Rousettus aegyptiacusEgyptian rousette

Ge­o­graphic Range

Rouset­tus ae­gyp­ti­a­cus is found through­out Africa south of the Sa­hara, in Egypt, and on the coast­lines of the Ara­bian Penin­sula (Grz­imek, 2003). Egypt­ian rousettes are most com­mon from lat­i­tudes 15 de­grees north through 37 de­grees south. How­ever, they have been found as far as 40 de­grees north in south­ern Turkey. They are also found on each of the Ca­nary Is­lands, west­ern North Africa, and through­out the Gulf of Guinea (No­gales et al., 2006; Nowak, 1999). (Grz­imek, 2003; No­gales, et al., 2006; Nowak, 1999)

Habi­tat

Egypt­ian rousettes are found through­out Africa’s trop­i­cal rain forests, trop­i­cal de­cid­u­ous forests, sa­vanna, and Turkey’s Mediter­ranean scrub forests. Egypt­ian rousettes have been found in arid bio­mes; how­ever, they pre­fer to re­main in habi­tats that pro­vide for­est cover, roost­ing op­por­tu­ni­ties, and abun­dant fruit tree growth (Kwiecin­ski and Grif­fiths, 1999). (Kwiecin­ski and Grif­fiths, 1999)

  • Other Habitat Features
  • caves
  • Range elevation
    0 to 4000 m
    0.00 to 13123.36 ft

Phys­i­cal De­scrip­tion

Egypt­ian rousettes are medium sized bats with dor­sal pelage rang­ing from dark brown to medium gray. Ven­tral pelage in both gen­ders is sev­eral shades lighter than dor­sal col­oration, with a col­lar of pale yel­low or or­ange fur often seen around the neck. There is no color dif­fer­ence be­tween gen­ders; how­ever, males have well-de­vel­oped stiff hairs along the throat that are more rec­og­niz­able than in fe­males (Kwiecin­ski and Grif­fiths, 1999). Short fur com­pletely cov­ers the head al­most to the end of the muz­zle, with the ex­cep­tion of the fore­head, where the fur is slightly longer. Ears are around the length of the muz­zle, with blunt tips and dark col­oration when com­pared to dor­sal pelage. Egypt­ian fruit bats have large eyes adapted for twi­light and night vi­sion. Wing mem­branes are dark brown with short fur ex­tend­ing to the prox­i­mal half of the fore­arm. A claw is pre­sent on both the first and sec­ond dig­its, while all other dig­its have car­ti­lage. Egypt­ian rousettes have five toes on both hind limbs, each with claws (Kwiecin­ski and Grif­fiths, 1999).

Males are typ­i­cally larger than fe­males with a total body length rang­ing from 14 to 19.2 cm, while fe­males range from 12.1 to 16.7 cm. Adults may weigh 80 to 170 g and have a wingspan close to 60 cm. The fore­arm varies be­tween 85 to 101.9 mm in males and 88.1 to 99 mm in fe­males (Kwiecin­ski and Grif­fiths, 1999; Grz­imek, 2003). (Grz­imek, 2003; Kwiecin­ski and Grif­fiths, 1999)

  • Sexual Dimorphism
  • male larger
  • Range mass
    80 to 170 g
    2.82 to 5.99 oz
  • Range length
    12.1 to 19.2 cm
    4.76 to 7.56 in
  • Average wingspan
    60 cm
    23.62 in

Re­pro­duc­tion

Rouset­tus ae­gyp­ti­a­cus is a polyg­a­mous species, with bian­nual breed­ing sea­sons. How­ever, in­stances of mo­noestry have been recorded at higher lat­i­tudes (Bay­demir and Al­bayrak, 2006; Bernard and Cum­ming, 1997). Egypt­ian rousettes ob­served in lat­i­tudes south of Egypt have two dis­tinct peak breed­ing sea­sons sep­a­rated by short pe­ri­ods. The first peak breed­ing sea­son is from spring through sum­mer, the sec­ond is fall through win­ter. Births occur to­wards the end of each breed­ing sea­son (Okia, 1987). (Bay­demir and Al­bayrak, 2006; Bernard and Cum­ming, 1997; Okia, 1987)

