Fruit Bat-Aza - Standard - Care

Standardized Guidelines for Fruit and Nectar Bat Care
List of Common and Scientific Names , Conservation Status, and numbers of Fruit Bats
held in North American Institutions, recorded in ISIS as of Jan 27,2006
Family - Pteropodidae
Scientific Name Common Name USFWS/CITES # Held in N.America
Cynopterus brachyotis Dog-faced fruit bat 19.25.0

Eidolon helvum Straw-colored fruit bat 220.197.18
Epomophorus wahlbergi Wahlberg’s epauletted bat 0.3.0
Pteropus conspicillatus Spectacled flying fox 0.1.0
Pteropus giganteus Indian fruit bat / II 59.66.8
Pteropus hypomelanus Island fruit bat / II 30.63.1
Pteropus poliocephalus Gray-headed flying fox / II 15.19.0
Pteropus pumilus Little golden-mantled fruit bat / II 22.23.0
Pteropus rodricensis Rodrigues fruit bat E / II 137.109.2
Pteropus vampyrus Large fruit bat / II 64.52.0
Pteropus voeltzkowi Pemba fruit bat / II 0.3.0
Rousettus aegyptiacus Egyptian fruit bat 146.156.21; 131 group
Rousettus lanosus Ruwenzori long-haired fruit bat 69.65.5
Family - Phyllostomatidae
Scientific Name Common Name USFWS/CITES # Held in N.America
Anoura geoffroyi Geoffroy’s tailless bat 1.4.0
Glossophaga soricina Long-tongued bat 402 group
Carollia perspicillata Short-tailed fruit bat 57.206.32; 3433 group
Artibeus jamaicensis Jamaican fruit bat 91.102.58; 1647 group
Artibeus lituratus Great Trinidadian fruit bat 2.0.0
Leptonycteris curasoae Southern Long-nosed bat 3.8.1
1. Abiotic Environmental Variables (address both exhibit and off-exhibit holding)

1.1 Temperature:
Fruit bats originate from tropical areas and do not tolerate low temperatures for extended
periods.1 Although researchers disagree on the optimal temperature in which to house bats,
laboratory studies have found that tropical bats do best in temperatures ranging between 70 and
90 degrees Fahrenheit (21-32˚C). According to Wilson (1988), Rasweiler (1975), and Barnard
(1991), the ideal temperature is 80 degrees Fahrenheit (27˚C). Bats do best when a constant
temperature is maintained.
In outdoor enclosures, supplemental heat is needed for temperatures below 70 degrees

Fahrenheit (24˚C) and, in regions where temperatures fall below 50 degrees Fahrenheit (10˚C)
for prolonged periods, indoor enclosures are necessary. If heat lamps or brood-rite heaters are
used they should be properly shielded from bats so that the animals cannot burn themselves or
chew through wires. In addition, we recommend creating multiple paths to heat sources to reduce
aggression between bats.
Shaded areas must always be provided in outdoor enclosures (e.g., by use of vegetation,
plywood structures, etc.). Shelter from rain & wind is also required.
1.2 Humidity: There are few data available regarding the proper humidity for bats.
Bats appear to do well at relative humidities of 60-90 percent (Rasweiler, 1975; Wilson, 1988).
Barnard suggests a relative humidity of 55 – 65% since high humidities can promote the growth
of potential pathogens. Low humidity appears to be a problem for some species. To maintain
proper humidity, we recommend using mister hoses when temperatures go above 85 degrees
(29˚C). Symptoms of low humidity include dry skin or wing membranes and/or cracked nails.
1.3 Illumination:
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1.3.1 Identify light intensity, spectral, and duration requirements.
Because there are few scientific studies describing circadian rhythms of bats, lighting is probably
best when it imitates natural photoperiods (Wilson, 1988; Rasweiler, 1975). Fruit bats do well on a
10/12 to 12/10 light/dark cycle (Wilson, 1988; MacNamara et al., 1980), and should never be kept
in 24 hour darkness or light.
1.3.2 Address the impact of and need for daily changes in light intensity and seasonal
changes in light intensity and duration.
Fruit bats do not need to be kept in a nocturnal exhibit. Many species are diurnal or crepuscular
and therefore active during the day. However, many species, especially nectar-feeding bats, are
more active at night and managers may prefer reversing the light cycle. Reverse red or blue
lighting is acceptable for use for display purposes, as are incandescent or florescent lights
(House and Doherty, 1975). If a reversed cycle is used, halogen lights, fluorescent tubes, and
blue filters make up a good lighting system. The day cycle lights are fluorescent and the nocturnal
lights are halogen with blue filters. Blue light can be used (rather than red) because it simulates
the color and intensity of moonlight and does not change the color of the bats. We strongly
recommend using a dim/anti-dim cycle to reduce the shock of immediate bright light or complete
darkness. During the fruit bat "day" cycle, shaded or darkened areas for roosting should be
provided. Access to full sunlight is beneficial for Pteropus and Eidolon. Fruit bats can see most
colors of light, including red.
1.4 Space
1.4.1. Identify what behaviors should be expected based on the behavioral repertoire
observed in the wild, which of these behaviors animals of a given size should be able to
exhibit within the enclosure/exhibit and how these behaviors influence the size and
configuration of enclosures. As appropriate address both horizontal and vertical space
as well as the appropriate level of complexity within that space. Adequate space should
be evaluated in terms of space that is useable for the taxon.
Fruit bats require ample space to perform normal maintenance behaviors and do best in
enclosures large enough to permit free flight (Wilson, 1988). USDA requires that flying mammal
species must be provided with sufficient unobstructed enclosure volume to enable movement by
flying and sufficient roosting space to allow all individuals to rest simultaneously [AWA Section 13;
9 CFR, Part 3, Section 3.128]. Bats that are unable to fly for a month or more may have muscle
atrophy and lose the ability to fly (Wilson, 1988).
Bats maintained for exhibition purposes should be housed in enclosures that promote
flight and roosting and allow for a broad spectrum of behaviors such as wing displays, grooming,
foraging, and social interactions Enclosures should be eight times the wing span long and four
times the wing span wide. .For example, an enclosure housing Island flying foxes (Pteropus
hypomelanus), a species with wing spans reaching 3.5 feet (1.07 m), should be at least 28 feet
(8.5 meters) long in order to encourage flight. Rectangular, octagonal, doughnut, circular or u-
shaped enclosures facilitate sustained flight better than square enclosures, but they are all
suitable for fruit bats.
Bats appear to be more comfortable in enclosures above human eye level, and will
usually move to the highest point in an exhibit. However, if an enclosure is too high, capturing the
bats for veterinary or husbandry purposes may be hindered. Generally, a cage height of six to
seven feet (1.83 to 2.13 meters) will suffice for most species. Larger bats, such as the pteropids,
could be housed in taller exhibits. In addition, greater space is needed for larger colonies. In free
flight enclosures or exhibits where high ceilings make hand capture difficult, it is beneficial to have
adjacent holding cages for management purposes.
The following space guidelines give details on the number of bats that can be housed in a
space based on wing span and a normal body weight range for the species:
1. Giant bats with a wingspread from 3.5 - 5 feet (800 – 1500 grams)
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a. For up to 6 animals, an enclosure 15 feet by 30 feet, 6 feet high or the equivalent area as long
as the minimum width is 2.5 times the largest wingspan. For each additional animal, increase
the enclosure size by 15 percent of the original floor area.
2. Large bats with a wingspread from 2.5 – 3.5 feet (300 – 800 grams)
3. Intermediate bats with a wingspread from 2 – 2.5 feet (200 - 350 grams)
4. Medium bats with a wingspread from 1.5 - 2 feet (80 - 200 grams)
5. Small bats with a wingspread less than 1.25 feet (< 80 grams)
a. For up to 10 animals, an enclosure 4 feet by 7.5 feet, 6 feet high or the equivalent area as long
All bat enclosures shall be large enough to permit aerial maneuvering within the enclosure. The
width of the unobstructed flight area shall be no less than two times the wingspread of the largest
bat in the enclosure, which is the approximate turning radius of the bat to allow for circular flight in
the exhibit. For mixed bat species exhibits the space for each species is proportional to the
number of bats in that size range and summing the space requirements for each group.
In the course of colony management, medical care may necessitate the use of non-flight
cages for bats with fractures or other health problems. A minimal acceptable enclosure height
should be no less than one and a half times the bats' body length to avoid contact with fecal
matter and spoiled food. The minimal acceptable length and width should be no less than one
and a half times the wing span to accommodate wing stretching and static flight behaviors (see
Appendix B, Ethogram, for definition of behaviors). Permanent non-flight cages may be
necessary for non-flighted bats with health problems that could damage themselves by having too
much space.
1.4.2. For social species indicate minimum inter-individual

distances that must be maintained and that will influence size of space. As
appropriate address need for temporary isolation from conspecifics.
Fruit bats vary from being very colonial with roost mates touching each other in the roost
as is the case with Egyptian fruit bats (Rousettus aegyptiacus) to more solitary species such as
the Little golden-mantled flying fox (Pteropus pumilus), which roosts in small dispersed groups.
Inter-individual roosting distances will also vary based on sex, with males requiring more space
and females roosting much closer together. Larger fruit bats and flying foxes that are over 200
grams should have a minimum inter-individual distance of one half wing span from roost mates in
the roost.
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Colonial species of fruit bats may have to be isolated from conspecifics for medical
reasons. This separation can be stressful, and where possible this should be avoided. Isolating
two bats instead of a single individual will allow for social interaction. Social interaction can also
be increased by placing isolated individuals in a howdy cage adjacent to their colony. As an
isolated individual improves they may also be placed in their normal enclosure during the day for
socialization, but isolated at night to monitor food consumption.
1.4.3. Identify appropriate furnishings to accommodate an array of

locomotory and foraging behaviors as well as resting and sleeping.
Each enclosure should have perching areas for tree roosting species with natural sunlight
or full spectrum lighting whereas crevice or cave-dwelling bat species will require dark areas or
night boxes. Each enclosure should allow all bats to roost simultaneously and to seek shelter
from the elements. They should also have climbing apparatus to facilitate exercise and landing
after flight.
Vinyl coated wire hung on the ceiling provides toeholds for bats. We particularly
recommend using 1” x 1” coated wire for larger species and 0.5” x 1” for smaller species, as it
mimics the network of branches that bats use for a variety of behaviors including breeding,
grooming, displaying, seeking food, roosting, and fleeing from aggressive encounters. Wire
ceilings also promote wing walking by the bats and increases activity in the exhibit. Bats can be
excluded from selected areas (e.g., over water or food dishes) by adding Plexiglas over the wire
to prevent roosting. Endurance net is soft alternate for hard wire for species that may abrade
their wing tips on the less flexible material.
Rough, naturalistic branches and vines should be provided as perches to help keep bats'
nails from overgrowing. However, there should be no points or sharp edges that could puncture
wings. Perches will need to be replaced occasionally; therefore, points of attachment should be
designed into the exhibit. Roosting areas such as perches, branches, or boxes should be
positioned far enough apart that animals can space themselves out during stress or flight
(MacNamara et al,1980). Vines, heavy ropes, or crawl ladders (e.g., wire "ladders" attached to
wall) should extend to the ground, especially if the exhibit contains solid or glass side walls. This
will aid bats who are trying to move from the floor to ceiling roosting areas, as bats can exhaust
and injure themselves attempting to fly up to their roost. Narrow gaps (e.g., one-half centimeter)
between pieces of furniture should be avoided because bats can get wings wedged in the gap.
Darkened roost boxes or other visual barriers are recommended to provide animals with
an avenue of escape from stressful situations, especially if aggression among individuals is
observed (Rasweiler, 1975). Consideration should be given to providing feeding stations
accessible from wire walls, vines, or perches; or, with sufficient clearance to allow bats to fly to
and away from them.
Nectar feeding bats like to roost in either the open, from the top of the cage, or in roost
boxes. Therefore, the top of the enclosure should be mesh and a roost box or two should be
provided. Roost boxes should be large enough for all bats to comfortably use and spread out.
The construction of roost boxes is important. They should be constructed of nontreated wood.
The inside of the front, the top and all four sides should be lined with polyethylene mesh or
grooved. The mesh or grooves provide an area for the bats to easily grasp and roost comfortably.
It is also necessary to have a long landing platform, which should also be meshed or grooved.
No light should be able to penetrate into the roost box. Two roost boxes (35 cm L x 21 cm W x 31
cm H with a landing platform of 12 cm) are suggested for a colony of thirty-three bats. It's
necessary for cave-dwelling species such as the Long-tongued bats (Glosophaga sorcina) to
have a dark place to retreat.
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1.4.4. Address the need for and appropriateness of visual, acoustic,
and olfactory barriers within the space. Consider options for minimizing “white
noise” from systems and the surrounding environment
Animals housed indoors should be protected as far as possible from disturbance by the visiting
public. Noise from environmental systems should be kept at a reasonable level. Fruit bats
should always be provided with an area in their exhibit where they can retreat and feel safe when
disturbed by loud noise, bright lights or any other sudden disturbance.
1.4.5. Identify appropriate substrates and nesting/bedding materials

if required.
As bats may occasionally descend to the ground, non-abrasive flooring should be used. Natural
substrates such as soil, grass, or mulch can be used with low densities of bats. We do not
recommend the use of sand or saw dust, as bats may ingest it. Flooring material should be easy
to clean and floors should have good drainage. Epoxy-based paints are useful in sealing cement
walls and floors.
1.4.6. Address mechanisms for the provision of change and variation in the
environment.
As with most mammals, exhibit “furniture” should be changed periodically, but not all at the same
time. These changes can include cleaning and repositioning of ropes, vines and visual barriers.
Environmental enrichment can also be a source of change and variation, and is important for fruit
bats. There are several enrichment options such as offering browse, giving the bats olfactory and
acoustic stimulation, or adding new props or visual barriers to the enclosure (Atkinson, 1993;
LeBlanc, 1999). Refer to Appendix E for a list of forages accepted by captive colonies of fruit
bats.
1.4.7. Address issues, such as scent marking, that may influence

how and how often space is cleaned.
Fruit bats use scent-marking to communicate (Dapson et al., 1977; Gustin and McCracken, 1987;
Quay, 1970; Scully et al., 2000), and they will scent mark their enclosure. Frequent disinfection
that removes all these scent marks may cause distress or aggression. Basic sanitation
procedures should be completed on a daily basis, such as sanitizing the food and water bowls,
and removing the food waste and fecal contamination in the enclosure. Appropriate design and
building materials for cages will greatly facilitate cleaning and sanitation (see Sections 1.4.9).
Raised, wire-floor cages allow feces and contaminated food to fall away from the animals and can
facilitate cleaning. Stainless steel dishes are preferable to those made from other materials (e.g.
plastic) because stainless steel disinfects well, and is less likely to be broken or scratched.
When disinfectants are used to clean dishes or exhibits, it is critical to rinse them well, as
disinfectants may be toxic to bats (Wilson, 1988). Appropriate disinfectants include diluted Clorox
(1:30 dilution) or quartenary ammonium compounds, although the latter does not kill certain
strains of pseudomonas. Ensure that all organic material is removed before applying
disinfectants. Allow adequate contact time (> 15 minutes), scrub in disinfectant well, then rinse
thoroughly. Do not expose bats to noxious vapors in enclosed, poorly ventilated areas or allow
bats to have contact with disinfectants.
Pest Control
A good pest control program should be in place and be maintained and monitored continuously.
The best control is to exclude pests. Cleanliness in and near bat enclosures is of the utmost
importance. Be sure the area around bat enclosures is free of places that harbor pests. Seal all
cracks and holes into the exhibit. Glue boards and snap traps can be used but must be shielded
from bats. Cockroaches are commonly found in and around fruit bat exhibits. Several methods for
5 18th October 2004

controlling cockroach populations are available. Pyrethrin-based insecticides and diatomaceous
earth can be used with care. Narrow-mouthed beer or soda bottles partially filled with fruit juices
can be used as traps. Tracking powder (e.g., Sevin Dust) has been used by some institutions to
control ants. However, we do not recommend it because bats will frequently descend to the floor
of the exhibit and can be adversely affected by the dust. 2 Snakes can be a problem for small bats,
especially in outdoor exhibits. Wire with small openings (e.g., one inch by one-half inch) will
exclude most, but not all, snake species. For outdoor exhibits where snakes may present a
problem, additional barriers may be needed.
There are no special concerns or requirements for sewage disposal; however,

requirements can vary by region and managers should be aware of local regulations.
1.4.8 Identify number of air or water changes/hour required

A highly efficient ventilation system is necessary to avoid the buildup of unpleasant odors,
especially in enclosures housing large colonies. A suggested rate of air change for larger colonies
is six to ten exchanges per hour with 25% fresh air. Because of the bacteriological breakdown of
bat guano, it is important that air leaving bat exhibits, especially that from large colonies, be
vented outside and not re-circulated or ducted into a public area (Wilson, 1988).
1.4.9 Identify necessary measures for safety and containment.

