Professional Documents
Culture Documents
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
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
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:
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)
2. Large bats with a wingspread from 2.5 – 3.5 feet (300 – 800 grams)
a. For up to 6 animals, an enclosure 11 feet by 21 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 20 percent of the original floor area.
3. Intermediate bats with a wingspread from 2 – 2.5 feet (200 - 350 grams)
a. For up to 6 animals, an enclosure 8 feet by 15 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 20 percent of the original floor area.
4. Medium bats with a wingspread from 1.5 - 2 feet (80 - 200 grams)
a. For up to 10 animals, an enclosure 6 feet by 12 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.
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
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.
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.
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.
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.
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.
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.
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
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.
1.4.10.5 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.
1.4.10.7 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.
1.5 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.
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.
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.
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.
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.
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:
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.
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:
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.
Section adapted from Dempsey, J. 2004. Fruit bats: Nutrition and dietary husbandry. Fact
Sheet 014. Nutrition Advisory Group Handbook.
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
Table 2. Comparison of the mineral content of cultivated and native fruits. Expressed on a
dry matter basis.
Nutrient Requirements
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.
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
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
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
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.
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.
* 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.
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.
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,
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.
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.
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.
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.
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.
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.
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.
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.
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.
Refer to Appendix F for a partial list of forages accepted by captive colonies of fruit bats.
6. Documentation
ENDNOTES (for Nutrition section 3.1 adapted from Nutrition Advisory Group Handbook)
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3 Ayaz K.M., E.C. Birney, and R. Jennes. 1976. Inability of bats to synthesize L-ascorbic acid.
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4 Bonaccorso, F.J. 1978. Foraging and reproductive ecology in a Panamanian bat community.
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9 Conklin-Brittain, N.L., E.S. Dierenfeld, R.W. Wrangham, M. Norconk, and S.C. Silver. 1999.
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26 Herbst, L.H. 1986. The role of nitrogen from fruit pulp in the nutrition of the frugivorous bat
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27 Herrera, L.G., E. Gutierrez, K.A. Hobson, B. Altube, W.G. Diaz, and V. Sanchez-Cordero.
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28 Hill, J.E., and J.D. Smith. 1984. Food habits and feeding. Pp. 60-72 in Bats: A Natural History.
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29 Iudica, C.A., F.J. Bonaccorso, and G. Richard. 1994. Sea water ingestion in Pteropus
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30 Klite, P.D. 1965. Intestinal bacterial flora and transit time of three neotropical bat species. J.
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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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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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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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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.
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.
APPENDIX A
Need to Update: List of Participants
1994 &/or 1995 AZA Chiropteran TAG Mid-year Meetings
The Lubee Foundation, Gainesville, Florida
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.