BEHAVIOR AND MAINTENANCE OF CAPTIVE WHITE-WINGED

VAMPIRE BATS, DIAEMUS YOUNGI

W. A. SCHurr, JR., F. MURADALI, N. MONDOL, K. JOSEPH, AND K. BROCKMANN

Natural Science Division, Southampton College, Long Island University,

Southampton, NY 11968 (WAS)
17 Winsor Road, Valsayn Park, Trinidad (FM)

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National Animal Disease Center, Caroni, North Bank Road, Trinidad (NM, KJ)
Department of Ecology and Systematics, Corson Hall, Cornell University, Ithaca, NY 07003 (KB)

Procedures for transportation and maintenance of the white-winged vampire bat, Diaemus
youngi, are documented for the first time. Contrary to previous reports, D. youngi has been
maintained on a diet of defibrinated bovine blood supplemented weekly with fresh chicken
blood. Gregarious by nature, D. youngi exhibits dominance-hierarchy behavior, and behav-
ioral patterns unreported in other species of bats. Although primarily arboreal with regard
to feeding behavior, D. youngi demonstrates the ability to feed terrestrially-a behavior
documented for the first time. Observations of our captive colony contradict anecdotal and
previously published information on feeding and behavior. Differences in feeding behavior
between D. youngi and the common vampire bat, Desmodus rotundus, appear to be related
to selection of prey (arboreal versus terrestrial prey, respectively). In places where these
vampire bats coexist, resource partitioning may serve to reduce competition.

Key words: Diaemus, Desmodus, vampire bats, captive maintenance, behavior

Reports on the maintenance and behavior unstudied. Similarly, relatively little is

of captive vampire bats (Phyllostomidae,
Desmodontinae) generally are restricted to
the common vampire bat, Desmodus rotun-
dus (Dickson and Green, 1970; Ditmars and
Greenhall, 1935; Greenhall, 1965; Trapido,
1946; Wimsatt, 1962a, 1962b; Wimsatt and
Guerriere, 1961). D. rotundus is common
in the wild (Ditmars and Greenhall, 1935;
Greenhall, 1965), easily maintained, and
long-lived in captivity (Dickson and Green,
1970; Trapido, 1946; Wimsatt, 1962a,
1962b; Wimsatt and Guerriere, 1961).
Wimsatt (1978) maintained a colony of D.
rotundus at Cornell University for >20
years and reported that at least one individ-
ual survived> 19 years. The hairy-legged
vampire bat, Diphylla ecaudata, is less
abundant compared with D. rotundus and
reportedly difficult to maintain in captivity
(Greenhall, 1976; Villa-R., 1967) although
Hoyt and Altenbach (1981) maintained
them successfully on a diet of blood from
live chickens. D. ecaudata remains largely
known about the biology of the white-
winged vampire bat, Diaemus youngi
(Greenhall and Schutt, 1996). As with D.
ecaudata, there is consensus that D. youngi
prefers avian blood (Gardner, 1977; Good-
win and Greenhall, 1961; Sazima and Uie-
da, 1980; Uieda et al., 1992).
There are few reports on the maintenance
of D. youngi in captivity, and bats in these
studies were not transported out of the
country of origin. Two adult females, cap-
tured and maintained in Brazil, were al-
lowed to feed on a diet of live chickens
(Uieda and de Araujo, 1987), and four
adults fed on chickens, Gallus gallus, do-
mestic pigeons, Columba livia, and collared
doves, Streptopelia decaocto (Uieda et al.,
1992). Lack of observational data on D.
youngi probably can be attributed to two
major factors: the relative rarity of D. youn-
gi in the wild and reports of difficulty main-
taining D. youngi in captivity. Goodwin and
Greenhall (1961) reported that D. youngi

Journal of Mammalogy, 80(1):71-81, 1999 71

72 JOURNAL OF MAMMALOGY
would not accept bovine blood in captivity,
whereas Pye (1967) reported that D. youngi
took small quantities of bovine blood but
did not survive without avian blood. Con-
trary to Goodwin and Greenhall (1961), we
found that captive D. youngi readily accepts
bovine blood. Bovine blood facilitates
maintenance of captive colonies, as it is rel-
atively easy to obtain in large quantities,
can be stored frozen for extended periods
of time, and eliminates the requirement for
a daily supply of live blood donors. It has
become a dietary staple of captive colonies
of D. rotundus (Dickson and Green, 1970;
Greenhall, 1965; Wimsatt and Guerriere,
1961).
We report here observations on the be-
havior and maintenance in captivity of 12
D. youngi. Many behavioral observations
are reported here for the first time, and this
is the first detailed account of a successful
maintenance program for captive D. youngi.
