In the reproductive biology of organisms, a continuum exists from “highly reproductive species” at one end to “survivor species” at the other end. Among other factors, the position of a species along this continuum affects its sensitivity to human exploitation and its vulnerability to extinction. Flying foxes are long-lived, seasonal breeders, with a rigid, well-defined breeding season that is largely or wholly genetically determined. Unlike opportunistic, highly reproductive species, such as rabbits or mice, female flying foxes are unable to produce viable young before their second or third year of life, and are then capable of producing just one young per year. Such a breeding strategy will be successful only if flying-foxes are long-lived and suffer naturally low mortality rates. In this paper,we assess the vulnerability of flying foxes to extinction, using basic parameters of reproduction observed in the wild, and in captive breeding colonies of P. poliocephalus, P. alecto and P. scapulatus, and survival rates that are likely to apply to Australian conditions. Our models show explicitly that flying-fox populations have a very low capacity for increase, even under the most ideal conditions.The implications of our models are discussed in reference to the long-term management and conservation needs of Australian flying foxes. We conclude that current death-rates of flying-foxes in NSW and Queensland fruit orchards are putting state populations at serious risk.

Flying-foxes, reproduction, culling, population decline, migration, endangered species
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.
,
The Distribution and Abundance of Animals
(pg.
31
-
54
)
Baker, J.R. and Baker, Z., 1936. The seasons in a tropical rain forest (New Hebrides). Pt. 3. Fruit Bats (Pteropodidae). Journal of the Linnean. Society (Zoology). 40: 123-142.
,
The seasons in a tropical rain forest (New Hebrides). Pt. 3. Fruit Bats (Pteropodidae)
,
Journal of the Linnean. Society (Zoology)
, vol.
40
(pg.
123
-
142
)
Benton, T.G. and Grant, A, 1999. Elasticity analysis as an important tool in evolutionary and population ecology. Trends in Ecology & Evolution 14(12): 467-471
,
Elasticity analysis as an important tool in evolutionary and population ecology
,
Trends in Ecology & Evolution
, vol.
14
(pg.
467
-
471
)
Boyce, M.S., 1984. Restitution or r- and K-selection as a model of density-dependent natural selection. Annual Review of Ecology and Systematics 15: 427-447.
,
Restitution or r- and K-selection as a model of density-dependent natural selection
,
Annual Review of Ecology and Systematics
, vol.
15
(pg.
427
-
447
)
Cole, L.C., 1954. The population consequences of life history phenomena. Quarterly Review of Biology 29: 103-37.
,
The population consequences of life history phenomena
,
Quarterly Review of Biology
, vol.
29
(pg.
103
-
37
)
De Kroon, H., Van Groenendael, J. and Ehrlen, J., 2000. Elasticities: A review of methods and model limitations Ecology. 81: 607-618.
,
Elasticities: A review of methods and model limitations
,
Ecology
, vol.
81
(pg.
607
-
618
)
Dolbeer, R. R., Fiedler, L. R. and Rasheed, H., 1988. Management of fruit bat and rat populations in the Maldive Islands, Indian Ocean. Proc. Vertebr. Pest. Conf. 13: 112-118.
,
Management of fruit bat and rat populations in the Maldive Islands, Indian Ocean
,
Proc. Vertebr. Pest. Conf.
, vol.
13
(pg.
112
-
118
)
Dukelow, W.R., Pow, C., Kennedy, J.H. and Martin, L., 1990. Stress effects on late pregnancy in the fluying fox Pteropus scapulatus. Zoological Science 7: 871-877.
,
Stress effects on late pregnancy in the fluying fox Pteropus scapulatus
,
Zoological Science
, vol.
7
(pg.
871
-
877
)
Eby, P., 1998. An analysis of diet specialization in frugivorous Pteropus poliocephalus (Megachiroptera) in Australian subtropical rainforests. Australian Journal of Ecology 23: 443-56.
,
An analysis of diet specialization in frugivorous Pteropus poliocephalus (Megachiroptera) in Australian subtropical rainforests
,
Australian Journal of Ecology
, vol.
23
(pg.
443
-
56
)
Eby, P, 1999. Low reproductive output in grey-headed flying foxes associated with a short period of food scarity. Bat Society Newsletter. 14: 17-20
,
Low reproductive output in grey-headed flying foxes associated with a short period of food scarity
,
Bat Society Newsletter
, vol.
14
(pg.
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 or resource concentration. Australian Zoologist. 31: 240-43
,
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 or resource concentration
,
Australian Zoologist
, vol.
31
(pg.