In general, enclosure surfaces should be smooth and non-porous, and all surfaces with which
bats will come into direct contact should be non-abrasive. Wall and floor surfaces must be able to
withstand a great deal of hosing and should be appropriately sealed. Galvanized steel cages and
wire should be avoided because bat urine corrodes tinned surfaces and may cause zinc toxicity if
ingested (Wilson, 1988). Therefore, if an enclosure contains wire, we recommend using vinyl
coated, Teflon sprayed, or non-galvanized wire 3. Polyethylene mesh and polypropylene knitted
netting (Endurance® netting)4 are also excellent caging material (Barnard, 2006). The size of the
openings in wire or mesh should be small enough to prevent animals from pushing a wing, wrist
or foot through it, but not so fine that animals' claws get stuck in the mesh (Wilson, 1988).
Enclosures with glass fronts present no special problems, although it may be necessary
to tape or soap the windows for a few days after animals are newly introduced to alert animals to
the presence of the glass. Piano wire has been used as a public barrier on some fruit bat exhibits,
with plastic coated wire being the easiest to clean. The disadvantages of piano wire include bats
getting caught in wire, especially when they are new to the exhibit, and urine and fecal material
passing through to the public viewing area. To minimize the second problem, place roosts at the
two ends of the exhibit, on sides adjacent to piano wire front. The bats will fly back and forth
between the roosts (in front of the viewing public) and, because bats tend to defecate while
turning at the ends of the exhibit, the amount of fecal material that is passed through the piano
wire front will be minimized.
For exhibits that are constructed as "bat caves” (e.g., the shot Crete wet mix design), the
vault must be semi-rough to facilitate roosting. Also, it is important to create many small concave
pockets in the vault to offer multiple territories and visual barriers favorable to the establishment
of a social structure such as harems. Finally, the vault must be uneven and characterized by
many elevation levels to simulate a natural environment.
For outdoor exhibits, use of a double enclosure will reduce the possibility of escape. This
is mandated by the Lacey Act for Pteropus species. The second enclosure should be more than
two inches from the first enclosure to minimize the risk of a bat's foot reaching through the first
enclosure and getting entangled with the second. Use small wire openings (one-half inch by one
half inch) or a Plexiglass barrier for pest control (see Section 1.4.7). Thin wire (e.g., chicken wire)
should be avoided as it may damage feet, especially with heavier bats (e.g., larger Pteropids).
1.4.10 Address issue of transport, identifying (in accordance with IATA)

Most bats can be shipped communally. Mothers with nursing infants should not be
shipped. Some international shipments may require use of a lift-out internal cage so that bedding
and fruit can be destroyed at the port of entry. Crates should be designed so that there is no
6 18th October 2004

leakage (e.g. of urine or feces) from the crate during shipment. Check with customs before
shipping and prepare the crates prior to shipment.
Zoos must follow IATA standards for all importations. Once within the United States, IATA
standards are still recommended by the Center for Disease Control (CDC), the United States
Department of Agriculture (USDA) Animal and Plant Health Inspection Service (APHIS), and
United States Department of Interior (USDI). However, the "secondary enclosure" technique,
described under Megabats, below, will be permitted (CDC and USDI, pers. comm.). Check with
appropriate agencies prior to shipment.
United States Regulations: A Centers for Disease Control (CDC) permit is required to import
fruit bats into the United States and to ship bats between states 5. (See Appendix E for copy of
permit). In 1995, CDC developed more stringent guidelines, so be sure to check with them prior
to shipping bats.
Canadian Regulations:
a) Shipments from the United States. There are no restrictions for bats being imported
from the United States; however, shipments require a health certificate and permission from the
Canadian Department of Interior.
b) Other foreign importation. According to Section Ten of the Animal Health Regulations,
administered by Agriculture Canada, an import permit must be issued by the Animal Health
Division of the Food Production and Inspection Branch of Agriculture Canada prior to importation.
To receive the permit, an application form must be completed by the importer and submitted to
the appropriate regional office. Upon their arrival in Canada, the bats will be confined for a thirty
day quarantine period in a previously approved area. If there is no health problem after this
period, the quarantine will be ended. Appendix D contains a copy of the import permit application
form (AGR 1551).
The introduction provides a table of bat species listed as endangered by the United
States Fish and Wildlife Service and CITES that can be referenced for shipping information.
1.4.10.1 Type of transport container:

The containers shown in Appendix C meet the requirements of the International Air
Transport Association (IATA). They are strong enough to withstand the rigors of air transport, and
they include appropriately constructed inner and outer components.
1. Megabats. Pteropus species must be transported in double cages. A vari-kennel with an

internal wire cage works well. Line the vented sides and the door with a breathable, opague
material like shade cloth or cheese cloth. This provides the bats with some privacy but still allows
for air flow. Care should be taken that there are no frayed edges in the material used for privacy
with which the bats could become entangled. Crates should be large enough so that when bats
are hanging their faces do not reach the floor.
2. Microbats.6 Microbats such as Glossophaga soricina, Carollia perspicillata, and Artibeus

jamaicensis can be transported in quart-sized cardboard cartons. These containers, which can be
purchased from paper goods companies, measure about four and one-quarter inches (10.8 cm)
in diameter at the top, three and one-half inches (8.9 cm) in diameter at the bottom, and are six
inches high. They should be moisture-resistant and have slip-on lids. To give the bats something
to hang on to, line the interior by stapling in one-eighth inch (3.2 mm) square mesh plastic
aquaculture Netting. Punch air holes in the sides with a cork borer. Up to three Carollia sized bats
can fit in one carton. Fourteen of these cardboard cartons can be placed inside a screen-lined
cardboard mouse shipping cage (with the partitions removed). For extra protection, the mouse
cages can be placed inside a strong protective container constructed of one inch (2.5 cm) square,
hollow aluminum tubing that is lined on the sides with one-quarter inch (6.2 mm) galvanized wire
mesh and has a plywood roof and floor.
1.4.10.2 Appropriate size of transport container
7 18th October 2004

See 1.4.10.1and Appendix C
1.4.10.3 Provision of food and water during transport

The bats should have access to food prior to shipment and it should not remain in the
containers for more than 15 hours.
1.4.10.4 Provision of bedding or substrate in transport container

Avoid using aromatic substrates such as pine bark or cedar chips because these
materials may cause respiratory distress to the occupants. Kraft paper (50 lb) can be used to line
the transport container.
1.4.10.5 Mechanism(s) for separating animal from urine and feces during transport
Some international shipments may require use of a lift-out internal cage so that bedding
and fruit can be destroyed at the port of entry. Crates should be designed so that there is no
leakage (e.g. of urine or feces) from the crate during shipment.
1.4.10.6 Identify appropriate temperature range during transport

Bats should be shipped when environmental temperatures correspond to a given species’
thermal requirement. Although some airlines will allow animal shipments when temperatures
range between 7.2º C and 29.4º C (45º F - 85º F) , regardless of the season, others find it easier
to refuse shipments during summer and winter months. When temperatures are close to the
allowable limits, it may be necessary to include heating or cooling packs (e.g., Hot Rod warm
packs or Uline cold packs) within a transport container.
When transporting bats by automobile during hot weather, NEVER leave them in a
parked car during rest stops unless the air conditioner is running. When transporting tropical
species in winter, the heater should be left on. Many animals have died in vehicles during
transport because simple precautions were not taken.
1.4.10.7 Consider appropriate light levels and how to minimize noise during
transport.
The appropriate transport container for shipping bats will have low light and minimize outside
noise. The mesh sides and door of a Vari-kennel transport container should be covered with a
breathable, opaque material to allow for ventilation, to reduce light levels in the container and to
give the bats privacy.
When fruit bats are shipped, a polite request to the airline staff to place live animals in locations
where loud noises are at a minimum and away from any other live animal shipments in the cargo
hold is advised.
1.4.10.8 Address appropriate group size or need for separation of individuals

during transport
Most bats can be shipped communally. Mothers with nursing infants should not be shipped.
When a group of bats of the same species is being shipped, the sexes should be separated. In
some cases, juveniles may have to be separated from adults. When transporting more than one
species at the same time, a separate container should be available for each species. Up to five
adult Rodrigues fruit bats (Pteropus rodricensis) can be shipped in an intermediate vari-kennel
(0.8 m x 0.56 m x 0.58 m), if the group is compatible. Three Large flying foxes (Pteropus
vampyrus) can be shipped in a large vari-kennel (0.91 m x 0.62 m x 0.66 m).
1.4.10.9 Consider need for handler/veterinarian access to animal during transport

Transport periods that are greater than 15 hours in length should be accompanied with
food and water instructions. In these cases it may be necessary to make arrangements with a
zoological facility close to the lay-over points prior to shipping for food and water provisions. A
contact number for the closest zoological facility should be provided and after 15 hours the
animals should be checked on by trained professionals. If needed, food and water should be
provided.
8 18th October 2004

1.4.10.10 Consider maximum duration of transport allowable before temporary
transfer to “normal housing” is required.
Bats should not remain in the containers for more than 24 hours.
1.4.10.11 Address appropriate timing of release, size and type of enclosure at

transport destination
Follow quarantine guidelines as required (see Section 3.2.1) and reintroduction guidelines (see
Section 5.3). Shipping crate should be placed directly into the quarantine space and the animals
allowed to exit on their own volition and at their own pace.
1.5 Water
1.5.1 For aquatic or semi-aquatic organisms identify acceptable water
quality parameters (pH, turbidity, salts, turnover rates, filtration,
disinfection, bacterial count, acceptable chemical residuals, etc).
1.5.2 For terrestrial and semi-aquatic organisms address appropriate
means of presentation of water, and appropriate placement of water
sources, acceptable bacterial levels, mineral levels, temperature as
appropriate
1.5.3 Address issues of depth and need for variation in depth and/or
current
Water sources should be checked for chemical residues, bacterial counts, mineral levels
and salts. Because water requirements are unknown for bats (Racey, 1970), water should be
offered at all times, even though some species (e.g., nectar-feeding bats) may rarely be seen
drinking. Some naturalistic bat exhibits have running sources of water such as waterfalls and
pools. This kind of water source is greatly used by Microchiropteran bats that glide over the water
surface for a drink. Care should be taken that fecal material and discarded pieces of food do not
accumulate in these areas, and the water source should be shallow enough to allow animals an
easy method of removing themselves from these features. Misters provide opportunities for bats
that are kept in outdoor exhibits to cool off if the temperature goes over 29.4 °C (85 °F).
2. Biotic Variables
2.1 Food and Water
2.1.1. Identify appropriate containers and protocols for the provision of food
and water
Standard glass, plastic, or ceramic bowls should be used for food and water bowls.
Several sources of water (e.g., bowls, pools, mister hoses, etc.) should be provided and water
should be changed daily. Large colonies and exhibits housing mixed bat species or aggressive
animals will need a greater number of water sources. Some of the larger fruit bat species are
capable of learning to drink from watering devices fitted with tubes. These devices have the
advantage of not becoming soiled by urine and feces. Some species or individuals may not be
able to utilize these devices, so bat caregivers need to monitor the situation to verify that all bats
are using them successfully. To minimize contamination of water sources by urine and feces,
place water source beneath areas of smooth ceiling where bats cannot hang.
For nectarivorous bats, dietary presentation is almost as important as the diet itself.
Nectar solution can be offered in a variety of hummingbird feeders. It is important to take into
consideration when choosing a feeder whether or not it will be accommodating and easily used
by the nectar-feeding bat. The feeder must not have bee guards or obstructions around the
opening. The opening must also be as close as possible to the nectar itself to ensure that the bat
will be able to reach the fluid with its tongue. The daily feeders used at OBC and Biodome are,
respectively, the Little Beginner Hummingbird Feeders by Perky-Pet Products Co. (2201 South
Wabash street, Denver, Co 80231. U.S.A.) and the Oriole Feeders by Opus Incorporated (P.O.
9 18th October 2004

Box 525, Bellingham, MA, 02019. U.S.A.). Open-topped feeders, such as a plate with an
extension in the middle hung from the ceiling, are favorites among the bats.
Yet another form of presentation includes open shallow dishes placed in hanging baskets.
More unique forms of presentation are a good source of daily enhancement but this should be
done with extra nectar so that it is guaranteed that the entire population receives enough to eat
each night. This can be done by using hummingbird feeders in the shape of artificial flowers
placed in a pot. Drowning is an important factor to consider when contemplating various ideas of
presenting food to smaller bats. To deter drowning, it is helpful to place a variety of marbles in
the open dishes to allow an area for footing should an individual accidentally fall into the dish.
2.1.2. Identify appropriate foodstuffs (see below) and feeding schedules

One daily feeding is sufficient for most species. Large colonies, however, should be fed
more frequently to ensure that all individuals will have access to food. Over feeding can cause
obesity in the colony, and this must also be monitored.
For nectarivorous bats, the liquid diets have a tendency to sour or ferment when left out
overnight especially in warm environments. The bats must be fed at the end of the light cycle to
minimize fermentation of the food prior to feeding.
2.1.3. Address the provision of variability in food type and presentation (e.g.
spatial and temporal dispersal of food resources)
Large fruit bat colonies may require twice a day feedings to ensure that all individuals will
have access to food. Captive fruit bats also require enrichment which can be offered to provide
variability in food type such as browse and flowers in addition to novel fruit items. This
enrichment can be presented so that bats have both a spatial and temporal dispersal of food
resources in their enclosure.
For nectarivorous bats, the feeders and fruit skewers should placed as far apart as
possible as well as in a different locations on a daily basis so that the bats are forced to actively
forage as they would in nature. There has been some territorial behaviors observed so it's
important to have the nectar and fruit as spread out as possible to ensure adequate consumption
by each individual.
2.1.4. Address opportunities for animals to process food in ways similar to

their wild counterparts, and consider mechanisms that enable animals
to work for food
All fruit bat species should be given whole fruit or nectar and allowed to process food in
ways similar to their wild counterparts as part of their enrichment program. Bats can also be
challenged to fly to feeding areas and to utilize their wings and feet to obtained food resources
such as Large flying foxes (Pteropus vampyrus) pulling up weighted plastic chain to reach novel
food items (Sequin et. al. 2000).
2.2 Social Considerations

2.2.1. Group Composition, including as appropriate
2.2.1.1. Suggested age and sex structure of social group
All fruit bat species maintained in North America can be maintained in single sex groups. Male
bachelor groups of fruit bats as with most mammals will have higher levels of aggression than
single sex female groups. In single sex groups of any size or sex, there may be minor dominance
struggles, especially among males. A common behavior that can be observed is male on male
mounting as part of normal dominance behavior.
10 18th October 2004

Bats that are part of an organized breeding program will require facilities to house multiple males,
and the breeding group, This does not necessitate that every zoo must maintaining all three
types of colonies (Male and female single sex groups and a breeding colony), but rather zoos
should trade animals to coordinate breeding programs. A sex ratio of 1.6 or 2.10 is suggested,
since problems with aggression can occur with maintaining too many males in a colony such as
decreased reproduction and males pulling infants off dams. A sex ratio of 3.12 Long-tongued bat
(Glossophaga soricina) is suggested for breeding with this nectar-feeding species.
Fruit bats are long lived in comparison to similar sized mammals, and breeding plans should
focus on maintaining healthy populations with appropriate age pyramids. Elderly male fruit bats
may have difficulty competing with younger males in single sex groups, and may have to be
maintained is single sex groups with males of a similar age class.
2.2.1.2. temporary isolation of parturient females and young, or of

males, and corresponding adequate and appropriate space for
animals when removed
Some fruit bat species have a post-partum estrus, and reproduction will occur if the
breeding males are not removed. Fruit bat species can breed even if they are housed in an
enclosure that is too small for them to fly. Parturient females should be left in maternity colonies,
and males should be returned to a bachelor group. Appropriate care must be taken in removing
breeding males from the colony, so as not to stress pregnant females or dams with pups, which
can cause abortions. Introductions of new bats into a maternity colony should also be avoided for
two to three months, to allow for a close bond to form between the dam and pup, and to minimize
aggressive encounters between newcomers and very young pups. If breeding males or
parturient females have to be temporarily isolated, then follow the space guidelines in section
1.4.1.
2.2.1.3. Seasonal separation of sexes, or for those species that are

truly solitary (=social interactions are primarily limited to
courtship and care of dependent offspring), seasonal
introduction of sexes
See Section 4, Reproduction, for more detailed information.
Wild bats are seasonal breeders, the timing of which is mostly species specific. However,
seasonality decreases in captivity, especially in indoor enclosures. Seasonality will also differ in
different climates.
Although male bats will attempt to copulate throughout the year (Martin et al., 1986), the
highest copulation activity in bats is during the mating season. When not in season, females will
avoid copulation and will generally fly away from interested males.
During their breeding season, males tend to become more territorial and aggressive.
Many, if not most, fruit bats form harems. In some species, males will be seen hovering around a
receptive female, showing interest by hooking the female with their thumbs. They will also "hook”
another male with a thumb to try to dislodge them and thereby keep them away from the female.
The colony hierarchy does not always hold for breeding, so do not assume that the dominant
male is the sire. Also, multiple males will breed one female so paternity in multiple male groups
can be difficult to ascertain.
Courtship in some of the larger bat species often involves male initiated allo-grooming,
especially around the genital area, followed by much bickering and then mating. Females are
generally resistant; they may fly to get away from an interested male. Females may choose with
whom they mate; however, more research needs to be done in this area.
2.2.1.4. nursery groups (groups of mothers with most recent young)