It should be noted that the daily mainte-
nance program we describe was developed
over the past 25 years by workers at the
National Animal Disease Center, Caroni,
Trinidad. Although this work primarily is
concerned with D. youngi, much of the
maintenance program described here (e.g.,
transportation and housing) can be applied
to D. rotundus. We speculate that captive
colonies of D. ecaudata require a diet com-
posed solely of avian blood. Several refer-
ences provided valuable information on the
care of captive D. rotundus (Dickson and
Green, 1970; Greenhall, 1976; Greenhall
and Schmidt, 1988; Wimsatt and Guerriere,
1961).
MATERIALS AND METHODS
Twelve adult D. youngi (10 males, 2 females)
were collected 17 Februrary-26 July 1993 by
mist netting from the following locations in
Trinidad (Guayaguayare, Mayaro; Ortoire, May-
aro; Matura, St. Andrew; Raghunan Road, Ca-
roni, Tabaquite, Caroni). All bats were main-
tained at the National Animal Disease Center,
Caroni, Trinidad, before transport to Cornell
University, Ithaca, New York, on 27 July 1993.
Vol. 80, No.1
Presence or absence of a cartilagenous zone in
the epiphyseal region of the fourth metacarpal-
phalangeal joint was used for determination of
relative age (Kunz and Anthony, 1982). For pur-
poses of identification and record keeping, fore-
arms of bats were fitted with individually num-
bered metal bands soon after capture.
Before and during transport, we complied
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with all rules and regulations concerning collec-
tion, export, and import of live animals. Regu-
lations differ from country to country and spe-
cial care should be taken to insure that all re-
quired permits are completed and available for
presentation during transit. Before our trip, we
contacted the United States Fish and Wildlife
Service's Office of Management Authority
(4401 North Fairfax Drive, Room 432, Arling-
ton, VA 22203), and we were instructed that
upon entering the United States with the bats,
we were to declare them and present United
States Customs Form 3177. Specimens of D.
youngi and D. rotundus, including several newly
captured individuals, were transported via com-
mercial airline from Port of Spain, Trinidad, to
New York City, and then by car to Ithaca, New
York. A wooden travel crate was constructed (90
by 90 by 90 cm), which securely held a pair of
wire-mesh, stainless steel, cages. Airholes (0.5
cm diameter) were drilled into the sides of the
crate, which were conspicuously labeled (e.g.,
Live Animals, Do Not Open, This End Up). On
several evenings before the trip, bats were
placed into the wire cages and the cages fitted
into the crate to acclimate the animals to the
transport container. The bottom of the cage con-
tained an aluminum tray onto which absorbant
material (newspaper) was placed. Before sealing
the crate for transport, bats were fed and placed
together into one of the wire cages (11 D. ro-
tundus were transported in the second wire
cage), and cages were fitted into the crate. The
crate lid was sealed with wood screws. All 23
of the bats survived transportation from their
country of origin; the crate was sealed for 18 h.
Captive D. rotundus and D. youngi were
maintained at Cornell University from July 1993
to September 1996. In September 1996, bats
were transported to the Burnett Park Zoo, Syr-
acuse, New York. At Cornell University, access
to the bat colony was restricted to research and
animal-care personnel. Specimens, separated by
genus, were housed in two, stainless-steel,
small-animal cages (56 by 51 by 43 cm). News-
1999 SCHurr ET AL.-DlAEMUS YOUNG1 IN CAPTIVITY
paper was used as an absorbent on the cage floor
and changed daily. A warm damp sponge was
used to remove blood and excreta from cage
walls, and cages were scrubbed with cage clean-
ing detergent on a weekly basis.
All research and animal-care personnel that
had any contact with our vampire bats had first
undergone a pre-exposure prophylaxis for rabies
virus. Although our D. youngi became tame and
did not attempt to bite us after their 1st week of
captivity, we continued to use thick, loose-fit-
ting, leather gloves when handling them.
Room temperature in our laboratory was 21-
24°C; relative humidity was variable at 20-S0%.
Bats were maintained on a cycle of 12 h of in-
direct flourescent ceiling light (0600-1800 h)
and 12 h of darkness. This schedule approxi-
mated the day-night light conditions in Trinidad.
We had few problems with illness in our col-
onies. Occasionally, however, bats presented
some or all of the following symptoms: lethargy,
dehydration, bloated abdomen, sneezing, and
vomiting of a previous blood meal. We treated
our bats in the following manner. At the first
sign of illness, the bat was isolated in a wire-
mesh travel cage draped in a clean towel to re-
duce drafts. Ambient temperature was increased
(ca. SoC) with an electric space heater. If dehy-
dration was observed, the bat was rehydrated
daily with a 1.0-ml subcutaneous injection of
0.9% saline solution. Animals were treated as
dehydrated if the loose skin covering the region
between their shoulder blades did not spring
back into place immediately after being gently
pinched by the handler. An antibiotic (10 mg
Baytril/kg-Bayer Corporation, Shawnee Mis-
sion, KS) was administered subcutaneously,
once per day. Subcutaneous injections were
made beneath the loose skin of the upper back.
