Seasonal variation in the reproductive tract of the Indian flying fox, Pteropus giganteus (Brünnich, 1782)

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Seasonal variation in the reproductive tract of Pteropus giganteus was studied at 13 sites of the Pothwar Plateau and Islamabad Capital Territory. Ninety-three dead specimens were collected from roosts, orchards and home-based small cultivations of fruit trees. In males, seasonal differences in testis length, testis width, cauda epididymidis length and penis length were recorded. Positive correlations were found between body weight, body length and all parameters of male reproductive tract. Significant monthly variations in testes conditions (viz. flaccid and turgid) were observed. In females, seasonal variation in reproductive tract weight, ovary length, ovary width, uterus length and uterus width were seen. In females, a strong relationship of body weight with total body length and female reproductive tract parameters were found. In flying foxes’ population of Pothwar, first mating was observed in December-January and second in June-July, suggesting two mating seasons in winter and summer.



AsdellS.A. (1964) Patterns of Mammalian Reproduction. Cornell University Press, New York.

BarclayR.M.R.HarderL.D. (2003) Life histories of bats: life in the slow lane. In: KunzT.H.FentonM.B. (Eds.) Bats Ecology. University of Chicago Press Chicago.

BatesP.J.J.HarrisonD.L. (1997) Bats of the Indian Subcontinent. Harrison Zoological Museum Publications, 258 pp.

BradburyJ.W. (1977) Lek mating behavior in hammer headed bat. Zeitscrift für Tierpsychologie, 45, 225-255.

CrichtonE.G.KrutzschP.H. (Eds.) (2000) Reproductive Biology of Bats. Academic Press, San Diego, CA, 528 pp.

de JongC.E.JonssonN.N.FieldH.SmithC.CrichtonE.C.PhilipsN.JohnstonS.D. (2005) Collection, seminal characteristics and chilled storage of spermatozoa from three species of free-range flying fox (Pteropus spp.). Theriogenology, 64, 1072-1089.

DeshmukhG.D.DhamaniA.A. (2011) Seasonal testicular changes during the annual reproductive cycle of Indian flying fox Pteropus giganteus giganteus (Brunnich). J. Cell and tissue Research, 11(3), 2943-2948.

DolbeerR.R.FiedlerL.R.RasheedH. (1988) Management of fruit bat and rat populations in the Maldive islands, Indian Ocean. In: A.C. Crabb & R.E. Marsh (Eds.) 13th Proc. Vertebr. Pest Conf., pp. 112-118. University of California, Davis.

DorlikarA.V.DhamaniA.A.ChardeP.N.MohiteA.S. (2013) Morphometric and histoarchitectural changes in the ovary of Pteropus giganteus (Brunnich) during various phases of reproductive cycle. International Journal of Molecular Zoology, 3, 8, DOI:10.5376/ijmz.2013.03.0008.

FalanruwM.V.C. (1988) On the status, reproductive biology and management of fruit bats of Yap. Micronesica, 21, 39-51.

FoxS.SpencerH.GemmaM. (2008) Analysis of twining in flying foxes (Megachiroptera) reveals superfoetation and multiple paternities. Acta Chiropterologica, 10, 271-278.

GrantG.S.BanackS.A. (1999) Harem structure and reproductive behavior of Pterpus tonganus in American Samoa. Australian Mammology, 21, 111-120.

HeidemanP.D. (2000) Environmental regulation of reproduction. In: CrichtonE.G.KrutzschP.H. (Eds.) Reproductive Biology of Bats, pp  469-499. Academic Press, San Diego, CA.

HeithausE.R.FlemingT.H.OpterP.A. (1975) Foraging patterns and resource utilization in seven species of bats in a seasonal tropical forest. Ecology, 56, 716-723.

HoodC.SmithJ. (1989) Sperm storage in a tropical nectar-feeding bat, Macroglossus minimus (Pteropodidae). Journal of Mammalogy, 70, 404-406.

IUCN (2010) Red list of threatened species, Version 2010.4. Available at:

JonesK.E.MaclarnonA. (2001) Bat life histories testing models of mammalian life history evolution. Evolutionary Ecology Research, 3, 465-476.

KingdonJ. (1974) East Affrican Mammals, Vol. II A: insectivores and bats. Academic Press, London, England, 392 pp.