Typically, parenting is done by the female. Allo-parenting is rare, but has been documented in
dog-faced fruit bats (Cynopterus brachyotis), straw-colored fruit bats (Eidolon helvum) and Little-
golden-mantled flying foxes (Pteropus pumilus). In the wild it is quite common to find maternity

colonies of many bat species, and zoos could replicate this behavior by removing males after
birth. Mothers carry and hold their infants in a variety of positions including front and back. They
will also "park" the infants for various lengths of time, especially during weaning and parent
feeding time (the length of time infants are parked appears to be species specific). Should the
infant fall or get knocked down, mother bats will pick their infants off the floor. In one case, a
female Large flying fox (P. vampyrus) at the Lubee Bat Conservancy had a pup that died
overnight, and the dam would carry the dead pup in her wings, and attempt to get the pup to
nurse by pushing the pup to the nipple until she lost her grip on the pup and it fell. This same
dam vocalized for her pup over a twenty-four hour period, and would investigate other pups in the
enclosure that made contact calls.
In large colonies of fruit bats or in crowded enclosures with other bat species, it has been noted
that lone infants that get separated from their dams become vulnerable. In overcrowded
situations, maternal neglect is not uncommon. Infanticide has been noted in fruit bat colonies, and
while the causes are unknown, it does not appear to be directly related to overcrowding or an
improper sex ratio (Carroll, 1988).
2.2.1.5. forced “emigration” of adolescents
In captivity, young male fruit bats must be removed before they reach sexually maturity. This date
varies on the species maintained, but in general it is several months after weaning. For the larger
fruit bats (e.g., Pteropus spp.) males may reach sexual maturity at twelve months while females
may reach sexual maturity when they become two years of old. Smaller species are weaned and
reach sexual maturity in less than six months. Be sure that juveniles are eating solid foods and
drinking water before removing them from the dam. Mother-juvenile fights are common,
especially during weaning, with the pup frequently trying to nurse its dam and then being
rebuffed. Pups will form lasting bonds with other bats in their crèche, and it is important to keep
them in these groups, especially when moving young males into older bachelor groups. Female
pups can stay in the maternity colony longer, and only need to be removed just prior to
introducing the breeding male for another round of breeding.
2.2.1.6. multigenerational groups (e.g. many primates, elephants)

Multigenerational single sex colonies may be formed in the wild such as male bachelor groups
and female maternity groups in fruit bats. In captivity, they can be maintained in single sex
groups.
2.2.1.7. groups deriving from cohorts (e.g., dolphin male pairs)

Young fruit bats that were creched together as pups may form long lasting bonds. Similar-sex
bats can be moved to single sex groups as part of normal population management.
2.2.1.8. all male groups

All fruit bat species maintained in North America can be maintained in single sex groups. Male
bachelor groups of fruit bats as with most mammals will have higher levels of aggression than
single sex female groups. In single sex groups of any size or sex, there may be minor dominance
struggles, especially among males. A common behavior that can be observed is male on male
mounting as part of normal dominance behavior.
2.2.1.9. daily and life stage variation in patterns of social affiliation

Bat behavior is formed largely through observational learning. Young bats are generally poor
flyers and will often crash into things as they learn how to fly. Static flying is common among
infant and juvenile bats (Carroll, 1979). Pups are very playful, and their need for environmental
enrichment is great (see Section 5.7). At weaning, a pup will first learn to eat solid food by licking
its mother's lips as she is feeding. Up to a year of age, pups will return to their mother's nipple
when they are frightened. Occasionally, a female will have an older pup on one nipple and a
younger pup on the other. Hall and Richards (2000) suggested that adult male flying foxes also
have a role in guiding weaned juveniles towards food resources in the wild.

2.2.2. Group Size, including
2.2.2.1. minimum and optimum group sizes
Because the majority of bat species are social, we recommend that no bats be maintained singly.
The only exception to this recommendation is if an animal requires special medical care. The
ideal number of individuals and sex ratio of bat colonies will be species specific and will depend
largely on the size of the enclosure. If bat densities are too high or low, the resulting stress may
lead to detrimental physiological and/or behavioral changes (Rasweiler, 1975). While managers
should observe colonies for excessive fighting and potential injuries, squabbles among bats are
common in the wild and in captivity and often result in harmless wing tears that heal quickly and
are of little concern. (For more information on wing tears, see Section 3.2.2). To help avoid undue
fighting among animals, be sure to place sufficient feeding and roosting stations in the enclosure
(see Section 1.4.3 for additional information on avoiding aggression).
2.2.2.2. inter-individual distances required
2.2.3. Conspecific groups, consider the need for/influence of adjacent

groups, or similar taxa, on territorial species
In general, bats do very well in mixed species exhibits, including housing bats with other
bat species and bats with other taxa. The primary concern with maintaining multiple bat species is
ensuring that the nutritional and spatial needs of each species are taken into consideration.
Enclosure size, environment (e.g., most appropriate feeding time and light cycle), and species
variability are important factors. It is also critical to offer enough food dishes of appropriate sizes
to minimize competition. Ideally, mixed species exhibits would house multiple colonies rather than
a colony of one species and one or two individuals of another (although this combination has
worked well in some instances). The size of the bat species may be a factor; however, more
information is needed. There are no documented cases of successful breeding between species
in zoos, but in Australia, Grey-headed flying foxes (Pteropus poliocephalus) have been observed
to hybridize with Black flying foxes (Pteropus alecto) in captivity (Luckoff pers. comm.).
2.2.4. Mixed species groups

2.2.4.1. identify appropriate species (consider relative size, diet,
mode of defensive behavior, potential for hybridization,
potential for disease transmission)
2.2.4.2. identify key environmental elements for each species
2.2.4.3. identify interspecific inter-animal distances required
2.2.4.4. address appropriateness of single-sexed groups
There is tremendous potential to increase the amount of space available to bats in zoos
and make exhibits more exciting by maintaining bats with other taxa. There are several potential
problems that should be taken into consideration before implementing a mixed taxa exhibit such
as looking at disease issues that could cross species, a lack of adequate space, competition and
aggression between species, and provision of enough food and a proper diet. An additional
concern when housing bats with terrestrial omnivores and carnivores is that they may consume
incapacitated bats that fallen to the ground. For more information on using bats in mixed species
exhibits, see Riger, Bear-Hull and Harmon (2001).
Bats in the genus Pteropus are prone to henipaviruses, and should not be exhibited with
primates, swine or hoofstock. Salmonella can also be a zoonotic disease threat with mixed
species, and veterinarians should screen animals for disease issues prior to managers mixing
different species. Managers should also avoid mixing bat species with species that require heavy
bedding material that would be soiled by bat guano, and create a possible culture for
Histoplasmosis. Bat species that are slated for possible reintroduction should also be excluded
from mixed species exhibits to minimize the risk of transferring captive wildlife diseases to wild
populations. The AZA Bat TAG suggests not mixing mega and microbat species, and to mix bat
species based on region, such as Asian flying foxes or South American fruit bats.
Rodrigues fruit bats (Pteropus rodricensis) have been successfully exhibited with the following
species, but it can be aggressive so any mixed species exhibits should be closely monitored:

Riverbanks Zoo – red billed hornbill Tockus erythrorhynchus
Brookfield Zoo – Prevost’s squirrel Callosciurus prevosti, giant Asian squirrel
Ratufa bicolor, small-clawed otter Ablonyx cinerea, boobook owl Ninox
novaeseelandiae, Matschie’s tree kangaroo Dendolagus matschiei,
brush-tailed bettong Bettongia penicillata
Lubee Bat Conservancy - Pteropid bats: Pteropus pumilus, Pteropus
hypomelanus, Pteropus vampyrus, Pteropus giganteus, Pteropus
conspiculattus, and Petaurus breviceps.
Oregon Zoo – Straw colored fruit bat Eidolon helvum, Egyptian fruit bat
Rousettus aegyptiacus, Jamaican fruit bat Artibeus jamaicensis
Philadelphia Zoo – Rousettus aegyptiacus
Organization for Bat Conservation – Short tailed fruit bat Carollia perspicillata,
Jamaican fruit bat Artibeus jamaicensis, Straw colored fruit bat Eidolon
helvum, Egyptian fruit bat Rousettus aegyptiacus, Ruwenzori fruit bat
Rousettus lanosus
Disney’s Animal Kingdom – Victoria crowned pigeon Goura victoria, Jungle
fowl Gallus sp., Pteropus vampyrus
 Additional species not mentioned above: Malayan tapir Tapirus indicus, white-
handed gibbon Hylobates lar, Brush-tailed porcupine Atherurus africanus, lesser
bushbaby Galago senegalensis (Riger et al. 2001).
2.2.4. Introductions
Identify appropriate introduction techniques and strategies for
conspecifics, and heterospecifics as appropriate
See section 5.3 for more information on introductions of conspecifics.
2.2.6. Human-animal interactions

2.2.6.1. identify acceptable forms of human/animal interaction
keeping in mind that animals should not show fear or
aggression during routine care, and interactions viewed
by the public should not encourage public interactions
with wild animals
All fruit bats will show stress when approached by a handler, and then try to escape. Stress levels
are greater for animals housed in small enclosures that have no room to escape. (For additional
information on the effects of stress see Widmaier and Kunz, 1993). When captured, most bats will
immediately urinate and defecate and many will try to bite. Vocalizations may accompany any of
these behaviors. Many large bat species will shiver and head bob when stressed. Long-term
stress may cause anorexia. Some bats, especially the micro fruit bats (e. g., Glossophaga
soricina), may go into stress-induced torpor. White-lines bats (Vampyrops helleri) and Long-
tongues bats (Glossophaga soricina) will exhibit the following behaviors when food deprived:
unfolding of the wings, no attempt to fly when approached or prodded, and no audible noises
emitted (Rasweiler, 1973).
2.2.6.2. Address both animal and keeper safety

In most States, a bat that bites a human must be killed to test it for rabies. For this reason, we
recommend that the public not have direct contact with bats. Only trained and qualified personnel
should handle bats. In addition, all personnel who come into contact with bats should be
immunized against rabies and have titers tested regularly. All bat bites should be reported to
County public health officials, per state guidelines.
Bats potentially are vectors of zoonotic diseases that can be passed to humans (see Constantine,
1988). Consult your local health department for additional information on health issues with bats.
Rabies and histoplasmosis are outlined in more detail below:

1. Rabies:
This virus manifests in itself in two forms: furious rabies and paralytic (or dumb rabies).
Furious rabies is characterized by agitation, thrashing, biting, and viciousness. Paralytic
rabies is characterized by apathy, depression, disorientation, and ends with coma and
death by respiratory failure. The furious form is rarely found in bats, and "outbreaks" of
rabies in bat colonies is extremely rare. Contrary to common belief (even by medical
"experts"), bats are not asymptomatic carriers of rabies. The statistics for the percentage
of bats that contract rabies varies depending on region and type of sampling technique
used for testing. No matter what the region or species, certain precautions should be
taken to protect animal handlers from contracting this disease.
a. All personnel working with bats should be vaccinated against rabies.
b. Personnel should have their rabies titers checked every one to two years and be
vaccinated as necessary.
c. The potential for bites should be minimized by wearing gloves, having only trained
personnel handle the animals, and by anesthetizing even for routine handling.
d. Establish a bite protocol, and record all bites, even if the skin is not broken.
e. Minimize the number of personnel in contact with the bats.
f. Greatest care must be taken with new, wild-caught imports.
2. Histoplasmosis: Histoplasmosis (Histoplasmosis capsulatum) is an airborne fungus

that affects the lungs of mammals. It is contracted from breathing dust in enclosed areas
such as caves or mines which contain infected bird or bat guano. The fungus occurs
primarily in the Mississippi Basin of the United States, the tropics and subtropics.
The risk of catching histoplasmosis from a captive colony of bats is small because these
colonies rarely reach the large numbers needed for the build-up of the fungus, and fecal
material is removed on a regular basis (i.e., not allowed to accumulate).
3. Henipavirus: This virus genus in the family Paramyxoviridae, contains two members,
Hendra virus (originally Equine morbillivirus, EMV) and Nipa hvirus. They are
characterized by their natural occurrence in Pteropid fruit bats (flying foxes), and their
recent emergence as zoonotic pathogens capable of causing illness and death in
domestic animals and humans. Hendra virus was discovered in September 1994 when it
caused the deaths of thirteen horses, and a trainer at a training complex in Hendra, a
suburb of Brisbane in Queensland, Australia. A serosurvey of wildlife in the outbreak
areas showed that pteropid fruit bats had a 47% seroprevalence for the virus. Virus
isolations from the reproductive tract and urine of wild bats indicated that transmission to
horses may have occurred via exposure to bat urine or birthing fluids (Halpin et al, 2000).
Nipah virus was identified in 1999 when it caused an outbreak of neurological and
respiratory disease on pig farms in peninsular Malaysia, resulting in 105 human deaths
and the culling of one million pigs (Field et al., 2001). Flying foxes are unaffected by
Hendra or Nipah virus infection. Bats in the genus Pteropus should not be mixed with
hoofstock, swine or primates.
3. Health and Nutrition

3.1. Diet
3.1.1. Identify existing standards for nutrient requirements for all life
stages if available
Foraging Ecology
Currently, it appears that the majority of data about foraging ecology is a description of
the bats' feeding or foraging habits in the wild. Some studies have involved collecting and
analyzing food items for various nutrients (Morrison, 1978; Morrison, 1980; Thomas, 1984).
Others examined excreta or stomach contents of bats in the wild (Stellar, 1986; Thomas, 1984).
Each of these methods has its limitations and none quantitatively defines the daily nutrient
requirements of fruit bats. Thus to date, there is little information available that describes the
nutrient levels required by fruit bats. The only fruit bats for which estimates of nutrient

requirements have been made are in the suborder Megachiroptera (Stellar, 1986; Thomas, 1984).
Few, if any, quantitative nutrient requirement data for frugivorous Microchiroptera are available.
Many species regularly consume fresh green forages; however, the extent to which forages
contribute to overall nutrition is unknown (Kunz and Ingalls, 1994; Kunz and Diaz, 1994; Marshall,
1985). Additionally, fruit and nectar-eating bats can consume other plant parts including flowers,
bracts, nectar, and pollen (Law, 1992; Lowry, 1989; Marshall, 1983). Since fruit is not a rich
source of protein, bats can consume protein rich plants and insects, especially during pregnancy,
to meet their protein requirements (Courts, 1998). Diets in the wild are influenced by seasonal
availability, and this can also be modeled in captivity.
Section adapted from Dempsey, J. 2004. Fruit bats: Nutrition and dietary husbandry. Fact
Sheet 014. Nutrition Advisory Group Handbook.
Digestive Tract Morphology and Physiology

Fruit bats are highly efficient in extracting the liquid portion of chosen foods. They have
fewer teeth than insectivores,10 and their teeth are broad and relatively flat for crushing fruits,
allowing the bats to squeeze out and swallow the juices. 23 The fibrous portion remaining is much
reduced in moisture and is spit out in tightly compressed pellets (ejecta). Seeds also may be
swallowed, but many of those from figs appear to pass whole into the feces. 23,48 To meet nutrient
needs, daily food intakes (on a wet basis) range as high as 2.5 times body mass. 14,15,16,26,31,48,63, 70
These large volumes of food are processed through the digestive tract rapidly, with transit times
ranging from 15-100 min.30,48,69
The gastrointestinal tract of the fruit bat species that have been studied is highly modified
compared to bat species with other feeding strategies. The stomach is large and more complex,
the small intestine is long and convoluted, the cecum is absent, and the large intestine is short and
nearly indistinguishable from the small intestine.30,68,69 There appear to be no areas of fermentation
analogous to the rumen of ruminants, and significant concentrations of gastrointestinal fermenting
anaerobes have not been found.30 Rather, it is thought that the enlarged, complex stomach and
long, convoluted intestine provide the space and surface area required for digestion and
absorption of nutrients from the large volumes of liquid consumed.
Nutrient Content of Foods

Upon comparison with established nutrient requirements of other mammals and the foods needed
to provide them, fruit, when consumed alone, would seem to constitute an inadequate diet.
Analyses of most cultivated fruits indicate that the concentrations of many nutrients are quite low.
However, there are few documented analyses of the nutrients in wild fruit species, and A.
jamaicensis has been reported to feed on fruits from over 70 genera. 23 Thus, it may be important
to distinguish between cultivated and wild fruits. Limited compositional data on proximate
fractions and minerals in both are presented in Tables 1 and 2. The cultivated fruits listed tend to
be lower in fiber, ash, and calcium (dry basis), with some higher in moisture, than the listed wild
fruits.8,19,25,31,59,60,66 Comparisons between cultivated and wild fruits in concentrations of vitamins
could not be made because published vitamin values for wild fruits were not found.
Fruit bats in the wild appear to meet their nutrient needs by consuming large quantities of a
mixture of native fruits, with some consumption of flower parts, leaves, and insects.
2,11,12,21,23,28,33,34,36,37,38,46-49,61,78
In captivity, fruit sources are limited to those that are cultivated and
readily available. As a consequence, it may be important to ensure that all essential nutrients are
supplied by using other nutritionally complete foods. To formulate such foods, it is important to
know which nutrients are essential to fruit bats and in what amounts they should be supplied.

