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Preprints & Manuscripts Submitted

Dantzer B, Mabry KE, Bernhardt JR, Cox RM, Francis CD, Ghalambor CK, Hoke KL, Jha S, Ketterson E, Levis NA, McCain KM, Patricelli GL, Paull SH, Pinter-Wollman N, Șafran RJ, Schwartz TS, Throop HL, Zaman L, Martin LB. Understanding organisms using ecological observatory networks. Submitted December 2022

Halhed A, Petrullo L, Boutin S, Dantzer B, McAdam AG, Wu M, Cottenie K. Spatiotemporal variability of core and non-core microbial taxa in North American red squirrel (Tamiasciurus hudsonicus) fecal microbiomes. Submitted December 2022

Devitz A-C, Dantzer B. Behavioral variation among fox squirrel (Sciurus niger) populations in urban and suburban environments. Resubmitted November 2022

Rubi TL, Prado JR, Knowles LL, Dantzer B. Patterns of genetic and epigenetic diversity across a range expansion in the white-footed mouse (Peromyscus leucopus). Resubmitted August 2022

Published Manuscripts

93) Dantzer B. 2023. Frank Beach Award Winner: The centrality of the hypothalamic-pituitary-adrenal axis in dealing with environmental change across temporal scales. Hormones & Behavior in press

92) Petrullo L, Boutin S, Lane JE, McAdam AG, Dantzer B. 2023. Phenotype-environment mismatch errors enhance lifetime fitness in wild red squirrels. Science

91) Sabol AC, Close WL, Petrullo L, Lambert CT, Keane B, Solomon NG, Schloss PD, Dantzer B. 2023. Sociality does not predict oral microbiome composition or diversity in free-living prairie voles. Animal Behaviour in press December 2022

90) Walmsley SF, Boutin S, Dantzer B, Lane JE, Coltman DW, McAdam AG. 2023. Benefits of living closer to kin vary by genealogical relationship in a territorial mammal. Proceedings of the Royal Society B 290, 20221569.

89) Petrullo L, Delaney DM, Boutin S, McAdam AG, Lane JE, Palme R, Dantzer B. 2022. The glucocorticoid response to environmental change is not specific to agents of natural selection in wild red squirrels. Hormones & Behavior 146, 105262.

88) Webber Q, Dantzer B, Lane JE, Boutin S, McAdam AG. 2022. Density-dependent plasticity in territoriality revealed using social network analysis. Journal of Animal Ecology 92, 207-221.

87) Gaidica M, Dantzer B. 2022. An implantable neurophysiology platform: broadening research capabilities in free-living and non-traditional animals. Frontiers in Neural Circuits 16, 940989.

86) Haines JA, Delaney DM, Wishart AE, McAdam AG, Coltman DW, Lane JE, Dantzer B, Boutin S. 2022. Sex-specific effects of capital resources on reproductive timing and success in red squirrels. Behavioral Ecology & Sociobiology 76, 142.

85) Santicchia F, Wauters LA, Dantzer B, Palme R, Tranquillo C, Preatoni D, Martinoli A. 2022. Native species exhibit physiological habituation to invaders: a reason for hope. Proceedings of the Royal Society B 289, 20221022.

84) Studd EK, Peers MJL, Menzies AK, Derbyshire R, Majchrzak YN, Seguin JL, Murray DL, Dantzer B, Lane JE,  McAdam AG, Humphries MM, Boutin S. 2022. Behavioral adjustments of predators and prey to wind speed in the boreal forest. Oecologia 200, 349-358.

83) Dantzer B, Newman AEN. 2022. Expanding the frame around social dynamics and glucocorticoids: from hierarchies within the nest to competitive interactions among species. Hormones & Behavior 144, 105204.

82) Martinig, AR, Karst HHJ, Siracusa ER, Studd EK, McAdam AG, Dantzer B, Delaney DM, Lane JE, Pokharel P, Boutin S. 2022. Animal personality: a comparison of standardized assays and focal observations in North American red squirrels. Animal Behaviour 190, 221-232.

