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Dr Daniel A. Warner Post-Doctoral
Fellow e-mail: dwarner@iastate.edu |
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In general, I am interested in how environmental and maternal factors influence variation in phenotypic traits, and their implications on the evolution of life-history traits. The majority of my research investigates how the embryonic environment and maternal factors modify various fitness-related traits in oviparous reptiles (specifically lizards), and, in turn, how these traits influence individual survival and reproduction (i.e., fitness). The current and past research projects that I have been involved with are briefly outlined below.
2003-2007: Adaptive significance of temperature-dependent sex determinationMy current research focuses on the ecology and evolution of temperature-dependent sex determination (TSD) in reptiles. Species with TSD typically do not have sex chromosomes, and offspring sex is determined by the temperatures embryos experience during development. This type of sex-determining mechanism is found in all crocodilians, most turtles, and several lizard species. The numerous independent occurrences of TSD within and among the major reptile groups raise several interesting questions about the evolution and adaptive significance of TSD.
The major component of my PhD research focuses on testing the differential fitness model proposed by Charnov and Bull (1977. Nature 266:828-830). This model provides an explanation for why environmental sex determination would be favoured over genetic sex determination. In terms of the adaptive significance of TSD, this model proposes that incubation temperatures differentially influence the fitness of male and female offspring, such that male-producing temperatures are best-suited for males and female-producing temperatures are best-suited for females.
One problem with testing the differential fitness model is that it is logistically difficult (if not impossible) to evaluate the long-terms effects of developmental temperatures on lifetime reproductive success (i.e., fitness), especially on extremely long-lived, late-maturing TSD reptiles, such as turtles and crocodilians. However, with the recent discovery of TSD in many short-lived agamid lizards in Australia, evaluating the long-term effects of incubation history on reproductive success now appears to be feasible.
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I am using the jacky dragon (Amphibolurus muricatus) to test the differential fitness model. Jacky dragons are common agamid lizards found in coastal heathland habitat in southeast Australia (see photo taken at Royal National Park, below). Jacky dragons have an interesting pattern of TSD, whereby low and high incubation temperatures produce females, and intermediate temperatures produce both sexes. These short-lived lizards can reach sexually maturity at an age of 9 months depending on how early they hatched in the previous season. Their nesting season occurs from October to February and they can produce up to four clutches of eggs within a single season.
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During the first year (2003-2004) of my PhD research, I initiated a long-term experimental test of four hypotheses for the adaptive significance of TSD; each hypothesis falls within the framework of the differential fitness model (see Warner and Shine. 2005. Evolution 59:2209-2221). The overall objective of this project is to evaluate the long-term effects of incubation temperature on male and female fitness. This project involves raising offspring to sexual maturity in semi-natural field enclosures to evaluate the sex-specific effects of incubation history on survival and actual reproductive success. Thus, I intend to continue this project for at least two more reproductive seasons in order to evaluate incubation and phenotypic influences on life-time reproductive success.
Research during my second season (2004-2005) focused on maternal influences on offspring sex ratios in the jacky dragon. I evaluated two types of maternal effects that likely play a role in sex determination; the first type is maternal nest-site selection behaviour, and the second is maternal allocation of endogenous steroid hormones into the yolks of eggs (in collaboration with Dr. Matthew Lovern, Oklahoma State University). I also conducted experiments to determine if the quality of the maternal diet and the operational sex ratio of the adult population influences sex allocation when eggs are incubated under a temperature that produces a 50:50 sex ratio.
During my third season (2005-2006), my research will focus on the nesting behaviour of females under natural field conditions. The objective of this study is to monitor field nest temperatures and determine the influence of natural incubation temperatures on offspring sex ratios under natural conditions. In addition, because environmental temperatures change over the course of the nesting season (October to February), I will evaluate how (or if) offspring sex ratios shift during the hatching season. This research will also determine the environmental characteristics that females consider when selecting a nest site.
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Female jacky dragon with radio transmitter digging nest
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Average clutch of eggs in nest before being buried
1998-2003: Environmental and maternal influences on phenotypes and survival of hatchlings lizards
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For my MS degree (advised by Dr. Robin Andrews, Virginia Tech), I examined the relative contributions of the incubation moisture environment, maternal yolk investment, and clutch to variation in offspring phenotypes and survival of the eastern fence lizard (Sceloporus undulatus). This research took place in southwest Virginia in Jefferson National Forest.