Table 1. Comparison of the macronutrient composition of cultivated and native fruits.
Expressed on a dry matter basis
Fruit Type Moisture Gross Protein Nitrogen Fat Fiber Carbohydrates Ash
% Energy % % % % % %
Kcal/g
Cultivated a,b
Apple 84.50 3.66f 1.03 0.16 2.00 3.55 95.46 1.55g
Banana 74.30 3.74f 4.08 0.63 2.06 2.06 91.03 3.11g
Cantaloupe 91.00 3.57f 8.88 1.38 2.69 3.33 85.55 5.68g
Orange 86.80 3.70f 7.58 1.21 0.54 3.26 88.25 3.47g
Native
Fig (F. ovalis)c 78.50 4.08 2.10 0.60 0.80 35.40 43.40 5.20
Fig (F. 83.70 3.60 3.50 0.56 2.10 5.20 84.70 4.50
sycomorous) d
Fig (Belize 72.08 -- 7.85 1.26 -- -- -- 7.96
native)e
Piper fruit 73 3.95 6.00 1.90 1.40 10.90 86.70 23.00
(P.amalago)c
Carob (C. 20.50 3.90 6.50 1.04 0.95 7.60 82.40 2.60
siliqua)d
A Data from Pennington and Church.60

B Fruit without skin or peel, includes seeds.
C Data from Herbst.25 Sum of values for protein, fat, fiber, carbohydrates, and ash deviate
significantly from 100%. Errors in reported values are suspected but are presented as published.
N=130 for F. ovalis, N=200 for P. amalago.
D Data from Korine et al.31 Samples analyzed were pooled from 20 specimens of each fruit
species.
E Data from Silver et al.66 Samples were pooled from multiple Ficus sp., N=6.
F Data provided by Debra A. Schmidt and Monty S. Kerley, Department of Animal Sciences,
Nutrition Laboratory, University of Missouri-Columbia. Values are for fruit pulp only, no skin, peel
or seeds.
G Data from USDA, Agricultural Handbook No. 8 Composition of Foods. 73
Table 2. Comparison of the mineral content of cultivated and native fruits. Expressed on a
dry matter basis.
Fruit Type Moisture Ca P Fe Se

% % % Ppm Ppm
Cultivated a,b
Apple 84.50 0.03 0.05 4.5 0.02
Banana 74.30 0.02 0.08 11.9 0.04
Cantaloupe 89.78 0.11 0.17 21.5 0.04
Orange 86.80 0.30 0.11 7.6 0.04
Native
Fig (F. pertusa)c ripe fruit 77.20 0.74 0.12 56.60 0.03
Fig (F. trigonata)cunripe 83.70 0.74 0.23 49.9 0.10
fruit
Fig (Belize native)d 72.08 2.04 0.18 58.1 --
Fig (Indonesia native)e -- 1.21 0.33 65.7 --
Fig (Uganda native)e -- 1.52 0.18 94.7 --
A Data from Pennington and Church.60

B Fruit without skin or peel, includes seeds.
C Data from Edwards.19 N=3 for F. pertusa, N=1 for F. trigonata.
D Data from Silver et al.66 Samples were pooled from multiple Ficus sp., N=5.
E Data from O’Brien et al.59 Indonesia N=20, Uganda N=10.
Nutrient Requirements

Despite differences in diet and feeding strategy among mammals, most species that have been
studied appear to have similar qualitative nutrient needs for normal tissue metabolism.
Presumably, these similarities also apply to fruit bats. Approximately fifty nutrients have been
identified as dietary essentials for mammals. That is, they must be obtained from the diet or
through the activity of gastrointestinal microbes because they cannot be endogenously
synthesized in quantities appropriate to meet animal needs. 72
Water
Water is the easiest and least expensive nutrient to provide in captivity. A supply of water is crucial
due to its involvement in a variety of essential functions within the body. The need for liquid water
is influenced by variables such as ambient air temperature and humidity, solar and thermal
radiation, metabolic rates, and food composition and intake. 64 Fruit bats have been maintained in
laboratory settings on mixed fruit diets without free access to liquid water and appear to obtain the
water they need from their high-moisture fruit diet and presumably from water released during
metabolism of absorbed energy sources (metabolic water). However, many species have been
observed to actively consume water both in captivity and in the wild. Wild species of Pteropus on
oceanic islands have been observed consuming sea water, and their kidneys appear well-adapted
to excreting salt.29 Therefore, considering the limited food choices and, in some cases, fluctuating
environmental conditions, it is prudent to provide ad libitum access to fresh water in captivity.
Protein and amino acids

Proteins contain nitrogen, are major constituents of the animal body, and are vital to all tissues.
Amino acids are the functional building blocks of proteins, and animals with relatively simple
stomachs and little or no capacity for fermentation require a dietary source of 10-12 amino acids
that cannot be synthesized metabolically at an adequate rate (dietary essentials). 64,72 Essential
amino acid requirements for bat species are not known nor has the amino acid composition of
many foods consumed by bats been determined, although Herbst 26 has reported relative
concentrations of arginine, histidine, isoleucine, leucine, lysine, methionine, phenylalanine,
threonine, and valine in fruit pulp protein of four plant species (Cecropia peltata, Chlorophora
tinctoria, Ficus ovalis, and Piper amalogo). It is not known whether the amino acid composition of
protein in juice squeezed out of the pulp and swallowed by fruit bats is the same as that in the
pulp.
Fruit, as the only food item, is considered by many researchers to constitute an
inadequate diet because of its low protein content when compared to other plant and animal food
sources, although the quality of that protein is relevant to such a conclusion. It has been argued
that both pteropodid and phyllostomid fruit bats must supplement their fruit diets with relatively
higher protein items, including a more complementary array of amino acids, such as might be
found in insects,48 pollen,35 or leaves32,33,38,78 in order to meet protein needs. However, other
researchers maintain that fruit bats can meet their protein requirements exclusively with
fruits,9,15,26,67,70 and fruit bats (Carollia brevicauda and Sturnira ludovici) in Costa Rica showed a
strong preference for fruit (Acristus arborescens) that was not infested with insect larvae.20
Further, it has been proposed that most fig wasps will have departed figs by the time they
become ripe and are consumed by bats, and residual corpses are unlikely to contribute
significantly to protein intake.8
Recent research supports the latter hypothesis, at least for maintenance of adults. These
studies have been conducted using diets that approximate the protein concentration of fruits (both
cultivated and wild) or by using wild fruits alone, thus suggesting that fruit bats have the ability to
meet their protein requirements for maintenance on very low protein diets. This may be
accomplished in part through minimal nitrogen excretion, 31 and by adjusting dry matter intake to
meet protein needs, regardless of dietary energy concentration. 63 A diet containing about 9%
crude protein (DM basis) was consumed by captive fruit bats (Pteropus hypomelanus, P. pumilus,
P. vampyrus) at 28% of body weight on an as fed basis or 7% as dry matter. 16 Studies of the
protein needs of A. jamaicensis (phyllostomids) weighing 39-50 g found that 0.28-0.30 g/day were
required for adult maintenance, or about 5% of dietary dry matter. 48,63 In another study, adult A.
jamaicensis bats weighing 36.8 g achieved nitrogen balance at protein intakes of 0.17 g/day on a
“high-energy” diet and 0.46 g/day on a “low-energy” diet. 15 Maintenance protein requirements for

adult Carollia perspicillata (phyllostomids) were estimated to be 0.14 g/day.14 Adult Rousettus
aegyptiacus (pteropodids) weighing 144-157 g achieved nitrogen balance at protein intakes of
0.36-0.83 g/day.15,31 It should be noted that when stable-N isotope analyses were used to quantify
the relative importance of plants (fruits) and animals (insects) as sources of assimilable N for five
species of free-ranging frugivorous New World bats (Artibeus jamaicensis, Uroderma bilobatum,
Dermanura phaetis, Sturnira lilium, and Carollia brevicauda), fruits were by far the most important
throughout the year. However, at the end of the rainy season and beginning of the dry season,
there was a trend in all species for a decline in the relative proportion of plant protein in the diet,
and in some individuals of S. lilium and C. brevicauda, insects became a significant source of
protein. It was suggested that fruit bat species may vary in their foraging strategy, with some
relying almost entirely on fruits throughout the year, whereas others may switch to insects when
fruits are less abundant.27
It is difficult to estimate exact requirements because quite different diets have been used
in published studies, and the factors to convert nitrogen to protein may have, in some cases,
been incorrect.9,26,43 Assuming appropriate amino acid concentrations in the protein, 4-6% protein
in the dry matter consumed and digested appears sufficient for maintenance. Requirements for
reproduction or growth may be higher but have not been experimentally determined. Presumably,
the essential amino acid composition of dietary proteins as compared to amino acid needs would
influence the amounts of protein required for any of the above functions.
Energy
Digestible energy and metabolizable energy are terms used to describe an animal’s energy
needs and to characterize the usable energy concentrations in food fed to that animal. Because
animal species vary in the way they digest and metabolize nutrient sources, the digestible energy
or metabolizable energy concentration in a food varies with the gross energy of that food, the
amount of food consumed, and the digestive and/or metabolic abilities of the animal to which the
food is fed. A more complete description of energy terms can be found in a National Research
Council publication.52
Due to their high concentrations in fruit, carbohydrates are quantitatively important
sources of energy in the diets of fruit bats. Dietary fat is a potentially important source of energy,
as well, but generally is a minor constituent of high-sugar fruits. Protein in the diet also can be
broken down for energy, but that is probably minimal on low-protein diets in which the balance of
essential amino acids is just adequate to meet amino acid requirements. Fruit bats presumably
have no difficulty meeting energy needs during periods of food abundance because they
consume large amounts of high carbohydrate fruits, both in captivity and in the wild. There
is controversy among researchers whether or not fruit bats must “over-ingest” energy in order to
meet protein requirements. This is due to the relatively high energy:protein ratio of most fruits.
High food (and energy) intakes were necessary to meet protein needs in pteropodids (Epomops
buettikoferi, Micropteropus pusillus, and Pteropus poliocephalus) when fed some low-protein
fruits but not when fed others.67,70 Carollia perspicillata (phyllostomids) did not need to consume
excessive energy to meet maintenance protein needs if appropriate wild foods were available and
selected.26 Estimates of metabolizable energy requirements for adult maintenance of A.
jamaicensis have been made by a number of researchers, and range from 12 to 34.3
kcal/day.15,48,63 The intermediate value of 17.2 kcal/day found by Reiter63 is equivalent to 43.9 kcal
of metabolizable energy/100 g body mass/day (mean body mass 39.2 g) or a metabolizable
energy density in ingested dry matter of 3.16 kcal/kg, a metabolizable energy concentration found
in many domestic fruits.
Estimates of metabolizable energy requirements for adult maintenance of C. perspicillata
weighing 18.5 g were 16.1 kcal/day/bat14 and were 40.1 kcal/day for R. aegyptiacus with a mean
body mass of 144 g,31 or 76.3 to 91.9 kcal/day for R. aegyptiacus with a mean body mass of
156.7 g.15
Care must be taken when interpreting published data on energy requirements of fruit
bats because species, experimental diets, and levels of permitted activity varied appreciably
among studies. Basal energy expenditures of bats appear to vary with species (particularly body
mass and foraging and roosting behavior), sexual dimorphism in body mass, precision of body
temperature regulation, tendency to enter torpor, and the environmental circumstances within

which they evolved.40,41 In addition, both dietary digestibility and metabolizability have usually
been estimated rather than directly determined. Some fruit bats eating very low protein diets may
consume large amounts of food to meet protein requirements, thus elevating energy intakes.
Researchers generally have noted little change in body mass, but studies of changes in body
composition appear not to have been made. With respect to energy needs for flight, requirements
are surely higher than for resting, and the amount of flight activity required in the wild will vary
with food abundance and distribution. In captivity, it is likely that energy requirements for resting
and for flight can be met by offering ad libitum quantities of nutritionally appropriate food.
Essential fatty acids

Apart from serving as sources of dietary energy and promoting fat-soluble vitamin
absorption, fats also provide essential fatty acids (n-6 and n-3 series). The most important dietary
fatty acids for the mammal species that have been studied are n-6 fatty acids, such as linoleic
and arachidonic, and n-3 fatty acids, such as ∞-linolenic and longer chain fatty acids in this
series. Qualitative requirements for essential fatty acids have been demonstrated in a number of
mammalian species although quantitative requirements are mostly undefined. Fruit bats are likely
to have dietary fatty acid requirements similar to other mammals. The recommendation for most
mammals is to include essential fatty acids in the diet at 1-2% of the total metabolizable energy
intake, with linoleic acid comprising the largest proportion of the total. However, the amounts of
total essential fatty acids and the proportions of each may vary with species, age, and physiologic
state.64, 72 Cultivated fruits generally have very low levels of fat and essential fatty acids. 60, 73
Essential fatty acid concentrations in the wild fruits eaten by bats are largely unknown.
Vitamins and minerals

Specific vitamin and mineral requirements have not been determined for fruit bats, and there is no
information on the nutrient requirements of their closest taxonomic relatives. Limited data are
available on a few primate species.50 The National Research Council (NRC) has established
vitamin and mineral requirements for a variety of domestic and laboratory animals. 50,53-56 NRC
nutrient requirements for rats and mice are shown in Table 3. 55 The range in body mass of
laboratory rats and mice is similar to that of several species of fruit bats, although relative intakes
of dietary dry matter and energy are typically higher for bats. Thus, when nutrient requirements
are expressed per unit of dietary dry matter, bats would be expected to consume larger amounts
of these nutrients per day than would rats or mice of comparable mass and physiologic state. As
a consequence, until specific requirements are determined, diets for captive bats may be
presumed sufficient if formulated to provide bioavailable 1 vitamin and mineral levels within the
ranges of nutrients for the listed laboratory species. It is known that some bat species lack the
ability to synthesize vitamin C and therefore require a dietary source. 3 Despite serum levels of 25-
hydroxyvitamin D <4 ng/ml in both captive and wild populations of the Egyptian fruit bat
(Rousettus aegyptiacus), small amounts of 1,25-dihydroxyvitamin D were found, serum mineral
concentrations were well regulated, and pathologic signs usually associated with vitamin D
deficiency were not evident.7 Others reported plasma concentrations of 1.5 ng 25-hydroxyvitamin
D/ml and 93-108 pg 1,25-dihydroxyvitamin D/ml in captive Pteropus hypomelanus and P.
vampyrus fed a diet containing 400 IU vitamin D3/kg DM and exposed to natural sunlight
immediately prior to the study.17
Qualitative information on some vitamin and mineral requirements of fruit bats is available
from observations of deficiencies or toxicities in captivity. Methodology must be carefully
considered when interpreting the quantitative implications of such results. One report has
described dilated cardiomyopathy in pteropodid fruit bats as a result of hypovitaminosis E. 24
Hemochromatosis (iron storage disease) has been reported in three species of pteropodid fruit
bats (Rousettus aegyptiacus, Pteropus giganteus, and P. poliocephalus), and was ascribed to
dietary iron overload.13 The diet contained 400 mg of iron/kg of dry matter, by analysis. The
principal unintended iron source was monodicalcium phosphate, which was used as a
phosphorus supplement and contained 11,860 mg iron/kg. Unusually high vitamin C
supplementation (estimated intake of 7,500 mg/kg dry diet) may have compounded the problem
by enhancing iron uptake.44,45 Studies of hematologic values and iron status of wild and captive

fruit bats (Rousettus aegyptiacus) found lower plasma iron concentrations in wild (175 ng/dl) than
in captive (286-316 ng/dl) subjects.75
Investigators studying the cause of nodular bone lesions in the same three species of
pteropodid fruit bats proposed fluoride toxicity as the probable cause. 18The diet contained shrimp
meal, which is naturally high in fluoride, dicalcium phosphate, which may contain significant
fluoride if a fertilizer-grade rather than a feed-grade product is used, fruit, and other supplements.
Although the concentration of fluoride in the total diet was not reported, the estimated amounts of
fluoride consumed by the bats were well above maximum tolerable levels reported for other
species.51 Recently, several more cases of suspected fluoride toxicosis have been reported in
pteropodid species (six Pteropus rodricensis, one Rousettus aegyptiacus) at one institution.71 The
affected bats exhibited periosteal hyperostosis of long bones, similar to the previous study, as
well as curled phalanges. The fluoride concentrations (by analysis) of all dietary items and water
sources, as well as other potential sources of fluoride contamination (e.g., exhibit rock-work,
flooring, and heating system condensate) were well below maximum tolerable levels for domestic
species. Signs of periosteal hyperostosis were not seen after municipal tap water was replaced
by distilled water for drinking.71 Quantitative fluoride requirements have not been established for
most species. However, excessive levels have been shown to cause bone and tooth lesions,
anorexia, lameness, necrosis of gastrointestinal mucosa, and cardiac failure. 51 The results of
these studies are not conclusive since requirements and tolerances of bats for fluoride have not
been established.
Table 3. Vitamin and mineral requirements of laboratory mice and rats in dietary dry
matter.a
Nutrient Mice Rats
Vitamins
A, IU/kg 2,600 2,600
D3, IU/kg 1,100 1,100
E (RRR-∞- tocopherol), mg/kg 24 20
K (phylloquinone), mg/kg 1.1 1.1
Thiamin, mg/kg 5.6 4.4
Riboflavin, mg/kg 7.8 4.4
Niacin, mg/kg 17 17
Panthothenic acid, mg/kg 18 11
Vitamin B6, mg/kg 9 7
Folic acid, µg/kg 555 1,100
Biotin, µg/kg 220 220
Vitamin B12, µg/kg 220 220
Choline, mg/kg 2,222 833
Minerals
Calcium, % 0.6 0.7
Phosphorus, % 0.33 0.4
Magnesium, % 0.06 0.07
Sodium, % 0.06 0.06
Potassium, % 0.22 0.4
Iron, mg/kg 39 83
Copper, mg/kg 7 9
Manganese, mg/kg 11 11
Zinc, mg/kg 11 28
Iodine, mg/kg 0.17 0.17
Selenium, mg/kg 0.17 0.4
a Requirements for most demanding life periods.55
Diets for Captive Frugivorous and Nectarivorous Bats

