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Many fruit growers see flying-foxes such as the Grey-headed Flying-fox as a threat to their livelihood. Understandably, when large numbers of the animals suddenly enter orchards and cause significant damage, growers start to talk of “population explosions” of the animals. In this paper we describe the seasonality, longevity and low reproductive potential of flying-foxes, and show that rapid expansion of their populations, at rates comparable to those of small rodents, simply is not possible. Sudden incursions of large numbers of flying-foxes into orchards reflect migration of animals into a food source from elsewhere. The “natural” longevity of flying-foxes and low reproductive rate of only one offspring per female per year reflects the environment in which these bats evolved - one in which they suffered a low annual mortality and could afford a low reproductive rate. We show that the major factor which determines the rate at which flying-fox populations increase or decline is mortality, and that an additional annual mortality of 10%, “imposed” (for example by “culling”) upon the animals' “natural” mortality, could lead the entire Grey-headed Flying-fox population into decline. Because of the animals' mobility, localised culling (as in the Melbourne Botanic Gardens) will not solve problems caused by the bats and, if continued, will act as a pteropucidal black hole, attracting to their deaths a continuing stream of animals from far afield. If the community wishes to protect flying-foxes, then it must recognise the genuine concerns of growers for their livelihoods, subsidise non-lethal modes of protecting crops, and provide support for research into cheaper and more effective means of protection.

Andrewartha, H. G. and Birch, L. C. 1954. The Innate Capacity for Increase in Numbers, Pp. 31-54 in The Distribution and Abundance of Animals, The University of Chicago Press, Chicago.
Collins, L. 1999. The impact of drought on the flying-foxes of the NSW North Coast. Australasian Bat Society Newsletter. 12: 31-32.
Dukelow, W. R., Pow, C., Kennedy, J. H. and Martin, L. 1990. Stress effects on late pregnancy in the flying fox Pteropus scapulatus. Zool. Sci. 7: 871-77.
Eby, P. 1995 The Biology and Management of Flying foxes in New South Wales, Species Management Report No. 18. NSW National Parks and Wildlife Service: Hurstville, New South Wales.
Eby, P. 1999. Low reproductive output in Grey-headed Flying-foxes associated with a short period of food scarcity. Australasian Bat Society Newsletter. 14: 17-20.
Eby P. Richards, G; Collins, L. and Parry-Jones, K. 1999. The distribution, abundance and vulnerability to population reduction of a nomadic nectarivore, the Grey-headed Flying-fox Pteropus poliocephalus in New South Wales, during a period of resource concentration. Aust. Zool. 31: 240-43.
Hall, L. S. and Richards, G. C. 1987. Crop protection and management of flying-foxes (Chiroptera: Pteropodidae). Aust. Mamm. 10: 137-39.
Hall, L. S., Martin, L., O'Brien, G., Kelly, R. and Luckhoff, H. 1991. Flying fox populations in crisis in southeast Queensland. Proc. Aust. Mamm. Soc. 37: 18.
Hassell, M. P. 1986. Detecting density dependence. Trends in Ecology and Evolution 1: 90-3.
Leslie, P. H. 1945. On the use of matrices in certain population mathematics. Biometrika 35: 183-212.
Lunney, D. and Moon, C., 1997. Flying foxes and their camps in rainforest remnants of north-east New South Wales. Pp. 247-77 in Australia's Ever Changing Forests III Ed by J. Dargavel. Centre for Resource and Environmental Studies, Australian National University: Canberra.
Martin, L. 1997. What regulates breeding season in Australian Flying foxes (genus Pteropus)? Proc. Aust. Soc. Reprod. Biol. 27: 100. (reprinted, 1999 in Australasian Bat Society Newsletter 12: 9-11).
Martin, L. and Bernard, R. T. F. 2000. Endocrine Regulation of Reproduction in Bats: the Role of Circulating Gonadal Hormones. Pp. 27-64 in Reproductive Biology of Bats, Ed by Crichton, E. G. and Krutzsch, P. H., Academic Press, London.
Martin, L., Towers, P. A., McGuckin, Little, L., Luckhoff, H. and Blackshaw, A. 1987. Reproductive biology of flying-foxes (Chiroptera: Pteropodidae). Aust. Mamm. 10: 115-18.
Martin, L., Kennedy, J. H., Little, L. Luckhoff, H., O'Brien, G. M., Pow, C. S. T., Towers, P. A., Waldon, A. K., and Wang, D. Y. 1996. The reproductive biology of Australian flying-foxes (genus Pteropus). Symposium Zoological Society of London, 67: 167-84.
McIlwee, A. P. and Martin, L. 2002. On the intrinsic capacity for increase of Australian flying-foxes ( Pteropus spp., Megachiroptera). Australian Zoologist 32: 76-100.
