snoring or even faint noises such 

as air movement through partially closed lips may suffice to 

attract a rabid bat.

Unprovoked Attacks by Flying Rabid Bats

Attacks on people and noisy pets by flying rabid bats 

are rare in the USA. Most reports involve the canyon bat 

(Fig. 10), which is abundant in the Southwest and prefers 

to shelter in rock crevices in cliffs. Fewer attacks are documented for the Western small-footed myotis (Fig. 11) and the 

California myotis (Fig. 12), both of which prefer to shelter in 

rock crevices, under loose tree bark, or other similar natural 

sites. All three species are strictly solitary and uncommonly 

take shelter in buildings. Flying attacks by rabid bats of other 

common North American species have been documented. 

The flying attacks are known to be swift and direct, and 

the bat is not seen until it has bitten the victim. That said, it 

seems superfluous to mention the only potentially preventive 

measure: do not make noise or do anything that may attract 

the animal’s attention. Most attacks have occurred in daylight, 

so bats flying in daytime should be suspect. Daylight attacks 

by rabid bats have been reported in Latin America (Haupt and 

Rehaag, 1921; Torres and de Quieroz Lima, 1935; Pawan, 

1936), but few case-specific details are available.

Translocation of Bats

Bats have been and continue to be accidentally and 

intentionally translocated by intra- and intercontinental ships, 

aircraft, and overland transport. Some of these bats may be 

incubating infectious diseases such as lyssaviruses. Bats are 

intentionally translocated to stock zoos or laboratory colonies 

and to supplement wild populations. Some bats intended for 

translocation have escaped en route to or at their destinations 

(Constantine, 2003). Only a small fraction of translocations 

are publicized, and they typically are highlighted only after 

information is released following accidents. Among various 

objections to translocations, especially when bats escape, 

are the potential introduction of foreign diseases that could 

impact humans and animals. For example, the recent introduction of West Nile virus into North America may have been 

due to the importation of thousands of foreign fruit bats for 

sale as pets and for exhibition (Constantine, 2003). In addition, a live little free-tailed bat was inadvertently transported 

by aircraft in a suitcase from South Africa to Los Angeles. 

This species is one of the very few known to support the 

experimental replication of the Ebola virus without exhibiting clinical signs of the disease (Swanepoel and others, 

1996). Increased regulation of the wildlife pet trade within 

the USA was urged following the accidental introduction of 

monkeypox virus, which was brought into the country by 

small African mammals intended for the pet market (Reynolds and others, 2007). The need for wildlife health certification for translocated and released wildlife has been elevated 

during recent years (Friend, 2006).

Vampire Bats and Rabies

Vampire bat populations, once dependent on native 

fauna for their blood meals, increased greatly following the 

introduction of domestic livestock throughout Latin America 

(Greenhall, 1988). Three different vampire bat species are 

involved in rabies transmission (Box 10). Rabies outbreaks 

in vampire bats (Fig. 24) have been observed to “migrate” 

throughout vampire populations, often along rivers, leaving 

a wake of death as the disease kills both the bats and livestock. Because vampire bats prefer feeding from cattle, the 

bats usually turn to other victims, including people, only after 

cattle die or are moved seasonally to distant pastures. Cattle 

depredations can be controlled by destroying vampire bat 

populations. This objective can be achieved through campaigns whereby petroleum jelly that contains an anticoagulant 

is applied to the backs of captured vampires (Fig. 25) that 

are then released to return home, where companion vampires 

lick off the jelly and perish as a consequence. Alternatively, 

the jelly is applied to wounds on bitten animals (Fig. 26) to 

poison vampire bats that return to the wounds on subsequent 

nights (Lord, 1988).
Points to Ponder: Interspecies Interactions in Potential Bat Rabies Transmission Settings 39

Figure 24. Common vampire bat. (D.G. Constantine)

Figure 26. Vampire bat-bitten burro.

(D.G. Constantine)

Figure 25. Application of anticoagulant 

jelly to the back of a vampire bat. 

(D.G. Constantine)
EXPLANATION

Reported range of the common vampire bat

 (Desmodus rotundus)

 Reported range of the hairy-legged vampire bat 

 (Diphylla ecaudata)

 One hairy-legged vampire bat was found in the USA

 Documented geographic locations of the white-winged 

 vampire bat (Diaemus youngi)

★

★

40 Bat Rabies and Other Lyssavirus Infections

Box 10 The Tripartite Vampire Bat Clan

Although most references to vampire bats refer to 

the common vampire, two less abundant vampire bat 

species exist that prefer the blood of birds, including 

poultry. These species, the white-winged vampire 

and the hairy-legged vampire, bite birds on the legs 

or the cloaca region from the underside of branches 

on which the birds perch (Greenhall, 1988), but 

these bats will also feed on the blood of mammals, 

including humans.

All three vampire species are distributed from Mexico 

to southern Brazil or Argentina and exhibit unique 

characteristics and behaviors. The white-winged 

vampire bat has white wing tips and a large, bulbous 

gland within each of its cheeks, from which it can 

emit through its open mouth a jet of liquid similar to 

the anal scent gland secretion sprayed by skunks. In 

contrast, the hairy-legged vampire bat is solitary and 

retiring rather than gregarious and confrontational. The 

Reported distribution of the three species of vampire bats: 

the common vampire bat (Greenhall and others, 1983), 

the hairy-legged vampire bat (Greenhall and others, 1984), 

and the white-winged vampire bat (Greenhall and 

Schutt, Jr., 1996).
The Tripartite Vampire Bat Clan 41

Hairy-legged vampire bat. 

(D.G. Constantine)

common vampire bat frequently regurgitates blood to 

supplement milk for its progeny older than 3 months 

of age. It also shares regurgitated blood with females 

that failed to secure meals through predation. Such 

food sharing among adult vampire bats is performed 

on a basis of reciprocity, requests from previously 

uncooperative bats being rejected.

Three species of vampire bats would be sufficient, but current research indicates 

that the common vampire may be divisible into additional species (Martins and 

others, 2007). In addition, the presumed extinct giant vampire bat may yet be found 

living (Ray and others, 1988).

White-winged vampire bat. (D.G. Constantine)

Common vampire bat. (D.G. Constantine)
42 Bat Rabies and Other Lyssavirus Infections

Attacks on People

Vampire bats may transmit numerous other known or 

potential diseases, in addition to rabies (Table 9) (Constantine, 1988b). Most vampire bat attacks on people are in undeveloped areas, where victims often do not recognize the cause 

of bites inflicted during sleep and darkness, or ascribe them to 

mythical agents such as witches, thus emphasizing the need 

for education. However, furious daytime attacks on people 

and animals have been reported during rabies outbreaks 

throughout Latin America (Pawan, 1936).

Physical exclusion of bats from the dwellings of remote 

native peoples most affected may be nearly impossible due 

to the primitive nature of these shelters and due to the lack of 

funding to construct more secure quarters. Infection would be 

preventable by pre-exposure antirabies vaccinations, which 

may be prohibitively expensive, or postexposure treatment 

with suitable vaccine and globulin products, which generally 

are neither affordable nor available. Under these circumstances, the potential for infection may be reduced by immediately and extensively washing the bite wound with soap and 

water, irrigating the wound with a virucidal agent, promptly 

testing the bat, and starting the patient on postexposure treatment. In view of the general unavailability of modern preventive measures, native peoples could benefit from revisiting the 

rabies-preventive measures that their predecessors taught the 

European explorers during the early 1500s: promptly cleanse 

vampire bat wounds and cauterize them with wood embers.

