Patrick J. Weatherhead

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Snake Research

The general areas of interest for the snake research done in my lab include mating systems and sexual selection, thermal ecology, the ecological role of snakes as avian nest predators, habitat selection, demography and population structure, and predator avoidance. Research on many of these topics is relevant both to basic behavior and ecology and to conservation biology. Field sites include locations in Canada, the Midwest and Texas.

Refereed Publications

78. Weatherhead, P.J., J.H. Sperry, G.L.F. Carfagno and G. Blouin-Demers. Latitudinal variation in thermal ecology of North American ratsnakes and its implications for the effect of climate warming on snakes. Journal of Thermal Biology, in press.

77. Weatherhead, P.J., G. Blouin-Demers and J.H. Sperry. Mortality patterns and the cost of reproduction in a northern population of ratsnakes, Elaphe obsoleta. Journal of Herpetology, in press.

76. Harvey, D.S. and P.J. Weatherhead. 2011. Thermal ecology of massasauga rattlesnakes (Sistrurus catenatus) near their northern range limit. Canadian Journal of Zoology 89: 60-68.

75. Harvey, D.S. and P.J. Weatherhead. 2010. Habitat selection as the mechanism for thermoregulation in a northern population of massasauga rattlesnakes (Sistrurus catenatus). Ecoscience 17: 411-419.

74. Sperry, J.H., G. Blouin-Demers, G.L.F. Carfagno and P.J. Weatherhead. 2010. Latitudinal variation in seasonal activity and mortality in ratsnakes (Elaphe obsoleta). Ecology 91:1860-1866.

73. Sperry, J.H. and P.J. Weatherhead. 2010. Ratsnakes and brushpiles: intended and unintended consequences of improving habitat for wildlife? American Midland Naturalist 163: 311-316..

72. Carfagno, G.L.F. and P.J. Weatherhead. 2009. Ratsnake response to bottomland flooding: implications for avian nest predation. Acta Herpetologica 4: 191-194.

71. Weatherhead, P.J., J.M. Knox, D.S. Harvey, D. Wynn, J. Chiucchi and H.L. Gibbs. 2009. Diet of Sistrurus catenatus in Ontario and Ohio: effects of body size and habitat. Journal of Herpetology 43: 693-697.

70. Sperry, J.H., L.K. Butler, L.M. Romero and P.J. Weatherhead. 2009. Effects of parasitic infection and radio-transmitters on condition, hematological characteristics and corticosterone concentrations in Texas ratsnakes. Journal of Zoology, 278: 100-107.

69. Sperry, J.H. and P.J. Weatherhead. 2009. Sex differences in behavior associated with sex-biased mortality in an oviparous snake species. Oikos 118: 627-633.

68. Weatherhead, P.J. and T. Madsen. 2009. Linking behavioral ecology to conservation objectives. Pages 149-171. In: Snakes: Ecology and Conservation (S.J. Mullin and R.A. Seigel, eds.), Cornell University Press.

67. Howes B.J., J.W. Brown, H.L. Gibbs, T.B. Herman, S.W. Mockford, K.A. Prior and P.J. Weatherhead. 2009. Directional gene flow patterns in disjunct populations of the black ratsnake (Pantheropis obsoletus) and the Blanding's turtle (Emydoidea blandingii). Conservation Genetics 10: 407-417.

66. Sperry, J.H. and P.J. Weatherhead. 2009. Does prey availability determine seasonal patterns of habitat selection in Texas ratsnakes? Journal of Herpetology 43: 55-64.

65. Sperry, J.H. and P.J. Weatherhead. 2008. Prey-mediated effects of drought on condition and survival of a terrestrial snake. Ecology 89: 2770-2776.

64. Blouin-Demers, G. and P.J. Weatherhead. 2008. Habitat use is linked to components of fitness through the temperature dependence of performance in ratsnakes (Elaphe obsoleta). Israeli Journal of Ecology and Evolution 54: 361-372.

