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Determining the population dynamics of the vulnerable Grey-headed Flying-fox, Pteropus poliocephalus requires accurate methods of estimating the age of wild animals. Traditionally, wild juvenile P. poliocephalus have been aged by comparing their forearm measurements to those of known-aged captive bred juveniles. To determine the degree of plasticity in the forearm growth rate and hence the reliability of this method, the forearm growth of two groups of captive-bred flying-foxes whose mothers were fed different diets was compared. The difference in diet, a protein supplement of pollen rather than milk powder, made no significant difference in the average length of the forearm at birth, but there were significant differences in the subsequent growth of each growth parameter tested, with the pollen supplemented maternal diet providing faster post-natal growth than the milk supplemented maternal diet. Maternal diet was also significantly correlated with a difference in the sex ratio with a majority of females being produced on the milk supplemented maternal diet and a majority of males being produced on the pollen supplemented maternal diet. There was no significant difference in the post-natal forearm growth of males and females independent of their mother's diet. The sigmoidal curve that best described the average post-natal growth rate of mother-reared P. poliocephalus was the Logistic Model Function. As the diet of wild flying-foxes varies from year to year it is likely that the growth rates of wild mother-reared flying-foxes vary from year to year along with the age of competent flight and independence. The plasticity in the forearm growth rate indicates that estimating the age of wild juvenile flying-foxes using growth rates of captive mother reared juveniles is unlikely to be an accurate method of ageing wild juvenile flying-foxes and other methods of ageing these animals should be investigated.

Altmann, J. and Alberts, S. 1987. Body Mass and growth rates in a wild primate population. Oecologia 72 (1): 15-20.
Anthony, E. L. P. 1988. Chapter 3: Age determination in bats. Pp 47-58 In Ecological and Behavioral Methods for the Study of Bats, edited by T.H. Kunz. Smithsonian Institution Press: Washington, D.C. USA.
Barclay, R. M.R. 1995Does energy or calcium availability constrain reproduction by bats? Symposium of the Zoology Society of London 67:245-258.
Case, T. J. 1978. On the evolution and adaptive significance of postnatal growth rates in the terrestrial vertebrates. The Quarterly Review of Biology 53: 243-282.
Cameron, E. Z. 2004. Facultative adjustment of mammalian sex ratios in support of the Trivers-Willard hypothesis: evidence for a mechanism. Proceedings of the. Royal Society of London-Series B: Biological Science 271: 1723-1728.
Del Prado, M., Delgado, G. and Villalpando, S. 1997. Maternal lipid intake during pregnancy and lactation alters milk composition and production and litter growth in rats. The Journal of Nutrition 127 (3): 458-462
Divljan, A. 2008. Population Ecology of the Grey-headed Flying-fox Pteropus poliocephalus: A study on the Age-Structure and the Effects of Mortality on a Vulnerable Species. Ph.D. Thesis. Biological Sciences, University of Sydney.
Gopalan, P. 2004. Food preference of the Grey-headed Flyingfox Pteropus poliocephalus. B.Sc. Thesis. Biological Sciences, University of Sydney.
Grosjean, P. 2001. Growth model of the reared sea urchin Paracentrotus lividus(Lamarck, 1816). Ph.D Thesis. Laboratoire de Biologie Marine. Universite Libre de Bruxelles.
Hall, L. S. and Richards, G. 2000. Flying-foxes: Fruit and blossom bats of Australia. University of New South Wales: Sydney. Australia.
Hayssen, V. and Kunz, T. H. 1996. Allometry of litter mass in bats: maternal size, wing morphology, and phylogeny. Journal of Mammalogy 77(2): 476-490
Herbst, H. 1986. The role of nitrogen from fruit pulp in the nutrition of the frugivorous bat, Carollia perspicillata.Biotropica 18(1): 39-44.
Hoying, K. M. and Kunz, T. H. 1998. Variation in size at birth and post-natal growth in the insectivorous bat Pipistrellus subflavus(Chiroptera: Vespertilionidae). Journal of Zoology London 245: 15-27.
Isaac, J. L., Krockenberger, A. K. and Johnson, C. N. 2005. Adaptive sex allocation in relation to life-history in the common brushtail possum, Trichosurus vulpecular.Journal of Animal Ecology 74(3): 552-558.
Jones, K. E., Purvis, A. and Gittleman, J. L. 2003. Biological correlates of extinction risk in bats. The American Naturalist 161: 601-614.
Koehler, C. E. and Barclay, R.M. 2000. Post-Natal Growth and Breeding Biology of the Hoary Bat ( Lasiurus cinereus). Journal of Mammalogy 81(1): 234-244.
Kunz, T. H. 1973. Population studies of the cave bat ( Myotis velifer) reproduction, growth and development. Occasional Papers of the Museum of Natural History, University of Kansas 15: 1-45.