Attacks on Livestock

In addition to vampire bat population control methods 

mentioned above, vampire-borne rabies in livestock can be 

limited by various other methods, the most effective of which 

is antirabies vaccination of livestock. Recent studies have 

revealed the existence of at least three rabies virus variants 

that are circulating among vampire bats in Latin America, and 

each variant has a specific geographical distribution that may 

differ from or overlap with that of other variants (De Mattos, 

C.A. and others, 2001; Vargas Pino, 2001). Consequently, 

antirabies vaccines tailored to each or all of these variants 

might increase their effectiveness.

Attacks on Domestic Dogs and Cats

Vampire bats may expose companion dogs or cats to 

rabies while seeking blood meals or under other circumTable 9. Diseases other than rabies that may be transmitted by vampire bats. 

[The feeding habits of vampire bats provide a mechanism for mechanical transmission of infectious agents to their live animal food sources.]

Disease Causative agent Agent type Primary hosts

Chagas disease Trypanosoma cruzi Protozoan Humans

Murrina (Surra) Trypanosoma evansi Protozoan Horses

Candidiasis Candida chiropterorum, etc. Fungus Mammals, birds

Salmonellosis Salmonella enterica serotype Typhimurium, etc. Bacterium Mammals

Brucellosis Brucella abortus, B. suis, etc. Bacterium Cloven-hooved mammals

Foot and mouth 

disease

Aphthovirus foot and mouth disease virus (FMDV) Virus Cloven-hooved mammals

Venezuelan equine 

encephalomyelitis

Alphavirus Venezuelan encephalomyelitis virus (VEEV) Virus Horses, humans
Disease Prevention and Control 43

stances (Box 9). Cats in particular may collect and deliver 

live or dead vampires, some captured as they attack the cat. 

Vampire bats can deliver exceptionally effective bites, and 

thus greater virus doses. They usually remove a divot of flesh 

about 1/8 inch (3 millimeters) in diameter with each bite, and 

they will do so either defensively for protection or to obtain 

blood for feeding. Rabies of vampire bat origin has adapted 

to and is being transmitted serially among dogs, cats, and 

some other bat species in Latin America (Carrieri and others, 

2001; Kotait and others 2001). Infected dogs or cats may 

then transmit rabies to humans or to other mammals. The 

prompt testing of a rabies-suspect vampire bat may indicate 

if it is infected. If the bat involved in an attack is unavailable or unsuitable for testing, or if testing is delayed, then the 

bat should be considered infected and known or potentially 

exposed subjects should be managed accordingly (see Disease 

Prevention and Control). This problem can be limited by 

vaccinating pets against rabies and confining them to prevent 

their contact with vampire bats. Control of stray and feral 

dogs and cats would mitigate their chance of infection and 

subsequent roles as virus sources.

Attacks on Other Bats

Vampire bats deliberately attack other mammals, including other bat species, for blood meals, and other bat species 

serially transmit vampire rabies variants (Lord and others, 

1973; Silva, 2001; Table 8). It is unknown if such changing 

of hosts will affect the nature of disease in the new host, such 

as changing a predominantly paralytic disease to a predominantly furious disease. Reports of furious rabies in various 

species of bats in Latin America are increasing, but more 

definitive evidence would delineate, qualify, and quantify 

relevant facts pertinent to devising appropriate responses 

or solutions.

Disease Prevention and Control

Preemptive actions to minimize exposure to rabid bats 

are possible (Table 10), and they can be highly effective 

when they are combined with sensible behaviors, such as 

avoiding hazardous situations and seeking treatment following potential exposure. When someone is potentially exposed 

to disease after being bitten by a bat or another mammal, 

the wound should immediately be scrubbed with water and 

soap and preferably irrigated with a virucidal agent, such 

as povidone-iodine solution. The victim should simultaneously take steps to initiate subsequent antirabies treatment. 

In addition, prompt testing of the bat may provide information valuable to treatment of the victim. However, if the bat 

is lost among a group of bats, one must consider whether it 

is possible or legal to capture and test all of the bats or to 

undertake treatment without further delay. Procrastination in 

seeking treatment can have a deadly outcome. The following sections provide general guidance and describe special 

circumstances when basic actions would be likely to mitigate 

the risk of disease or the severity of the infection.

Humans

Information about human rabies prevention is available 

from the Recommendations of the Advisory Committee on 

Immunization Practices (Centers for Disease Control and 

Prevention, 2008), which reviews rabies biologics, pre-exposure vaccination, postexposure prophylaxis, serologic testing, 

adverse reactions, and precautions. Additionally, current 

information referable to all lyssavirus laboratory services or 

biologics may be sought from local to international health 

department channels. The World Health Organization (WHO), 

Geneva, Switzerland, has designated various laboratories 

as WHO Collaborating Centers for lyssavirus reference 

and research.

Persons who work with rabies virus or with known or 

potentially infected animals can reduce their risk of infection 

by avoiding bites or other exposures and by receiving preexposure vaccinations followed by periodic tests for rabies 

antibody levels. Potentially virus-contaminated skin breaks, 

such as bites or scratches, should be scrubbed at once with 

soap and water, followed by irrigation with a virucidal agent. 

In these situations, tetanus prophylaxis and bacterial infection 

control may also be beneficial. Rapid initiation of laboratory 

tests on an animal source may indicate the species and virulence of lyssavirus, if present. The immediate initiation of 

antirabies globulin and vaccine treatments may be a lifesaving measure. However, some people have received treatment 

immediately and still died of rabies, having lost the race 

between the virus and immune-system response (Centers for 

Disease Control and Prevention, 2008). Nevertheless, rabies 

vaccines and globulin products approved for use in the USA 

are usually highly effective and reliable. It is important to 

note that tests to identify, and treatment for, exposures other 

than rabies or Australian bat lyssavirus may not be available 

or entirely satisfactory (Table 1). 

Wildlife Rehabilitators

Among the many people who become wildlife rehabilitators are those who care for disabled bats. Because 1 in 10 

disabled bats submitted for testing in the USA proves to have 

rabies, bat rehabilitators may avoid infection while handling 

bats by receiving inoculations of rabies vaccine. Rehabilitators may not emphasize the rabies hazard associated with bats 

during presentations given to school children or other audiences, because they may be motivated to care for and protect 

the reputation of bats and feel securely vaccinated. Complete 

presentations on the ecology of bats and their role in consequent rabies mortality may enhance audience appreciation 

of bats and educate them about the transmission of diseases 

carried by bats. 
44 Bat Rabies and Other Lyssavirus Infections

Some rehabilitators may become embarrassed or fear 

consequent negative effects on their careers and on bats after 

they repeatedly submit rabid bats for testing. However, they 

need to know if a bat that has bitten them is infected so that 

they can receive vaccine boosters. Some rehabilitators ask 

friends to submit bats that bite, along with false histories, 

to distant laboratories for testing. Such actions may deprive 

the original discoverers of the bats of test results, and this 

knowledge is critical for the informed initiation of potentially 

lifesaving antirabies treatment.

