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Dr Daniel A. Warner

Post-Doctoral Fellow
Department of Ecology,
Evolution and Organismal Biology
Iowa State University
Ames, Iowa 50011, USA

e-mail: dwarner@iastate.edu

http://www.public.iastate.edu/~fjanzen/

Education

Publications

Abstracts and Presentations

Grants and Awards



Past Research

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 determination

My 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.

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.


2jackies

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.

nesting
Female jacky dragon with radio transmitter digging nest
nest
Average clutch of eggs in nest before being buried


1998-2003:  Environmental and maternal influences on phenotypes and survival of hatchlings lizards

 

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.
 






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.

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Education

Iowa State University, Ames, Iowa, USA.
Bachelor of Science in Animal Ecology.  May 1998.
Advisor:  Dr. Fredric Janzen

Virginia 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

goanna
Hatchling Varanus rosenbergi from Royal National Park


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Publications

Some papers are available as pdf files. To read these you will need Adobe Acrobat Reader.
If you have any difficulties downloading files please contact Mel.

 
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 undulatusHerpetological 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 undulatusHerpetologica 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. (doi: 10.1111/j.1471-8286.2006.01644.x)
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.


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Abstracts and Presentations
 
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)

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Grants and Awards

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.
Last updated: June 2006

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