When formulating diets for captive fruit bats, foods available and appropriate for meeting probable
nutrient needs must be identified. Bats that are primarily frugivorous or primarily nectarivorous are
commonly fed similar diets. Nutritionally complete nectar formulas are sometimes included in
diets for species that consume large amounts of nectar in the wild. Cultivated fruits are readily
accepted by both groups. However, the concentrations of several nutrients in cultivated fruits are
typically low or imbalanced, and an additional food source must generally be provided to ensure
that nutrient concentrations in the total diet will be within the range required by other nonruminant
mammalian species.50,53-56 The preferred method for ensuring dietary adequacy is to provide
nutritionally complete feeds such as low-fiber, dry primate diets or dry diets formulated for
frugivorous birds. These feeds should be formulated to meet the currently known requirements of
the species for which they have been designed. The use of individual vitamin and mineral
compounds as nutrient supplements is not recommended unless they are provided in a
preparation specifically formulated for fruit bats. Over- or under-dosing, when using individual
vitamin or mineral supplements, is common, as are the consequent toxicity and/or deficiency
signs. The nutritionally complete feed used should be as finely ground as possible to promote
adequate consumption. Mixing fruit nectars, such as peach or apricot, with the ground feed will
soften and/or suspend the diet and encourage intake, since fruit bats tend to be attracted to fruit
nectars. Fruit also may be added to this mixture. However, it is advisable to chop the fruit into
very small pieces or to mix ingredients in a blender so bats cannot preferentially consume fruit
only. Because this mixture may gel if not mixed correctly and will ferment in a hot environment, its
condition should be assessed periodically and the mixture replaced when necessary. Behavioral
enrichment in the form of hanging whole fruits may be used to encourage natural feeding
behavior, providing fruit intakes relative to intakes of the complete diet are carefully monitored
and are not excessive. Microchiropteran and megachiropteran fruit bats are sometimes offered
insects in addition to fruit, nutritionally complete feeds, and nectar formulas. A number of “home
made” nectar formulas used successfully in captivity have been previously described. 62 However,
these formulations contain a variety of supplements that can be dangerous if misused.
Nutritionally complete nectar powders formulated for hummingbirds are currently available with
varying protein concentrations and may be appropriate for nectarivorous bats. Megachiroptera
also are sometimes offered leafy vegetables and browse, both of which are readily
accepted.34,36,37 Preliminary research suggests that certain browses are consumed
preferentially,32,33,38 but the reasons for these choices are not entirely clear. 2
The total diet should be formulated so that the items consumed will provide the probable nutrient
requirements. Formulas and nutrient specifications for three diets that have been used
successfully for 5 years, with generally minor modifications, are shown in Table 4.
Table 4. Formulas of and calculated nutrient concentrations in three sample fruit bat diets.
(Percent by weight, as fed)
Ingredient Diet A a Diet B b Diet C c
Fruits, whole d 5.2 42.0 --
Fruits, chopped d 30.5 13.2 17.0
Vegetables, chopped e 7.3 1.65 1.0
Starchy vegetables, chopped f 2.0 1.65 2.0
Leafy green vegetables, chopped g 10.4 -- 7.0
PMI Prolab High Protein monkey 13.7 2.99 --
diet 5045 h
Marion Jungle Biscuit i -- -- 22.0
Calcium Phosphate, dibasic 0.30 0.41 --
Peach nectar, canned 6.0 38.1 22.0
Orange juice, frozen, reconstituted 6.0 -- --
Water 18.6 -- 29.0
100.0 100.0 100.0
Nutrient Calculated Concentration, DM basis

Dry matter, % 25.2 20.6 29.0

Crude protein, % 16.2 6.5 18.3
Crude fat, % 3.8 1.8 6.4
Vitamin A, IU/kg 23,419 6,240 10,484
Vitamin D3, IU/kg 3,595 958 2,877
Vitamin E, mg/kg 49 20 252
Vitamin C, mg/kg 588 752 444
Thiamin, mg/kg 10.6 4.0 6.5
Riboflavin, mg/kg 6.2 3.2 7.8
Niacin, mg/kg 72 32 59
Pantothenic acid, mg/kg 37 15 21
Vitamin B6, mg/kg 12 8 9
Folic acid, µg/kg 6,144 1,907 4,458
Biotin, µg/kg >103 >28 >301
Vitamin B12, µg/kg 26 7 23
Choline, mg/kg >980 >261 >1,079
Calcium, % 0.85 0.63 0.85
Phosphorous, % 0.61 0.52 0.58
Magnesium, % 0.17 0.09 0.15
Sodium, % 0.19 0.06 0.21
Potassium, % 1.16 0.90 0.84
Iron, mg/kg 411 352 140
Copper, mg/kg 14 7 15
Manganese, mg/kg 84 30 38
Zinc, mg/kg 90 29 78
Iodine, mg/kg 0.98 0.26 >0.8
Selenium, mg/kg 0.13 0.05 0.28
a Fed to Rodriguez fruit bats (Pteropus rodricensis) at Brookfield Zoo for 5 years (Ca phos.,
dibasic, added at 3 yr).
b Fed to Neotropical fruit bats (Artibeus jamaicensis) at Brookfield Zoo for 5 years.
c Fed to Rodriguez fruit bats (Pteropus rodricensis) at Philadelphia Zoo for 5 years.
d Whole fruits include apple, banana and orange; chopped fruits include apple, banana,
blueberries, currants, grapes, mango, and raisins.
e Vegetables (nonstarchy type) consisted of carrots, either steamed or raw.
f Starchy vegetables consisted of sweet potatoes, either steamed or raw.
g Leafy green vegetables included celery, romaine lettuce, and spinach.
h Purina Mills, LLC, 1401 S. Hanley Rd., St. Louis, MO 63144 (800-227-8941).
i Marion Zoological Inc., 2003 E. Center Circle, Plymouth, MN 55441 (800-327-7974).
Summary
Although specific nutrient requirements for frugivorous and nectarivorous bats remain virtually
unknown information exists which provides practical guidelines for formulating diets for captive
bats. This information must be reviewed critically, applied sensibly, and compared with what is
known for other mammalian species until more specific research is conducted. Opportunities for
research in bat nutrition abound and, in view of the diversity of species, the information derived
would improve not only our understanding of the nutrient requirements and dietary husbandry of
bats but also of other species with similar dietary habits and feeding strategies.
(*note- end of section adapted from Nutrition Advisory Group Handbook)

3.1.2. Provide sample recommended diet(s) for all life stages based on
nutritional requirements and identify body condition norms as
determined from wild animals, if possible
PRELIMINARY TARGET NUTRIENT LEVELS (drv matter basis)

Daily Diet
Nutrient Composition
Crude protein (%) 2.0- 15.0*
Fat (%) 5.0-9.0
Vitamin A (IU/g) 4.0- 14.0
Vitamin D2 & D3 (IU/g) 0.2-2.0
Vitamin E (mg/kg) 11.0-56.0
Calcium (%)* * 0.5- 1 .O
Phosphorus (%) * * 0.4-0.9
*Reported levels of protein consumed by free-ranging and captive Megachiropteran and
Microchropteran fruit- and nectar-feeding bats fall in this range (Morrison, 1980; Rasweiler, 1977;
Reiter, 1993; Stellar, 1986; Thomas, 1984).
**For Mammals, Calcium:Phosphorus ratios of 1: 1 to 2: 1 are recommended (Robbins, 1993).
Because we have yet to delineate the target nutrient levels, we are providing diets that, on face
value, appear to meet the above indicated nutrient levels. No examination of micro-nutrient levels
has occurred. These diets have been provided by several institutions that consider their programs
successful. Please note that these are presented only as suggestions and have not been
subjected to thorough review. To provide suggestions for easily adaptable diets, the data have
been modified slightly and presented as food groups rather than specific food items. The total
quantity to feed is not given and is dependent on many factors. However, as a rough guideline, an
average active adult bat may consume a total of approximately 10- 15% of body weight per day
(dry matter basis); or 50-120% of body weight on an as fed basis. If the animal is lactating this
may increase to 1.5 times that per day. Care should be taken that the animals do not become
obese when fed ad libitum.
Comparison of the Approximate Nutrient Content of Three Working

Diets with Preliminary Target Nutrient Levels (dry matter basis)
Target Zoo Zoo Zoo
Nutrient Levels A B C*
Crude protein (%) 2.0-15.0 12.3 19.0 4.9
Fat (%) 1.0--5.0 7.2 3.0 4.4
Vitamin A (IU/g) 4.0-14.0 26.0 14.9 8.1
VitaminD2& D3
(IU/g) 0.2-2.0 6.2 3.2 0. I**
Vitamin E (mg/kg) 11.0-56.0 133.1 50.2 29.8
Calcium (%) 0.5- 1.0 0.4 0.8 0.1 ***
Phosphorus (%) 0.4-0.9 0.3 0.4 0.1
* Values reported are averages because daily diet composition varies at this Zoo.
**Animals at this facility are housed under full spectrum lights
***Animals have ad libitum access to a mineral wheel
Working Diet Composition

We are aware of a number of nutritional concerns about some of the diets being used in captive
feeding regimes today. We hope to address these in the final document.
Food items: The proportions represented below are derived from the three working diets
presented above. It is possible to achieve the nutrient levels of the Zoo diets outlined above by
offering a diet consisting of the following food items. For adaptability to different institutional

situations, we are presenting the diets as food groups in percent to be included in the total diet.
This is percent contribution of each item (or food group) by weight, as fed.
COMPARISON OF THE PERCENT CONTRIBUTION OF FOOD GROUPS IN THREE

WORKING DIETS (AS FED BASIS)
Zoo Zoo Zoo
Food group A B C*
Fruit(%) 74.1 16.0 90.4
Vegetables (%) - 7.0 -
Starchy vegetables (%) - 7.0 -
Leafy, green vegetables (%) - 14.0 2.3
Fruit juice/nectar (%) - - 6.5
Water (%) - 41.0 -
Nutritionally complete products (%) 23.3** 7.5*** -
Protein supplements (animal-based) (%) - - 0.7
Home-made protein/ vitamin/mineral
supplements (%) 2.6 4.5 0.1
* Values reported are averages because daily diet composition varies at this Zoo.
**This consists of 14.2% canned primate diet and 9.1% canned feline diet.
*** Total contribution is from high protein monkey chow.
Food groups: The diet can comprise one or more of the following foods in the quantities outlined
above.
Fruit: apple, banana, grapes, pear, papaya, dried figs, raisins, melon, kiwi, etc.
Vegetables: carrots, green beans, etc.
Starchy vegetables: sweet potatoes, corn, etc.
Leafy, green vegetables: lettuce, spinach, kale, collard, mustard, etc.
Please notice that all of the working diets outlined above use home-made supplements in
conjunction with commercially-made, nutritionally complete products. If one or more of the
products are not included in each of the diets as outlined, the diet will no longer reflect the
nutrients presented in the previous section. Many of the home-made recipes are difficult, and
sometimes expensive, to make. Therefore, in the final nutrition chapter we intend to provide
recommendations for use of easily attainable, nutritionally complete diets that will allow
formulated diets to meet target nutrient levels.
In addition to specialized housing, Long-tongued bats have specialized dietary requirements. A

nutritionally complete diet for nectarivorous bats is composed of a liquid diet of fruit juice/nectar
and fresh fruits such as cantaloupe and banana. Table 1 present a successful diet currently in
use at the Biodome and the Organization for Bat Conservation’s facility (OBC).
Table 1. Liquid diet for nectarivorous bats
Food components Quantity

Tropical Passion Oasis unsweetened (Juice)* 960 ml
Corn oil 20 ml
Purina High Protein Monkey Chow 5045 15 g
Nektar-plus† 55 g
Nectar powder mixture 30 g
Sources of diet components
* A.Lassande Inc., Rougemont (Quebec), Canada
† Nekton, Pforzheim, West Germany

Nectar powder mixture
Quantity
Components
Mixed cereal 500 g
Fructose 500 g
Powder milk 300 g
Wheat germ oil 160 g
Vitadol Plus§ 40 g
§ Upjohn Company, Animal Health Division, Orangeville, Ontario, Canada.
3.1.3. As appropriate address the influence of the following variables on

dietary requirements
Age (infant, juvenile, reproductive adult, senescent adult, etc.)
body size
reproductive status
seasonal changes in ambient temperature
seasonal changes in body condition
seasonal changes in nutritional requirements
activity levels
health status
3.1.4. Address issues of palatability, texture, processing, etc. that will
encourage species-appropriate appetitive behaviors.
Fruit bats squeeze the juices out of their food and spit out the dry, fibrous portion.
Blending or puréeing fruits forces the bats to ingest large quantities of fiber that may impair
absorption of nutrients (George, 1990). Furthermore, because fruit bats are unable to concentrate
the fiber in blended diets, rectal prolapse may occur (Snell, 1994). Trupkiewicz et al. (in prep.)
reported frequent rectal prolapses in a zoo colony of Pteropus rodricensis that may have been
related to diet.
There remains a large amount of information to be collected in order to finalize recommendations

for target nutrient levels and appropriate feeding procedures for Megachiropteran and
Microchiropteran fruit bats. Much of this information will become available through our proposed
pre-survey and following in-depth survey of diets and feeding practices in institutions currently
housing fruit bats. The information still needed includes, but is certainly not limited to, the
following: Nutritional deficiencies/toxicities, feeding behavior in captivity, quantities of food to offer
(factors include: age, species, size of housing, number in group, etc.), form of diet offered (e.g.,
chopped vs. whole fruits, etc.), problems with obesity, seasonality (e.g., feeding different diets
based on season), feeding schedules, feeding location, use and potential nutrient contribution of
fresh green forages and other plant parts (see Appendix F for a partial list of forages accepted by
captive colonies of fruit bats), behavioral enrichment for captive fruit bats.
3.2. Medical management – guidelines should address and specify

acceptable husbandry practices for the taxon at different life stages
with respect to
3.2.1. Quarantine and hospitalization
We strongly recommend consulting with veterinarians and others experienced in
quarantine procedures when receiving bats. In addition, the Centers for Disease Control (CDC)
can provide quarantine information and recommendations.
Ideally, wild-caught animals should be held for one month prior to shipment from their
country of origin. This has the benefit of allowing detection and removal of any sick animals, and
it allows the animal to adapt to captivity prior to shipment 7. During the holding period prior to

shipment, the bats should be examined, preferably by a veterinarian, and treated for ecto- and
endoparasites.
When captive- born bats are received from another institution, ideally they should be
quarantined for 90 days. However, this may vary depending on the source of animals and their
previous standard of care and medical testing. Minimally, they should be quarantined for 30 days.
Wild-caught bats arriving from another institution should be quarantined for a minimum of six
months. Quarantine conditions should closely mimic the bats' natural ambient conditions (torpor
can lengthen the disease incubation time). All bats should be examined as soon as possible after
shipment for evidence of disease. An additional exam should be performed before the animals
are removed from quarantine. Any bats that die during quarantine must have a complete
necropsy (see Section 3.2.2). If a bat dies during quarantine and the cause is unknown, the
quarantine period should be extended to allow sufficient time for detection of disease in the
remaining animals. All Pteropus species should be screened for henipavirus.
Following completion of quarantine, it is recommended that wild-caught animals placed
on exhibit be isolated from other animals for at least one year, and housed in cages that protect
the public from bat urine and feces (e.g., plexiglass-fronted cages). Animals to be introduced to a
mixed collection should have been in isolation at least one year, and the group should be free of
evidence of infectious diseases. It is preferable that animals to be introduced into mixed exhibits
be first or second generation removed from wild-caught animals. The same is also true for
education animals.
Except for the 90 day quarantine period following receipt of the bats, the other
requirements (e.g., one year isolation and first and second generations) can be completed at
other accredited facilities.
3.2.1.1. Identify problems arising from isolation of social taxa

and suggest possible mechanisms for avoiding these
problems
3.2.2. Preventive medicine (testing, vaccinations, parasite control, etc.)
Ideally, all bats should be individually identifiable (see Section 5.1) and each bat should have a
separate medical records.
Inoculations
Recent studies suggest that captive bats can be vaccinated for rabies (Peters and Isaza,
2005; Lollar, 2004). However, vaccination of insectivorous bats constitutes extra-label
use of the rabies vaccine.
Recommended Routine Medical Procedures

Ideally, each bat should be examined once a year. When applicable, this examination
could include collection of blood for hematology, plasma biochemical analysis, and
plasma for freezing (freezing temperature equals -70˚F). When colony size renders
individual exams impractical, a few individuals could be selected from the colony for
examination.
Life Span
For their size, bats have very long life spans. Short-tailed leaf-nosed bats (Carollia
perspicillata) can live up to 12 years of age, and have a shorter life span than the larger
tropical species of megachiroptera. Straw-colored fruit bats (Eidolon helvum) can live up
to 22 years and the record age of an Indian flying fox (P. giganteus) is 31 years (Jones,
1982; Richarz and Limbrunner, 1993). More data need to be collected on this issue.
Necropsy Protocol
All bats that die should be necropsied. A gross necropsy alone is inadequate without
submission of representative tissues (brain, skeletal muscle, heart, lung, liver, stomach,

pancreas, small and large intestines, kidneys, gonads, bone and bone marrow) for
pathology. A section of brain from new or wild-caught bats that have died in captivity
should always be submitted for rabies testing. Accurate necropsy records should be
made and mortality rates should be tracked on a yearly basis (including neonatal deaths).
Parasites
1. Screening. All parasites collected should be submitted for identification. Fecal
examination for parasites should be done every six months for individuals and every
month on a sample collected from the colony.8 These tests should be performed by
people capable of identifying the parasites likely to be found in bats.
2. Commonly observed parasites and treatments.