Cave Explorers

Persons such as speleologists or biology students who 

contemplate entering bat harborages, including caves or mine 

tunnels, may be exposed to the health hazards associated 

with this activity and can take steps to decrease the risk of 

exposure. Applicable health hazards, precautions, and necessary preparations are detailed in a 38-page book chapter by 

Constantine (1988a). Topics covered include: 

1. risks associated with exposure to pathogenic agents 

including rabies, rabies-related viruses, other viruses of 

bats, and histoplasmosis; 

2. risks associated with exposure to bat urine, guano, and 

ectoparasites; 

3. risks associated with oxygen deficit; 

4. first aid and rescue concerning hazardous underground 

atmospheric gases such as ammonia, carbon dioxide, 

methane, carbon monoxide, and hydrogen sulfide; 

5. risks associated with the deliberate contamination of bats 

or bat roosts with anticoagulants or poisonous gases; 

6. recommended personal protective equipment; and 

7. bat health problems caused by roost visitations.

Table 10. Preemptive actions for minimizing human exposures to lyssaviruses.

Potential source for exposure Preemptive actions

All bat species

“Home deliveries” of bats by domestic cats Vaccinate and confine cats. Control stray and feral cats.

Colonies of bats in manmade structures

Schools Prevent bat entry.

Other public contact sites Prevent bat entry or post warning signs.

Colonies of bats in caves or mine tunnels Prohibit entry to all but immunized persons with justifiable need for entry.

Flying bats observed entering buildings at night Avoid contact and promptly facilitate ability of flying bat to exit building. Safely collect 

and submit non-flying bats for testing.

Unprovoked attacks by day-flying bats Upon seeing flying bat, remain quiet and motionless until bat departs.

Vampire bats

Colonies in human-inhabited areas Destroy colonies using appropriate techniques.

Attacks on people Educate susceptible human populations about vampire bat rabies and initiate appropriate 

protective measures, including vaccination and post-exposure treatment programs.

Attacks on livestock Vaccinate livestock, employ protective husbandry of livestock, and institute vampire bat 

control programs.

Attacks on dogs and cats Vaccinate pets, control their outdoor movements, and house pets indoors.

Attacks on other bats Consider tropical bat bites, and other interactions between vampires and other bat 

species, as rabies exposures unless quickly proven otherwise.
References Cited 45

Relevant preparation prior to cave exploration can save 

lives later. For example, carbon dioxide, which originates 

from various sources within underground caverns, including the exhalations of bats, is heavier than air and displaces 

oxygen, so it accumulates at lower underground levels where 

one may sit down to rest and soon become too weak to rise. 

Myotis bats show extraordinary resistance to anoxia (Britton and Kline, 1945), and Brazilian free-tailed bats can live 

in 100 times the ammonia concentration tolerated for five 

minutes by a human (Studier and others, 1967; National 

Institute for Occupational Health Standards, 1983). Therefore, 

the presence of live bats does not provide assurance that the 

atmosphere is safe for people.

Studies have shown that inhaled rabies virus enters nerve 

cells (olfactory receptor cells) that extend into the nasal cavity and are freely exposed to air as it is inhaled through the 

nose (Constantine and others, 1972). Accordingly, the virus 

essentially enters the brain as one inhales air. Airborne rabies 

transmission in the presence of massive Brazilian free-tailed 

bat aggregations within bat caves has been demonstrated 

(Constantine, 1967b), and similar aerosol transmission of 

other lyssaviruses and pathogens may be likely in similarly 

congested bat harborages, especially in tropical and subtropical zones. Fortunately, most explorers are repelled by these 

environments or undertake necessary preparations to ensure 

their safety and survival prior to entering. Pre-exposure rabies 

vaccination (Centers for Disease Control and Prevention, 

2008) for all explorers of caverns known to harbor bats is an 

essential place to begin, but it may provide little or no protection against lyssaviruses other than rabies (Hanlon and others, 

2005).

Animals

Recommendations for the prevention and control of animal rabies are available in the Compendium of Animal Rabies 

Prevention and Control (National Association of State Public 

Health Veterinarians, 2008). Information includes responses 

to known or potential exposures of domestic pets, such as 

dogs, cats, and ferrets, to rabid animals. 

The prompt testing of bats that expose pet mammals is 

indicated, and if the bat is unavailable or unsuitable for testing, then it is prudent to assume that the bat is infected and 

that the animal was exposed to the virus. Exposed healthy 

pets with up-to-date vaccinations should be revaccinated 

immediately, kept under the owner’s control, and observed for 

45 days. Immediate euthanasia of an unvaccinated pet would 

ensure that disease is not transmitted to other animals or to 

humans. If euthanasia is not the first course of action for an 

unvaccinated pet, then it can be placed in strict isolation for 

6 months and vaccinated upon entry into isolation or 1 month 

before release from isolation. 

Domestic cats that are permitted to hunt and that bring 

home their victims account for the great majority of contacts 

between bats and humans and resulting human lyssavirus 

mortality. Because cat owners and their children are the foremost victims of lyssavirus transmission from pets, the relative 

ease and cost effectiveness of pet management would be 

likely to significantly decrease the problem (Box 9).

Clearly, much remains to be learned about lyssaviruses. For example, in an experimental case, a cat developed 

paralysis of its rear legs 17 days after it was inoculated with 

rabies virus from a big brown bat. The cat underwent gradual 

recovery until about 2 years later, when it suffered a relapse, 

became debilitated, and was euthanized, whereupon the virus 

was recovered from the cat’s central nervous system (Murphy 

and others, 1980). This study discloses another method by 

which rabies virus can survive incognito over time, supplementing known incubation periods similar to dormancy in 

hibernating animals during the winter.

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Glossary

A

active immunity The internal capability of 

the body to combat disease agents; develops 

either following infection by a disease agent 

or by vaccination as the immune system 

responds by producing antibodies against the 

agent. (See also passive immunity.)

African bats Bat species native to the African continent. 

Americas Lands of the New World (Western 

hemisphere), consisting of North, Central, and 

South America and their associated islands.

amphibians A group of poikilothermic

(“cold-blooded”), scaleless, vertebrate

animals that usually begin life in the water as 

gill-breathing tadpoles and that later develop 

lungs and spend part of their time on land. 

amplification host A host in which infectious agents multiply to high concentrations, 

(See also intermediate host, reservoir host.)

anoxia A condition characterized by the 

absence of oxygen. For living organisms, this 

condition involves the absence of a supply of 

oxygen to an organ or a tissue.

antibody A protein formed in the body of a 

vertebrate that is used by the immune system 

to identify and neutralize the effects of foreign 

invading proteins, called antigens, such as 

bacteria and viruses. (See also maternal antibody, passive immunity.)

arthropods The largest group of animals, 

comprising more than 80 percent of described 

living animal species, that includes insects, 

spiders, and crustaceans. Arthropods are covered by a hard exoskeleton and are characterized by a segmented body, jointed legs, and 

many pairs of limbs. 

B

bats Mammals in which the forelimbs have 

developed as wings, making them the only 

mammals in the world naturally capable of 

flight. There are estimated to be about 1,100 

species of bats worldwide, accounting for 

about 20 percent of all mammal species. (See 

also African bat.)

insectivorous bats Those that primarily 

feed on insects while in flight

fruit-eating bats Those that feed almost 

entirely on fruit and on some green vegetation. (See also frugivorous.)

true vampire bats Those that feed 

on blood obtained from animals 

and humans.

biological transmission Disease transfer 

requiring an amplification host (often an 

arthropod) for development or multiplication 

of the disease agent (or both) necessary for 

infection of another host.

biologics A range of pharmaceutical products including vaccines, blood components, 

and therapeutic proteins used to treat a variety 

of medical conditions.

birds Warm-blooded vertebrates with 

wings and feathers, although the wings are 

poorly developed in some species and they 

are flightless. Birds belong to the taxonomic

class Aves.

blood serum The pale fluid that remains 

after blood has clotted.

burro A term used in the Western USA to 

refer to a donkey.