KofronC. (2007) Reproduction of the dusky fruit bat Penthetor lucasi (Pteropodidae) in Brunei, Borneo. Mammalia, 71, 166-171.

KoilrajJ.A.AgoramoorthyG.MarimuthuG. (2001) Copulatory behavior of Indian flying fox Pteropus giganteus. Current Science, 80, 15-16.

KretschmannK.HayesR. (2004) Old World fruit bats I (Pteropus). In: HutchinsM.EvansA.JacksonJ. (Eds.) Animal Life Encyclopedia, Vol. 13: mammals II. 2nd ed., pp.  319-332. Grzimek’s Detroit, Gale.

KrutzschP.H. (1979) Male reproductive patterns in non hibernating bats. J. Reprod. Fert., 1979, 333-344.

KrutzschP.H. (2000) Anatomy, physiology and cyclicity of the male reproductive tract. In: CrichtonE.G.KrutzschP.H. (Eds.) Reproductive Biology of Bats, pp.  91-156. Academic Press, San Diego, CA.

KunzT.H.FentonM.B. (Eds.) (2003) Bat Ecology. University of Chicago Press, IL, 784 pp.

MarshallA.J. (1949) Pregestational changes in the giant fruit bat (Pteropus giganteus), with special reference to an asymmetric endometrial reaction. Proceedings of the Linnean Society of London, 161(1), 26-36.

MarshallA.G. (1985) Old World phytophagus bats (Megachiroptera) and their food plants survey. Zoological Journal of the Linnean Society, 83, 351-369.

MartinL.TowersP.A.McGuckinM.A.LittleL.LuckhoffH.BlackshawA.W. (1987) Reproductive biology of flying-foxes (Chiroptera: Pteropodidae). Aust. Mammal., 10, 115-118.

MathurV.PriyaY.S.KumarH.ElangovanV. (2012) Reproductive behaviour and population dynamics of Indian flying fox Pteropus giganteus. Journal of Threatened Taxa, 7(4), 2699-2704.

MatthewsL.H. (1941) Notes on genitalia and reproduction of African bats. Proceeding of the Zoological Society of London B, 111, 289-346.

McCannC. (1941) Further observations on the flying fox (Pteropus giganteus Brunnich) and the fulvus fruit-bat (Rousettus leschenaultia Desm). J. Bomb. Nat. Hist. Soc., 42, 587-592.

McCleanJ.A.SpeakmanJ.R. (1999) Energy budgets of lactating and non lactating brown long eared bats suggest females use metabolic compensation during lactation. Funct. Ecol., 13, 373-383.

MelvilleD.F.O’BrienG.M.CrichtonE.G.TheilemannP.McKinnonA.JohnstonS.D. (2012) Reproductive seasonality and the effect of the GnRH agonist deslorelin as a contraceptive in captive male black flying-foxes (Pteropus alecto). Theriogenology, 77, 652-661.

MiguelA.RicardoL.G.OmarL.W.EdithH.P.RíosA.RosadoA. (2005) Male reproductive cycle of Mexican big-eared bats, Corynorhinus mexicanus (Chiroptera: vespertilionidae). The Southwestern Naturalist, 50(4), 453-460.

MutereF. (1980) Eidolon helvum Revisited. In: D. Wilson & A. Gardner (Eds.) Proceedings of the Fifth International Bat Res. Conference. Lubbock, Texas, U.S.A., pp. 145-150. Texas Tech. Press.

NelsonJ.E.W. (1963) The biology of the flying-fox (genus Pteropus) in south-eastern Queensland. Ph.D. Thesis. Univ. Queensland, St. Lucia.

NeuweilerG. (1969) Verhaltensbeobachtungen an einer Indischen Flughundkolonie (Pteropus giganteus). Zeitscrift fur Tierpsychologie, 26, 166-199.

NowakR. (1999) Walker’s Mammals of the World. The Johns Hopkins University Press, London.

O’BrienG.M. (1993) Seasonal reproduction in flying foxes, reviewed in the context of other tropical mammals. Reproduction, Fertility and Development, 5(5), 499-521.

O’BrienG.M.NankervisR.F. (1994) Coital behavior of male Pteropus scapulatus (little red flying foxes) in captivity. Physiology & Behavior, 56(30), 471-477.

PiersonE.D.RaineyW.E. (1992) The Biology of flying foxes of genus Pteropus. Biological Reprodution, 90, 1-17.