a) Ectoparasites: Emtoparasites include mites, lice, chiggers, fleas, ticks and
flies (Whitaker, 1988). No ectoparasites should be tolerated and several
ectoparasites are potential vectors for blood borne diseases. Imported animals
should be examined and treated in the country of origin. Many ectoparasite
species will spontaneously leave the wild caught bat. Some ectoparasites spend
most of their time in the environment, and therefore any treatment program
should include changing props, substrate and so forth, as well as disinfecting,
and in large cages, fumigating.
Pest strips work well to deter ectoparasites confined to the host (e.g.,
mites, chiggers and lice). However, animals in outdoor exhibits (where use of
pest strips is not feasible), and those with heavy ectoparasite loads can be
treated with ivermectin (200-400 pg/kg PO, IM, SC) every 10 to 14 days for three
treatments. Repeated treatment may be necessary to kill the parasites that
subsequently hatch from the protected eggs. Ivermectin has a narrow therapeutic
index. Care should therefore be taken in calculating dosages. Signs of toxicity
range from lethargy and decreased appetite to full paralysis. These signs occur
within the first 24 to 48 hours. Treatment is supportive; there is no antidote.
Ivermectin for injection must be diluted with propylene glycol, not water. This is
because it is an oily substance and does not mix well with water. There is an oral
liquid preparation for horses that can be diluted with water and administered
orally.
Alternatively, some ectoparasites are susceptible to pyrethrin-based
sprays. Take care not to use excessive amounts (e.g., soaking the animal), and
do not use in confined, poorly ventilated areas. Rather than spray, it may be
better in smaller bats to moisten a towel, and rub into the coat. Whenever using a
treatment for the first time, use in a few individuals first and observe.
b) Endoparasites (see Coggins, 1988 and Constantine, 1993):

The major endoparasite of concern is Toxocara pteropodis.
Round worms:
Ivermectin 200 to 400 μg/kg PO. Extreme care must be taken when
diluting ivermectin, especially with smaller bats.
Fenbendazole 75 to 100 mg/kg PO, repeat in 10 days.
Tapeworms:
Praziquantel 7.5 to 15 mg/kg IM, PO.
Trematodes:
Prauquantel: high dosages (50 mg/lkg) are not very effective.
c) Hemoparasites: Hemoparasites (Hepatocystis sp.) have been observed in

imported Pteropus pumilus from the Philippines. These parasites spontaneously
disappeared from the blood over a 12 to 14 month period. No present treatment
is recommended.

d) Protozoa: Metronidazole 75 to 100 mg/kg PO.
Behavioral Signs of Illness

Behavioral signs of illness include: decreased appetite, wings constantly wrapped to the
body (even when disturbed), lethargy, reluctance to move 9, not flying or abnormal flying,
and animal found on floor (note: some healthy individuals will sleep on floor or retrieve
food from floor).
Common Injuries and Their treatment

1. Band constrictions: Pay attention immediately after placement and check on a
periodic basis (see Section 5.1). Remove the band immediately if swelling and/or
ulceration occur.
Treat severe injuries with antibiotics.
2. Other thumb injuries (e.g., tom nails, simple fractures or abrasions as a result
of contact with rough surfaces): Treat with antibiotics if deemed necessary.
3. Wing tears generally heal well by themselves. Wing tears that involve the
leading or supporting wing edges require suturing.
4. Bandaging that incorporates the wing web may obstruct inflow of arterial blood
causing subsequent necrosis (i.e., ischemic necrosis).
5. Fractures (wings and legs): Take special care not to pull too hard when
removing bats from wire and perches. Rough handling in young bats may cause injuries
at growth plates of long bones.
6. Dry skin and feet may be caused by low humidity (see Section 1.2).
7. Wing bone injuries: These injuries need to be carefully evaluated by a
veterinarian to determine appropriate treatment.
8. Swollen joints: Swollen joints may be caused by septic (bacterial) arthritis.
Treatment is based on culture and antibiotic sensitivity, as well as radiographs to confirm
severity of lesions.
9. Infected marking glands on chest: These may respond to antibiotics. In severe
cases, it may be necessary to excise the lesion surgically.
10. Torn lower lips. These may need to be sutured.
Physiological Reference Values

1. Heart and respiratory rates: Rapid, except during torpor. 10
2. Hematology and blood reference values: Available data are limited. The TAG
encourages zoos to submit data from their collections to ISIS. In addition, more research
is required to obtain normal reference values from free-ranging bats. Data that are
available are sometimes compromised by poor blood collection and handling techniques.
The following tips are recommended to facilitate collection: In a healthy bat, 10% of blood
volume can safely be collected. This is approximately 1% of body weight in grams (1
gram = 1 ml). For example, in a
450 gm bat, 4.5 mls of blood can be collected for plasma biochemical determination.
Blood should be collected into a heparinized syringe, immediately centrifuged, and the
plasma refrigerated prior to analysis.
3.2.3. Management of hereditary diseases or disorders
Major Disease Problems and Their Treatments

1. Viral. No treatment is available for viral diseases.
a. Rabies (see Constantine, 1988 and 1993): Rabies is not a major disease
problem for bats, although, when contracted, the virus will kill animals. The primary
concern is for public health (see Section 2.2.6.2). All bats should be vaccinated for rabies,
although this constitutes an extra-label use of the vaccine (Peters and Isaza, 2005; Lollar,
2004).
b. Henipaviruses

c. Other viruses: further research required.
2. Bacterial. Treatment is based on culture and sensitivity.
a. Salmonella
b. Other
Antibiotics used in bats include: fruit-flavored oral preparations, Baytril , Trimethoprimsulfa
combination, Amoxicillin, Clavamox, Chloramphenicol palmitate, and Keflin. Injectables include:
TMZ, Amikacin, Enrofloxacin.
3. Parasitic (see Section 3.2.2).
4. Nutritional
Diets for long-term maintenance of bats need to be perfected (see Section 3.)
a. Dilated cardiomyopathy - probably vitamin E deficiency.
b. Vitamins C and B 12
c. Metabolic bone disease.
5. Toxic
a. Zinc (galvanized wire)
b. Lead
c. Fluoride - P. rodricensis appears to be sensitive to low level exposure to
fluoride. Cumulative effects of fluoride exposure may result in toxicity (J.
Trupkiewicz pers. com.). At the Philadelphia Zoo, multicentric hyperostosis (bone
lesions) that is consistent with fluorosis has been diagnosed during post-mortem
examination of five P. rodricensis. Similar lesions have been identified in other
species of captive pteropids exposed to high levels of dietary fluoride (Duncan et
al., 1996).
d. Some plants
3.2.4. appropriate capture, restraint and immobilization techniques and

training for routine and non-routine procedures
Methods of Capture, handling, and restraint
Chiropteran wing bones are delicate and can be easily broken during capture. If using a net,
make sure it is wider than the bat's wingspan to avoid damage to wing bones and membranes.
Mist nets can also be used to capture bats. Be sure to have experienced personnel and adequate
staff on hand when using mist nets. It is important to remove bats from nets immediately so they
do not get overly entangled. To remove bats that are hanging on wire or perches, be sure to
unhook their nails rather than pulling on the bat. This will avoid injury to toes. In large, free-flight
aviaries, it may be useful to erect a food trap. This is accomplished by feeding animals in a small
enclosed area on a regular basis so that when it becomes necessary to capture bats, the area
can be sealed as the animal is eating.
Appropriate techniques for handling small mammals are generally adequate for bats, although
care should always be taken to immobilize the wings. Do not hold any bat by the wing tips during
restraint and always fold wings close to the body. To avoid permanent damage, special care
should be taken not to hyperextend the joints. While we do not know if holding bats head-up for
prolonged periods presents a risk to bats, we recommend trying to hold them head down (or at
least horizontally) as a precaution.
It is a good idea to wear a long sleeved shirt and gauntlet gloves 11 when handling bats, especially
the larger fruit bats. To reduce keeper injuries, welders’ gloves can be useful in handling large fruit
bats. However, use of them may cause injuries to the animals. If used, extreme care should be
taken not to apply excessive pressure. When weighing fruit bats, place the bat in a cloth bag
before putting it on the scale to reduce stress.
Immobilization and Anesthesia

The following recommendations are for a one pound fruit bat. The dose will need to be higher per
kilogram for smaller bats and lower per kilogram for larger bats. The preferred method of

anesthesia is isoflurane in oxygen. Mask down animal at 5% isoflurane until the animal is relaxed,
then decrease to 2.5 %. Maintain oxygen flow at 2 liters per minute. If gas anesthesia is
unavailable, a mixture of ketamine and xylazine can be utilized. For 30 minutes of immobilization,
combine equal volumes of ketamine (100mg/ml) and xylazine (20mg/ml) to achieve a
concentration of 50mg/ml ketamine and 10mg/ml xylazine and administer. Give .2mls/kg of
mixture for a 500 gram bat.
3.2.5. Management of neonates and geriatric animals

Neonatal Examinations
Births, as well as birthing problems and neonatal deaths, should be recorded. The neonate
should be visually assessed soon after birth to make sure it is suckling, has normal activity for its
age, and there are no gross congenital abnormalities. Although not practical or advisable for most
situations,12 neonatal bats can be examined, weighed and measured immediately after birth and
during development. Information from such a procedure can help to establish baseline
information for the individual, as well as for the species. Growth curves are invaluable for
valuating health status of lactating females and hand-raised bats. For more information on
neonatal exams, see Barnard, 1994.
When examining neonates and suckling bats, it is recommended that the mother be anesthetized
(see Section 3.2.4) to decrease the likelihood of injury to both parent and pup. The pup can be
removed from the nipple by gently placing a smooth probe (e.g., small rubber spatula) at the
comer of the mouth of the pup until it releases its grip on the teat. Force should not be used and
care should be taken to avoid damaging the milk teeth. Rubber spatulas work well to insure teeth
and mouth do not get damaged. The pup is either placed on the mother before recovery, or
placed in a small cage with the awake mother. Placement of the pup on the physically restrained
mother is stressful, may result in injury, and is generally unnecessary.
3.2.6. Management during pregnancy
4. Reproduction
Fruit bats share many of the general reproductive characteristics present in other mammalian
species. They reproduce sexually, produce live young, and females nurse pups from one or two
mammae located on either side of the thorax. Fruit bats have a rather low reproductive rate,
especially considering their size. Most field studies on bats, however, refer to sporadic
observation collected only a few times a year. Thus, much of the information on bat reproduction
is anecdotal. Some of the information in this chapter was collected through a TAG bat
reproduction survey that was sent to all AZA institutions housing bats. Tables 1 and 2, taken from
this survey, contain information on development in captive born bats and a summary of bat
reproduction data, respectively. For additional information, see Galindo et al, 1995.

Age-Specific Fecundity
Generally, females of !arger species do not give birth until they are between one and two years
old (Asdell, 1964; Nelson, 1965; Thomas and Marshal, 1984). Pteropus poliocephalus females
reach sexual maturity at one and a half years, while males do not reach effective fertilization until
they are two and a half years, even though they are considered sexually mature at 18 months of
age (Martin et al., 1986). The males of some other Pteropus species also do not reach effective
fertilization until two and a half years, even though they are sexually mature at 18 months (Martin
et al., 1986). Smaller species are able to reproduce as early as four to eight months of age
(Miclkeburgh, et al., 1992; Galindo et al., 1995), with males not becoming mature until between
one and two years of age. Cynopterus brachiotis females are sexually mature at four to six
months, while the males are mature at one year of age (Heideman, pers. comm., 1992). Many
species of fruit bats show a post partum estrus (Galindo et al., 1995; Mickleburgh et al., 1992).
4.1. Identify seasonal changes in physiology and behavior associated

with reproduction and address management implications of such
changes
Breeding Seasons
Beck (1973) reports a monoestrous cycle (one single estrus per breeding season [McDonald,
1987) in Pteropus giganteus, polyestrous cycles (multiple estrus cycles per breeding season
[McDonald, 1987]) in Rousettus aegyptiacus and three pregnancy peaks per year
in Cynopterus brachyotis. There has been one reported case of possible sperm storage in P.
giganteus in a captive environment, where one female gave birth after nine months of being in a
single sex female group (Galindo et al., 1995). In general, larger species of fruit bats seem to be

monoestrous, while smaller species are generally polyestrous. In captive Pteropus poliocephalus,
mating increases in February, peaking in April (Martin et al., 1986).
Female G. soricina exhibit menstruation and seems to have a reproductive cycle of 22-26
days (Rasweiler, 1972). Mating activity should normally occur during these 3 or 4 weeks. In
Central America, Flemming et al. (1972) reported that ovulation and mating may be biannual.
Females give birth throughout the year, although the majority of births at the Biodome occur in the
warmest period of the year (July and August).
Behavior
Although male bats will attempt to copulate throughout the year (Martin et al., 1986), the highest
copulation activity in bats is during the mating season. When not in season, females will avoid
copulation and will generally fly away from interested males. As part of the mating ritual, the
interested male may flick his wings towards the female and vocalize loudly. He will then attempt
to groom the female while he orients himself to her dorsal surface. He will attempt to clasp her
with his wings and thumbs, grasping the thickened nape of her neck with his teeth. During a
successful copulation, the female cooperates by hanging from the male's legs (Baker, 1991;
Martin et al.; West, 1986). Intromission is short and is accompanied by a quick ejaculation (Baker,
1991; Martin et al., 1986). Copulation is generally ventral/dorsal; however, Eidolon helvum have
been observed copulating in the ventral/ventral position (Galindo, 1995). 13
Breeding success of G. soricina seems not to be affected by the environment and outside
disturbance. Indeed, a few studies showed that G. soricina can mate and conceive despite being
housed in small cages and subject to daily handling (Rasweiler 1972, 1974). According to
Rasweiler (1974), a low male/female sex ratio may be advisable to ensure an efficient breeding
success with G. soricina.
Gestation
As a general rule, bats carry only one young per gestation period. Twining is present occasionally,
though survivability of both the pups and the dam is very low. At least one of the pups may need
to be hand-raised to relieve the dam of the stress of raising twins. In the wild, however, smaller
species such as Artibeus jamaicensis, Eidolon helvum and others have been seen to do quite
well with twins (Skinner and Smithers, 1990). Gestation in Pteropodid bats has been estimated at
105 days to 210 days. Examples include Pteropus rodricensis at 140- 180 days, Rousettus
aegyptiacus at 105- 107 days, and Epomophorus wahlbergi at 180-210 days (West and
Redshaw, 1987; Baker, 1991; Kunz et al., 1994; Carroll, 1988; West, 1985-86; Falanrow, 1988;
Sowler, 1984).
Eidolon helvum are essentially monoestrous, and are thought to have two distinct breeding
strategies in the wild. One strategy is to have a four month gestation with conceptions in the
autumn and births in the spring (Anderson, 1912). Another is to copulate in June or July and,
utilizing a delayed implantation strategy, have gestation begin in November, with the young being
born between March and April. This strategy usually coincides with the rainy season (Fayenuwo
and Halstead, 1974; Funmilayo, 1979; Galindo et al., 1995). Births of Pteropus rodricensis in
captivity occur mainly during October through April, with a peak between late March and April.
They show synchronized births and can potentially have two birth periods a year with an interval
of 260 days (Carroll, 1988).
There are some suggestions that Epomophorus wahlbergi have conceptions that occur from May
to December, with the peak occurring May through July. Births peak in November and December
(Sowler, 1984), thus making gestation roughly six to eight months. Rousettus aegyptiacus breed
two times per year in East Africa and year-round in North Africa (Kingdom, 1974).
Parturition
There is little behavioral change in the weeks and days prior to parturition. Female bats
will pant immediately prior to and during parturition. For birthing, she will alternate hanging with all
four limbs and clinging to the wire hanging from just her thumbs. In the Pteropodidae, infants are
born head first and, as soon as the head is exposed, the female will bring her feet up and catch
the pup as it emerges. The Microchiroptera are usually born rump or feet first.