C

carnivores Animals whose diet consists 

mainly of meat.

clinical signs Readily observable indications of a disease or injury.

conspecific An organism of the same 

species.

cornea The transparent covering on the 

front of the eye.

D

dewlap A flap of skin containing stored fat 

that hangs beneath the lower jaw of animals 

including mammals, birds, reptiles, or 

amphibians.

DNA (deoxyribonucleic acid) A nucleic 

acid found mainly in the chromosomes that 

contains the hereditary information of organisms and some viruses.

donkey A pony-sized domesticated member 

of the horse family.

Glossary 53
E

Ebola The common name for the virus that 

causes the fatal disease Ebola hemorrhagic 

fever. Ebola has caused a number of serious and highly publicized outbreaks since its 

discovery in Africa in 1976.

exsanguination The fatal process of total 

blood loss, or “bleeding to death.”

F

flaccid paralysis Paralysis with a loss of 

muscle tone.

flying fox Large, fruit-eating bats with a foxlike appearance of the head. These bats are 

also commonly referred to as “fruit bats.”

frugivorous An animal that feeds primarily 

or exclusively on fruit.

furious rabies A manifestation of rabies 

that in animals is characterized by periods of aggressive behavior towards inanimate objects and towards other animals 

and humans.

G

genet Small, largely nocturnal catlike 

carnivores with short legs, long bodies and 

tails, rounded ears, and pointed snouts; found 

in forests, savannahs, and grasslands of Africa 

and parts of Europe.

genotype The genetic makeup of a biological organism.

genus A taxonomic group of similar species; similar genera are grouped into families.

guano A term used here for the excrement 

of bats, but also including the manure of sea 

birds used as a fertilizer and other manure 

resembling the appearance of such excrements.

H

harp trap A large boxlike device effective 

for catching bats. A series of vertical wires 

extend upwards from the collection chamber. 

Bats in flight collide with the wires and then 

fall into the collection chamber below.

heterothermous An animal with the capability for regulating its internal body temperature 

above that of its environment during periods 

of activity. When the animal is at rest, however, its body temperature drops to that of its 

surrounding environment. (See also homoiothermous, poikilothermous.)

hibernation A state of inactivity in animals 

characterized by a lower body temperature, 

slower breathing, and a reduced metabolic 

rate. Hibernation allows animals to conserve 

energy during the winter when food may 

be unavailable.

histoplasmosis A potentially fatal disease 

that primarily effects the lungs and is caused 

by the fungus Histoplasma capsulatum. 

Spores of H. capsulatum, which are found in 

soil contaminated with bat or bird droppings, 

can become airborne when the contaminated 

soil is disturbed. Inhaling the spores can result 

in infection.

homoiothermous An animal with a body 

temperature that is constant and largely 

independent of the temperature of its surroundings. (See also heterothermous, 

poikilothermous.)

host An organism that harbors or nourishes 

microbes, viruses, and parasites. (See also

intermediate host, reservoir host.)

hyperimmune sera Typically, blood serum

with an especially high concentration of antibodies produced by repeated antigen injections into healthy donor hosts. 

hypothermia A condition in which an animal’s core body temperature drops below that 

required for normal physiological functions.

I

incubation period The time from the initiation of infection until the first recognized 

clinical signs.

insectivorous An animal that feeds primarily on insects.

intermediate host An organism in or on 

which a parasite develops for part of its life 

cycle. (See also host, reservoir host.)

J

jaguar The largest wild cat of the Americas. Typically found in thick wooded and 

arid shrubby environments from border areas 

of Texas and Arizona south to about 40° 

South latitude. A fierce predator, the jaguar is 

distinctly marked by black spots within the 

center of the rosettes that pattern its hide.

54 Bat Rebies and Other Lyssavirus Infections 
K

L

lyssavirus A genus of viruses of the family 

Rhabdoviridae.

M

mammals Warm-blooded vertebrate

animals that possess hair during some part of 

their life and suckle their young.

maternal antibody An antibody

transferred from a mammalian mother 

through the placenta into her fetus. 

(See also passive immunity.)

mechanical transmission Disease transfer 

involving contact with surfaces contaminated 

by pathogens. Mechanical transmission 

from bites, scratches, contact with inanimate 

objects, and other means allows disease-causing levels of the pathogen to enter the host. 

Megachiroptera One of two taxonomic

suborders of bats (the Chiroptera) and comprised of a single family, the Pteropodidae, or 

Old World fruit bats.

melancholia A nonspecific depression characterized by low levels of enthusiasm and low 

levels of eagerness for physical activity.

Microchiroptera The taxonomic suborder 

of bats (the Chiroptera) that contains all bats 

other than the fruit bats (flying foxes). It 

should be noted that some fruit bats (Megachiroptera) are smaller in size than some bat 

species within the Microchiroptera. 

mites A medically important group of 

arthropods of the taxonomic order Acarina, 

but excluding ticks. These external parasites, 

closely related to spiders, cause a variety 

of infestations, often of the skin of humans 

and animals, and of the hair and feathers 

of animals.

monkeypox A viral disease related to smallpox that is transmissible between animals 

and humans and that primarily occurs in 

central and western Africa. The exotic pet 

trade resulted in the 2003 introduction of this 

disease into the USA.

mucosal surfaces Moist surfaces of 

orifices and internal parts of the body that 

cover, protect, and provide secretory and 

absorptive functions.

mule The usually sterile offspring of a male 

donkey and a female horse.

N

O

omnivorous An animal with a diet of both 

plants and animals.

P

paralytic rabies A manifestation of rabies 

that in animals is characterized by signs of 

posterior paralysis and that generally lacks the 

violent symptoms of furious rabies.

passive immunity Immunity achieved 

following the receipt of antibodies from 

an immune donor. Protection by passive 

immunity lasts for a relatively short time. 

For example, in humans, antibodies passed 

from mother to fetus confer passive immunity 

to the baby for the first 4 to 6 months of life. 

(See also active immunity.)

pathogen Typically, a mircoorganism capable of inducing disease, but broadly includes 

all disease-inducing agents.

placenta A temporary organ rich in blood 

vessels that transfers oxygen and nutrients 

from the mother to the fetus and that permits 

the release of carbon dioxide and waste products from the fetus. The placenta is expelled 

during the birth process.

plague An infectious disease of animals 

and humans caused by the bacterium Yersinia 

pestis. Although millions of people in Europe 

died from plague during the Middle Ages, 

modern antibiotics provide effective treatment 

of this disease when administered promptly 

following development of symptoms.

poikilothermous An animal with a body 

temperature that varies with the temperature 

of its environment. (See also heterothermous, 

homoiothermous.) 

poultry A general term for farmed domestic 

birds, including chickens, turkeys, ducks, 

and geese.

prevalence The total number of cases of a 

disease in a population at a given time.

prodromal Early nonspecific symptoms, 

such as fever, headache, malaise, and nausea, 

indicating the start of a disease before specific 

symptoms occur.

putative Assumed or accepted as true.

Glossary 55
Q

R

reptiles A group of poikilothermic 

(“cold-blooded”), air-breathing, vertebrate animals with scaled bodies. (See also

poikilothermous.)

reservoir host An organism that maintains 

a disease agent in nature and that provides 

a source of infection to susceptible hosts. 

(See also intermediate host.)