In the evening, the bat was allowed to feed on
a live chicken. This treatment regime proved ex-
tremely successful, and all bats treated in this
manner recovered within S days. At that time,
treatment was discontinued, and the bat was re-
introduced into the colony.
Our feeding regime for D. youngi was iden-
tical to one that had been developed and suc-
cessfully employed to maintain bats in Trinidad
(Muradali et aI., 1993). Specifically, bats were
fed bovine blood six times per week and offered
live chickens once per week. Contrary to pre-
vious reports (Goodwin and Greenhall, 1961;
Pye, 1967), our D. youngi readily accepted de-
73
fibrinated bovine blood. That blood was ob-
tained monthly at a local slaughterhouse from
freshly killed domestic cattle. A clean, plastic,
7.S-1 container was used to collect blood. To pre-
vent clots during storage or feeding, blood was
fibrinated immediately by manual agitation. A
metal kitchen colander was used to skim off
clots, and the newly defibrinated blood was
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stored for transport in 3.78-1 plastic jars. In the
laboratory, the blood was transferred to individ-
ual 473-ml glass bottles. We stored enough in
each bottle to feed each bat 30 mi. That volume
was more than sufficient because it was about
twice the amount normally consumed by our
bats. Bottles of blood were frozen at -20°C until
use-keeping the bottle's lid loose to prevent
glass bottles from cracking when frozen. On the
day that the blood was used, a bottle was thawed
slowly, and blood was poured into a plastic ice-
cube tray and served at room temperature. We
filled one ice-cube tray depression (ca. 30 ml)
with blood for each bat. Because D. youngi
feeds arboreally in the wild (Gardner, 1977;
Goodwin and Greenhall, 1961; Sazima and Uie-
da, 1980; Uieda et aI., 1992), we elevated the
feeding tray ca. 30 cm off the cage floor with a
wooden block. Bats were fed in the late after-
noon or evening, and the tray was removed and
cleaned the next morning.
In our study of feeding capacity, the average
daily intake of blood was determined, over a 30-
day period, for isolated D. youngi. Eight indi-
viduals were used (four times each) in 32 trials.
Thirty milliliters of blood was offered in a spill-
proof feeding container. The next day, the
amount of blood remaining was subtracted from
the starting amount to yield the amount con-
sumed. A correction factor of 2 ml was applied
to account for volume loss due to evaporation
and blood that clung to the bat during feeding.
Although our bats were placed into a stain-
less-steel feeding cage with the chickens, >SO
arboreal feeding sessions were carried out in a
1.8 by 3.0-m screened enclosure, with chickens
roosting on a branch. During preparation for
those sessions, a chicken was placed onto the
upper section of a branch (inverted L-shape, ca.
1.3 m in length, 4 cm in diameter), the lower
section of which was attached to a stand con-
structed with two sections of S- by lO-cm pine
studs. With the room lights lowered or turned
off, a single D. youngi was placed on the lower
74 JOURNAL OF MAMMALOGY
part of the branch (ca. 15 cm off the floor) in a
head-up position.
One evening per week, two specimens of D.
youngi were placed in a stainless-steel feeding
cage (55 by 50 by 42.5 cm). A single subadult
chicken (9-10 weeks old) was placed in the
cage. Bats were permitted to feed on the chicken
overnight.
For observations of terrestrial feeding behav-
ior a large wire-mesh cage (51 by 84 by 37 cm)
was used. Twenty terrestrial feeding sessions
were observed, during which a single bat was
placed into the cage with one adult or subadult
chicken. Because there were no perches in the
cage, the chickens stood on the cage floor. Room
lights were turned off and a small flashlight fit-
ted with a red-gel filter was used to observe ter-
restrial feeding behavior. Sex and age of chick-
ens and the identification number of the bat used
were noted. Behavior during those sessions was
recorded in a notebook.
RESULTS AND DISCUSSION
Wimsatt et al. (1973) designed a cage for
vampire bats with a novel and time-saving
sanitation system, but satisfactory housing
can be provided with less elaboration. Basic
requirements for cages are that they prevent
bats from escaping, allow bats to move
about and roost comfortably, and be easy to
clean. Because excreta is tar-like and cor-
rosive, housing constructed of stainless
steel or plastic is advisable.
In a previous study (Uieda and de Ar-
aujo, 1987), D. youngi was maintained at
temperatures similar to those in our labo-
ratory. In Trinidad, our bats were caged in
a open-air facility where temperatures gen-
erally were 21-29°C. Greenhall (1965)
maintained D. rotundus and D. youngi in
Trinidad at similar temperatures. Although
it has been reported that vampire bats re-
quire high relative humidity (60% mini-
mum) in captivity (Joermann, 1988), D.
youngi does not appear sensitive to low hu-
midity, and we have maintained D. youngi
and D. rotundus at relative humidities of
20-50%. This agrees with previous data on
maintenance of captive specimens of D. ro-
tundus (Wimsatt and Guerriere, 1961) and
Vol. 80. No.1
indicates that captive vampire bats may not
be as sensitive to relative humidity as other
species of bats kept in captivity (e.g., ves-
pertilionids). Racey (1972) reported that
low humidities can be injurious to wing
membranes. Although wing injuries have
not been noted in our laboratory, humidity
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should be given careful consideration in
studies where bats are required to fly exten-
sively.