81) Petrullo L, Ren T, Wu M, Boonstra R, Palme R, Boutin S, McAdam AG, Dantzer B. 2022. Glucocorticoids coordinate changes in gut microbiome composition in wild North American red squirrels. Scientific Reports 12, 2605.

80) Tranquillo C, Villa F, Wauters LA, Dantzer B, Palme R, Preatoni D, Martinoli A, Santicchia F. 2022. Physiological stress and spatio-temporal fluctuations of food abundance and population density in Eurasian red squirrels. Hystrix 33, 58-64.

79) Hacket Pain A, Foest JJ, Pearse IS, … Dantzer B… Ascoli D (89 total authors). 2022. MASTREE+: time-series of plant reproductive effort from six continents. Global Change Biology 28, 3066-3082.

78) Santicchia F, Wauters LA, Tranquillo C, Villa F, Dantzer B, Palme R, Preatoni D, Martinoli A. 2022. Invasive alien species as an environmental stressor and its effects on coping style in a native competitor, the Eurasian red squirrel. Hormones & Behavior 140, 105127.

77) Prado JR, Rubi TL, Baumgartner J, Hoffman S, Dantzer B, Knowles LL. 2022. Postglacial colonization in the Great Lakes Region by the white-footed mouse (Peromyscus leucopus): conflicts between genomic and field data. Journal of Mammalogy 103, 243-254.

76) Dantzer B, Boutin S, Lane JE, McAdam AG. 2022. Integrative studies of the effects of mothers on offspring: an example from wild North American red squirrels. In Patterns of Parental Behavior: from Animal Science to Comparative Ethology and Neuroscience (Edited by Gabriela González Mariscal).

75) McAdam AG, Webber QMR, Dantzer B, Lane JE, Boutin S. 2022. Social effects on annual fitness in red squirrels. Journal of Heredity, esab051

74) Kittendorf A, Dantzer B. 2021. Urban fox squirrels exhibit tolerance to humans but respond to stimuli from natural predators. Ethology 127, 697-709.

73) Westrick SE, van Kesteren F, Boutin S, Lane JE, McAdam AG, Dantzer B. 2021. Maternal glucocorticoids have minimal effects on HPA axis activity and behavior of juvenile wild North American red squirrels. Journal of Experimental Biology 224, jeb236620.

72) Hare AJ, Newman AEM, Dantzer B, Lane JE, Boutin S, Coltman DW, McAdam AG. 2021. An independent experiment does not support stress-mediated kin discrimination through red squirrel vocalizations. Animal Behaviour 176, 185-192.

71) Cooke, S.J., J.N. Bergman, C.L. Madliger, R.L. Cramp, J. Beardall, G.P. Burness, T.D. Clark, B. Dantzer, E. de la Barrera, N.A. Fangue, C.E. Franklin, A. Fuller, L.A. Hawkes, K.R. Hultine, K.E. Hunt, O.P. Love, H.A. MacMillan, J.W. Mandelman, F.C. Mark, L.B. Martin, A.E.M. Newman, A.B. Nicotra, G.D. Raby, S.A. Robinson, Y. Ropert-Coudert, J.L. Rummer, F. Seebacher, A.E. Todgham, S. Tomlinson, S.L. Chown.  2021. One hundred research questions in conservation physiology for generating actionable evidence to inform conservation policy and practice. Conservation Physiology  9, coab009.

70) Underhill, V, Pandelis GG, Papuga J, Sabol AC, Rife A, Rubi T, Hoffman SMG, Dantzer B. 2021. Personality and behavioral syndromes in two Peromyscus species: presence, lack of state-dependence, and lack of association with home range size. Behavioral Ecology and Sociobiology 75, 9.

69) Siracusa ER, Boutin S, Dantzer B, Lane JE, Coltman DW, McAdam AG. 2021. Familiar neighbors, but not relatives, enhance fitness in a territorial mammal. Current Biology 31, 1-8.

68) Martinig A, Mathot K, Lane JE, Dantzer B, Boutin S. 2021. Selective disappearance does not underlie age-related changes in trait repeatability in red squirrels. Behavioral Ecology 32, 306-315.