A typical Pteropus birth and placental delivery may last only a few minutes to two hours
(Baker, 1991). There are usually few warning signs to show when a female is about to give birth.
The first signs are manifested two to three weeks prior to parturition. The bulk of the fetus shifts
from its usual sprawled lateral position to a more centralized, ventral position on her abdomen,
directly above the vaginal area, which seems to swell slightly. Hours prior to parturition, the
expectant mother separates herself from others in the group and starts fanning herself. When she
is ready to deliver, she hangs from her thumbs, as though she is attempting to void, and strains
hard. As the pup emerges head first, it is positioned so that the wings are folded around the head.
There is little blood loss. At the actual moment of birth, the female adopts a horizontal position,
grooming and licking the genitalia and the emerging infant (West, 1986). There is usually a pause
between expulsion of the head and rest of the body (Martin et al., 1986) which will generally last
from ten minutes to several hours (Martin et al., 1986). During this time, the pup has its eyes
open, twitches its nose, and occasionally will vocalize. The mother opens up her wings to make a
"net" to protect the infant from falling. The offspring hangs on to the mother with its feet and
thumbs and is guided to a nipple by the mother. The mother, and occasionally other cage mates,
will then eat the placenta (Martin et al., 1986). Kunz et al. (1994) have documented alloparenting
in Pteropus rodricensis.
Infant Development
Fruit bats grow and mature quickly. Smaller species of fruit bats are born with their eyes closed
and ears folded back, both opening after ten days (Lombard, 1961). Artibeus jamaicensis have an
average birth weight of four grams, and wean at an average age of three and a half months
(Galindo et al., 1995). All fruit bats are born with some fur, well developed claws to cling to the
mother's ventral side, and deciduous recurved milk teeth (West, 1985-86; Nelson, 1965; Pook,
1977). Pteropus rodricensis have short, snubbed muzzles, small wings and large feet (Baker,
1991; West, 1985-86) with a body weight of 45 grams. Forearm length is 43% of adult length
(West, 19486). Pteropus poliocephalus are an average of 76 grams at birth (Hood, 1989).
Milk cannot be expressed manually after about six months. However, many species of fruit bats
have been seen still on the mother's nipple after one year of age (Galindo et al., 1995; West,
1985-86; Baker, 1991). From observations in the field and in captivity, many species show a post
partum estrus (Galindo et al, 1995; Mickleburgh et al., 1992). At birth, G. sorcina males are
slightly larger than females (Walker 1975). Nursing lasts approximately 1 month before the young
bat can fly.
Neonatal/ Infant Mortality

According to the TAG'S published survey on reproduction of frugivorous bats in North American
Zoos (Galindo et al, 1995), 1370 individuals were born within the last ten years in responding
institutions. Of those born, 458 (33%) were either stillborn or died before the age of two months.
Some of the major causes of neonatal death included: environmental stress (overcrowding,
extremes in temperature, capturing for medical procedures), cage mate inflicted trauma, hazards
within the enclosure (drowning in pools, eaten by other species housed with the bats), and
medical inflictions (cesarean section deaths, infections). The survey also reported that there were
higher instances of neonatal deaths in larger species (100 grams and over) to pups born to first
time mothers than those born to experienced mothers. Cannibalism was also seen in many
instances where overcrowding was an issue. Birthing dams have been seen to partially consume
stillborn pups.
In Rodrigues fruit bats, pup mortality was investigated through radiographic analysis, by
Cooper and West (1988) who noted a preponderance of cranial injuries, including dislocated
mandibular symphyses, and depression fractures of the skull. These injuries are typical of flying
fox mothers who immediately reject the pup after birth. First time mothers, obese females, and
mothers under stress are more prone to reject pups than well adapted females who have a
history of successful births (LeBlanc, pers. comm.).
4.2. Address hormonal tracking as a mechanism for identifying

reproductive state, and assessing feasibility of introduction for
solitary species

4.3. Address timing of introductions for individuals of solitary species
4.4. Address provision of and describe facilities for parturition and as
appropriate, management of females during isolation or denning
4.5. Address what, if any, circumstances might warrant hand-rearing and
identify acceptable hand-rearing and reintroduction protocols
Hand-rearing
Hand-rearing fruit bats is similar to hand-rearing other small mammals. Be sure to wrap the infant
in a towel or hang it on a stuffed animal for its comfort and security. Initially, infants should be fed
with milk replacer every two hours from six a.m. until midnight. The milk replacer should be a high
fat (e.g., 43% dry weight) product that utilizes vegetable oil, not soy14 or butterfat.
Some examples of acceptable formulas are Similac, Puppylac, and Just Born. Feedings can be
decreased gradually over time with pureed and then diced fruits being introduced until infants are
weaned (large Pteropus species are usually weaned by six months; smaller fruit bats will be
weaned earlier). Many suckling fruit bats remain on the nipple until they are weaned and should
be given continuous access to a nipple. For additional information on hand-rearing, See Barnard,
1995; Young, 1987; and George, 1988.15
Fostering
The Lubee Bat Conservancy has successfully fostered a Little-golden mantled flying fox pup to its
grandmother who was in another enclosure, and lactating. The Durrell Wildlife Conservation
Trust has also successfully fostered a Rodrigues fruit bat.
4.6. Recommend means and duration of contraception for taxon; include

all acceptable alternatives and identify the benefits and drawbacks of
each
Contraception
Little or no information is available on reversible chemical contraceptive methods in bats.
The most commonly used method of reversible contraception in mammals in AZA institutions are
MGA implants. However, the efficacy of these in bats has yet to be determined. Barnard and
Soloway (1992) describe a method of male castration as a form of irreversible contraception.
Recently, Lafortune et al. (2004) showed that vasectomies are possible in bats. Unlike castration,
a vasectomy has the potential to be reversed.
Husbandry techniques, such as maintaining single-sex groups, are recommended as the most
simple and inexpensive method of contraception. Downfalls such as lack of space to house
separate groups of the same species can be overcome. One way to acquire additional space for
single sex colonies is to house bats with other taxa (see Section 2.2.4). Another method is for
institutions to trade the same species of bat, so each institution receives all of one sex.
5. Behavior management
5.1. Identify procedures that have been successful in managing the taxon
for routine husbandry. Routine husbandry may include such
activities as shifting from one area to another, tolerating close visual
inspection, tolerating close proximity to caretakers, etc.
Record Keeping
All bats should be included in ISIS records. We further recommend that all neo-natal deaths,
stillborns, and aborted fetus' be included in ISIS records. While this is not always an easy task,
especially with large colonies of microbats, it is an important tool in determining population
demographics.
Individual Identification Methods

All known identification methods have advantages and disadvantages and tend to be species
specific. The following is a summary of selected identification methods used in bats (taken from
Kunz, in prep.)

1. Transponders. The AZA Chiropteran TAG recommends all fruit bats be permanently
identified using the Trovan Electronic Identification System (i.e., transponders), although due to
the development of ISO universal readers and transponders other systems such as AVID and
Digital Angel (Formerly Destron) are acceptable. 16 To insert a transponder, pinch and pull the
skin, insert needle under the skin, and inject transponder right side up. The best placement is in
the posterior dorsal region where it will not interfere with movement. Some managers have
placed transponders between the shoulder blades; however, this location may interfere with
muscles and shoulder blade movements associated with flight. Transponders can migrate, and
several observations of the transponders occurring in the patagium have been noted at the Lubee
Bat Conservancy, even when they were injected below the scapulas. Advantages of this
technique include relative permanence and few health/injury risks. The disadvantage of
transponders is that the individual must be in-hand to read the identification.
2. Bead-chain necklace. Many field researchers have used bead-chain necklaces quite
successfully and they seem to work well with some species. The advantages of bead-chain
necklaces are that they can be seen from a distance and come in many color and/or number
combinations. Disadvantages are that beads have been reported to fade, they can break, be
abrasive, harbor mites and ticks, food can cake around necklace, and the public sometimes
reacts negatively to the "jewelry."
3. Thumb bands. Some species do fairly well with thumb bands, while others, such as
Rousettus aegyptiacus and Pteropus rodricensis are able to remove them. Use colored bands17
that are placed on the animal loosely enough to turn easily and not cut off circulation. Aluminum
bands are not recommended for fruit bats as they can be compressed if bats chew on them. The
advantages of colored thumb bands are that they are fairly easy to read from a distance and
there are many color and/or number combinations available. Multicolored band combinations as
well as numbered bands should be read from distal end of thumb in toward the wing. Stainless
steel bands are also available and work well with some species. However, stainless steel bands
cannot be read from a distance (as colored bands can). Therefore, we recommend using
transponders for close identification needs and colored bands for remote identification.
4. Forearm bands. Extreme care must be taken when using forearm bands because
they can restrict circulation in the propatagium. They may also injure the underlying muscle and
tendons if bats chew and bend the band. If used, band combinations or numbers should be read
from the distal end of thumb.
5. Ear notching. This technique is not recommended because it disfigures the animal
and there are relatively few combinations available.
6. Ear tags. Ear tags can cake with food and can tear out of the ear. In addition, they are
difficult to read from a distance.
7. Tattooing Wing tattooing is a short-term identification method because the tattoo

does not last long. Another disadvantage is the need to catch the animal in order to read the
tattoo. Tattooing of the ear with light colored species such as Egyptian fruit bats (R. aegyptiacus)
and Wahlberg’s epauletted fruit bats (Epomophorus wahlbergi) is permanent (LeBlanc, pers.
comm.).
8. Freeze branding. This method of identification has not been successful to date.
9. Nail polish. Nail polish, when placed on the individual's back, is fairly useful for short
term identification purposes, and many color variations are available. The disadvantage of this
method is the frequent need to replace polish. In addition, while nail polish makes for easy
identification because it is easy to see, it may be offensive to the visiting public.
All methods of identification, but especially bands, should be checked regularly.

5.2 Identify procedures that have been successful in managing the taxon for non-routine
husbandry. Non-routine husbandry includes regular weighing, sample collection, blood
pressure monitoring, injections, crating, swimming/climbing into a stretcher, walking into
a small stall/funneling space, transport, etc.
See section 3.2.4, and appendix for SOP’s.
5.3 Identify procedures that have been successful in facilitating introductions.

These may include separation of individuals from group, stationing, tolerance
while feeding, “howdy” units, visitation gates, etc.
Be careful not to overextend quarantine of individual animals or their social behavior may be
adversely affected. If possible, quarantine bats in groups rather than singly.
Introducing and removing bats from a colony is generally not problematic. New animals
can be introduced immediately into a group rather than maintained in a traditional "howdy" cage
(i.e., a small cage placed within the main exhibit cage for the purpose of letting animals become
familiarized). However, the use of a howdy cage for introducing a group of bats into a large free-
flight exhibit can help new bats get acclimated to the environment and possibly reduce
aggression between the new group and established colony. Browse and enrichment can be
utilized to offer new animals sanctuary from aggressive cage residents by offering more visual
barriers in the enclosure, and distracting current cage residents with food. Other visual barriers
that can be utilized are corrugated vinyl roofing, cardboard or cotton bed sheets.
Introducing new bats into an existing colony should take place at the start of a keeper
shift, so as to allow maximum observation time. Newly introduced animals should be watched off
and on for 24-48 hours to ensure that the animal(s) is eating and has adjusted well. Juvenile and
adult bats will frequently perform static flying when placed in a new exhibit as they grow
accustomed to the new environment (Carroll, 1978).
The stress of capture of microchiropteran fruit bats (e. g., Carollia perspicillata & and
Artibeus jamaicensis) , can induce torpor and/or a period of fasting that can last up to 24 hours.
To help prevent this problem, we recommend that removal of individuals be planned for after the
normal feeding period.
Initially there will be many confrontations among the bat colony as the group's social
structure is challenged and perhaps altered. New males may take over existing males' territories
or they will be dominated by the cage residents. Females tend to accommodate to new members
more easily than males. Removing bats from colonies causes minimal stress.
If an animal needs to be treated for medical purposes, it is best to provide veterinary care
while leaving the animal in the main enclosure. If it is absolutely necessary to remove the animal
for treatment, be sure to minimize the amount of time it is separated from the colony and follow
recommendations for introductions when reintroducing the animal to the colony. For bats that
have to be isolated for protracted periods, their scent marks can be captured by placing a sealed
muslin cloth on the ceiling of the isolation cage as a visual barrier, and then moving the scent-
marked cloth into the regular pen. Since the scent of this bat was maintained in the enclosure,
the bat may have an easier time being reintroduced to its cage mates.
There appears to be no major effect of removing animals from a colony except that, if the
individual removed is high ranking in the social order, there may be some jostling while a new
hierarchy is established. This may affect reproduction, but usually just for a short time.
If maintaining the colony for genetic management, the breeding male should be removed
immediately after the first young are born. If it is known that a female is pregnant, the male can be
removed earlier, although care should be taken not to stress the entire colony while attempting to
capture males. New males or females being added to the colony should be introduced two to
three months after the colony's females give birth. At this stage, the infants will be large enough to
fend for themselves should the newcomers act aggressively.
5.4 Identify facility design considerations, husbandry training
techniques, and implementation plan that can be used to elicit desired
behaviors in a way that is safe for both caretakers and animals.

5.5 Identify those techniques that have been shown to be most
effective.
5.6 Identify technical skills and competencies needed by staff
5.7 Appropriate methods of enrichment for the taxon should be

identified if not included in categories 1-3 above.
Environmental Enrichment
Environmental enrichment is important for bats, and keepers and managers should spend some
time developing new ways to stimulate their bat colonies. Methods of environmental enrichment
that are known to be successful include providing naturalistic foods, varying the food
presentation, supplying olfactory stimulation, adding props to the enclosure such as ropes,
branches, mirrors and so forth, providing adequate flight space, and supplying bats with browse
(for additional information on environmental enrichment for fruit bats, see Atkinson, 1993; LeBlanc
and Seyjagat, 2000).
Refer to Appendix F for a partial list of forages accepted by captive colonies of fruit bats.
Notes on Individual Species and Unusual Behaviors

Epomophorus wahlbergi will rock back and forth (laterally) with their wings wrapped around them
when resting (Wickler and Seibt, 1976).
Rousettus aegyptiacus and other small bats will occasionally sleep on their backs on the floor of
the cage (usually propped against a wall).
There are usually high levels of aggression between males in a breeding colony of Cynopterus
brachyotis.
In outdoor enclosures (and indoor enclosures where browse is provided), many bat species will
crawl to the ground in search of green foliage.
6. Documentation
ENDNOTES (for Nutrition section 3.1 adapted from Nutrition Advisory Group Handbook)
1 Ammerman, C.B., D.H. Baker, and A.J. Lewis (eds.). 1995. Bioavailability of Nutrients for
Animals. Academic Press, San Diego, CA.
2 Ange, K.D., S. Rhodes, and S.D. Crissey. 2001. Browse consumption and preference in the
Rodriguez fruit bat (Pteropus rodricensis). AAZK Animal Keepers’ Forum 28(12):475-482.
3 Ayaz K.M., E.C. Birney, and R. Jennes. 1976. Inability of bats to synthesize L-ascorbic acid.
Nature 260:626-628.
4 Bonaccorso, F.J. 1978. Foraging and reproductive ecology in a Panamanian bat community.
Bull. Florida State Mus. Biol. Sci. 24:359-408.
5 Bonaccorso, F.J., and S.R. Humphrey. 1979. Population and community ecology. Pp. 409-441
in Baker, R.J., J.K. Jones, Jr., and D.C. Carter (eds.). Biology of Bats of the New World Family
Phyllostomatidae, Part III. Mus. Texas Tech. Univ. Spec. Publ. 16.
6 Bradbury, J.W. 1977. Social organization and communication. Pp. 1-72 in Wimsatt, W.A. (ed.).
Biology of Bats, Vol. III. Academic Press, New York, NY.
7 Cavaleros, M., R. Buffenstein, F.P. Ross, and J.M. Pettifor. 2003. Vitamin D metabolism in a
frugivorous nocturnal mammal, the Egyptian fruit bat (Rousettus aegyptiacus). Gen. Comp.
Endocrinol.133:109-117.

8 Conklin, N.L., and R.W. Wrangham. 1994. The value of figs to a hind-gut fermenting frugivore: a
nutritional analysis. Biochem. Sys. Ecol. 22(2):137-151
9 Conklin-Brittain, N.L., E.S. Dierenfeld, R.W. Wrangham, M. Norconk, and S.C. Silver. 1999.
Chemical protein analysis: a comparison of kjeldahl crude protein and total protein ninhydrin
using wild, tropical vegetation. J. Chem. Ecol. 25:2601-2622.
10 Constantine, D.G. 1986. Insectivorous bats. Pp. 650-655 in Fowler, M.E. (ed.). Zoo and Wild
Animal Medicine, 2nd Ed. W.B. Saunders Company, Philadelphia, PA.
11 Courts, S.E. 1996. Insectivory in captive Livingstone’s and Rodrigues fruit bats Pteropus
livingstonii and Pteropus rodricensis (Chiroptera: Pteropodidae): A behavioural adaptation for
obtaining dietary protein. J. Zool., London. 242:404-410.
12 Courts, S.E. 1998. Dietary strategies of Old World fruit bats (Megachiroptera, Pteropodidae):
how do they obtain sufficient protein? Mammal Rev. 28(4):185-194.
13 Crawshaw, G., S. Oyarzun, E. Valdes, and K. Rose. 1995. Hemochromatosis (iron storage
disease) in fruit bats. Pp. 136-147 in Proc. Annu. Meet. Nutr. Advisory Group of Am. Zoo Aquar.
Assoc.
14 Delorme, M., and D.W. Thomas. 1996. Nitrogen and energy requirements of the short-tailed
fruit bat (Carollia perspicillata): fruit bats are not nitrogen constrained. J. Comp. Physiol. B
166:427-434.
15 Delorme, M., and D.W. Thomas. 1999. Comparative analysis of the digestive efficiency and
nitrogen and energy requirements of the phyllostomid fruit bat (Artibeus jamaicensis) and the
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16 Dierenfeld, E.S., and J. Seyjagat. 2000. Intake and diet digestibility in three species of captive
pteropid bats. Pp. 73-80 in Nijboer, J., J.–M. Hatt, W. Kaumanns, A. Beynen, and U. GansloBer
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17 Dierenfeld, E.S., and J. Seyjagat. 2000. Plasma fat-soluble vitamin and mineral concentrations
in relation to diet in captive pteropodid bats. J. Zoo Wildl. Med. 31:315-321.
18 Duncan, M., G.J. Crawshaw, K.G. Mehren, K.P.H. Pritzker, M. Mendes, and D.A. Smith. 1996.
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19 Edwards, M.S. 1995. Comparative Adaptations to Folivory in Primates. Ph.D. Dissertation.

Michigan State University, E. Lansing, MI.
20 Engriser, E.M. 1995. The effect of insect larvae infestation on fruit choice in phyllostomid fruit
bats: an experimental study. Biotropica 27:523-525.
21 Fenton, M.B., and T.H. Kunz. 1977. Movements and behavior. Pp. 351-364 in Baker, R.J., J.K.
Jones, Jr., and D.C. Carter (eds.). Biology of Bats of the New World Family Phyllostomatidae,
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22 Fleming, T.H. 1988. Evolution and ecology of Phyllostomid bats. Pp. 3-35 in Schaller, G.B.
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Chicago, IL.
23 Gardner, A. L. 1977. Feeding habits. Pp. 293-350 in Baker, R.J., J.K. Jones, Jr., and D.C.
Carter (ed.). Biology of Bats of the New World Family Phyllostomatidae, Part II. Mus. Texas Tech.
Univ. Spec. Publ.13.