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After completing my MS degree (January 2001), I extended this research in collaboration with Travis Robbins (University of South Florida). We investigated the effects of flutctuations in the moisture environment during incubation on phenotypes and survival of hatchling Sceloporus undulatus from a population in west-central Florida.
2001-2003: Population biology and fisheries management of marine arthropods
After finishing my MS degree, I moved to Florida and lined up a job at the Florida Marine Research Institute with the Crustacean Fisheries Research Group (supervised by Dr. Theresa Bert) in St. Petersburg. I was involved with various research projects on the population biology and fishery management of stone crabs (Menippe adina & M. mercenaria), blue crabs (Callinectes sapidus), shrimp (penaeid species), and horseshoe crabs (Limulus polyphemus).
1996-1998: Population biology of the six-lined racerunner in northwestern Illinois
As an undergraduate student I worked in the laboratory of Dr. Fredric Janzen (Iowa State University) for about 3 years. During this time I undertook an independent research project to study the demography and population biology of six-lined racerunners (Cnemidophorus sexlineatus) in northwest Illinois. I was also involved with various other projects that focused on the demographics of western hognosed snakes (Heterodon nasicus), and nesting patterns and TSD in painted turtles (Chrysemys picta) and snapping turtles (Chelydra serpentina). During my time in the Janzen lab, I spent a summer (1997) working with John Tucker on his demographic studies of red-eared slider turtles (Trachemys scripta elegans) in west-central Illinois.
Iowa State University, Ames, Iowa, USA.
Bachelor of Science in Animal Ecology. May 1998.
Advisor: Dr. Fredric JanzenVirginia Polytechnic Institute and State University, Blacksburg, Virginia, USA.
Master of Science in Biology. January 2001.
Advisor: Dr. Robin Andrews
The University of Sydney, Sydney, New South Wales, Australia.
PhD candidate in Biology, July 2003 - May 2007.
Advisor: Prof. Richard Shine
Iowa State University, Ames, Iowa, USA.
Post-doctoral Fellowship, Department of Ecology, Evolution and Organismal Biology, July 2007
Hatchling Varanus
rosenbergi
from Royal National
Park
| 1. | Warner, D. A. 1997. An overview on the evolution of the family Iguanidae. Journal of the International Iguana Society (Iguana Times) 6:57-65. |
| 2. | Tucker, J. K., and D. A. Warner. 1997. Sternotherus odoratus (common musk turtle). Herpetological Review 28:209. |
| 3. | Tucker, J. K., and D. A. Warner. 1998. Rana blairi (plains leopard frog). Herpetological Review 29:108. |
| 4. | Tucker, J. K., and D. A. Warner. 1998. Apalone spinifera (spiny softshell) reproduction. Herpetological Review 29:234. |
| 5. |
Warner, D. A. 1998. A preliminary
report on
a population of Cnemidophorus sexlineatus in
northwestern
Illinois. Bulletin of the Chicago
Herpetological Society
33:6-8. |
| 6. |
Warner, D. A.
1998.
Overcrowding effects on larval red-eyed treefrogs
(Agalychnis callidryas).