240
-
43
)
Ferson, S. and Akcakaya, H.R., 1990. Modelling fluctuations in age-structured populations. RAMAS/age User Manual. Applied Biomathematics. Setauket, New York.
Garnett, S., Whybird, O. and Spencer, H., 1999. The conservation status of the spectacled flying fox Pteropus conspicillatus.Australian Zoologist 31: 38-54.
,
The conservation status of the spectacled flying fox Pteropus conspicillatus
,
Australian Zoologist
, vol.
31
(pg.
38
-
54
)
Hall, L.S., 2000. Recent historical changes to the distribution, and declines in the abundance of the grey-headed flying fox in south-east Queensland. 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 p51-55.
Hall, L.S. and Richards, G.C., 1987. Crop protection and management of flying foxes (Chiroptera: Pteropodidae). Australian Mammalogy 10: 137-39.
,
Crop protection and management of flying foxes (Chiroptera: Pteropodidae)
,
Australian Mammalogy
, vol.
10
(pg.
137
-
39
)
Hall, L. and Richards, G.C., 2000.Flying foxes: Fruit and blossom bats of Australia. UNSW Press, Sydney.
,
Flying foxes: Fruit and blossom bats of Australia
Hall, L.S., Martin, L., O’Brien, G., Kelly, R. and Luckhoff, H., 1991. Flying fox populations in crisis in southeast Queensland. Proceedings of the Australian Mammal Society 37: 18.
,
Flying fox populations in crisis in southeast Queensland
,
Proceedings of the Australian Mammal Society
, vol.
37
pg.
18
Harvey, P.H., Read A.F. and Promislow, D.E.L., 1989. Life-history variation in placental mammals unifying the data with theory. Oxford Surveys in Evolutionary Biology 6: 13-31
,
Life-history variation in placental mammals unifying the data with theory
,
Oxford Surveys in Evolutionary Biology
, vol.
6
(pg.
13
-
31
)
Hassell, M.P., 1986. Detecting density dependence. Trends in Ecology and Evolution 1: 90-93.
,
Detecting density dependence
,
Trends in Ecology and Evolution
, vol.
1
(pg.
90
-
93
)
Heideman, P.D. and Heaney, L.R., 1989. Population cbiology and estimates of abundance of fruit bats (Pteropodidae) in Phillipine submontane rainforest. Journal of Zoology (London) 218: 565-586.
,
Population cbiology and estimates of abundance of fruit bats (Pteropodidae) in Phillipine submontane rainforest
,
Journal of Zoology (London)
, vol.
218
(pg.
565
-
586
)
Heppell, S.S., Caswell, H. and Crowder, L.B., 2000. Life histories and elasticity patterns: Perturbation analysis for species with minimal demographic data. Ecology 81(3): 654-665
,
Life histories and elasticity patterns: Perturbation analysis for species with minimal demographic data
,
Ecology
, vol.
81
(pg.
654
-
665
)
Leslie, P.H., 1945. On the use of matrices in certain population mathematics. Biometrika 35: 183-212.
,
On the use of matrices in certain population mathematics
,
Biometrika
, vol.
35
(pg.
183
-
212
)
Leslie, P.H. and Ranson, R.M., 1940. The mortality, fertility and rate of natural increase of the vole (Microtis agrestis) as observed in the laboratory. Journal of Animal Ecology. 9: 27-52.
,
The mortality, fertility and rate of natural increase of the vole (Microtis agrestis) as observed in the laboratory
,
Journal of Animal Ecology
, vol.
9
(pg.
27
-
52
)
Lotka, A.J., 1922. The stability of the normal age distribution. Proceedings of the National Academy of Science 24: 199-216.
,
The stability of the normal age distribution
,
Proceedings of the National Academy of Science
, vol.
24
(pg.
199
-
216
)
Lotka, A. J., 1925. [cited by Andrewartha and Birch (1954)] Elements of physical biology. Williams and Wilkins, Baltimore
,
Elements of physical biology
Macarthur, R.H. and Wilson, E.O., 1967.The theory of island biogeography. Princeton University Press, Princeton.
,
The theory of island biogeography
Marshall, A.J., 1948. The breeding cycle of an equatorial bat (Pteropus giganteus of Ceylon). Proc. Linn. Soc. Lond. 159: 103-111.
,
The breeding cycle of an equatorial bat (Pteropus giganteus of Ceylon)
,
Proc. Linn. Soc. Lond.
, vol.
159
(pg.
103
-
111
)
Martin, L., 1997. What regulates breeding season in Australian Flying foxes (genus Pteropus)? Proceeding of the Australian Society for Reproductive Biology 27: 100. [reprinted 1999 in The Australasian Bat Society Newsletter 12: 9-11.]