63. Carfagno, G.L.F. and P.J. Weatherhead. 2008. Energetics and space use: intraspecific and interspecific comparisons of movements and home ranges of two Colubrid snakes. Journal of Animal Ecology 77: 416-424. [PDF]

62. Blouin-Demers, G., L.P.G. Bjorgan and P.J. Weatherhead. 2007. Changes in habitat use and movement patterns with body size in black ratsnakes (Elaphe obsoleta). Herpetologica 63: 421-429. [PDF]

61. Blouin-Demers, G. and P.J. Weatherhead. 2007. Allocation of offspring size and sex by female black ratsnakes. Oikos 10: 1759-1767. [PDF]

60. Row, J.R., G, Blouin-Demers and P.J. Weatherhead. 2007. Demographic effects of road mortality in black ratsnakes (Elaphe obsoleta). Biological Conservation 137: 117-124. [PDF]

59. Gibbs, H.L., S.J. Coreyl, G. Blouin-Demers, K.A. Prior, and P.J. Weatherhead. 2006. Hybridization between mtDNA-defined phylogeographic lineages of ratsnakes (Pantherophis sp.). Molecular Ecology 15:3755-3767 [PDF]

58. Richardson, M.L., P.J. Weatherhead, and J.D. Brawn. 2006. Habitat use and activity of prairie kingsnakes (Lampropeltis calligaster calligaster) in Illinois. Journal of Herpetology 40:424-428 [PDF]

57. Carfagno, G.L.F. and P.J. Weatherhead. 2006. Intraspecific and interspecific variation in use of forest-edge habitat by snakes. Canadian Journal of Zoology 84:1440-1452. [PDF]

56.  Weatherhead, P.J., K.J. Kissner, and S.J. Sommerer. 2006. Prenatal sex ratios and expression of sexually dimorphic traits in three snake species. Journal of Experimental Biology 305A: 603-609. [PDF]

55. Carfagno, G.L.F., E.J. Heske, and P.J. Weatherhead. 2006. Does mammalian prey abundance explain forest-edge use by snakes? Ecoscience 13:293-297. [PDF]

54. Harvey, D.S. and P.J. Weatherhead. 2006. A test of the hierarchical model of habitat selection using eastern massasauga rattlesnakes (Sistrurus c. catenatus). Biological Conservation 130:206-216. [PDF]

53. Harvey, D.S. and P.J. Weatherhead. 2006. Conservation implications of hibernation site selection by eastern massasauga rattlesnakes (Sistrurus c. catenatus) near the northern limit of their range. Journal of Herpetology 40:66-73. [PDF]

52. Kissner, K.J., P.J. Weatherhead, and H.L. Gibbs. 2005. Experimental assessment of ecological and phenotypic factors affecting male mating success and polyandry in northern watersnakes. Behavioral Ecology and Sociobiology 59:207-214. [PDF]

51. Kissner, K.J. and P.J. Weatherhead. 2005. Phenotypic effects on survival of neonatal northern watersnakes. Journal of Animal Ecology 74:259-265. [PDF]

50. Blouin-Demers, G., H.L. Gibbs and P.J. Weatherhead. 2005. Genetic evidence for sexual selection in black ratsnakes, Elaphe obsoleta. Animal Behaviour 69: 225-234. [PDF]

49. Weatherhead, P.J. and G. Blouin-Demers. 2004. Long-term effects of radiotelemetry on black ratsnakes. Wildlife Society Bulletin 32: 900-906. [PDF]

48. Blouin-Demers, G., P.J. Weatherhead and J.R. Row. 2004. Phenotypic consequences of nest-site selection in black rat snakes (Elaphe obsoleta). Canadian Journal of Zoology 82: 449-456. [PDF]

47. Brown, G.P. and P.J. Weatherhead. 2004. Sexual abstinence and the cost of reproduction in adult male water snakes, Nerodia sipedon. Oikos 104: 269-276. [PDF]

46. Weatherhead, P.J., G. Blouin-Demers, and K.M. Cavey. 2003. Seasonal and prey-size dietary patterns of black ratsnakes (Elaphe obsoleta obsoleta). American Midland Naturalist 150: 275-281. [PDF]

45. Blouin-Demers, G., P.J. Weatherhead, and H.A. McCracken. 2003. A test of the thermal coadaptation hypothesis with black rat snakes (Elaphe obsoleta) and northern water snakes (Nerodia sipedon). Journal of Thermal Biology 28: 331-340. [PDF]

44. Blouin-Demers, G. and P.J. Weatherhead. 2002. Implications of movement patterns for gene flow in black rat snakes (Elaphe obsoleta). Canadian Journal of Zoology 80: 1162-1172. [PDF]

43. Weatherhead, P.J., M.R. Prosser, H.L. Gibbs, and G.P. Brown. 2002. Male reproductive success and sexual selection in northern water snakes determined by microsatellite DNA analysis. Behavioral Ecology 13: 808-815. [PDF]