S

salivary gland A gland that produces saliva.

sanguivorous An animal that feeds on 

blood; for example, certain bats, leeches, 

and insects.

serial transmission The sequential spreading of a disease agent among multiple susceptible hosts. (See also host.)

skunk Several species of carnivores

within the family Mephitidae; specifically, the striped, hooded, spotted, and 

hog-nosed skunk. 

species A population of organisms whose 

members are able to breed among themselves 

and produce fertile offspring. More precise 

determinations of species are based on similarity of DNA. (See also strain, variant.)

speleologist A person who explores caves.

strain A genetically or biochemically distinguishable subtype of a microorganism.

symptomatic Showing symptoms or signs of 

a disease or injury.

T

taxonomy The systematic principles and 

procedures of grouping and arranging organisms into a hierarchical order.

temperate zone Latitudes of the globe that 

lie between the tropics and the polar circles 

that exhibit seasonal temperature patterns.

torpid Dormant, as observed for hibernating 

animals. (See also hibernation.)

tropical Geographically, the zone on either 

side of the equator, bounded by the Tropic 

of Cancer (23°27’ North latitude) and the 

Tropic of Capricorn (23°27’ South latitude); 

climatically hot and humid for the majority of 

the year.

U

umbilical cord A tube that connects a developing embryo or fetus to the placenta. 

V

vaccine A biological preparation containing an antigen, a molecule that stimulates 

the immune system, that is used to establish 

immunity to a disease.

variant A group of organisms of the same 

species that exhibit subtle yet characterizable 

differences from one another.

vector An animal, including insects, that 

carries a disease-causing organism from an 

infected to a healthy animal or plant, thus 

causing the latter to become infected.

vertebrates All animals that have spinal 

columns: mammals, birds, reptiles, amphibians, and fishes.

virulence The degree or ability of a 

pathogenic organism to cause disease. 

(See also pathogen.)

virus A microscopic infectious agent that 

can cause a disease, such as rabies in animals and potato virus in potato, tomato, and 

tobacco plants. A virus can reproduce only by 

invading living cells, and it exists outside of 

living cells as an inactive particle. 

W

X

Y

Z

Zebu An Asian ox, domesticated and 

differentiated into many breeds, and physically distinguished from European cattle 

by the presence of a large fleshy hump 

over the shoulders, a loose skin prolonged 

in a dewlap and folds under the belly, and 

large droopy ears.

56 Bat Rebies and Other Lyssavirus Infections 
Appendix A 57

Appendix A. Bat Species Reported Infected with Rabies in the USA from 1953 

to 2007

Bat species (common name) Family

Big brown bat Vespertilionidae

Big free-tailed bat Molossidae

Brazilian free-tailed bat Molossidae

California leaf-nosed bat Phyllostomidae

California myotis Vespertilionidae

Canyon bat Vespertilionidae

Cave myotis Vespertilionidae

Eastern red bat Vespertilionidae

Evening bat Vespertilionidae

Fringed myotis Vespertilionidae

Gray myotis Vespertilionidae

Greater bonneted bat Molossidae

Hoary bat Vespertilionidae

Keen’s myotis Vespertilionidae

Little brown myotis Vespertilionidae

Long-eared myotis Vespertilionidae

Bat species (common name) Family

Long-legged myotis Vespertilionidae

Northern myotis Vespertilionidae

Northern yellow bat Vespertilionidae

Pallid bat Vespertilionidae

Peter’s ghost-faced bat Mormoopidae

Pocketed free-tailed bat Molossidae

Seminole bat Vespertilionidae

Silver-haired bat Vespertilionidae

Southeastern myotis Vespertilionidae

Southern yellow bat Vespertilionidae

Spotted bat Vespertilionidae

Townsend’s big-eared bat Vespertilionidae

Tricolored bat Vespertilionidae

Western red bat Vespertilionidae

Western small-footed myotis Vespertilionidae

Western yellow bat Vespertilionidae

Yuma myotis Vespertilionidae
58 Bat Rabies and Other Lyssavirus Infections

Appendix B. Bat Species Reported Infected with Rabies in the 

Americas through 2006 

Bat species (common name) Family Country where infection was identified

Argentine brown bat Vespertilionidae Brazil

Big brown bat Vespertilionidae Canada, Cuba, Mexico, United States

Big free-tailed bat Molossidae Brazil, Mexico, United States

Big-eared brown bat Vespertilionidae Chile

Black bonneted bat Molossidae Argentina, Brazil

Black mastiff bat Molossidae Brazil, Mexico, Venezuela

Black myotis Vespertilionidae Argentina, Belize, Brazil, Panama, Peru

Black-winged little yellow bat Vespertilionidae Mexico

Brazilian brown bat Vespertilionidae Colombia

Brazilian free-tailed bat Molossidae Argentina, Brazil, Chile, Mexico, United States

Broad-eared free-tailed bat Molossidae Brazil, Mexico

Brown fruit-eating bat Phyllostomidae Peru

Brown mastiff bat Molossidae Brazil

Bulldog bat Noctilionidae Panama

California leaf-nosed bat Phyllostomidae United States

California myotis Vespertilionidae Canada, United States

Canyon bat Vespertilionidae United States

Cave myotis Vespertilionidae Mexico, United States

Chilean myotis Vespertilionidae Chile

Cinnamon dog-faced bat Molossidae Brazil

Cinnamon myotis Vespertilionidae Belize, Guatemala

Common mustached bat Mormoopidae Mexico, Trinidad

Common sword-nosed bat Phyllostomidae Brazil

Common tent-making bat Phyllostomidae Panama

Common vampire bat Phyllostomidae Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, 

Ecuador, El Salvador, Guatemala, Mexico, Nicaragua, 

Paraguay, Peru, Trinidad, Venezuela

Davy’s naked-backed bat Mormoopidae Mexico, Trinidad

Diminutive serotine Vespertilionidae Brazil

Eastern red bat Vespertilionidae Canada, Mexico, United States

Evening bat Vespertilionidae United States

Fringed myotis Vespertilionidae United States

Fringe-lipped bat Phyllostomidae Brazil

Fruit-eating bat Phyllostomidae Mexico, Peru
Appendix B 59

Bat species (common name) Family Country where infection was identified

Geoffroy’s tailless bat Phyllostomidae Brazil

Gray myotis Vespertilionidae United States

Gray short-tailed bat Phyllostomidae Mexico

Great fruit-eating bat Phyllostomidae Belize, Bolivia, Brazil, Guatemala, Mexico, Peru, Trinidad

Great stripe-faced bat Phyllostomidae Brazil

Greater bonneted bat Molossidae Brazil, United States

Greater bulldog bat Noctilionidae Mexico

Greater spear-nosed bat Phyllostomidae Brazil, Peru

Hairy-legged vampire bat Phyllostomidae Brazil, Mexico

Hoary bat Vespertilionidae Argentina, Brazil, Canada, Mexico, United States

Jamaican fruit-eating bat Phyllostomidae Belize, Bolivia, Brazil, Grenada, Mexico, Panama, Trinidad