Bulletin of the
Chicago Herpetological Society
33:212-214. |
| 7. |
Tucker, J. K., and D. A. Warner. 1999. Microgeographic variation in response of red-eared slider (Trachemys scripta elegans) embryos to similar incubation environments. Journal of Herpetology 33:549-557. |
| 8. |
Kolbe, J. J., L. J. Harmon, and D. A. Warner. 1999. New state record lengths and associated natural history notes for some Illinois snakes. Transactions of the Illinois State Academy of Science 92:133-135. |
| 9. | Warner, D. A. 2000. Ecological observations on the six-lined racerunner (Cnemidophorus sexlineatus) in northwestern Illinois. Transactions of the Illinois State Academy of Science 93:239-248. |
| 10. | Tucker, J. K., and D. A. Warner. 2000. Mud accumulation in nesting aquatic turtles. Chelonian Conservation and Biology 3:753-755. |
| 11. | Andrews, R. M., T. Mathies, and D. A. Warner. 2000. Effect of incubation temperature on morphology, growth, and survival of juvenile Sceloporus undulatus. Herpetological Monographs 14:420-431. |
| 12. | Warner, D. A., and R. M. Andrews. 2002. Laboratory and field experiments identify sources of variation in phenotypes and survival of hatchling lizards. Biological Journal of the Linnean Society 76:105-124. |
| 13. | Warner, D. A., and R. M. Andrews. 2002. Nest-site selection in relation to temperature and moisture by the lizard Sceloporus undulatus. Herpetologica 58:399-407. |
| 14. | Warner, D. A., and R. M. Andrews. 2003. Consequences of extended egg retention by the eastern fence lizard (Sceloporus undulatus). Journal of Herpetology 37:309-314. |
| 15. | Warner, D. A., and J. J. Kolbe. 2003. Porthidium nasutum (hog-nosed pitviper) prey. Herpetological Review 34:377. |
| 16. |
Warner, D. A. 2003.
Environmental
and maternal influences on eggs and hatchlings of the eastern
fence lizard (Sceloporus undulatus). Bulletin
of
the Chicago Herpetological Society
38:129-136. |
| 17. |
Warner, D. A.
2003.
The horseshoe crab in Florida. Florida
Wildlife 57(4):28-29. |
| 18. |
Warner, D. A. 2003.
Horseshoe crabs:
living fossils. Sea Stats, Florida Marine
Research
Institute Publication. 4pp. |
| 19. | Warner, D. A., A. L. McMillen-Jackson, T. M. Bert, and C. R. Crawford. 2004. The efficiency of a bycatch reduction device used on skimmer trawls in the Florida shrimp fishery. North American Journal of Fisheries Management 24:853-864. |
| 20. |
Warner,
D. A., and R. Shine.
2005. The adaptive significance of
temperature-dependent sex
determination: experimental tests with a short-lived
lizard.
Evolution
59:2209-2221. |
| 21. | Warner, D. A., J. K. Tucker, N. I. Filoramo, and J. B. Towey. 2006. Claw function of hatchling and adult red-eared slider turtles (Trachemys scripta elegans). Chelonian Conservation and Biology 5:317-320. |
| 22. |
Warner,
D. A., and R. Shine.
2006. Morphological variation does
not influence locomotor
performance within a cohort of hatchling lizards
(Amphibolurus
muricatus, Agamidae). Oikos
114:126-134. |
| 23. |
Allsop,
D. J., D. A. Warner, T.
Langkilde, W. Du, and R. Shine. 2006.
Do operational sex
ratios influence sex allocation in viviparous lizards
with
temperature-dependent sex determination? Journal of Evolutionary Biology
19:1175-1182. |
| 24. |
Opell,
B.
D., J. E. Bond, and D. A. Warner. 2006. The effects of
capture
spiral composition and orb-web orientation on prey
interception. Zoology
109:339-345. |
| 25. |
Radder,
R., D. Warner, J. Cuervo, and R. Shine. 2007. The
functional
significance of residual yolk in hatchling lizards
(Amphibolurus muricatus,
Agamidae). Functional Ecology
21:302-309. |
| 26. |
Warner,
D., J. Thomas,
and R. Shine. 2007. A simple and reliable method
for attaching
radio-transmitters to lizards. Herpetological Conservation
Biology 1:129-131. |
| 27. |
Warner,
D.
A., and R. Shine. 2007. Reproducing lizards modify
sex
allocation in response to operational sex ratio. Biology Letters 3:47-50. |
| 28. |
Warner,
D. A., M. Lovern, and R. Shine.
2007. Maternal nutrition
affects reproductive output and sex
allocation in a lizard with environmental
sex determination. Proceedings of the Royal
Society of London
Series B 274:883-890. |
| 29. |
Shine,
R., D. Warner, and R. Radder. 2007. Windows
of sexual
lability during embryonic development in two lizard
species with environmental
sex determination. Ecology 88:1781-1788. |
| 30. |
Robbins,
T. R., and D. A. Warner.
In review. Fluctuations in
the incubation moisture
environment: does the pattern or magnitude influence
egg survival
and hatchlings phenotypes in a lizard? Oecologia |
| 31. |
Leung, A. N.,
T. T. L. Phu, D.
Lister, J. Poolice, L. Caon, R. Jones, N. Chowdhary, and
D.