Kunz, T. H. and Anthony E. L. P. 1982. Age estimation and post-natal growth in the bat Myotis lucifugus.Journal of Mammalogy 63(1): 23-32.
Kunz, T. H. and Robson, S. K. 1995. Postnatal growth and development in the Mexican Free-tailed Bat ( Tadarida brasiliensis mexicana): birth size, growth rates, and age estimation. Journal of Mammalogy 76(3): 769-783
Kunz, T. H. and Stern, A. A. 1995. Maternal investment and post-natal growth in bats. Symposium of the Zoology Society of London 67: 123-138.
McIlwee, A.P. and Martin, L. 2002. On the intrinsic capacity for increase of Australian flyng-foxes ( Pteropus spp., Megachiroptera). Australian Zoologist 32(1): 76-100.
Marshall, A. G. 1983. Bats, flowers and fruit: evolutionary relationships in the Old World. Biological Journal of the Linnean Society 20: 115-135.
Messer, M. and Parry-Jones, K. 1997. Milk composition in the Grey-headed Flying-fox, Pteropus poliocephalus(Pteropodidae: Chiroptera). Australian Journal of Zoology 45: 65-73.
Mickleburgh, S. P., Hutson, A. M. and Racey, P. A. 1992. Old World Fruit Bats: An Action Plan for their Conservation. I.U.C.N./S.S.C., Gland, Switzerland.
Morris, P. 1972. A review of mammalian age determination methods. Mammal Review 2(3): 69-104.
Nelson, J. E. 1965a. Behaviour of Australian Pteropodidae (Megachiroptera). Animal Behaviour 13: 544-557.
Nelson, J. E. 1965b. Movements of Australian Flying Foxes (Pteropodidae: Megachiroptera). Australian Journal of Zoology 13: 53-73
Oftedal, O. T. 1984. Milk composition, milk yield and energy output at peak lactation: a comparative review. Symposium Zoological Society, London 51: 33-85
Parry-Jones, K. 1993. The Movements of Pteropus poliocephalus in New South Wales. Ph.D. Thesis. Biological Science, University of New South Wales.
Parry-Jones, K. A. and Augee, M. L. 1991. Food Selection by Grey-headed Flying Foxes ( Pteropus poliocephalus) occupying a summer colony site near Gosford, New South Wales. Wildlife Research 18: 111-24.
Parry-Jones, K. and Augee, M. 2001. Factors affecting the occupation of a colony site in Sydney, New South Wales by the Grey-headed Flying-fox Pteropus poliocephalus(Pteropodidae). Austral Ecology 26: 47-55.
Ratcliffe, F. N. 1931. The Flying Fox ( Pteropus) in Australia. CSIR Bulletin 53: (Government Printer: Melbourne).
Read, A. F. and Harvey, P. H. 1989. Life history differences among the eutherian radiations. Journal of Zoology (London) 219: 329-351.
Rose, R. W., Morahan, T. M., Mulchay, J. E. and Ratkowsky, D. A. 2003. Milk Composition and growth in wild and captive Tasmanian bettongs, Bettongia gaimardi(Marsupialia). Journal of Comparative Physiology B. Biochemistry. Systems & Environmental Physiology 173 (2): 125-133.
Rose, R. W. and Flowers, K. 2005. Milk composition and growth in wild and captive Tasmanian pademelons, Thylogale billardierii(Marsupialia). Australian Journal of Zoology 53 (4): 241-248.
Rosenfeld, C. S. and Roberts, R. M. 2004. Maternal diet and other factors affecting offspring sex ratio: a review. Biology of Reproduction 71(4): 1063-1070.
Stace, P. 1996. Protein content and amino acid profiles of honeybee-collected pollens. Bees 'N Trees Consultants: Lismore, NSW, Australia. http://www.honeybee.com.au/Library/Pollenindex.html.
Steller, D. C. 1986. The dietary energy and nitrogen requirements of the Grey-Headed Flying-Fox Pteropus poliocephalus(Temminck) (Megachiroptera). Australian Journal of Zoology 34: 339-350.
Stern, A., Kunz, T., Studier, E. and Oftedal, O. 1997. Milk composition and lactational output in the greater spear-nosed bat, Phyllostomus hastatus. J. Comparative Physiology B, Biochemistry. Systems & Environmental Physiology 167: 389-398.
Studier, E. H. and Kunz, T. H. 1995. Accretion of nitrogen and minerals in suckling bats, Myotis velifer and Tadarida Braziliensis. Journal of Mammalogy 76 (1): 32-42.
Swartz, S. M., Bennett, M. B. and Carrier, D. R. 1992. Wing bone stresses in free flying bats and the evolution of skeletal design for flight. Nature (London) 359: 726-729.