Fe­males give birth to and raise a sin­gle pup with each breed­ing ef­fort, though twins have been recorded oc­ca­sion­ally (Ko­rine et al., 1994). Es­ti­mated ges­ta­tion length is 3.5 to 4 months, with syn­chro­nized births oc­cur­ring in breed­ing colonies (Kwiecin­ski and Grif­fiths, 1999; Okia, 1987). New­born bats are born with de­cid­u­ous teeth and are al­tri­cial - com­pletely naked apart from a thin layer of down along the head and back. Pups are weaned at 6 to 10 weeks with no dif­fer­ence in growth be­tween sexes. Young reach full adult weight and size around 9 months old, about the same time young be­come in­de­pen­dent from their moth­ers (Kwiecin­ski and Grif­fiths, 1999). Sex­ual ma­tu­rity is reached on av­er­age at 15 months, how­ever, fe­males have been noted to be sex­u­ally ma­ture at 5 to 7 months ("Egypt­ian Fruit Bat", 2006). ("Egypt­ian Fruit Bat", 2006; Ko­rine, et al., 1994; Kwiecin­ski and Grif­fiths, 1999; Okia, 1987)

  • Breeding interval
    Breeding occurs twice yearly throughout most of their range, although one breeding effort per season may occur in northern latitudes.
  • Breeding season
    Breeding occurs typically from April to August and October to February
  • Range number of offspring
    1 to 2
  • Average number of offspring
    1
  • Range gestation period
    3.5 to 4 months
  • Average gestation period
    4 months
  • Range weaning age
    6 to 10 weeks
  • Range time to independence
    8 to 9 months
  • Average time to independence
    9 months
  • Range age at sexual or reproductive maturity (female)
    5 to 16 months
  • Average age at sexual or reproductive maturity (female)
    15 months
  • Range age at sexual or reproductive maturity (male)
    14 to 18 months
  • Average age at sexual or reproductive maturity (male)
    15 months

After birth Egypt­ian rousette pups are cared for solely by their moth­ers for around 9 months. They are ca­pa­ble of fly­ing at 63 to 70 days after birth and stay with their mother until they have reached adult weight and size. Once ma­ture, males leave ma­ter­nity colonies to join bach­e­lor groups, fe­males join ma­ter­nity colonies (Kwiecin­ski and Grif­fiths, 1999). (Kwiecin­ski and Grif­fiths, 1999)

  • Parental Investment
  • altricial
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female

Lifes­pan/Longevity

The recorded max­i­mum lifes­pan of Rouset­tus ae­gyp­ti­a­cus is 22 years in the wild, with a max­i­mum of 25 years in cap­tiv­ity for both males and fe­males. In wild pop­u­la­tions, av­er­age lifes­pan is typ­i­cally 8 to 10 years due to fac­tors such as pre­da­tion, vi­t­a­min D de­fi­ciency, cal­cium-phos­pho­rus im­bal­ance, and os­teo­pro­lif­er­a­tion. Longevity in cap­tive bats is also de­pen­dent on en­vi­ron­men­tal tem­per­a­tures and neu­ro­log­i­cal im­pair­ment (Kwiecin­ski and Grif­fiths, 1999). (Kwiecin­ski and Grif­fiths, 1999)

  • Average lifespan
    Status: wild
    9 years
  • Range lifespan
    Status: captivity
    25 (high) years
  • Average lifespan
    Status: captivity
    22 years
  • Typical lifespan
    Status: wild
    8 to 10 years
  • Typical lifespan
    Status: captivity
    25 (high) years
  • Average lifespan
    Status: captivity
    22 years

Be­hav­ior

Egypt­ian rousettes are found in groups of var­i­ous sizes, rang­ing from small colonies of 20 to 40 mem­bers to large colonies of 9,000. These colonies pre­fer to roost dur­ing the day in dark, slightly humid en­vi­ron­ments such as cave sys­tems and ruins, though small colonies have been seen roost­ing in trees (Grz­imek, 2003). Dur­ing breed­ing sea­sons, males and fe­males sep­a­rate; males form­ing bach­e­lor groups and fe­males form­ing ma­ter­nity colonies. Egypt­ian rousettes roost in close con­tact with other mem­bers of the colony to re­duce the in­flu­ence of tem­per­a­ture fluc­tu­a­tions. Close con­tact among mem­bers of the colony also al­lows com­mu­ni­ca­tion with each other through­out the day dur­ing roost­ing. Egypt­ian rousettes are more ac­tive dur­ing the late af­ter­noon and into the night, when groom­ing is more fre­quent (Kwiecin­ski and Grif­fiths, 1999). In­di­vid­u­als leave the day roost close to sun­set each evening to for­age, re­turn­ing be­fore sun­rise. Dur­ing sum­mer months longer daylength de­lays emer­gence from the roost, sug­gest­ing a cir­ca­dian rhythm syn­chro­nized to light-dark cy­cles. Due to the change in light in­ten­sity, Egypt­ian rousettes have dif­fer­ent pat­terns of ac­tiv­ity be­tween sum­mer and win­ter sea­sons. Sum­mer sea­sons are char­ac­ter­ized by longer feed­ing in­ter­vals than win­ter sea­sons, de­spite re­duced dark pe­ri­ods and de­layed roost emer­gence (Kwiecin­ski and Grif­fiths, 1999). (Grz­imek, 2003; Kwiecin­ski and Grif­fiths, 1999)