A. Warner. 2006. Diurnal skinks (Lampropholis delicata)
seek shelter in the presence of scent from a nocturnal predator and
prey.
Herpetofauna: in press. |
| 32. |
Warner,
D. A. 2007. Amphibolurus muricatus. Predation.
Herpetological
Review: in press. |
| 33. |
Schwartz,
T. S., D. A. Warner, L. B. Beheregaray, and M. Olsson. 2007.
Microsatellite
loci for Australian agamid lizards. Molecular
Ecology Notes: in press.
( |
| 34. |
Warner,
D. A., X. Bonnet,
K. A. Hobson, and R. Shine. In review. Multi-clutching
lizards fuel
reproduction with recently-acquired energy. Ecology. |
| 35. |
Warner,
D. A., and R. Shine. 2007. Fitness of
juvenile lizards depends on seasonal
timing of hatching, not
offspring body size. Oecologia: in press. |
| 36. |
Radder,
R. S., D. A. Warner, and R. Shine. 2007.
Compensating for a bad start:
catch-up growth in juvenile lizards
(Amphibolurus muricatus, Agamidae).
Journal of
Experimental Zoology: in press. |
| 37. |
Warner,
D. A., T. Uller, and R. Shine. In review. Fitness effects of the
seasonal
timing of hatching may drive the evolution of
temperature-dependent sex determination
in short-lived lizards.
Journal of Evolutionary Biology. |
| 1. | Warner, D. A. 1997. A survey of a northern population of six-lined racerunners (Cnemidophorus sexlineatus). Department of Zoology and Genetics Summer Intern Symposium, Iowa State University, Ames, Iowa, USA. (oral presentation) |
| 2. | Warner, D. A. 1998. A survey of an isolated population of the six-lined racerunner (Cnemidophorus sexlineatus). Department of Biochemistry and Biophysics Spring Symposium, Iowa State University, Ames, Iowa, USA. (poster presentation) |
| 3. | Warner, D. A. 1998. A survey of a population of six-lined racerunners (Cnemidophorus sexlineatus) in northwestern Illinois. Chicago Herpetological Society monthly meeting. Field Museum of Natural History, Chicago, Illinois, USA. (invited seminar) |
| 4. | Warner, D. A., and R. M. Andrews. 2000. Phenotypes and survival of hatchling lizards. Virginia Academy of Science annual meeting, Radford University, Radford, Virginia, USA. May 2000. (oral presentation) |
| 5. | Warner, D. A., and R. M. Andrews. 2000. Clutch and incubation effects on phenotypes of hatchling lizards. Joint meeting of the American Society of Ichthyologists and Herpetologists, Society for the Study of Amphibians and Reptiles, and the Herpetologists’ League, La Paz, Baja California Sur, Mexico. (oral presentation) |
| 6. | Warner, D. A. 2000. Phenotypes and survival of hatchling lizards. Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, USA. (seminar) |
| 7. | Warner, D. A., and R. M. Andrews. 2001. Environmental and maternal contributions to phenotypic variation and survival of the lizard Sceloporus undulatus. Society for Integrative and Comparative Biology annual meeting, Chicago, Illinois, USA. (oral presentation) |
| 8. | Warner, D. A. 2001. Phenotypes and survival of hatchling lizards. Ecology, Evolution, and Systematics Seminar Series. Department of Biology, Virginia Polytechnic Institute and State University, Blacksburg, Virginia, USA. (MS defense seminar) |
| 9. | Warner, D. A. 2001. Phenotypes and survival of hatchling lizards. Department of Biology, University of South Florida, Tampa, Florida, USA. (invited seminar) |
| 10. | Warner, D. A., and R. M. Andrews. 2001. Clutch and incubation effects on phenotypes and survival of hatchling lizards. Florida Ecological and Evolutionary Symposium. Archbold Biological Station, Florida, USA. (oral presentation) |
| 11. | Warner, D. A., and R. M. Andrews. 2001. Phenotypes and survival of hatchling lizards. Joint meeting of the Society for the Study of Amphibians and Reptiles, and the Herpetologists’ League, Indianapolis, Indiana, USA. (oral presentation) |
| 12. | Warner,
D.