Keen’s myotis Vespertilionidae Canada, United States

Lesser long-nosed bat Phyllostomidae Mexico

Lesser spear-nosed bat Phyllostomidae Peru

Little big-eared bat Phyllostomidae Brazil, Panama, Peru

Little brown myotis Vespertilionidae Canada, United States

Little yellow-shouldered bat Phyllostomidae Brazil

Long-eared myotis Vespertilionidae Canada, United States

Long-legged myotis Vespertilionidae United States

Mexican long-nosed bat Phyllostomidae Mexico

Northern ghost bat Emballonuridae Trinidad

Northern myotis Vespertilionidae Canada, United States

Northern yellow bat Vespertilionidae Mexico, United States

Pale spear-nosed bat Phyllostomidae Belize, Guatemala, Mexico

Pallas’ long-tongued bat Phyllostomidae Mexico, Peru

Pallas’ mastiff bat Molossidae Belize, Brazil, Colombia, Panama, Peru, Trinidad

Pallid bat Vespertilionidae Mexico, United States

Patagonian dwarf bonneted bat Molossidae Argentina

Peters’ ghost-faced bat Mormoopidae Mexico, United States

Pocketed free-tailed bat Molossidae United States

Pygmy round-eared bat Phyllostomidae Brazil

Seba’s short-tailed bat Phyllostomidae Colombia, Peru, Trinidad

Seminole bat Vespertilionidae Mexico, United States
60 Bat Rabies and Other Lyssavirus Infections

Bat species (common name) Family Country where infection was identified

Silver-haired bat Vespertilionidae Canada, United States

Silver-tipped myotis Vespertilionidae Brazil

Sinaloan mastiff bat Molossidae Belize, Brazil, Guatemala, Honduras

Small big-eared brown bat Vespertilionidae Argentina

Southeastern myotis Vespertilionidae United States

Southern dog-faced bat Molossidae Panama

Southern yellow bat Vespertilionidae Brazil, Mexico, Paraguay, United States

Spotted bat Vespertilionidae Canada, United States

Tailed tailless bat Phyllostomidae Brazil

Thomas’ mastiff bat Molossidae Panama

Townsend’s big-eared bat Vespertilionidae United States

Tricolored bat Vespertilionidae Canada, Mexico, United States

Tropical big-eared brown bat Vespertilionidae Brazil

Wagner’s bonneted bat Molossidae Cuba

Wagner’s mustached bat Mormoopidae Mexico

Waterhouse’s leaf-nosed bat Phyllostomidae Mexico

Western red bat Vespertilionidae Argentina, Brazil, Chile, United States

Western small-footed myotis Vespertilionidae Canada, United States

Western yellow bat Vespertilionidae United States

White-lined broad-nosed bat Phyllostomidae Peru

White-winged vampire bat Phyllostomidae Brazil, Mexico, Trinidad

Woolly false vampire bat Phyllostomidae Brazil

Yuma myotis Vespertilionidae Canada, United States
Appendix C 61

Appendix C. Common and Scientific Names for Species Cited 

[Common and scientific names from Wilson and Reeder (2005)]

Common name Scientific name

African shrews Crocidura sp.

African straw-colored fruit bat Eidolon helvum

Angolian free-tailed bat Mops condylurus

Argentine brown bat Eptesicus furinalis

Assassin bug Arilus cristatus

Big brown bat Eptesicus fuscus

Big-eared brown bat Histiotus macrotus

Big free-tailed bat Nyctinomops macrotis

Black bonneted bat Eumops auripendulus

Black flying fox Pteropus alecto

Black mastiff bat Molossus rufus

Black myotis Myotis nigricans

Black-winged little yellow bat Rhogeessa tumida

Brazilian brown bat Eptesicus brasiliensis

Brazilian free-tailed bat Tadarida brasiliensis

Broad-eared free-tailed bat Nyctinomops laticaudatus

Brown fruit-eating bat Artibeus concolor

Brown mastiff bat Promops nasutus

Bulldog bat Noctilio sp.

Burro Equus asinus

California leaf-nosed bat Macrotus californicus

California myotis Myotis californicus

Canyon bat Parastrellus hesperus 1

Cat (domestic) Felis catus

Cattle (domestic) Bos taurus

Cave myotis Myotis velifer

Chilean myotis Myotis chiloensis

Cinnamon dog-faced bat Cynomops abrasus

Cinnamon myotis Myotis fortidens

Common mustached bat Pteronotus parnellii

Common pipistrelle Pipistrellus pipistrellus

Common sword-nosed bat Lonchorhina aurita

Common name Scientific name

Common tent-making bat Uroderma bilobatum

Common vampire bat Desmodus rotundus

Crocidura shrew Crocidura sp.

Daubenton’s bat Myotis daubentonii

Davy’s naked-backed bat Pteronotus davyi

Dermestid beetle Dermestes carnivorus

Diadem leaf-nosed bat Hipposideros diadema

Diminutive serotine Eptesicus diminutus

Dog (domestic) Canis lupus

Donkey Equus asinus

Eastern red bat Lasiurus borealis

Egyptian rousette Rousettus egyptiacus

Egyptian slit-faced bat Nycteris thebaica

Evening bat Nycticeius humeralis

Ferret Mustela putorius

Flying fox Pteropus sp.

Fringed myotis Myotis thysanodes

Fringe-lipped bat Trachops cirrhosus

Fruit-eating bat Artibeus sp.

Gambian slit-faced bat Nycteris gambiensis

Genet Genetta tigrina

Geoffroy’s tailless bat Anoura geoffroyi

Giant vampire bat Desmodus draculae

Goat (domestic) Capra hircus

Gray fox Urocyon cinereoargenteus

Gray-headed flying fox Pteropus poliocephalus

Gray myotis Myotis grisescens

Gray short-tailed bat Carollia subrufa

Great fruit-eating bat Artibeus lituratus

Great stripe-faced bat Vampyrodes caracciolae

Greater bonneted bat Eumops perotis

Greater bulldog bat Noctilio leporinus
62 Bat Rabies and Other Lyssavirus Infections

Common name Scientific name

Greater horseshoe bat Rhinolophus ferrumequinum

Greater spear-nosed bat Phyllostomus hastatus

Greater tube-nosed bat Murina leucogaster

Hairy-legged vampire bat Diphylla ecaudata

Hamster Mesocricetus auratus

Hoary bat Lasiurus cinereus

Horse (domestic) Equus caballus

Indian flying fox Pteropus giganteus

Jackal Canis adustus

Jaguar Panthera onca

Jamaican fruit-eating bat Artibeus jamaicensis

Keen’s myotis Myotis keenii

Lesser dawn fruit bat Eonycteris spelaea

Lesser long-nosed bat Leptonycteris yerbabuanae

Lesser mouse-eared myotis Myotis blythii

Lesser spear-nosed bat Phyllostomus elongatus

Little big-eared bat Micronycteris megalotis

Little brown myotis Myotis lucifugus

Little free-tailed bat Chaerephon pumila

Little red flying fox Pteropus scapulatus

Little yellow-shouldered bat Sturnira lilium

Long-eared myotis Myotis evotis

Long-legged myotis Myotis volans

Lyle’s flying fox Pteropus lylei

Mexican long-nosed bat Leptonycteris nivalis

Mongoose Galerella sanguinea

Mouse-eared myotis Myotis myotis

Nathusius’ pipistrelle bat Pipistrellus nathusii

Noctule Nyctalus noctula

Northern bat Eptesicus nilssonii

Northern ghost bat Diclidurus albus

Northern myotis Myotis septentrionalis

Northern yellow bat Lasiurus intermedius

Pale spear-nosed bat Phyllostomus discolor

Pallas’ long-tongued bat Glossophaga soricina

Pallas’ mastiff bat Molossus molossus

Common name Scientific name

Pallid bat Antrozous pallidus

Particolored bat Vespertilio murinus

Patagonian dwarf bonneted bat Eumops patagonicus

Peter’s ghost-faced bat Mormoops megalophylla

Peter’s lesser epauletted fruit 

bat Micropteropus pusillus

Pig (domestic) Sus scrofa

Pocketed free-tailed bat Nyctinomops femorosaccus

Pond myotis Myotis dasycneme

Pygmy round-eared bat Lophostoma brasiliense

Queensland tube-nosed fruit bat Nyctimene robinsoni

Red fox Vulpes vulpes

Rhesus monkey Macaca mulatta

Rusty-bellied brush-furred rat Lophuromys sikapusi

Schreibers’ long-fingered bat Miniopterus schreibersii

Seba’s short-tailed bat Carollia perspicillata

Seminole bat Lasiurus seminolus

Serotine bat Eptesicus serotinus

Sharp-nosed tomb bat Taphozous georgianus

Sheep (domestic) Ovis aries

Shrew Crocidura sp.