Home Range

Home range varies in Rouset­tus ae­gyp­ti­a­cus. This species has been noted to mi­grate south­ward in the north­ern re­gions of their ge­o­graphic range and for­age great dis­tances away from day roosts. Males ap­pear to de­fend ter­ri­to­ries, but their size has not been char­ac­ter­ized (Kwiecin­ski and Grif­fiths, 1999). (Kwiecin­ski and Grif­fiths, 1999)

Com­mu­ni­ca­tion and Per­cep­tion

Egypt­ian rousettes are one of 3 mem­bers of the Rouset­tus genus to use both vi­sual ori­en­ta­tion and echolo­ca­tion. Echolo­ca­tion in this species is pro­duced through a se­ries of crude, short clicks of the tongue against the side of the mouth (Roberts, 1975; Hol­land et al., 2004). These short, im­pul­sive-paired clicks as­sist in nav­i­ga­tion in the dark. The fre­quency range is usu­ally 12 to 70 kHz, with click struc­ture and du­ra­tion most sim­i­lar to dol­phins. This form of echolo­ca­tion has evolved in­de­pen­dently from the echolo­ca­tion sys­tem used by other echolo­cat­ing bats, such as ves­per­til­ion­ids (Roberts, 1975; Hol­land et al., 2004; Hol­land and Wa­ters, 2007). (Hol­land and Wa­ters, 2007; Hol­land, et al., 2004; Roberts, 1975)

Food Habits

Egypt­ian rousettes are fru­gi­vores, pre­fer­ring to eat the pulp and juice of very ripe fruit. They typ­i­cally take fruit from trees, such as lilac, mul­berry, carob, sycamore, fig, and baobob (Kwiecin­ski and Grif­fiths, 1999; Ko­rine et al., 1996; Grz­imek, 2003). Egypt­ian rousettes con­sume 50 to 150% of total body mass in fruit each night. After se­lect­ing fruit, Egypt­ian rousettes take their fruit and roost close to the feed­ing tree. Only the pulp and juice is con­sumed; after thor­ough chew­ing, the seeds are spit out. Egypt­ian rousettes feed while hold­ing the fruit close to the body, as a means of pro­tect­ing the fruit from being pil­fered by other bats. Steal­ing fruit is com­mon, and these bats are ag­gres­sive when feed­ing (Kwiecin­ski and Grif­fiths, 1999). (Grz­imek, 2003; Ko­rine, et al., 1996; Kwiecin­ski and Grif­fiths, 1999)

  • Plant Foods
  • fruit

Pre­da­tion

Be­cause of their fru­giv­o­rous diet, Egypt­ian rousettes are some­times con­sid­ered a threat to fruit crops dur­ing the grow­ing sea­son. Plan­ta­tion own­ers and farm­ers often hunt Egypt­ian rousettes or hire bounty hunters dur­ing the flow­er­ing and fruit­ing sea­sons to re­duce the risk of crop dam­age and profit loss (Fu­jita and Tut­tle, 1991). Egypt­ian rousettes, like many mem­bers of the Fam­ily Pteropo­di­dae, are hunted for meat as well. As a re­sult roost pop­u­la­tion den­si­ties have de­clined (Fu­jita and Tut­tle, 1991). Nat­ural preda­tors have not been de­scribed but are likely to in­clude aer­ial preda­tors, such as fal­cons, and cave preda­tors, such as snakes and mustelids. Their noc­tur­nal habits, cave roost­ing, flight, and cryp­tic col­oration all help them to avoid pre­da­tion. (Fu­jita and Tut­tle, 1991)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