Ectoparasite infestation was observed in
newly captured vampire bats, but this prob-
lem disappeared within a month. As re-
ported in previous studies (Dickson and
Green, 1970; Wimsatt and Guerriere, 1961),
this problem disappeared gradually over the
course of several weeks, probably due to
the disruption of ectoparasite lifecycles in
captive conditions (Wimsatt and Guerriere,
1961). Information on ectoparasites and en-
doparasites of D. rotundus exists (Mendez,
1988), but data are fragmentary with regard
to D. youngi (Mendez, 1988).
Cultures of the bovine blood were tested
(Cornell University, Department of Pathol-
ogy) and indicated presence of bacteria that
may have caused occasional gastrointestinal
distress observed in several of our bats.
Tests to determine presence of Salmonella
were negative.
Data on the demeanor of vampire bats in
captivity is variable. Some authors have
found, as we have, that D. rotundus does
not tame in captivity (Dickson and Green,
1970), while others report that D. rotundus
tames readily (J. S. Altenbach, pers. comm.;
Novick, 1963; Wimsatt and Guerriere,
1961). D. youngi has been described as ag-
gressive in captivity (Uieda and de Araujo,
1987), while we have found the opposite to
be true. Like D. rotundus, D. youngi ap-
pears to be gregarious by nature and can be
housed in a group dwelling. There have
been no problems maintaining mixed-sex
populations, although D. youngi does ap-
pear to exhibit dominance hierarchies. The
following behaviors were noted in Trinidad,
when a new animal was introduced into an
established group and at Cornell University,
1999 SCHUTT ET AL.-D1AEMUS YOUNG1 IN CAPTIVITY
the day after their arrival. In both instances,
a male that had been captured recently was
approached by another male. These bats are
hereafter referred to as the newcomer and
the dominant male, respectively. The bats
faced each other and both displayed a rep-
ertoire of threat behavior. After rising up
into an elevated stance, a distinctive hissing
was accompanied by other audible vocali-
zations varying in frequency and intensity.
Forelimbs, held in a closed-wing position,
were used for parrying. That behavior,
which was accompanied by feints, lunges,
and head bobbing, lasted ca. 20 s. In the
Trinidad episode, the newcomer then
crouched quietly, partially opened his
mouth and exposing a pair of enlarged oral
glands. The dominant male then inserted his
tongue into the mouth of the newcomer for
several seconds. At Cornell University, af-
ter a similar face-off (in this instance there
was no tongue insertion), the dominant
male approached the newcomer from the
rear. With both bats vocalizing loudly, the
dominant male partially extended a wing,
wrapped it around the back of the newcom-
er, and climbed over it. This took several
seconds, after which time the bats ceased
their vocalizations and moved away from
each other. That stylized fighting behavior
was observed on a number of occasions in
our lab especially during sessions with pairs
of bats feeding on live chickens. Unlike the
violent and bloody clashes reported in stud-
ies on D. rotundus during roost-defense en-
counters (Wilkinson, 1985, 1988), those be-
tween D. youngi have not resulted in blood-
shed.
Diaemus youngi possesses two large oral
glands whose function is unknown (Good-
win and Greenhall, 1961; Greenhall, 1988).
When disturbed (such as we have seen dur-
ing dominance-hierarchy behavior or at
times during handling), D. youngi opens its
mouth, and these large glands can be clear-
ly observed filling the caudolabial part of
the oral cavity. A peculiar hissing vocaliza-
tion, described by Goodwin and Greenhall
(1961) as a "pssst," is accompanied by
75
emission of a fine spray of liquid, which has
a strong musky odor. We have observed
musk spraying only during threat response.
We suggest that this aerosol may be used to
discourage predators and may serve a role
in territory marking and individual recog-
nition. Enlarged oral glands and musk
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spraying have not been reported in D. ro-
tundus or D. ecaudata. There have been
studies on anatomy and histochemistry of
the salivary glands of D. rotundus (Di-
Santo, 1960) and the salivary antihemostat-
ic factors of D. rotundus and D. youngi
(Hawkey, 1966, 1967). However, there
have been no studies on the anatomy of sal-
ivary glands in D. youngi or D. ecaudata.