Welbergen, J.A. 2004. The social organization of the Greyheaded Flying-fox, Pteropus poliocephalus. Ph. D. Thesis. University of Cambridge.
Williams, R. and George, H. 1983. Hand-rearing Greyheaded Fruit Bats and observations on the growth of captive bred animals. Proceedings of the 4th Annual Conference of Australasian Soc. Zookeepers WPZ Dubbo.
Wills, R.B., Lim, J.S. and Greenfield, H. 1987. Composition of Australian foods. 40. Temperate Fruits. Food Technology in Australia 39(11): 520-522.
Zullinger, E. M., Ricklefs, R. E., Redford, K. H. and Mace G. M. 1984. Fitting sigmoidal equations to mammalian growth curves. Journal of Mammalogy 65(4): 607-636.
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References

Altmann, J. and Alberts, S. 1987. Body Mass and growth rates in a wild primate population. Oecologia 72 (1): 15-20.
Anthony, E. L. P. 1988. Chapter 3: Age determination in bats. Pp 47-58 In Ecological and Behavioral Methods for the Study of Bats, edited by T.H. Kunz. Smithsonian Institution Press: Washington, D.C. USA.
Barclay, R. M.R. 1995Does energy or calcium availability constrain reproduction by bats? Symposium of the Zoology Society of London 67:245-258.
Case, T. J. 1978. On the evolution and adaptive significance of postnatal growth rates in the terrestrial vertebrates. The Quarterly Review of Biology 53: 243-282.
Cameron, E. Z. 2004. Facultative adjustment of mammalian sex ratios in support of the Trivers-Willard hypothesis: evidence for a mechanism. Proceedings of the. Royal Society of London-Series B: Biological Science 271: 1723-1728.
Del Prado, M., Delgado, G. and Villalpando, S. 1997. Maternal lipid intake during pregnancy and lactation alters milk composition and production and litter growth in rats. The Journal of Nutrition 127 (3): 458-462
Divljan, A. 2008. Population Ecology of the Grey-headed Flying-fox Pteropus poliocephalus: A study on the Age-Structure and the Effects of Mortality on a Vulnerable Species. Ph.D. Thesis. Biological Sciences, University of Sydney.
Gopalan, P. 2004. Food preference of the Grey-headed Flyingfox Pteropus poliocephalus. B.Sc. Thesis. Biological Sciences, University of Sydney.
Grosjean, P. 2001. Growth model of the reared sea urchin Paracentrotus lividus(Lamarck, 1816). Ph.D Thesis. Laboratoire de Biologie Marine. Universite Libre de Bruxelles.
Hall, L. S. and Richards, G. 2000. Flying-foxes: Fruit and blossom bats of Australia. University of New South Wales: Sydney. Australia.
Hayssen, V. and Kunz, T. H. 1996. Allometry of litter mass in bats: maternal size, wing morphology, and phylogeny. Journal of Mammalogy 77(2): 476-490
Herbst, H. 1986. The role of nitrogen from fruit pulp in the nutrition of the frugivorous bat, Carollia perspicillata.Biotropica 18(1): 39-44.
Hoying, K. M. and Kunz, T. H. 1998. Variation in size at birth and post-natal growth in the insectivorous bat Pipistrellus subflavus(Chiroptera: Vespertilionidae). Journal of Zoology London 245: 15-27.
Isaac, J. L., Krockenberger, A. K. and Johnson, C. N. 2005. Adaptive sex allocation in relation to life-history in the common brushtail possum, Trichosurus vulpecular.Journal of Animal Ecology 74(3): 552-558.
Jones, K. E., Purvis, A. and Gittleman, J. L. 2003. Biological correlates of extinction risk in bats. The American Naturalist 161: 601-614.
Koehler, C. E. and Barclay, R.M. 2000. Post-Natal Growth and Breeding Biology of the Hoary Bat ( Lasiurus cinereus). Journal of Mammalogy 81(1): 234-244.
Kunz, T. H. 1973. Population studies of the cave bat ( Myotis velifer) reproduction, growth and development. Occasional Papers of the Museum of Natural History, University of Kansas 15: 1-45.
Kunz, T. H. and Anthony E. L. P. 1982. Age estimation and post-natal growth in the bat Myotis lucifugus.Journal of Mammalogy 63(1): 23-32.
Kunz, T. H. and Robson, S. K. 1995. Postnatal growth and development in the Mexican Free-tailed Bat ( Tadarida brasiliensis mexicana): birth size, growth rates, and age estimation. Journal of Mammalogy 76(3): 769-783
Kunz, T. H. and Stern, A. A. 1995. Maternal investment and post-natal growth in bats. Symposium of the Zoology Society of London 67: 123-138.
McIlwee, A.P. and Martin, L. 2002. On the intrinsic capacity for increase of Australian flyng-foxes ( Pteropus spp., Megachiroptera). Australian Zoologist 32(1): 76-100.