PuddicombeR. (1981) A behavioral study of the grey headed flying fox Pteropus poliocephalus (Megachiroptera). B.Sc. Honors Thesis. University of New England: Armidale, New South Wales, Australia.

RaceyP.A. (1982) Ecology of bat reproduction. In: KunzT.H. (Ed.) Ecology of Bats, pp.  57-104. Plenum Press, New York, NY.

RaceyP.A.SpeakmanJ.R. (1987) The energy costs of pregnancy and lactation in heterothermic bats. Symp. Zool. Soc. Lond., 57, 107-125.

RaceyP.A.SwiftS.M. (Eds.) (1995) Ecology, evolution and behaviour of bats, Symposia of Zoological Society of London, No. 67. Oxford University Press, London, England, 446 pp.

RatcliffeF.N. (1931) The flying fox (Pteropus) in Australia. Bulletin of CSIRO, 53, 1-80.

RobertsT.J. (1997) Mammals of Pakistan. Revised Ed. Oxford Univ. Press, Karachi.

TidemannC.R.KelsonS.L.JamiesonG. (1997) Flying-fox damage to orchard fruit in Australia – incidence, extent and economic impact. Australian Biologist, 10, 179-186.

TowersP.A.MartinL. (1985) Some aspects of female reproduction in the grey-headed flying fox P. poliocephalus (Megachiroptera: Pteropodidae). Aust. Mammal., 8, 257-263.

VardonM.J.TidemannC.A. (1998) Reproduction, growth and maturity in the black flying-fox, Pteropus alceto (Meagchiroptera: Pteropodidae). Aust. J. Zool., 46, 329-344.

WelbergenJ.A. (2010) Growth, bimaturation, and sexual size dimorphism in wild gray-headed flying foxes (Pteropus poliocephalus). Journal of Mammalogy, 91(1), 38-47.

WelbergenJ.A. (2011) Fit females and fat polygynus males: seasonal body mass changes in the grey-headed flying fox. Oecologia, 165, 629-637.

WilesG.J. (1987) Current research and future management of Marianas fruit bats (Chiroptera:Pteropodidae) on Guam. Aust. Mammal., 10, 93-95.

WimsattW.A. (1979) Reproductive asymmetry and unilateral pregnancy in Chiroptera. Journal of Reproduction and Fertility, 56, 345-357.

WongC.H.TayT.W.MarinaM.T.ZubaidA.KamisA.B. (2002) Reproductive biology of male Cynopterus brachyotis in a fragmented forest and an agriculture habitat in Peinsuala Malaysia. Malaysian Applied Biology, 31, 5-61.

XiaopingZ.HaiyanL.QizhiO.ZhongC. (2007) Wild fulvous fruit bats (Rousettus leschenaultii) exhibit human-like menstrual cycle. Biology of Reproduction, 77, 358-364.


  • Specimen collection sites in four districts of Pothwar plateau.

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  • Illustration of ventral view of flying fox showing various anatomical measurements taken during the study duration. Abbreviations: C., calcar; Cpt., chiropatagium; E.L., ear length; E.Wd., ear width; F.A.L., forearm length; F.Cr., keel of calcar; Fr.L., femur length; H.C.L., hind foot claw length; H.L., hind foot length; Met. II-V, metacarpalia; PhI. II-V, phalanges; Plpt., plagiopatagium; Prpt., protopatagium; T.B.L., total body length; Tb.L., tibia length; Tmp., thumb; U.A.L., upper arm length; Upt., uropatagium; W.S., wing span.

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  • Female and male dead specimens of P. giganteus (A) collected in different seasons and (B) monthly variations in opportunistic sampling.

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  • Monthly variations in body weight (A) and total length (B) of both sexes of P. giganteus.

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  • Mean seasonal variations in (A) ovary length, (B) ovary width, (C) uterus length, (D) uterus width and (E) weight of reproductive tract of P. giganteus.

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  • Mean monthly variations in (A) testicular volume, (B) testis weight, (C) penis length, (D) cauda epididymidis length and (E) prostate gland length of male P. giganteus.

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  • Male reproductive tracts of P. giganteus (A) external view of penis and testes, (B) appearance of prostate gland and (C) seminal vesicles (vascular extended) and cauda epididymidis. This figure is published in colour in the online version.

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