Our vampire bats spent considerable time
roosting. During that time, they clung to
each other with claws and thumbs of their
hind limbs. Hind limbs not in contact with
conspecifics generally were used to grasp a
branch or the cage ceiling. Number of bats
roosting together varied but most often,
they clung together in one dense mass. Dur-
ing roosting, we frequently observed an in-
dividual apparently biting the bat hanging
adjacent and ventral to it. Upon examina-
tion, however, we learned that those were
not actually bites but rather one bat took a
loose fold of skin from a cage mate (usually
from the upper back or dorsal surface of the
neck, the nape) into its mouth (Fig 1a).
There, the skin fold was held securely for
several seconds. Bats on the receiving end
of that mouth grasp did not appear dis-
turbed. Upon examination of those animals,
neither wounds nor hair loss were observed.
This behavior has not been reported in other
studies on D. youngi in the wild (Sazima
and Uieda, 1980; Uieda et aI., 1992) or in
captivity (Uieda and de Araujo, 1987; Uie-
da et al., 1992).
Mouth grasping is most reminiscent of
behavior seen in non-fiying neonatal bats as
they cling onto their mother's body with
their mouths, thumbs, and hindlimbs. We
suggest that mouth grasping is a component
of dominance-hierarchy behavior. A. Stem
(pers. comm.) reported similar behavior in
76 JOURNAL OF MAMMALOGY
FIG. I.-a) Mouth-grasping behavior by Diae-
mus youngi. While roosting or during domi-
nance-hierarchy behavior, individuals often use
their mouths to grasp cage mates by the loose
skin between the shoulders or on the nape of the
neck. b) D. youngi mounting a young female
chicken dorsally prior to feeding. Rather than
becoming agitated, female chickens mounted in
this fashion assume a crouched posture similar
to that observed during copulatory behavior; we
suggest that the position and body weight of the
bat trigger submissive behavior in female chick-
ens.
Pteropus vampyrus during dominance
fights and copulations in several species of
Pteropus. Additionally, mouth grasping ap-
pears to assist D. youngi in stabilizing its
body during roosting, especially during
shifts in position of hind limbs. Mouth-
grasping behavior was not observed in our
D. rotundus nor has it been reported in oth-
er microchiropterans (K. E Koopman, pers.
comm.).
In previous studies on D. youngi in cap-
tivity (Uieda and de Araujo, 1987; Uieda et
Vol. 80, No.1
al., 1992), live birds were used as the sole
source of blood. Although our bats readily
accepted defibrinated bovine blood, we be-
lieve that citrated blood (Wimsatt and Guer-
riere, 1961), also would be acceptable. Be-
cause these bats feed primarily on avain
blood in the wild, we chose to provide them
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with a weekly supplement of blood from a
live chicken. While feeding from an ele-
vated ice-cube tray, bats hung from the cage
ceiling by their hind limbs and extended
their upper bodies toward the tray (until
their mouths were just above the level of
the blood). As the bat fed, the tongue was
extended into the blood and withdrawn at a
rate of ca. 4 licks/so Feeding usually lasted
ca. 15 min and fighting was never observed
during those sessions.
Observations on feeding capacity of D.
rotundus in captivity (Wimsatt and Guerri-
ere, 1961) indicated that average daily in-
take of blood by two groups was 15.3 and
15.6 mI. In our study, mean daily intake for
D. youngi was 16.4 ± 3.6 (SD) mlIday, but
there were differences in the method of
those two studies (e.g., the study of D. ro-
tundus was carried out over a full year but
animals were only fed 6 days/week). Wil-
kinson (1984) noted that D. rotundus did
not necessarily feed every night and some
bats received blood regurgitated by other
individuals. Blood sharing has not been re-
ported in D. youngi.
Sazima and Uieda (1980) reported on the
feeding behavior of D. youngi on free-rang-
ing poultry (chickens, Guinea fowl, and tur-
keys), but were unable to observe the actual
biting behavior and the prey's reaction. Uie-
da and de Araujo (1987) documented some
aspects of maintenance of D. youngi and D.
ecaudata in captivity but did not go into
great detail concerning feeding behavior.
Uieda et al. (1992) studied arboreal feeding
on chickens and smaller birds (pigeons and
doves). They described two feeding pos-
tures used by bats, hanging and quadrupe-
dal, and stressed the importance of diameter
of branch as a limiting factor in attacks by
D. youngi on small birds.
1999 SCHUTT ET AL.-DlAEMUS YOUNG1 IN CAPTIVITY
The following description characterizes a
typical arboreal feeding session in our lab-
oratory and in Trinidad (Muradali et aI.,
1993). Holding on with its thumbs and hind
feet, D. youngi clings quietly with its ven-
tral body surface held tightly to the branch.
During this period, which lasts :::; 10 min,
the bat raises its head occasionally, sniffs at
the air with its mouth partially open, and
observes the chicken perched above it. At
this time, the bat moves to the underside of
the branch and slowly climbs closer to the
perching chicken. D. youngi advances one
limb at a time (with the grip on the branch
maintained by the three non-moving limbs)
always keeping the branch between itself
and the underside of the chicken. After po-
sitioned beneath the chicken's digits (ca. 1
min), the bat situates itself so that its head
is close to a single digit of the roosting bird.