A., and R. M. Andrews. 2002. Nest site selection
in relation to temperature and moisture by the lizard
Sceloporus undulatus.
Joint meeting of the American
Society of Ichthyologists and Herpetologists,
Society for the
Study of Amphibians and Reptiles, and the
Herpetologists’
League, Kansas City, Missouri, USA.
(poster presentation) |
| 13. | Warner, D. A.
2003.
Sources of variation in phenotypes and survival of
hatchling fence lizards
(Sceloporus
undulatus). Monthly meeting of The Mansota
Herpetological Society. Marie Selby Gardens, Sarasota, Florida,
USA.
(invited seminar) |
| 14. | Robbins, T. R, and D. A. Warner.
2003. The influence of fluctuating moisture conditions during
incubation
on phenotypes of hatchling Sceloporus
undulatus. Joint meeting
of the American Society of
Ichthyologists and Herpetologists, Society for
the Study of
Amphibians and Reptiles, and the Herpetologists' League, Manaus,
Amazonas, Brazil. (oral presentation by T.R.
Robbins) |
| 15. |
Warner, D. A., and
T. R.
Robbins. 2003. The importance of morphology,
performance,
and behavior to survival of hatchling lizards: a
preliminary evaluation.
Joint meeting of the American
Society of Ichthyologists and Herpetologists,
Society for
the Study of Amphibians and Reptiles, and the
Herpetologists’
League, Manaus, Amazonas, Brazil.
(poster presentation) |
| 16. |
Warner,
D. A., and
T. R. Robbins. 2003.
Associations among the incubation
environment, phenotypes, and survival of
hatchling fence lizards
(Sceloporus undulatus). Australian
Society of
Herpetologists. Mary River Park, Northern Territory,
Australia.
(oral presentation) |
| 17. |
Warner,
D. A., and T. R. Robbins. 2004. Associations among
the
incubation environment, phenotypes and survival of hatchling
fence lizards
(Sceloporus undulatus). Meeting of the
Australasian Society
for the Study of Animal Behaviour
and the Australasian Society
for the Study of Evolution,
Adelaide, South Australia, Australia.
(oral
presentation) |
| 18. |
Warner,
D. A. 2004. The
adaptive significance of temperature-dependent
sex determination
in an Australian agamid lizard. The University
of Sydney,
New South Wales, Australia. (Introductory Postgraduate
Seminar) |
| 19. |
Warner,
D. A. 2004. The
adaptive significance of temperature-dependent
sex determination
in an Australian agamid lizard. Evolution meetings,
Fort
Collins, Colorado, USA. (oral presentation) |
| 20. |
Warner,
D. A. 2005. The adaptive significance of
temperature-dependent
sex determination in the jacky
dragon. Macquarie University, Sydney,
Australia.
(Invited Departmental Seminar) |
| 21. |
Warner,
D. A. 2005. The
adaptive significance of temperature-dependent
sex determination
in an Australian agamid lizard. Fifth World Congress
of
Herpetology. Stellonbosch, South Africa. (Invited paper
at Sex Allocation Symposium) |
| 22. |
Warner,
D. A. 2005. The
adaptive significance of temperature-dependent
sex determination
in a short lived lizard (Amphibolurus muricatus:
Agamidae). Joint Ichthyology and Herpetology meeting.
Tampa,
Florida, USA. (oral presentation) |
| 23. |
Warner,
D. A. 2005. The adaptive significance of
temperature-dependent
sex determination in the jacky
dragon. Department of Zoology, Oklahoma
State
University. Stillwater, Oklahoma, USA. (Departmental
Seminar) |
| 24. |
Warner,
D. A. 2006.
Maternal diet affects offspring sex in a lizard with
temperature-dependent
sex determination. Australian Society of
Herpetologists. Healesville, Victoria,
Australia. (oral
presentation) |
| 25. |
Warner,
D. A. 2006.
Maternal diet influences offspring sex ratios in a lizard
with
temperature-dependent sex determination. Workshop on Sex Allocation
in
Reptiles Workshop. Wollongong, New South Wales, Australia. (oral
presentation) |
| 26. |
Warner,
D. A. 2006. The
adaptive significance of temperature-dependent sex determination
in
a short-lived lizard. Ecological Society of Australia. Wellington,
New
Zealand. (oral presentation) |
| 27. |
Warner,
D. A. 2006. The adaptive significance of temperature-dependent
sex determination.