Marshall, A. G. 1983. Bats, flowers and fruit: evolutionary relationships in the Old World. Biological Journal of the Linnean Society 20: 115-135.
Messer, M. and Parry-Jones, K. 1997. Milk composition in the Grey-headed Flying-fox, Pteropus poliocephalus(Pteropodidae: Chiroptera). Australian Journal of Zoology 45: 65-73.
Mickleburgh, S. P., Hutson, A. M. and Racey, P. A. 1992. Old World Fruit Bats: An Action Plan for their Conservation. I.U.C.N./S.S.C., Gland, Switzerland.
Morris, P. 1972. A review of mammalian age determination methods. Mammal Review 2(3): 69-104.
Nelson, J. E. 1965a. Behaviour of Australian Pteropodidae (Megachiroptera). Animal Behaviour 13: 544-557.
Nelson, J. E. 1965b. Movements of Australian Flying Foxes (Pteropodidae: Megachiroptera). Australian Journal of Zoology 13: 53-73
Oftedal, O. T. 1984. Milk composition, milk yield and energy output at peak lactation: a comparative review. Symposium Zoological Society, London 51: 33-85
Parry-Jones, K. 1993. The Movements of Pteropus poliocephalus in New South Wales. Ph.D. Thesis. Biological Science, University of New South Wales.
Parry-Jones, K. A. and Augee, M. L. 1991. Food Selection by Grey-headed Flying Foxes ( Pteropus poliocephalus) occupying a summer colony site near Gosford, New South Wales. Wildlife Research 18: 111-24.
Parry-Jones, K. and Augee, M. 2001. Factors affecting the occupation of a colony site in Sydney, New South Wales by the Grey-headed Flying-fox Pteropus poliocephalus(Pteropodidae). Austral Ecology 26: 47-55.
Ratcliffe, F. N. 1931. The Flying Fox ( Pteropus) in Australia. CSIR Bulletin 53: (Government Printer: Melbourne).
Read, A. F. and Harvey, P. H. 1989. Life history differences among the eutherian radiations. Journal of Zoology (London) 219: 329-351.
Rose, R. W., Morahan, T. M., Mulchay, J. E. and Ratkowsky, D. A. 2003. Milk Composition and growth in wild and captive Tasmanian bettongs, Bettongia gaimardi(Marsupialia). Journal of Comparative Physiology B. Biochemistry. Systems & Environmental Physiology 173 (2): 125-133.
Rose, R. W. and Flowers, K. 2005. Milk composition and growth in wild and captive Tasmanian pademelons, Thylogale billardierii(Marsupialia). Australian Journal of Zoology 53 (4): 241-248.
Rosenfeld, C. S. and Roberts, R. M. 2004. Maternal diet and other factors affecting offspring sex ratio: a review. Biology of Reproduction 71(4): 1063-1070.
Stace, P. 1996. Protein content and amino acid profiles of honeybee-collected pollens. Bees 'N Trees Consultants: Lismore, NSW, Australia. http://www.honeybee.com.au/Library/Pollenindex.html.
Steller, D. C. 1986. The dietary energy and nitrogen requirements of the Grey-Headed Flying-Fox Pteropus poliocephalus(Temminck) (Megachiroptera). Australian Journal of Zoology 34: 339-350.
Stern, A., Kunz, T., Studier, E. and Oftedal, O. 1997. Milk composition and lactational output in the greater spear-nosed bat, Phyllostomus hastatus. J. Comparative Physiology B, Biochemistry. Systems & Environmental Physiology 167: 389-398.
Studier, E. H. and Kunz, T. H. 1995. Accretion of nitrogen and minerals in suckling bats, Myotis velifer and Tadarida Braziliensis. Journal of Mammalogy 76 (1): 32-42.
Swartz, S. M., Bennett, M. B. and Carrier, D. R. 1992. Wing bone stresses in free flying bats and the evolution of skeletal design for flight. Nature (London) 359: 726-729.
Welbergen, J.A. 2004. The social organization of the Greyheaded Flying-fox, Pteropus poliocephalus. Ph. D. Thesis. University of Cambridge.
Williams, R. and George, H. 1983. Hand-rearing Greyheaded Fruit Bats and observations on the growth of captive bred animals. Proceedings of the 4th Annual Conference of Australasian Soc. Zookeepers WPZ Dubbo.
Wills, R.B., Lim, J.S. and Greenfield, H. 1987. Composition of Australian foods. 40. Temperate Fruits. Food Technology in Australia 39(11): 520-522.
Zullinger, E. M., Ricklefs, R. E., Redford, K. H. and Mace G. M. 1984. Fitting sigmoidal equations to mammalian growth curves. Journal of Mammalogy 65(4): 607-636.
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