On most occasions, the bat positioned its
head next to the posteriorly directed hallux
(digit 1). It appears that feeding from this
digit keeps the bat concealed under its prey
to a greater degree than would occur if an
anteriorly directed digit (digits 2-4) was
fed upon. Moving its mouth to the digit, the
bat's tongue is extended and withdrawn (in
'a piston-like manner), and the potential bite
area is licked at a rate of ca. 2 licks/so
It is not known if saliva of vampire bats
contains anesthetic or enzymatic compo-
nents to aid in preparation of bites or if lick-
ing softens the keratinized reticulate scales
that cover the surface of the digit. The bite-
preparation period lasts from 10 s to 2 min.
Bites were never inflicted without this prep-
aration period. In all of our observations,
immediately prior to the bite, D. youngi
turns its head so that the midline of the head
is perpendicular to the long axis of the digit
being bitten. The actual bite is never initi-
ated quickly or violently but occurs when
licking stops and the bat applies its partially
opened mouth to the site. At this time, it
appears that the jaws close, driving the ra-
zor-sharp upper incisors through the epi-
dermis and into the dermis. This position is
maintained for 1-2 s until one of the fol-
77
lowing events occurs to complete the bite.
Either a series of quick backward steps are
made by the bat or the chicken shifts its
position slightly on the branch and the digit
is pulled away from the bat's mouth. In the
majority of bites, chickens show little, if
any indication that they have been disturbed
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by the bite.
Bites appear as small crater-shaped in-
dentations that extend into the dermis-a
layer richly supplied by blood vessels. Bites
inflicted by D. rotundus appear similar and
have been well characterized (Greenhall,
1988). In D. rotundus, canine teeth are used
to shave the fur around a wound site. The
lobed lower incisors secure a firm hold on
the skin while the concave upper incisors
remove a divot of flesh. D. youngi may use
its canines in a similar manner to shave
feathers, but this has not been reported and
we did not observe this behavior.
After the bite is inflicted, D. youngi re-
turns to the wound and resumes the licking
behavior at a rate of ca. 4 licks/so Within
30 s of the bite, blood flows freely from the
wound. Blood is conducted into the bat's
mouth via two grooves on the ventral sur-
face of the tongue (Uieda, 1986) through a
gap between the two pairs of lower incisors,
and above a V-shaped groove in the lower
lip. Most feeding sessions last ca. 15 min
and may stop temporarily if the bird
changes positions or becomes alarmed.
When this occurs, the bat may back off,
concealing itself under the branch, until the
bird calms. On other occasions, the bat re-
mains in position and appears to hide under
the body of the bird. After the bird settles
down, the bat re-establishes its position and
begins feeding. At no time (during either
arboreal or terrestrial feeding behavior) did
we observe a feeding bat make a second
bite on the same chicken. Interruptions in
feeding always were followed by a resump-
tion of feeding from the same wound. We
have regularly observed periods of urina-
tion, lasting several seconds, commencing
at ca. 5 min into the feeding session. While
urinating, one hind limb usually is extended
78 JOURNAL OF MAMMALOGY
laterally and slightly downward, preventing
the bat from soiling itself.
On several occasions, bites made on the
terminal digital pad of digit 1 (ventral to the
proximal portion the claw) were not licked
during feeding. Instead, the protruding dig-
ital pad was taken into the bat's mouth as
it fed. On two occasions, we attempted to
dislodge the bat from the bird's digit and
found that it took a considerable amount of
pulling for the bat to release its hold. Our
attempts to dislodge the bat did not disturb
the chicken, and no further wounds were
apparent on them after the bat was pulled
free. We suggest that suction (rather then
the use of teeth) was used during this pe-
culiar feeding behavior. This observation
lends support to the hypothesis that vampire
bats may use their mouths to form a seal
around the wound when feeding (Greenhall,
1988). Suction and wound licking appear to
be two methods for obtaining blood from a
wound. Because suction was observed only
when D. youngi fed on the distal digital
pad, it appears that these feeding methods
may be dependent on location of wounds.
After ca. 15 min, D. youngi concluded its
feeding session by releasing its thumbs
from their hold on the branch. After hang-
ing briefly by its hind limbs, the bat re-
leased its grip on the branch. In many in-
stances, flight was initiated by this method,
but on a few occasions D. youngi dropped
directly to the ground before scrambling off
to hide.
Because of the design of cages, cages
used during weekly supplements of diet
with chicken blood were not used to ob-
serve bat-bird interactions but solely for
feeding purposes. In all cases, subadult
chickens placed into the feeding cage were
removed and euthanized after 4 h. In a pre-
vious study (Uieda and de Araujo, 1987),
bats and chickens were kept together con-
tinuously. Those chickens exhibited de-
creased activity and weight loss, and a
number of them died within 7-10 days. We
examined locations of bites on carcasses of
chickens and observed that most bites were
Vol. 80, No.1
located on digits or the posterior side of the
intertarsal joint. Some bites, however, were
made on the comb, neck, or abdomen, and
a single bite was found on the proximal pre-
patagium of the wing.