42. Prosser, M.R., P.J. Weatherhead, H.L. Gibbs, and G.P. Brown. 2002. Genetic analysis of the mating system and opportunity for sexual selection in northern water snakes (Nerodia sipedon). Behavioral Ecology 13: 800-807. [PDF]

41. Weatherhead, P.J., G. Blouin-Demers, and K.A. Prior. 2002. Synchronous variation and long-term trends in two populations of black rat snakes. Conservation Biology 16: 1602-1608. [PDF]

40. Blouin-Demers, G., and P.J. Weatherhead. 2002. Habitat-specific behavioural thermoregulation by black rat snakes (Elaphe obsoleta obsoleta). Oikos 97: 59-68. [PDF]

39. Blouin-Demers, G., K.A. Prior, and P.J. Weatherhead. 2002. Comparative demography of black rat snakes (Elaphe obsoleta) in Ontario and Maryland. Journal of Zoology 256: 1-10. [PDF]

38. Blouin-Demers, G. and P.J. Weatherhead. 2001. Thermal ecology of black rat snakes (Elaphe obsoleta) in a thermally challenging environment. Ecology 82: 3025-3043. [PDF]

37. Blouin-Demers, G. and P.J. Weatherhead. 2001. An experimental test of the link between foraging, habitat selection and thermoregulation in black rat snakes Elaphe obsoleta obsoleta. Journal of Animal Ecology 70: 1006-1013. [PDF]

36. Blouin-Demers, G. and P.J. Weatherhead. 2001. Habitat use by black rat snakes (Elaphe obsoleta obsoleta) in fragmented forests. Ecology 82: 2882-2896. [PDF]

35. Prior, K.A., G. Blouin-Demers, and P.J. Weatherhead. 2001. Sampling biases in demographic analyses of black rat snakes (Elaphe obsoleta). Herpetologica 57: 460-469.

34. Gibbs, H.L. and P.J. Weatherhead. 2001. Insights into population ecology and sexual selection in snakes through the application of DNA-based genetic markers. Journal of Heredity 92: 173-179. [PDF]

33. Blouin-Demers, G. and P.J. Weatherhead. 2000. A novel association between a beetle and a snake: parasitism of Elaphe obsoleta by Nicrophorus pustulatus. Ecoscience 7: 395-397. [PDF]

32. Parent, C. and P.J. Weatherhead. 2000. Behavioral and life history responses of eastern massasauga rattlesnakes (Sistrurus catenatus catenatus) to human disturbance. Oecologia 125: 170-178. [PDF]

31. Lougheed S.C., H.L. Gibbs, K.A. Prior, and P.J. Weatherhead. 2000. A comparison of RAPD versus microsatellite DNA markers in population studies of the massasauga rattlesnake. Journal of Heredity 91: 458-463. [PDF]

30. Kissner, K.J., G. Blouin-Demers, and P.J. Weatherhead. 2000. Sexual dimorphism in malodorousness of musk secretions of snakes. Journal of Herpetology 34: 491-493.

29. Blouin-Demers, G., P.J. Weatherhead, C.M. Shilton, C.E. Parent, and G.P. Brown. 2000. Use of inhalant anesthetics in three snake species. Contemporary Herpetology ISSN 1094-2246. [PDF]

28. Blouin-Demers, G., K.J. Kissner, and P.J. Weatherhead. 2000. Plasticity in preferred body temperature of young snakes in response to temperature during development. Copeia 2000: 841-845. [PDF]

27. Brown, G.P. and P.J. Weatherhead. 2000. Thermal ecology and sexual size dimorphism in northern water snakes, Nerodia sipedon. Ecological Monographs 70: 311-330. [PDF]

26. Blouin-Demers, G., K.A. Prior and P.J. Weatherhead. 2000. Patterns of variation in spring emergence by black rat snakes (Elaphe obsoleta obsoleta). Herpetologica 56: 175-188.

25. Lougheed, S.C., H.L. Gibbs, K.A. Prior and P.J. Weatherhead. 1999. Hierarchical patterns of genetic population structure in black rat snakes (Elaphe obsoleta obsoleta) as revealed by microsatellite DNA analysis. Evolution 53: 1995-2001.

24. Brown, G.P. and P.J. Weatherhead. 1999. Demography and sexual size dimorphism in northern water snakes, Nerodia sipedon. Canadian Journal of Zoology 77: 1358-1366.

23. Weatherhead, P.J., G.P. Brown, M.R. Prosser and K.J. Kissner. 1999. Factors affecting neonate size variation in northern water snakes, Nerodia sipedon. Journal of Herpetology 33: 577-589.