24 Heard, D.J., C.D. Buergelt, P.S. Synder, A.K. Voges, and E.S. Dierenfeld. 1996. Dilated
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25 Herbst, L.H. 1983. Nutritional analyses of the wet season diet of Carollia perspicillata
(Chiroptera: Phyllostomidae) in Parque Nacional Santa Rosa, Costa Rica. M.S. Dissertation.
University of Miami, Miami, FL.
26 Herbst, L.H. 1986. The role of nitrogen from fruit pulp in the nutrition of the frugivorous bat
Carollia perspicillata. Biotropica 18:39-44.
27 Herrera, L.G., E. Gutierrez, K.A. Hobson, B. Altube, W.G. Diaz, and V. Sanchez-Cordero.
2002. Sources of assimilated protein in five species of New World frugivorous bats. Oecologia
133:280-287.
28 Hill, J.E., and J.D. Smith. 1984. Food habits and feeding. Pp. 60-72 in Bats: A Natural History.
University of Texas Press, Austin, TX.
29 Iudica, C.A., F.J. Bonaccorso, and G. Richard. 1994. Sea water ingestion in Pteropus
hypomelanus. Bat Research News 35(4):102 (Abstr.).
30 Klite, P.D. 1965. Intestinal bacterial flora and transit time of three neotropical bat species. J.
Bacteriology 90:375-379.
31 Korine, C., Z.Arad, and A. Arieli. 1996. Nitrogen and energy balance of the fruit bat Rousettus
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32 Kunz, T.H., P.V. August, and C.D. Burnett. 1983. Harem social organization in cave roosting
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33 Kunz, T.H., and C.A. Diaz. 1995. Folivory in fruit-eating bats, with new evidence from Artibeus
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34 Kunz, T.H., and D.M. LeBlanc. 1996. Leaf-eating habits of “fruit bats”. P. 2 in Pteropus, Newsl.
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35 Law, B.S. 1992. The maintenance nitrogen requirements of the Queensland blossom bat
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36 LeBlanc, D., A. Field, and J. Roberts. 1993. Browsing: encouraging folivory in flying foxes. The
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37 LeBlanc, D. 1999. Bat enrichment survey. AAZK Animal Keepers’ Forum 26(7):267-285.
38 Lowry, J.B. 1989. Green-leaf fractionation by fruit bats: is this feeding behaviour a unique
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39 Luft, S., E. Curio, and B. Tacud. 2003. The use of olfaction in the foraging behavior of the
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40 McNab, B.K., and M.I. Armstrong. 2001. Sexual dimorphism and scaling of energetics in flying
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41 McNab, B.K., and F.J. Bonaccorso. 2001. The metabolism of New Guinean pteropodid bats. J.
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42 Mikich, S.B. 2002. The diet of fruit-eating bats (Mammalia, Chiroptera, Phyllostomidae) in a
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43 Milton, K., and F.R. Dentzis. 1981. Nitrogen-to-protein conversion factors for tropical fruit
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44 Monsen, E. 1982. Ascorbic acid: an enhancing factor in iron absorption; Chapter 5. in Kies, C.
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45 Morris, E.R. 1987. Iron. Pp. 79-142 in Mertz, W. (ed.). Trace Elements in Human and Animal
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46 Morrison, D.W. 1978. Foraging ecology and energetics of the frugivorous bat Artibeus
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47 Morrison, D.W. 1979. Apparent male defense of tree hollows in the fruit bat, Artibeus
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48 Morrison, D.W. 1980. Efficiency of food utilization by fruit bats. Oecologia 45:270-273.
49 Morrison, D.W. 1980. Foraging and day-roosting dynamics of canopy fruit bats in Panama. J.
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50 National Research Council. 1978. Nutrient Requirements of Nonhuman Primates, Publ. No.
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51 National Research Council. 1980. Mineral Tolerance of Domestic Animals. National Academy
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52 National Research Council. 1981. Nutritional Energetics of Domestic Animals and Glossary of
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54 National Research Council. 1986. Nutrient Requirements of Cats, Publ. No. 3682. National
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56 National Research Council. 1998. Nutrient Requirements of Swine, 10th Rev. Ed. National
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bat feeding preferences in Panama. Biotropica 32:489-501.
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76 Wilson, D.E. 1979. Reproductive patterns. Pp. 317-318 in Baker, R.J., J.K. Jones, Jr., and
D.C. Carter (eds.). Biology of Bats of the New World Family Phyllostomatidae, Part III. Mus.
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78 Zortea, M., and S.L. Mendez. 1993. Folivory in the big fruit-eating bat, Artibeus lituratus
(Chiroptera, Phyllostomidae) in eastern Brazil. J. Trop. Ecol. 9:117-120.
ENDNOTES (for all other sections except Nutrition section 3.1)
1. Only two species, Pteropus poliocephalus and Rousettus aegyptiacus extend from tropical
regions into cool temperate regions.
2. At least one zoo has reported bat deaths due to tracking powder (K. Whitman, pas. comm.).
3. Available from Valentine Incorporated. Phone: (800) GET STUFF
4. Netting is available from Endurance Net, Inc. PO Box 127, Roebling, NJ 08554; Phone (800)
808-6387; www.EnduranceNetInc.com; email: EnduranceNet@aol.com
5. For more information about CDC permits contact: Department of Health & Human Services,
Public Health Service, Center for Disease Control, Office of Biosafety, Atlanta GA 30333. Phone:
(404) 329-3883.
6. Microbat shipping information was provided by Dr. John J. Rasweiler, The New York Hospital -
Cornell Medical Center.
7. In addition, if an animal is not adapting to captivity, the possibility of releasing them remains.
8. Ideally, samples should be collected from individuals. In large collections, however, a survey
sample number of specimens should be collected.
9. Lethargy and reluctance to move can also be caused by torpor, which is normal for
chiropterans.
10. Many factors affect heart rate. It is necessary to calculate respiration for each species. Both
heart rate and respiration vary with body size.
11. Gauntlet gloves & capture equipment can be obtained through Fuhrman Diversified, 2912
Bayport Blvd., Seabrook, TX 77586-1501. Phone: (281) 474-1388; email: fdi@flash.net
12. It is impractical to attempt to examine neonates of large colonies. In addition, it may lead to
mother bats abandoning their infants. This process should only be undertaken when deemed
necessary in order to gather specific scientific data for a research program.
13. In addition, some researchers believe that Pteropus pumilus may show lek behavior (R.
Spears, pen. comm.). However, there is little evidence to support this claim.
14. Some fruit bat rehabilitators believe that soy-based milk can be lethal to flying foxes (H.
Luckoff, pers. corn.)
15. Additional information can be obtained by writing to Susan Barnard, 6146 Fieldcrest Dr.
Morrow, GA 30260; Helen George, Wombaroo Food Products, ass Vale Road, Beaumont,
Australia 2577. Phone: (044) 65- 1328 and/or Helen Luckhoff, 52 Cardiff Road, Darru,
Queensland, Australia 4076.
Teats for handrearing flying foxes can be obtained from: S. Grzegorski, Australian Wildlife Teats,
60 Sutherland St., Calliope, Australia, 4680.
16. Trovan transponders can be obtained by contacting Infopet Identification Systems, Inc., 415
W. Travelers Trail, Burnsville, MN 55337, Phone: (612) 890-2080.
AVID transponders can be obtained by contacting AVID Identification Systems, Inc., 3185 Hamner
Ave, Norco, CA 92860-1983, Phone: (951) 371-7505
17. One recommended brand is L&M Bird Leg Bands, Inc. PO Box 2636, San Bernadino, CA
92406, Phone: (909) 882-4649; another is Gey Band & Tag Co, Inc. (now National Band & Tag
Co.) PO Box 72430, Newport, KY 41072-0430; Phone: (859) 261-2035; email:
tags@nationalband.com; another is A.C. Hughes, 1 High Street, Hampton Hill, Greater London
TW12 lNA, U.K. Phone: (081) 797- 1366.

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(Sundevall), with special reference to male calling. Z. Tierpsychol., 40: 345-376.
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Definitions
Animal welfare:
The degree to which an animal can cope with challenges in its environment as determined by a
combination of measures of health (including pre-clinical physiological responses) and measures
of psychological well-being:
 Good health represents the absence of diseases or physical and physiological conditions
that result (directly or indirectly) from inadequate nutrition social conditions, or other
environmental conditions to which an animal fails to cope successfully.
 Psychological well-being is dependent on there being the opportunity for animals to perform
strongly motivated behaviours, especially those that arise in response to aversive stimuli.
 Enhanced psychological well-being is conditional on the choices animal have to respond
appropriately to variable environmental conditions, physiological states, developmental
stages and social situations, and the extent to which they can develop and use their
cognitive abilities.
Stress:
a) A non-specific response of the body to any demand made on it; b) an environmental effect that
is likely to or does reduce the Darwinian fitness of the organism; c) the alarm system in a
homeostatic organism, d) a cumulative response, the result of an animal’s interaction with its
environment.
Eustress:
The outcome when a stressor is met by effective coping strategies and transformed thereby from
distress (negative) to eustress (positive).
Distress. The state of being unable to defend against or control effects of an acute or chronic
environmental stressor. A stressor may be either physical or emotional.
Suggested Readings
Beckoff, M., ed. 1998 Encyclopedia of Animal Rights and Animal Welfare. Greenwood Press,
Westport, Conneticut.
Gibbons, Edward F., Jr., Barbara S. Durrant, and Jack Demarest, eds. 1995. Conservation of
Endangered Species in Captivity: An Interdisciplinary Approach. State University of New York
Press, Albany, New York.
Kleiman, Devra G., Mary E. Allen, Katerina V. Thompson, Susan Lumpkin, eds. 1996 Wild
Mammals in Captivity: Principles and Techniques. University of Chicago Press, Chicago, Illinois.

Moller, Anders P., Manfred Milinski, Peter J. B. Slater, eds. 1998. Stress and Behavior,
Advances in the Study of Behavior, vol. 27. Academic Press, San Diego, California.
Norton, Bryan G., Michael Hutchins, Elizabeth F. Stevens, and Terry L. Maple, eds. 1995. Ethics
on the Ark. Smithsonian Institution Press, Washington, D. C.
Shepherdson, David J., Jill D. Mellen, Michael Hutchins, eds. 1999. Second Nature:
Environmental Enrichment for Captive Animals. Smithsonian Institution Press, Washington, D. C.
Stanford, S. Clare, Peter Salmon, eds. 1993. Stress: From Synapse to Syndrome. Academic
Press, San Diego, California.
Veterinary Standards Committee, American Association of Zoo Veterinarians. 1999. Guidelines

for Zoo and Aquarium Veterinary Medical Programs and Animal Hospitals.
APPENDIX A
Need to Update: List of Participants
1994 &/or 1995 AZA Chiropteran TAG Mid-year Meetings
The Lubee Foundation, Gainesville, Florida
Keith Atkinson, The Lubee Foundation

Bruce Bohmke, Phoenix Zoo
Bruce Brewer, Brookfield Zoo
Bryan Carroll, Jersey Wildlife Preservation Trust
Michel Delorme, Biodome De Montreal
Jan Dempsey, St. Louis Zoo
Ellen Dierenfeld, Wildlife Conservation Park/ Bronx Zoo
Nina Fascione, Defenders of Wildlife
Adriana Galindo, University of Florida
Tim Gross, University of Florida
Roger Haagenson, Lubee Foundation
Darryl Heard, University of Florida
Scott Heinrichs, Lincoln Park Zoo
Karen Killmar, San Diego Zoo
Tom Kunz, Boston University
Norma Lewis, Birmingham Zoo
Dan Moms, Omaha's Henry Door1 y Zoo
Terry O’Connor, Woodland Park Zoo
Dennis Pate, Metro Washington Park Zoo
Harry Reeve, Busch Gardens Tampa
Peter Riger, Wildlife Conservation Park/Bronx Zoo
Alan Rost, Jacksonville Zoo
John Seyjagat, The Lubee Foundation
Kim Tropea, The Wildlife Conservation Park/Bronx Zoo
Kim Whitman, Philadelphia Zoological Garden
Steven Wing, Folsom Children's Zoo
Roland Wirth, Munchen, Germany
Husbandry Manual Editors (in addition to those at the TAG Meetings):

Sue Barnard, Zoo Atlanta
Sue Crissey, Brookfield Zoo
Brock Fenton, York University
Peter Myroniuk, Melbourne Zoo
Dominic Wormell, Jersey Wildlife Preservation Trust

Janette Young, Jersey Wildlife Preservation Trust
APPENDIX B
PARTIAL ETHOGRAM OF BAT BEHAVIORS
(from Carroll, 1979)
Autogrooming: licking or nibbling the fur or scratching with one hind foot. Forelimbs are not used
in grooming.
Allogrooming: seen quite frequently among all sex/age classes, although adult males confine
allogrooming to adult females. Involves groomer licking groomee's fur.
Hanging Alert: animal is hanging stationary, but is awake and alert.
Play Chase: seen among immature. Two or more animals fly from one location to another and
rapidly leave again.
Play Wrestle: usually seen among immatures. Involves close ventral contact between individuals
while giving inhibited bites to neck area of the other animal. More than two animals may be
involved.
Roosting: animal is asleep, suspended by one or both feet from a perch.
Scent Marking: rubbing neck and chin glands against perches or the walls of cage. Seen most
frequently in adult males.
Static Flying: flapping the wings as if to fly but retaining hold of the perch with the feet. Seen
mainly among infants who cannot yet fly.
Walking on the Ground: animal will descend to ground to search for fallen food. Limb sequence
is typical for quadrupedal gait and claw of the thumb is used to pull the animal along. While is
may be difficult for some species, bats can take off from the ground.
Wing Fan: seen during hanging alert. Is a gentle fanning with half-folded wing, may be
thermoregulatory.
Wing Shake: aggressive action. One or both wings held with arms outstretched from the body.
Wings shaken with thumb pointing at addressee. Claw can be used in physical encounters and is
capable of inflicting injury.
Wing Flick: short flick of the distal half of wing achieved by moving the digits. Seen sometimes
during courtship by males.

APPENDIX C

Shipping Container: Vari-Kennek (Doskocil) with a wire mesh cage
Large- Pteropus vampyrus (39”L x 26.5”W x 28.5”H)
Medium- Pteropus rodricensis (27”L x 20”W x 19”H)
Small- (19”L x 12”W x 10”H)

APPENDIX E

APPENDIX F
Partial List of Forages Accepted by Captive Colonies of Fruit Bats

alder
ash
aspen
bamboo
banana
beech
black locust
box elder
crabapple
dogwoods
elm
ficus
forsythia
grape
hackberry
honeysuckle (Japanese)
Kentucky coffee tree
kudzu
mangrove (Florida)
maple (not red)**
mulberry (not red)
ornamental pear
poplar (not tulip poplar)
sweetgum
torch ginger
** Although red maple has been fed in some institutions, there have been reports of health
incidents possibly related to the ingestion of red maple. Therefore, it is not recommended.

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Standardized Guidelines for Fruit and Nectar Bat Care

Cynopterus brachyotis Dog-faced fruit bat 19.25.0

1. Abiotic Environmental Variables (address both exhibit and off-exhibit holding)

In outdoor enclosures, supplemental heat is needed for temperatures below 70 degrees

1 18th October 2004

2 18th October 2004

1.4.2. For social species indicate minimum inter-individual

3 18th October 2004

1.4.3. Identify appropriate furnishings to accommodate an array of

4 18th October 2004

1.4.5. Identify appropriate substrates and nesting/bedding materials

1.4.7. Address issues, such as scent marking, that may influence

5 18th October 2004

There are no special concerns or requirements for sewage disposal; however,

1.4.8 Identify number of air or water changes/hour required

1.4.9 Identify necessary measures for safety and containment.

1.4.10 Address issue of transport, identifying (in accordance with IATA)

6 18th October 2004

1.4.10.1 Type of transport container:

1. Megabats. Pteropus species must be transported in double cages. A vari-kennel with an

2. Microbats.6 Microbats such as Glossophaga soricina, Carollia perspicillata, and Artibeus

1.4.10.2 Appropriate size of transport container

7 18th October 2004

1.4.10.3 Provision of food and water during transport

1.4.10.4 Provision of bedding or substrate in transport container

1.4.10.6 Identify appropriate temperature range during transport

1.4.10.8 Address appropriate group size or need for separation of individuals

1.4.10.9 Consider need for handler/veterinarian access to animal during transport

8 18th October 2004

1.4.10.11 Address appropriate timing of release, size and type of enclosure at

9 18th October 2004

2.1.2. Identify appropriate foodstuffs (see below) and feeding schedules

2.1.4. Address opportunities for animals to process food in ways similar to

2.2 Social Considerations

10 18th October 2004

2.2.1.2. temporary isolation of parturient females and young, or of

2.2.1.3. Seasonal separation of sexes, or for those species that are

2.2.1.4. nursery groups (groups of mothers with most recent young)

11 18th October 2004

2.2.1.6. multigenerational groups (e.g. many primates, elephants)

2.2.1.7. groups deriving from cohorts (e.g., dolphin male pairs)

2.2.1.8. all male groups

2.2.1.9. daily and life stage variation in patterns of social affiliation

12 18th October 2004

2.2.3. Conspecific groups, consider the need for/influence of adjacent

2.2.4. Mixed species groups

13 18th October 2004

2.2.6. Human-animal interactions

2.2.6.2. Address both animal and keeper safety

14 18th October 2004

2. Histoplasmosis: Histoplasmosis (Histoplasmosis capsulatum) is an airborne fungus

3. Health and Nutrition

15 18th October 2004

Digestive Tract Morphology and Physiology

Nutrient Content of Foods

16 18th October 2004

A Data from Pennington and Church.60

Fruit Type Moisture Ca P Fe Se

A Data from Pennington and Church.60

17 18th October 2004

Protein and amino acids

18 18th October 2004

19 18th October 2004