Martin, L., 1999. On seasonal breeding in Australian Flying-foxes, with August observations on a maternity camp of Pteropus scapulatus in north-west Queensland. The Australasian Bat Society Newsletter 12: 5-9.
,
On seasonal breeding in Australian Flying-foxes, with August observations on a maternity camp of Pteropus scapulatus in north-west Queensland
,
The Australasian Bat Society Newsletter
, vol.
12
(pg.
5
-
9
)
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, editors, Crichton, E.G. and Krutzsch, P.H., Academic Press, London.
,
Reproductive Biology of Bats
(pg.
27
-
64
)
Martin, L. and McIlwee, A.P., 2002. The reproductive biology and intrinsic capacity for increase of the Grey-headed Flying-fox (Pteropus poliocephalus: megachiroptera), and the implications of culling. Australian Zoologist (in press)
Martin, L., Towers, P.A., McGuckin, M.A., Little, L., Luckhoff, H. and Blackshaw, A., 1987. Reproductive biology of flying-foxes (Chiroptera: Pteropodidae). Australian Mammalogy, 10: 115-118.
,
Reproductive biology of flying-foxes (Chiroptera: Pteropodidae)
,
Australian Mammalogy
, vol.
10
(pg.
115
-
118
)
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: 167-184.
,
The reproductive biology of Australian flying-foxes (genus Pteropus)
,
Symposium Zoological Society of London
, vol.
67
(pg.
167
-
184
)
McCoy, M., 1990. Pollination of eucalypts by flying foxes in northern Australia, pp. 33-37 in Slack, J.M., Flying-fox workshop proceedings, New South Wales Department of Agriculture and Fisheries, Sydney.
Mills L.S., Doak, D.F. and Wisdom, M.J., 1999. Reliability of conservation actions based on elasticity analysis of matrix models. Conservation Biology. 13(4): 815-829
,
Reliability of conservation actions based on elasticity analysis of matrix models
,
Conservation Biology
, vol.
13
(pg.
815
-
829
)
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). Australian Journal of Zoology 13: 53-73.
,
Movements of Australian flying foxes (Pteropodidae: megachiroptera)
,
Australian Journal of Zoology
, vol.
13
(pg.
53
-
73
)
O’Brien, G.M., 1993. Seasonal reproduction in flying-foxes, reviewed in the context of other tropical mammals. Reproduction Fertility and Development, 5: 499-521.
,
Seasonal reproduction in flying-foxes, reviewed in the context of other tropical mammals
,
Reproduction Fertility and Development
, vol.
5
(pg.
499
-
521
)
O’Brien, G. M., Martin, L. and Curlewis, J. D., 1993. Effect of photoperiod on the annual cycle of testis growth in a tropical mammal, the little-red flyingfox Journal of Reproduction and Fertility 98: 121-127.
,
Effect of photoperiod on the annual cycle of testis growth in a tropical mammal, the little-red flyingfox
,
Journal of Reproduction and Fertility
, vol.
98
(pg.
121
-
127
)
Parry-Jones, K, 2000. Historical declines since the early 1900's, and current mortality factors and abundance of the grey-headed flying fox in NSW. 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 Pp.56-65.
Pianka, E.R., 1970. On r- and K-selection. American Naturalist 104: 592-597.
,
On r- and K-selection
,
American Naturalist
, vol.
104
(pg.
592
-
597
)
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.
Pritchard, K., 2001. George (1978-2001). Friends of Bats Newsletter (Ku-rin-gai Bat Conservation Soc.) 61: 7.
,
George (1978-2001)
,
Friends of Bats Newsletter (Ku-rin-gai Bat Conservation Soc.)
, vol.
61
pg.
7
Promislow, D.E.L. and Harvey, P.H., 1990. Living fast and dying young: A comparative analysis of life history variation in mammals. Journal of Zoology, (London) 232: 417-437.
,
Living fast and dying young: A comparative analysis of life history variation in mammals
,
Journal of Zoology, (London)
, vol.
232
(pg.
417
-
437
)
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., 1931. The flying fox (Pteropus) in Australia. CSIR Bulletin 53: 1-81.
,
The flying fox (Pteropus) in Australia
,
CSIR Bulletin
, vol.
53
(pg.
1
-
81
)
Ratcliffe, F., 1932. Notes on the fruit bats (Pteropus spp) of Australia. Journal of Animal Ecology 1: 33-57.
,
Notes on the fruit bats (Pteropus spp) of Australia
,
Journal of Animal Ecology
, vol.
1
(pg.