Silver-haired bat Lasionycteris noctivagans

Silver-tipped myotis Myotis albescens

Sinaloan mastiff bat Molossus sinaloae

Skunk Mephitis sp., Spilogale sp.

Small big-eared brown bat Histiotus montanus

Southeastern myotis Myotis austroriparius

Southern dog-faced bat Cynomops planirostris

Southern yellow bat Lasiurus ega

Spectacled flying fox Pteropus conspicillatus

Spotted bat Euderma maculatum

Stone martin Martes foina

Tailed tailless bat Anoura caudifer

Thomas’ mastiff bat Molossus currentium

Townsend’s big-eared bat Corynorhinus townsendii

Tricolored bat Perimyotis subflavus 2

Tropical big-eared brown bat Histiotus velatus
Appendix C 63

Common name Scientific name

Wagner’s bonneted bat Eumops glaucinus

Wagner’s mustached bat Pteronotus personatus

Wahlberg’s epauletted fruit bat Epomophorus wahlbergi

Waterhouse’s leaf-nosed bat Macrotus waterhousii

Water mongoose Atilax paludinosusis

Western red bat Lasiurus blossevillii

Western small-footed myotis Myotis ciliolabrum

Western yellow bat Lasiurus xanthinus

Common name Scientific name

Whiskered myotis Myotis mystacinus

White-lined broad-nosed bat Platyrrhinus lineatus

White-winged vampire bat Diaemus youngi

Woolly false vampire bat Chrotopterus auritus

Yellow-bellied sheathtail bat Saccolaimus flaviventris

Yuma myotis Myotis yumanensis

Zebu Bos indicus

1

 The canyon bat was formerly the Western pipistrelle (Pipistrellus hesperus). The species has been reclassified as the only member of the genus Parastrellus

(Hoofer and others, 2006) and is currently referred to as the canyon bat. 

2 The tricolored bat was formerly the Eastern pipistrelle (Pipistrellus subflavus). This species has been reclassified as the only member of the genus Perimyotis

(Menu, 1984; Hoofer and others, 2006) and is currently referred to as the tricolored bat.
64 Bat Rabies and Other Lyssavirus Infections

Appendix D. Lyssavirus Infections in Bats Reported in Africa, Asia, Australia, 

and Europe

Common name Family Food habits Geographic location Lyssavirus species Citation

African straw-colored 

fruit bat

Pteropodidae Frugivorous Nigeria

Senegal

Lagos

Lagos

Boulger and Porterfield, 1958

Markotter and others, 2006a

Black flying fox Pteropodidae Frugivorous Australia Australian Warrilow and others, 2003

Common pipistrelle Vespertilionidae Insectivorous Germany

France

Unknown

Unknown

WHO CCRSR,1 1988

Bruyère and Janot, 2000

Daubenton’s bat Vespertilionidae Insectivorous Denmark

Germany

Switzerland

United Kingdom

EBL-2

EBL-2

EBL-2

EBL-2

Johnson and others, 2006

Johnson and others, 2006

Johnson and others, 2006

Johnson and others, 2006

Egyptian rousette Pteropodidae Frugivorous Egypt Lagos Markotter and others, 2006b

Egyptian slit-faced bat Nycteridae Insectivorous Zimbabwe Duvenhage Foggin, 1988

Gambian slit-faced bat Nycteridae Insectivorous Senegal Lagos Markotter and others, 2006a

Gray-headed flying fox Pteropodidae Frugivorous Australia Australian Warrilow and others, 2003

Greater horseshoe bat Rhinolophidae Insectivorous Turkey Unknown Tunçman, 1958

Greater tube-nosed bat Vespertilionidae Insectivorous Russia Irkut Botvinkin and others, 2003

Indian flying fox Pteropodidae Frugivorous India Unknown Pal and others, 1980

Lesser mouse-eared 

myotis

Vespertilionidae Insectivorous Kyrghyzstan Aravan Botvinkin and others, 2003

Little red flying fox Pteropodidae Frugivorous Australia Australian Warrilow and others, 2003

Mouse-eared myotis Vespertilionidae Insectivorous Germany Unknown Hentschke and Hellman, 1975

Nathusius’ pipistrelle 

bat

Vespertilionidae Insectivorous Germany EBL-1 WHO CCRSR,1 1986

Noctule Vespertilionidae Insectivorous Ukraine

Yugoslavia

EBL-1

Unknown

Khozinski and others, 1990; 

Selimov and others, 1990

Nicolíc and Jelesíc, 1956
Appendix D 65

Common name Family Food habits Geographic location Lyssavirus species Citation

Northern bat Vespertilionidae Insectivorous Russia Unknown Khozinski and others, 1990; 

Selimov and others, 1990

Particolored bat Vespertilionidae Insectivorous Ukraine EBL-1 Kappeler, 1989

Peter’s lesser 

epauletted fruit bat

Pteropodidae Frugivorous Central African 

Republic

Lagos Sureau and others, 1980

Pond myotis Vespertilionidae Insectivorous Denmark

Netherlands

EBL-2

EBL-2

WHO CCRSR,1 1986

Van der Poel and others, 2003

Schreiber’s 

long-fingered bat

Vespertilionidae Insectivorous Russia WCBV Botvinkin and others, 2003

Serotine bat Vespertilionidae Insectivorous Russia

Czech Republic.

Denmark

France

Germany

Hungary

Netherlands

Poland

Spain

Ukraine

Unknown

Unknown

EBL-1

EBL-1

EBL-1

Unknown

EBL-1

EBL-1

EBL-1

Unknown

Amengual and others, 1997

Matouch, 1999

Amengual and others, 1997

Amengual and others, 1997

Amengual and others, 1997

Kerekes, 1999

Amengual and others, 1997

Amengual and others, 1997

Amengual and others, 1997

Selimov and others, 1986

Spectacled flying fox Pteropodidae Frugivorous Australia Australian Warrilow and others, 2003

Wahlberg’s epauletted 

fruit bat

Pteropodidae Frugivorous South Africa Lagos Crick and others, 1982

Whiskered myotis Vespertilionidae Insectivorous Tajikistan Khujand Kuzmin and others, 2003

Yellow-bellied sheathtail batEmballonuridae Insectivorous Australia Australian Warrilow and others, 2003

1. World Health Organization Collaborating Centre for Rabies Surveillance and Research, Tübingen, Germany.
66 Bat Rabies and Other Lyssavirus Infections

Appendix E. Course of Bat Lyssavirus Infections in Humans

Case 3. Exposure in North America 

(Irons and others, 1957)

The victim was exposed to airborne rabies in a cave occupied by millions of Brazilian free-tailed bats.