McGuckin, M.A, and Blackshaw, A. W. 1987. Seasonal changes in spermatogenesis (including germ cell degeneration) and plasma testosterone concentrations in the Grey-headed fruit bat, Pteropus poliocephalus. Aust. J. Biol. 40: 211-20.
Nelson J. E. 1963. The biology of the flying-fox (genus Pteropus) in south-eastern Queensland. PhD Thesis, The University of Queensland.
Nelson, J. E. 1965. Movements of Australian flying-foxes (Pteropodidae: megachiroptera). Aust. J. Zool. 13: 53-73.
O'Brien, G. M. 1993. Seasonal reproduction in flying-foxes, reviewed in the context of other tropical mammals. Reprod. Fertil. Develop. 5: 499-521.
Parry-Jones, K. 2000. Submission to NSW scientific committee regarding the status of the Grey-headed Flying-fox Pteropus poliocephalus. in Proc. Workshop to Assess the Status of the Grey-headed Flying Fox in NSW. edited by G. Richards. http://www.batcall.csu.edu.au/abs/ghff/ghffproceedings.pdf.
Pierson, E. D. and Rainey, W. E. 1992. The Biology of flying-foxes of the Genus Pteropus: A Review. Pp 1-17 In Pacific Island Flying-foxes: Proceedings of an International Conservation Conference. Biological Report 90 (23) US Department of the Interior, Fish and Wildlife Service, Washington, DC.
Puddicombe, R. 1981. A behavioural study of the Grey-headed Flying-fox Pteropus poliocephalus (Megachiroptera). Bsc Honours Thesis, University of New England, Armidale.
Pulliam, H. R. 1996. Sources and sinks: empirical evidence and population consequences. Pp45-69 in Population Dynamics in Ecological Space and Time. ed. Rhodes, O. E. Jnr, Chesser, R. K. and Smith, M. H. University of Chicago.
Ratcliffe, F. N. 1931. The flying-fox ( Pteropus) in Australia. CSIR Bull. 53: 1-81.
Ratcliffe, F. N. 1932. Notes on the fruit bats ( Pteropus spp.) of Australia. J. Anim. Ecol. 1: 32-57.
Richards, G. C. 2000. The extent of flying-fox culling in orchards and industry attitudes to the NPWS permit system. Submission to NSW scientific committee regarding the status of the Grey-headed Flying-fox Pteropus poliocephalus. Proc. Workshop to Assess the Status of the Grey-headed Flying Fox in NSW. edited by G. Richards. http://www.batcall.csu.edu.au/abs/ghff/ghffproceedings.pdf.
Rigden, P., Page, J. and Chapman, J. 2000 To Net or Not to Net? Flying Fox Control in Orchards Through Netting Protection. Queensland Department of Primary Industries.
Stearns, S. C. 1992 The evolution of life histories. Oxford University Press, Oxford.
Tidemann, C. R. 1999. Biology and management of the Grey-headed Flying-fox P. poliocephalus. Acta Chiropterol. 1: 151-64.
Tidemann, C. 2000. Referenced in Rigden, et al. (2000) as “Personal communication”.
Tidemann, C. R. 2000a. Expert opinion. Booth v Bosworth Federal Court Proceeding No Q163 of 2000, p.3.
Tidemann, C. R. and Vardon, M. J. 2001. The Spectacled Flying-fox, Pteropus conspicillatus, in Queensland: a review of available information. An unpublished document (copy obtained by subpoena) submitted to the Sustainable Ecosystems and the Rainforest Cooperative Research Centre, Atherton Queensland, for this institute's Report for Environment Australia, prepared by Westcott, D. A., Dennis, A. J., McKeown, A. Bradford, M. and Margules, C. R. The Spectacled Flying Fox Pteropus conspicillatus, in the Context of the World Heritage Values of the Wet Tropics World Heritage Area.
Tidemann, C., Eby, P., Parry-Jones, K. and Vardon, M. 1999. Grey-headed Flying Fox. Pp. 31-35 in The Action Plan for Australian Bats ed Duncan, A., Baker, B. and Montgomery, N. Natural Heritage Trust, Canberra.
Towers, P. A. and Martin, L. 1985. Some aspects of female reproduction in the Grey-headed Flying-fox Pteropus poliocephalus (Megachiroptera: Pteropodidae). Aust. Mammal. 8: 257-63.
Towers, P. A. and Martin, L. 1995. Peripheral plasma progesterone concentrations in pregnant and nonpregnant Grey-headed flying-foxes ( Pteropus poliocephalus) and Little red flying-foxes ( P. scapulatus). Reprod. Fertil. Dev. 7: 1163-76.