33
-
57
)
Richards, G.C., 1990. The spectacled flying-fox, Pteropus conspicillatus in North Queensland: Part 2, Diet, feeding ecology and see dispersal. Australian Mammalogy, 13: 25-31.
,
The spectacled flying-fox, Pteropus conspicillatus in North Queensland: Part 2, Diet, feeding ecology and see dispersal
,
Australian Mammalogy
, vol.
13
(pg.
25
-
31
)
Richards, G.C., 2000. Submission to NSW scientific committee regarding the status of the Grey-headed flying-fox Pteropus poliocephalus: The extent of flying fox culling in orchards and industry attitudes to the NPWS permit system. Proc. Workshop to Assess the Status of the Grey-headed Flying Fox in NSW.
Ross, C.R., 1987. The intrinsic rate of natural increase and reproductive effort in primates. Journal of Zoology 214: 119-220.
,
The intrinsic rate of natural increase and reproductive effort in primates
,
Journal of Zoology
, vol.
214
(pg.
119
-
220
)
Saether, B.E., Ringsby, T.H. and Roskaft, E., 1996. Life history variation, population processes and priorities in species conservation: Towards a reunion of research paradigms. Oikos 77: 217-226.
,
Life history variation, population processes and priorities in species conservation: Towards a reunion of research paradigms
,
Oikos
, vol.
77
(pg.
217
-
226
)
Saether, B.E. and Bakke, O., 2000. Avian life history variation and contribution of demographic traits to the population growth rate. Ecology 81(3): 642-653.
,
Avian life history variation and contribution of demographic traits to the population growth rate
,
Ecology
, vol.
81
(pg.
642
-
653
)
Sibly, R.M., Collett, D., Promislow, D.E.L., Peacock, D.J. and Harvey, P.H., 1997. Mortality rates of mammals. Journal of Zoology, London 243: 1-12.
,
Mortality rates of mammals
,
Journal of Zoology, London
, vol.
243
(pg.
1
-
12
)
Stearns, S.C., 1992.The evolution of life histories. Oxford University Press, Oxford.
,
The evolution of life histories
Tidemann, C.R., 1999. Biology and management of the grey-headed flying-fox P. poliocephalus. Acta Chiropterologica 1: 151-164.
,
Biology and management of the grey-headed flying-fox P. poliocephalus
,
Acta Chiropterologica
, vol.
1
(pg.
151
-
164
)
Tidemann, C.R., 2000. 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 supoena) submitted to The Sustainable Ecosystems and the Rainforest Cooperative Research Centre, Atherton Queensland, for this institutes Report for Environment Australia, prepared by Westcott, D.A., Dennis, A.J., McKeown, A., Bradford, M. and Marqules, C.R. The Spectacled Flying Fox Pteropus conspicillatus in the Context of 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.
,
The Action Plan for Australian Bats
(pg.
31
-
35
)
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: 257-263.
,
Some aspects of female reproduction in the grey headed flying-fox Pteropus poliocephalus (Megachiroptera: Pteropodidae)
,
Aust. Mammal.
, vol.
8
(pg.
257
-
263
)
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.
,
Ecology of Bats
(pg.
105
-
150
)
Vardon, M.J. and Tidemann, C.R., 1995. Harvesting of flying foxes (Pteropus spp.) in Australia: Could it promote the conservation of endangered Pacific Island species? in Conservation through sustainable use of wildlife ed Grigg, G., Hale, P. and Lunney, D. University of Queensland.
Vardon, M.J. and Tidemann, C.R., 1998. Reproduction, growth and maturity in the black flying fox, Pteropus alecto (Megachiroptera: Pteropodidae). Australian Journal of Zoology. 46: 329-344.
,
Reproduction, growth and maturity in the black flying fox, Pteropus alecto (Megachiroptera: Pteropodidae)
,
Australian Journal of Zoology
, vol.
46
(pg.
329
-
344
)
Vardon, M.J. and Tidemann, C.R., 2000. The black flying fox (Pteropus alecto) in north Australia: juvenile mortality and longevity. Australian Journal of Zoology. 48: 91-97.
,
The black flying fox (Pteropus alecto) in north Australia: juvenile mortality and longevity
,
Australian Journal of Zoology
, vol.
48
(pg.
91
-
97
)
Williams, G. C., 1966.Adaptation andNatural Selection. Princeton University Press, Princeton, New Jersey.
,
Adaptation andNatural Selection
Whybird, O., Clague, S. and Clague, C., 2000. Spectacled Flying Fox – the need and justification for listing. Report to the Endangered Species Monitoring Program, Environmental Protection Agency, Queensland.
This content is only available as a PDF.