Day 1 of symptoms, about 6 weeks later: The victim had 

visual problems that precluded driving. The victim reported 

a night of fitful sleep and nightmares.

Day 2: The victim had convulsive seizures while trying to 

drink water and the throat muscles became paralyzed. The 

victim also had breathing difficulties and acute anxiety.

Day 3: Symptoms grew worse, and supportive therapy was 

ineffectual.

Day 4: The victim died.

Case 4. Exposure in North America 

(Humphrey and others, 1960)

The victim picked up an injured silver-haired bat that 

was being attacked by dogs, placed the bat in a tree, and was 

bitten on the left forefinger. The victim was given antiserum 

3 days later and started on vaccine 4 days after being bitten. 

Day 1 of symptoms, 26 days after the bite: The victim’s inner 

left arm became numb to the first two fingers. The victim’s 

abdomen and spine became painful. 

Day 8: The symptoms worsened, and numbness extended to 

the victim’s arms. The victim was unable to urinate, then 

suffered overflow incontinence. 

Day 9: The victim had poor limb muscle control and pain in 

many parts of the body. The victim was able to eat breakfast. 

Day 10: The victim had severe leg and abdominal pain yet ate 

well. 

Day 11: Symptoms became worse. The victim could not pass 

gas but ate well. 

Day 12: The victim could not swallow mucus and felt heaviness in the chest. The victim died.

Case 1. Exposure in South America 

(Hurst and Pawan, 1932) 

The victim was awakened by being bitten on a toe of the 

left foot, whereupon a vampire bat was observed flying away 

from the foot.

Day 1 of symptoms, 27 days later: The victim felt a burning sensation and cramps in the left foot and was unable 

to flex the foot at the ankle. The left foot was also numb to 

touch. The symptoms later spread to the victim’s leg, which 

became weak. 

Day 3: The victim’s right foot became numb. Symptoms grew 

worse over several days.

Day 7: The victim had flaccid paralysis and numbness in the 

left leg and right foot and numbness over the trunk to the 

level of eighth to ninth rib. The victim was unable to pass 

urine freely and had been constipated for the past 8 days. 

The victim could answer questions intelligently.

Day 8: The victim’s symptoms became worse. The victim 

experienced slight breathing distress, some arm weakness, 

great thirst, and had a fever of 105°F (40.5°C). 

Day 9: The victim experienced acute breathing distress, 

profuse sweating over the entire body, arm incoordination, 

and ascending numbness. Swallowing became impossible. 

The victim became semiconscious and died.

Case 2. Exposure in North America 

(Sulkin and Greve, 1954)

The victim picked up a grounded bat and was bitten on the 

forearm. The bat was discarded.

Day 1 of symptoms, 16 days later: The victim experienced 

aching of the forearms and developed a fever of 103°F 

(39.4°C). 

Day 2: The victim felt generally achy, became nauseated and 

vomited, and one arm became paralyzed. 

Day 7: The victim was unable to swallow and later became 

semicomatose to comatose. 

Day 9: The victim’s blood pressure decreased, and the 

victim died.
Appendix E 67

Case 5. Exposure in North America 

(Humphrey and others, 1960)

The victim was awakened by a bat biting his left ear lobe. 

A cat later ate the bat. 

Day 1 of symptoms, 21 days later: The victim felt tingling and 

pain in the area of the left ear and had pain and tremors that 

spread to the neck and the left arm. 

Days 5 and 6: After further deterioration, the victim had 

marked tremors in the left arm, prolonged muscle spasms of 

the legs and neck similar to those seen in cases of tetanus, 

and hydrophobic spasms when offered water. 

Day 7: The victim died.

Case 6. Exposure in Europe 

(Lumio and others, 1986)

The victim was bitten by a Daubenton’s bat that was acting 

abnormally. The bat was released. 

Day 1 of symptoms, 51 days later: The victim’s right palm 

was numb and felt pain in the neck.

Day 2: The victim’s right cheek was numb and painful. 

Day 9: The victim was unable to walk and felt generally weak. 

The victim’s right arm and neck were numb, and the chest 

was painful.

Day 10: The victim developed muscle spasms of the jaw and 

the legs, prolonged rapid and deep breathing, throat spasms, 

facial grimacing, disturbed speech, swallowing difficulties, 

and dilated pupils. The victim exhibited overwhelming fear 

and was given antirabies serum and sedatives. Symptoms 

progressed to delirium with shouting, motor hyperexcitability, hypersalivation, and convulsions. 

Day 11: The victim died.

Case 7. Exposure in Africa 

(Meredith and others, 1971)

The victim was awakened by a bat bite on the lip. The bat 

was destroyed. 

Day 1 of symptoms, about 1 month later: The victim had 

headache, dizziness, and muscle aches in the neck and back.

Days 2 and 3: The victim sweated continuously and had 

muscle spasms in the face, arm, and leg. The victim became 

increasingly confused, irritable, and aggressive, had difficulty sleeping, and had nightmares. The victim’s swallowing became progressively more difficult.

Day 4: All of the victim’s reflexes became markedly exaggerated. When the victim attempted to drink, tremendous 

spasms of the mouth and throat muscles prevented swallowing and caused impaired breathing. Despite sedatives, the 

victim was tense, anxious, sweated profusely, and became 

increasingly violent and difficult to control. Upon further 

attempts to drink, the victim cried out in pain and had pharyngeal muscle spasms as well as spasms of the upper torso 

and arms. 

Day 5: The victim went “completely berserk,” frothing at the 

mouth and kicking at attendants, and had to be forcibly restrained. The victim had spontaneous uncontrollable spasms 

at decreasing intervals until the victim died during a seizure.
68 Bat Rabies and Other Lyssavirus Infections

Case 8. Exposure in India 

(Veeraraghavan, 1955)

The victim chased boys from a grounded bat that they 

were pelting with stones. The victim then picked up the bat 

to place it in a tree, and the bat bit the victim’s forearm, from 

which it was dislodged with great difficulty. The bat was 

discarded. 

Day 1, about 11 weeks later: The victim’s symptoms were not 

recorded. 

Day 2 (perhaps later): The victim had severe general sickness, 

was restless, had difficulty swallowing, and regurgitated 

fluid that was in the mouth. 

Days 3 and 4: The victim’s condition deteriorated. The victim 

became very excited and restless and would shout, but could 

not explain why. 

Day 5: The victim died.

Publishing support was provided by the USGS Columbus Publishing 

Service Center.

Layout by Elizabeth Ciganovich.

Illustrations by Elizabeth Ciganovich and Rosemary Stenback.

Case 9. Exposure in Australia 

(Allworth and others, 1996)

The victim evidently became infected consequent to caring for fruit bats and experiencing a scratch or bite on an 

unknown date. 

Day 1 of symptoms: The victim had pain and numbness in an 

upper limb. 

Day 4: The victim’s muscular control became uncoordinated. 

The victim had difficulty focusing, had shoulder pain, and 

vomited. 

Day 6: After the victim’s throat muscles became paralyzed, 

the victim was unable to swallow and breathe. The victim’s 

condition continued to deteriorate. 

Day 11: The victim became comatose. 

Day 16: The victim died.
Back cover photos (D.G. Constantine)

Top: A California leaf-nosed bat departing from 

a mine tunnel in the Sonoran Desert, California.

Bottom: A diadem leaf-nosed bat, Queensland, Australia. 
Constantine—Bat Rabies and Other Lyssavirus Infections—Circular 1329

Printed on recycled paper