Descriptions of terrestrial feeding behav-
ior by D. youngi are lacking. Uieda et al.
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(1992) mentioned a single observation of
quadrupedal posture employed by D. youn-
gi during feeding on a domestic pigeon, but
it is unclear if that occurred during arboreal
or terrestrial feeding. The following is a de-
scription of a typical terrestrial feeding ses-
sion in our laboratory. After the bat is
placed into the cage, it uses a hopping gait
to position itself under the posterior part of
the bird's body behind the feet. If the bird
moves about the cage, the bat hops after it,
trying to maintain its position under and be-
hind the bird. After the bird is stationary,
the bat approaches the posterior side of the
metatarsus. Raising itself into an elevated
quadrupedal stance, similar to that de-
scribed for D. rotundus (Altenbach, 1979,
1988; Sazima, 1978), a bite-preparation
phase is initiated. In most instances, bites
are made on the posterior or lateral surface
just distal to the intertarsal joint. Larger
scales (scutes or scutella) are partially or
fully removed during bites to the metatar-
sus. Bite infliction is similar in terrestrial
and arboreal feeding in that the bite is com-
pleted if the chicken moves away from the
stationary bat or the bat steps backward
away from the bird's leg. After blood be-
gins to flow from the wouQd, the bat ap-
proaches the wound and begins to feed. If
the bird moves around the cage, the bat dis-
engages feeding and follows the bird by
hopping behind and underneath it. Feeding
resumes when the bird assumes a stationary
pose.
During a number of terrestrial feeding
sessions, we noted agitated and aggressive
behavior by male chickens toward a bat.
That behavior consisted of rapid pacing
back and forth in the feeding cage by the
bird accompanied by pecks directed toward
the bat. Inevitably, that behavior resulted in
1999 SCHUTT ET AL.-D1AEMUS YOUNG1 IN CAPTIVITY
the bat retreating to the cage ceiling where
it remained until removed. D. youngi were
observed feeding on male chickens on
many occasions, but they seem to prefer fe-
male chickens.
In ca. 10% of terrestrial feeding sessions
observed, the bat did not feed from the
chicken's leg or digit but leaped or climbed
onto the chicken's back to feed on the head
region. In those instances, the bat posi-
tioned itself with its body facing cranially
with regard to the bird. That position was
maintained by partially spreading (abduct-
ing) forelimbs while grasping onto dorsal
feathers with both thumbs and digit claws
of the hind limb.
Male chickens that were mounted dor-
sally by D. youngi quickly became agitated,
often dislodging bats from their back. When
young female chickens were mounted dor-
sally, however, rather than becoming agi-
tated, they assumed a crouched posture
(Fig. 1b) similar to that observed during
copulatory behavior. They remained in this
position even when moving around the
cage. We suggest that the forces applied to
the female bird's back by the bat triggers
this innate submissive response behavior.
Attacks on poultry roosting in trees are
frequent in Trinidad, but there have been no
reports of attacks on birds confined in cages
on the ground although the mesh of these
cages would allow entrance by D. youngi.
Our feeding experiments, however, showed
that D. youngi are quite capable of efficient
terrestrial feeding. Examination of stomach
contents of D. youngi has, in fact, shown
them to feed on mammalian blood (e.g.,
cattle and pigs-Greenhall, 1988). Our ob-
servations suggest that although D. youngi
is capable of feeding terrestrially, it will not
do so as long as arboreal prey is available.
One possibility is that arboreal feeding
evolved to reduce competition between D.
youngi and the terrestrially feeding D. ro-
tundus where their ranges overlap, although
Schutt and Altenbach (1997) suggest that
arboreal feeding is a primitive behavior in
vampire bats.
79
According to Koopman (1988), there is
disagreement with regard to the recognition
of Desmodus and Diaemus as seperate gen-
era. Handley (1976) for example, included
Diaemus within the genus Desmodus. Enu-
merating 18 morphological characters,
Koopman (1988:12) was "inclined to rec-
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ognize Diaemus as a valid genus along with
Diphylla and Desmodus." In our studies,
we observed a number of behavioral char-
acters, related to feeding and roosting be-
havior (e.g., mouth grasping), which sup-
port that view.
The locomotor morphology of D. rotun-
dus was studied in detail by Altenbach
(1979, 1988) and Schutt et al. (1997). Al-
tenbach (1988) described the quadrupedal
stance of this genus as elevated. This stance
apparently allows "immediate movement in
any axis parallel to the surface or move-
ment away from it without repositioning of
either pectoral or pelvic limbs" (Altenbach,
1988:72). The elevated posture appears to
be an adaptation for rapid terrestrial loco-
motion in D. rotundus, allowing it, for ex-
ample, to initiate flight by jumping verti-
cally into the air. Jumping functions to get
the heavily loaded bat airborne. It also aids
in predator avoidance (e.g., snakes) and
may prevent the bat from being stepped on
by ungulate prey while feeding.