Final PhD seminar and Postgraduate Excellence
Prize seminar. School of Biological
Sciences, University of Sydney,
Australia. (seminar) |
| 1997 | Study Abroad Scholarship (Ecuador and Galápagos Islands), International Field Trips in Biology Program, Iowa State University. |
| 1997 | Department of Zoology and Genetics Summer Internship, Iowa State University. |
| 1997 | Undergraduate Research in Herpetology Grant, Chicago Herpetological Society. |
| 1998 | J.N. “Ding” Darling Conservation Foundation Scholarship, Department of Animal Ecology, Iowa State University. |
| 1998 | Gary C. White Award, Department of Animal Ecology, Iowa State University. |
| 1998 | Study Abroad Scholarship (Costa Rica), International Field Trips in Biology Program, Iowa State University. |
| 1998 | Undergraduate Research in Herpetology Grant, Chicago Herpetological Society. |
| 1999 | Graduate Research in Herpetology Grant, Chicago Herpetological Society. |
| 1999 | Grant-in-Aid of Research, Sigma Xi. |
| 1999 | Graduate Research Development Project Grant, Graduate Student Assembly, Virginia Polytechnic Institute and State University. |
| 2000 | Honorable Mention for best student paper, annual meeting of The Virginia Academy of Sciences. |
| 2000 | Travel Grant, Graduate Student Assembly, Virginia Polytechnic Institute and State University. |
| 2001 | Master of Science Research Award in Biology, The Society of the Sigma Xi, Virginia Polytechnic Institute and State University. |
| 2001 | Henri Seibert Award for best student paper in ecology and evolution, Society for the Study of Amphibians and Reptiles. |
| 2003 | International Postgraduate Research Scholarship (IPRS), Australian Department of Education, Science, and Training. |
| 2003 | International Postgraduate Award (IPA), The University of Sydney. |
| 2003 | Grant-in-Aid of Research, Sigma Xi. |
| 2004 |
Gaige
Fund Award, American Society of Ichthyologists and
Herpetologists. |
| 2004 |
Travel
Grant, Australasian Society for the Study of
Animal Behaviour. |
| 2004 |
Postgraduate
Research Support Scheme, The University
of Sydney. |
| 2004 |
Norman
Wettenhall Foundation Grant. |
| 2004 |
Joyce
Vickery Scientific Research Fund, Linnean Society of New South
Wales. |
| 2004 |
Research
Grant, Australian Society of Herpetologists.
(honourable mention) |
| 2004 |
Travel
Grant, Australian Society of Herpetologists.
(declined) |
| 2005 |
Graduate
Student Research Grant, Chicago
Herpetological Society. |
| 2005 |
Grant-in-Aid,
Society for Integrative and Comparative
Biology. |
| 2005 |
Mary
Ethyl Read Research Grant, Royal Zoological
Society of New South Wales. |
| 2005 |
Student
Research Grant, Ecological Society of Australia. |
| 2005 |
Student
Travel Grant, Society for the Study of Amphibians and Reptiles. |
| 2005 |
Student
Travel Grant, American Society of Ichthyologists and
Herpetologist. |
| 2005 |
Postgraduate
Research Support Scheme, The University
of Sydney. |
| 2005 |
James
Kentley Memorial Scholarship, The University of
Sydney. |
| 2006 |
Student
Travel Grant, Ecological Society of
Australia. |
| 2006 |
Student
Travel Grant, Australian Society of Herpetologists. |
| 2006 |
Student
Research Grant, Australian Society of Herpetologists. |
| 2006 |
Postgraduate
Research Support Scheme, The University of Sydney. |
| 2006 |
Early Career
Researchers Support Program. Australian Research Council,
Environmental Futures
Network. |
| 2006 |
Best student
paper award, Meeting of the Australian Society of
Herpetologists. |
| 2006 |
Postgraduate
Excellence Award, School of Biological
Sciences, University of Sydney. |