Tuttle, M. D. and Stevenson, D. 1982. Growth and survival in bats. Pp. 105-150 in Ecology of Bats ed Kunz, T. H., Plenum Press, New York.
Vardon, M. J. and Tidemann, C. R. 1998. Reproduction, growth and maturity in the black flying-fox, Pteropus alecto (Megachiroptera: Pteropodidae). Aust. J. Zool. 46: 329-44.
Vardon, M. J. and Tidemann, C. R. 2000. The black flying-fox ( Pteropus alecto) in north Australia: juvenile mortality and longevity. Aust. J. Zool. 48: 91-7.
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References

Andrewartha, H. G. and Birch, L. C. 1954. The Innate Capacity for Increase in Numbers, Pp. 31-54 in The Distribution and Abundance of Animals, The University of Chicago Press, Chicago.
Collins, L. 1999. The impact of drought on the flying-foxes of the NSW North Coast. Australasian Bat Society Newsletter. 12: 31-32.
Dukelow, W. R., Pow, C., Kennedy, J. H. and Martin, L. 1990. Stress effects on late pregnancy in the flying fox Pteropus scapulatus. Zool. Sci. 7: 871-77.
Eby, P. 1995 The Biology and Management of Flying foxes in New South Wales, Species Management Report No. 18. NSW National Parks and Wildlife Service: Hurstville, New South Wales.
Eby, P. 1999. Low reproductive output in Grey-headed Flying-foxes associated with a short period of food scarcity. Australasian Bat Society Newsletter. 14: 17-20.
Eby P. Richards, G; Collins, L. and Parry-Jones, K. 1999. The distribution, abundance and vulnerability to population reduction of a nomadic nectarivore, the Grey-headed Flying-fox Pteropus poliocephalus in New South Wales, during a period of resource concentration. Aust. Zool. 31: 240-43.
Hall, L. S. and Richards, G. C. 1987. Crop protection and management of flying-foxes (Chiroptera: Pteropodidae). Aust. Mamm. 10: 137-39.
Hall, L. S., Martin, L., O'Brien, G., Kelly, R. and Luckhoff, H. 1991. Flying fox populations in crisis in southeast Queensland. Proc. Aust. Mamm. Soc. 37: 18.
Hassell, M. P. 1986. Detecting density dependence. Trends in Ecology and Evolution 1: 90-3.
Leslie, P. H. 1945. On the use of matrices in certain population mathematics. Biometrika 35: 183-212.
Lunney, D. and Moon, C., 1997. Flying foxes and their camps in rainforest remnants of north-east New South Wales. Pp. 247-77 in Australia's Ever Changing Forests III Ed by J. Dargavel. Centre for Resource and Environmental Studies, Australian National University: Canberra.
Martin, L. 1997. What regulates breeding season in Australian Flying foxes (genus Pteropus)? Proc. Aust. Soc. Reprod. Biol. 27: 100. (reprinted, 1999 in Australasian Bat Society Newsletter 12: 9-11).
Martin, L. and Bernard, R. T. F. 2000. Endocrine Regulation of Reproduction in Bats: the Role of Circulating Gonadal Hormones. Pp. 27-64 in Reproductive Biology of Bats, Ed by Crichton, E. G. and Krutzsch, P. H., Academic Press, London.
Martin, L., Towers, P. A., McGuckin, Little, L., Luckhoff, H. and Blackshaw, A. 1987. Reproductive biology of flying-foxes (Chiroptera: Pteropodidae). Aust. Mamm. 10: 115-18.
Martin, L., Kennedy, J. H., Little, L. Luckhoff, H., O'Brien, G. M., Pow, C. S. T., Towers, P. A., Waldon, A. K., and Wang, D. Y. 1996. The reproductive biology of Australian flying-foxes (genus Pteropus). Symposium Zoological Society of London, 67: 167-84.
McIlwee, A. P. and Martin, L. 2002. On the intrinsic capacity for increase of Australian flying-foxes ( Pteropus spp., Megachiroptera). Australian Zoologist 32: 76-100.
McGuckin, M.A, and Blackshaw, A. W. 1987. Seasonal changes in spermatogenesis (including germ cell degeneration) and plasma testosterone concentrations in the Grey-headed fruit bat, Pteropus poliocephalus. Aust. J. Biol. 40: 211-20.
Nelson J. E. 1963. The biology of the flying-fox (genus Pteropus) in south-eastern Queensland. PhD Thesis, The University of Queensland.
Nelson, J. E. 1965. Movements of Australian flying-foxes (Pteropodidae: megachiroptera). Aust. J. Zool. 13: 53-73.