In contrast, D. youngi exhibits a more
crouched stance than D. rotundus. The
body of D. youngi is held so that its center
of mass is closer to the support (the ground
or branch). D. youngi has never been re-
ported to perform flight-initiating jumps in
the wild or in captivity. Additionally, we
never observed this behavior during force-
platform experiments analyzing terrestrial
locomotion (Schutt et al., 1997). We sug-
gest that D. youngi, with its arboreal feed-
ing habits, is under less threat from terres-
trial predators than D. rotundus. Unlike the
terrestrially feeding D. rotundus, D. youngi
is in no danger of being crushed by large-
ungulate prey. During terrestrial feeding
sessions with D. youngi, chickens were ob-
served to suddenly squat down on top of a
80 JOURNAL OF MAMMALOGY
feeding bat. No ill effects have been ob-
served in bats in those situations, and on
most occasions there was no interruption in
feeding.
We believe that the stance of D. youngi
is less upright than that of D. rotundus for
two additional reasons: climbing efficiency
and crypsis. As a climber and arboreal
hunter, D. youngi has decreased the tenden-
cy to topple from its support (the branch)
by several means: 1) keeping its center of
mass close to the support (this is especially
valauble during vertical climbing behavior);
2) climbing underneath the branch; 3) using
its thumbs and hind feet to exert a torque
that resists the toppling moment (for a re-
view of the biomechanics of climbing see
Cartmill, 1985). Additionally, by clinging
with its body under and close to the branch,
D. youngi is better hidden from the sight of
its avian prey. D. youngi, initiates flight by
simply dropping from its arboreal feeding
position and has no need for the flight-ini-
tiating jumps that are characteristic of the
terrestrially feeding D. rotundus.
In summary, specimens of D. youngi and
D. rotundus captured in Trinidad have been
maintained successfully in captivity. The only
difference in maintenance programs for these
two genera is a once per week supplement of
live chicken blood for D. youngi. Many dif-
ferences in behavior, however, have been not-
ed between D. youngi and D. rotundus. We
believe that these differences are related to
their mode of prey selection. D. rotundus ex-
hibits a number of unique adaptations for ter-
restrial feeding (e.g., speed, flight-initiating
jumps). Although D. youngi is quite capable
of terrestrial feeding, there have been no re-
ports of this mode of feeding in the wild, and
it appears that D. youngi is better adapted for
arboreal feeding. Blood feeding in mammals
is restricted to three genera of vampire bats,
and these exhibit many unique adaptations
that allow them to maintain a sanguivorous
lifestyle (e.g., highly modified digestive and
excretory systems, quadrupedal locomotion,
altruistic behavior). In Trinidad where D. ro-
tundus and D. youngi coexist, competition
Vol. 80, No.1
between these genera may be reduced
through resource partitioning.
ACKNOWLEDGMENTS
This study was made possible by funding
from the American Museum of Natural History
(Theodore Roosevelt Memorial Fund. Coleman
Postdoctoral Fellowship); the American Society
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of Mammalogists (grant-in-aid). Sigma Xi. and
Cornell University. Special thanks to the Min-
istry of Agriculture, Lands and Marine Re-
sources. Trinidad. The authors thank the follow-
ing individuals in Trinidad: the vampire-bat
crew (A. Johnson and P. Wallace). D. Booda. C.
James, S. Johnstone. N. Gyan. H. Nelson, S.
Ramdass. J. Latchman, Ali family. and especial-
ly G. Ramsawak and O. Ramsawak (our hosts
at Mount Saint Benedict's). We thank individu-
als at Cornell University: J. W. Hermanson. J.
Bertram, D. McCleam. H. Evans. D. Cullinane,
Y. H. Chang. C. Coen. P. Faure. R Quimby. L.
Carbone. H. Bartlett. D. Woodin. C. Grant, and
L. L. Lafrance. We thank the Department of
Mammalogy at the American Museum of Nat-
ural History, especially N. B. Simmons, R.
MacPhee, D. Lunde. T. Conway. and K. R
Koopman. Thanks also to J. S. Altenbach. R.
Dugal. R. Adamo. R. "Tuna" Sinclair. W. Uie-
da. A. Stem. T. Kunz, M. Schutt. J. Schutt. W.
R. Schutt. and Bloomfield College (especially J.
Noonan. I. Anderson. M. LaBare, and P. Russo).
This paper is dedicated to our friend. A. M.
Greenhall, who pioneered the modem study of
vampire bats. Without his encouragement. this
work would not have been undertaken.
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Associate Editor was Troy L. Best.