Egypt­ian rousettes are pol­li­na­tors of many noc­tur­nally flow­er­ing trees through­out pa­le­otrop­i­cal forests. Due to their fru­giv­o­rous diets, they not only pol­li­nate the flow­ers of fruit trees but also as­sist as the pri­mary agent in seed dis­per­sal for many tree species (Fu­jita and Tut­tle, 1991). Egypt­ian rousettes are car­ri­ers for both ec­topar­a­sites and en­dopar­a­sites through­out their ge­o­graphic range. Ec­topar­a­sites that use Egypt­ian rousettes as hosts in­clude: Spin­turnix lat­er­alis, An­cy­s­tro­pus lele­upi, An­cy­s­tro­pus zele­bori, An­cy­s­tro­pus lat­er­alis, An­cy­s­tro­pus zele­borii, Liponys­sus gluti­nosus, Liponys­sus longi­manus, Eu­camp­sipoda africanum, Nyc­teri­bosca africana, Brachy­tarsina al­lu­audi, Eu­camp­sipoda hyrtlii, Nyc­teri­bosca di­versa, Thaumap­sylla bre­vi­cep, Nyc­teribia pedic­u­laria, Nyc­teribia schmidlii, Eu­camp­sipoda africana, Afrocimex lele­upi, Thaumap­sylla bre­vi­ceps, Ar­chaeop­sylla met­allescens, and Alec­toro­bius cam­i­casi (Kwiecin­ski and Grif­fiths, 1999). En­dopar­a­sites that are found in Rouset­tus ae­gyp­ti­a­cus in­clude the he­mo­sporid­ian Plas­mod­ium rous­setti and the round­worm Nyc­teri­do­coptes rousetti. Out­breaks of ra­bies are also found in pop­u­la­tions of Egypt­ian fruit bats (Kwiecin­ski and Grif­fiths, 1999). (Fu­jita and Tut­tle, 1991; Kwiecin­ski and Grif­fiths, 1999)

  • Ecosystem Impact
  • disperses seeds
  • pollinates
Mu­tu­al­ist Species
  • com­mon lilacs (Sy­ringa vul­garis)
  • mul­berry species (Morus)
  • lo­quats (Eri­obotrya japon­ica)
  • carob (Cer­a­to­nia sili­qua)
  • fig species (Ficus)
Com­men­sal/Par­a­sitic Species
  • stre­blid flies (Nyc­teri­bosca species)
  • sar­cop­tid mites (Nyc­teri­do­coptes rousetti)
  • malar­ial plas­mod­ium (Plas­mod­ium rous­setti)
  • par­a­sitic mites (Spin­turnix lat­er­alis)
  • par­a­sitic mites (An­cy­s­tro­pus species)
  • par­a­sitic mites (Liponys­sus species)
  • par­a­sitic flies (Eu­camp­sipoda species)
  • bat bugs (Afrocimex lele­upi)
  • fleas (Thaumap­sylla bre­vi­ceps)
  • fleas (Ar­chaeop­sylla met­allescens)
  • par­a­sitic mites (Alec­toro­bius cam­i­casi)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Through­out their range, Egypt­ian rousettes pol­li­nate many com­mer­cially valu­able fruit trees. This species also works to dis­perse seeds away from the par­ent plant, al­low­ing seed ger­mi­na­tion and spread in un­pre­dictable con­di­tions (Fu­jita and Tut­tle, 1991; Izhaki et al., 1995). (Fu­jita and Tut­tle, 1991; Izhaki, et al., 1995)

  • Positive Impacts
  • food
  • pollinates crops

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Rouset­tus ae­gyp­ti­a­cus is per­ceived by many fruit farm­ers to be a pest of fruit crops grown com­mer­cially for human pro­duc­tion through­out its range, though lit­tle ev­i­dence sup­ports this and cases are often ex­ag­ger­ated (Al­bayrak et al., 2008). (Al­bayrak, et al., 2008)

Con­ser­va­tion Sta­tus

Rouset­tus ae­gyp­ti­a­cus is con­sid­ered a species of least con­cern by the IUCN. How­ever, due to poach­ing and a neg­a­tive in­ter­ac­tions with com­mer­cial farm­ing, there has been a de­cline in roost­ing sites (Al­bayrak et al., 2008). (Al­bayrak, et al., 2008)

Con­trib­u­tors

Rachel Cohen (au­thor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor), Rad­ford Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

echolocation

The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

rainforest

rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.

scrub forest

scrub forests develop in areas that experience dry seasons.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

Ref­er­ences

2006. "Egypt­ian Fruit Bat" (On-line). Rosa­mond Gif­ford Zoo. Ac­cessed April 20, 2010 at http://​www.​rosamondgiffordzoo.​org/​assets/​uploads/​animals/​pdf/​EgyptianFruitBat.​pdf.