O'Brien, G. M. 1993. Seasonal reproduction in flying-foxes, reviewed in the context of other tropical mammals. Reprod. Fertil. Develop. 5: 499-521.
Parry-Jones, K. 2000. Submission to NSW scientific committee regarding the status of the Grey-headed Flying-fox Pteropus poliocephalus. in Proc. Workshop to Assess the Status of the Grey-headed Flying Fox in NSW. edited by G. Richards. http://www.batcall.csu.edu.au/abs/ghff/ghffproceedings.pdf.
Pierson, E. D. and Rainey, W. E. 1992. The Biology of flying-foxes of the Genus Pteropus: A Review. Pp 1-17 In Pacific Island Flying-foxes: Proceedings of an International Conservation Conference. Biological Report 90 (23) US Department of the Interior, Fish and Wildlife Service, Washington, DC.
Puddicombe, R. 1981. A behavioural study of the Grey-headed Flying-fox Pteropus poliocephalus (Megachiroptera). Bsc Honours Thesis, University of New England, Armidale.
Pulliam, H. R. 1996. Sources and sinks: empirical evidence and population consequences. Pp45-69 in Population Dynamics in Ecological Space and Time. ed. Rhodes, O. E. Jnr, Chesser, R. K. and Smith, M. H. University of Chicago.
Ratcliffe, F. N. 1931. The flying-fox ( Pteropus) in Australia. CSIR Bull. 53: 1-81.
Ratcliffe, F. N. 1932. Notes on the fruit bats ( Pteropus spp.) of Australia. J. Anim. Ecol. 1: 32-57.
Richards, G. C. 2000. The extent of flying-fox culling in orchards and industry attitudes to the NPWS permit system. Submission to NSW scientific committee regarding the status of the Grey-headed Flying-fox Pteropus poliocephalus. Proc. Workshop to Assess the Status of the Grey-headed Flying Fox in NSW. edited by G. Richards. http://www.batcall.csu.edu.au/abs/ghff/ghffproceedings.pdf.
Rigden, P., Page, J. and Chapman, J. 2000 To Net or Not to Net? Flying Fox Control in Orchards Through Netting Protection. Queensland Department of Primary Industries.
Stearns, S. C. 1992 The evolution of life histories. Oxford University Press, Oxford.
Tidemann, C. R. 1999. Biology and management of the Grey-headed Flying-fox P. poliocephalus. Acta Chiropterol. 1: 151-64.
Tidemann, C. 2000. Referenced in Rigden, et al. (2000) as “Personal communication”.
Tidemann, C. R. 2000a. Expert opinion. Booth v Bosworth Federal Court Proceeding No Q163 of 2000, p.3.
Tidemann, C. R. and Vardon, M. J. 2001. The Spectacled Flying-fox, Pteropus conspicillatus, in Queensland: a review of available information. An unpublished document (copy obtained by subpoena) submitted to the Sustainable Ecosystems and the Rainforest Cooperative Research Centre, Atherton Queensland, for this institute's Report for Environment Australia, prepared by Westcott, D. A., Dennis, A. J., McKeown, A. Bradford, M. and Margules, C. R. The Spectacled Flying Fox Pteropus conspicillatus, in the Context of the World Heritage Values of the Wet Tropics World Heritage Area.
Tidemann, C., Eby, P., Parry-Jones, K. and Vardon, M. 1999. Grey-headed Flying Fox. Pp. 31-35 in The Action Plan for Australian Bats ed Duncan, A., Baker, B. and Montgomery, N. Natural Heritage Trust, Canberra.
Towers, P. A. and Martin, L. 1985. Some aspects of female reproduction in the Grey-headed Flying-fox Pteropus poliocephalus (Megachiroptera: Pteropodidae). Aust. Mammal. 8: 257-63.
Towers, P. A. and Martin, L. 1995. Peripheral plasma progesterone concentrations in pregnant and nonpregnant Grey-headed flying-foxes ( Pteropus poliocephalus) and Little red flying-foxes ( P. scapulatus). Reprod. Fertil. Dev. 7: 1163-76.
Tuttle, M. D. and Stevenson, D. 1982. Growth and survival in bats. Pp. 105-150 in Ecology of Bats ed Kunz, T. H., Plenum Press, New York.
Vardon, M. J. and Tidemann, C. R. 1998. Reproduction, growth and maturity in the black flying-fox, Pteropus alecto (Megachiroptera: Pteropodidae). Aust. J. Zool. 46: 329-44.
Vardon, M. J. and Tidemann, C. R. 2000. The black flying-fox ( Pteropus alecto) in north Australia: juvenile mortality and longevity. Aust. J. Zool. 48: 91-7.
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