22. Brown, G.P. and P.J. Weatherhead. 1999. Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon). Behavioral Ecology and Sociobiology 47: 9-16. [PDF]

21. Brown, G.P. and P.J. Weatherhead. 1999. Growth and sexual size dimorphism in northern water snakes (Nerodia sipedon). Copeia 1999: 723-732.

20. Prosser, M.R., H.L. Gibbs and P.J. Weatherhead. 1999. Microgeographic population genetic structure in the northern water snake, Nerodia sipedon sipedon detected using microsatellite DNA loci. Molecular Ecology 8: 329-333.

19. Weatherhead, P.J., G.P. Brown, M.R. Prosser and K.J. Kissner. 1998. Variation in offspring sex ratios in the northern water snake, Nerodia sipedon. Canadian Journal of Zoology 76: 2200-2206.

18. Gibbs, H.L., K.A. Prior, P.J. Weatherhead and G. Johnson. 1997. Genetic structure of populations of the threatened eastern massasauga rattlesnake, Sistrurus c. catenatus: evidence from microsatellite DNA markers. Molecular Ecology 6: 1123-1132. [PDF]

17. Prior, K.A., H.L. Gibbs and P.J. Weatherhead. 1997. Population genetic structure in the black rat snake: implications for management. Conservation Biology 11: 1147-1158. [PDF]

16. Brown, G.P. and P.J. Weatherhead. 1997. Effects of reproduction on survival and growth of female northern water snakes, Nerodia sipedon. Canadian Journal of Zoology 75: 424-432.

15. Weatherhead, P.J. and G.P. Brown. 1996. Measurement versus estimation of condition in snakes. Canadian Journal of Zoology 74: 1617-1621.

14. Prior, K.A. and P.J. Weatherhead. 1996. Habitat features of black rat snake hibernacula in Ontario. Journal of Herpetology 30: 211-218.

13. Scribner, S. and P.J. Weatherhead. 1995. Locomotion and antipredator behaviour in three species of semi-aquatic snakes. Canadian Journal of Zoology 73: 321- 329.

12. Weatherhead, P.J., F.E. Barry, G.P. Brown and M.R.L. Forbes. 1995. Sex ratios, mating behavior and sexual size dimorphism of the northern water snake, Nerodia sipedon. Behavioral Ecology and Sociobiology 36: 301-311.

11.  Prior, K.A. and P.J. Weatherhead. 1994. Response of free-ranging eastern massasauga rattlesnakes to human disturbance. Journal of Herpetology 28: 255-257.

10. Gibbs, H.L., K.A. Prior and P.J. Weatherhead. 1994. Genetic analysis of populations of threatened snake species using RAPD markers. Molecular Ecology 3: 329-337.

9.  Weatherhead, P.J. and I.C. Robertson. 1992. Thermal constraints on swimming performance and escape response of northern water snakes (Nerodia sipedon). Canadian Journal of Zoology 70: 94-98.

8.  Robertson, I.C. and P.J. Weatherhead. 1992. The role of temperature on microhabitat selection by northern water snakes (Nerodia sipedon). Canadian Journal of Zoology 70: 417-422.

7.  Barry, F.E., P.J. Weatherhead and D.P. Philipp. 1992. Multiple paternity in a wild population of northern water snakes, Nerodia sipedon. Behavioral Ecology and Sociobiology 30: 193-199.

6.  Weatherhead, P.J. and K.A. Prior. 1992. Preliminary observations of habitat use and movements of the eastern massasauga rattlesnake (Sistrurus c. catenatus). Journal of Herpetology 26: 447-452.

5.  Weatherhead, P.J. and I.C. Robertson. 1990. Homing to food by black rat snakes (Elaphe obsoleta). Copeia 1990: 1164-1165.

4.  Weatherhead, P.J. and D.J. Hoysak. 1989. Spatial and activity patterns of black rat snakes (Elaphe obsoleta) from radiotelemetry and recapture data. Canadian Journal of Zoology 67: 463-468.

3.  Weatherhead, P.J. 1989. Temporal and thermal aspects of hibernation of black rat snakes (Elaphe obsoleta) in Ontario. Canadian Journal of Zoology 67: 2332-2335.

2.  Weatherhead, P.J. and M.B. Charland. 1985. Habitat selection in an Ontario population of the snake, Elaphe obsoleta. Journal of Herpetology 19: 12-19.

1.  Weatherhead, P.J. and F.W. Anderka. 1984. An improved radio transmitter and implantation technique for snakes. Journal of Herpetology 18: 264-269.