Al­bayrak, I., N. Asan, T. Yorul­maz. 2008. The Nat­ural His­tory of the Egypt­ian Fruit Bat, Rouset­tus ae­gyp­ti­a­cus, in Turkey (Mam­malia: Chi­roptera). Turk­ish Jour­nal of Zo­ol­ogy, 32: 11-18.

Bay­demir, N., I. Al­bayrak. 2006. A Study on the Breed­ing Bi­ol­ogy of Some Bat Species in Turkey (Mam­malia: Chi­roptera). Turk­ish Jour­nal of Zo­ol­ogy, 30: 103-110.

Bernard, R., G. Cum­ming. 1997. African Bats: Evo­lu­tion of Re­pro­duc­tive Pat­terns and De­lays. The Quar­terly Re­view of Bi­ol­ogy, 72: 253-274.

Fu­jita, M., M. Tut­tle. 1991. Fly­ing Foxes (Chi­roptera: Pteropo­di­dae): Threat­ened An­i­mals of Key Eco­log­i­cal and Eco­nomic Im­por­tance. Con­ser­va­tion Bi­ol­ogy, 5: 455-463.

Grz­imek, B. 2003. Grz­imek's An­i­mal Life En­cy­clo­pe­dia. Farm­ing Hills, Michi­gan: Gale Vir­tual Li­brary.

Hol­land, R., D. Wa­ters. 2007. The Ef­fect of Fa­mil­iar­ity on Echolo­ca­tion in the Megachi­ropteran Bat Rouset­tus ae­gyp­ti­a­cus. Be­hav­ior, 144: 1053-1064.

Hol­land, R., D. Wa­ters, J. Rayner. 2004. Echolo­ca­tion Sig­nal Struc­ture in the Megachi­ropteran Bat Rouset­tus ae­gyp­ti­a­cus Ge­of­frey 1810. The Jour­nal of Ex­per­i­men­tal Bi­ol­ogy, 207: 4361-4369.

Izhaki, I., C. Ko­rine, Z. Arad. 1995. The Ef­fect of Bat (Rouset­tus ae­gyp­ti­a­cus) Dis­per­sal on Seed Ger­mi­na­tion in East­ern Mediter­ranean Habi­tats. Oe­colo­gia, 101: 335-342.

Ko­rine, C., Z. Arad, A. Arieli. 1996. Ni­tro­gen and En­ergy Bal­ance of the Fruit Bat Rouset­tus ae­gypt­la­cus on Nat­ural Fruit Diets. Phys­i­o­log­i­cal Zo­ol­ogy, 69: 618-634.

Ko­rine, C., I. Izhaki, D. Makin. 1994. Pop­u­la­tion Struc­ture and Emer­gence Order in the Fruit-bat (Rouset­tus aeyyp­ti­a­cus: Mam­malia, Chi­roptera). The Zo­o­log­i­cal So­ci­ety of Lon­don, 232: 163-174.

Kwiecin­ski, G., T. Grif­fiths. 1999. Rouset­tus egyp­ti­a­cus. Mam­malian Species, 611: 1-9.

No­gales, M., J. Ro­driguez-Lu­engo, P. Mar­rero. 2006. Eco­log­i­cal Ef­fects and Dis­tri­b­u­tion of In­va­sive Non-Na­tive Mam­mals on the Ca­nary Is­lands. Mam­mal Re­view, 36: 49-65.

Nowak, R. 1999. Walker's mam­mals of the world. Bal­ti­more: John Hp­kins Uni­ver­sity Press.

Okia, N. 1987. Re­pro­duc­tive Cy­cles of East African Bats. Jour­nal of Mam­mal­ogy, 68: 138-141.

Roberts, L. 1975. Con­fir­ma­tion of the Echolo­ca­tion Pulse Pro­duc­tion Mech­a­nism of Rouset­tus. Jour­nal of Mam­mal­ogy, 56: 218-220.