67) Dantzer B, McAdam AG, Humphries MM, Lane JE, Boutin S. 2020. Decoupling the effects of food and density on life history plasticity of wild animals using field experiments: insights from the steward who sits in the shadow of its tail, the North American red squirrel. Journal of Animal Ecology 89, 2397-2414.

66) Menzies A, Studd E, Majchrzak Y, Peers M, Boutin S, Dantzer B, Lane J, McAdam A, Humphries MM. 2020. Body temperature, heart rate, and activity patterns of two boreal homeotherms in winter: homestasis, allostasis, and ecological coexistence. Functional Ecology 34, 2292-2301.

65) Haines JA, Nason SE, Skurdal AMM, Bourchier T, Boutin S, Taylor RW, McAdam AG, Lane JE, Kelley AD, Humphries MM, Gorrell JC, Dantzer B, Coltman DW, Hämäläinen A. 2020. Sex- and context-specific associations between personality and a measure of fitness but no link with life history traits. Animal Behaviour 167, 23-39.

64) Westrick SE, Taylor RW, Boutin S, Lane JE, McAdam AG, Dantzer B. 2020. Attentive red squirrel mothers have faster-growing pups and higher lifetime reproductive success. Behavioral Ecology & Sociobiology 74, 72.

63) Romeo C, Wauters L, Santicchia F, Dantzer B, Palme R, Martionli A, Ferrari N. 2020. Complex relationships between physiological stress and endoparasite infections in natural populations. Current Zoology 66, 449-457.

62) Gaidica M, Dantzer B. 2020. Quantifying the autonomic response to stressors – one way to expand the definition of “stress” in animals. Integrative & Comparative Biology 60, 113-125.

61) Studd, EK, Menzies AK, Siracusa ER, Dantzer B, Lane JE, McAdam AG, Boutin S, Humphries MM. 2020. Optimisation of energetic and reproductive gains explains behavioral responses to environmental variation across seasons and years. Ecology Letters 23, 841-850.

60) Cooke SJ, Madliger CL, … Dantzer B, … Todgham AE. 2020. Reframing conservation physiology to be more inclusive, integrative, relevant and forward-looking: reflections and a horizon scan. Conservation Physiology 8, coaa016.

59) Dantzer B, van Kesteren F, Westrick SE, Boutin S, McAdam AG, Lane JE, Gillespie R, Majer A, Haussmann M, Monaghan P. 2020. Maternal stress promotes offspring growth without inducing oxidative stress or shortening telomeres in wild red squirrels. Journal of Experimental Biology 223, jeb212373.

58) Sabol AC, Lambert CT, Keane B, Solomon NG, Dantzer B. 2020. How does individual variation in sociality influence fitness in prairie voles? Animal Behaviour 163, 39-49

57) Hendrix J, Fisher D, Martinis AR, Boutin S, Dantzer B, Lane JE, McAdam AG. 2020. Territory acquisition mediates the influence of predators and climate on juvenile red squirrel survival. Journal of Animal Ecology. 89, 1408-1418.

56) Martinig AR, McAdam AG, Dantzer B, Lane JE, Coltman DW, Boutin S. 2020. Immigrant male red squirrels experience benefits whereas females pay a fitness cost after settlement. Ecology Letters 23, 430-438.

55) Santicchia F, Wauters LA, Dantzer B, Westrick SE, Ferrari N, Romeo C, Palme R, Preatoni DG, Martinoli A. 2020. Relationships between personality traits and the physiological stress response in a wild mammal. Current Zoology 66, 197-204.

54) Rubi TL, Knowles LL, Dantzer B. 2020. Museum epigenomics: characterizing cytosine methylation in historic museum specimens. Molecular Ecology Resources 20, 1161-1170.

53) Westrick SE, van Kesteren F, Palme R, Boonstra R, Lane JE, Boutin S, McAdam AG, Dantzer B. 2019. Stress activity is not predictive of coping style in North American red squirrels. Behavioral Ecology and Sociobiology 73, 113.

52) Guindre-Parker S, McAdam AG, van Kesteren F, Palme R, Boonstra R, Boutin S, Lane JE, Dantzer B. 2019. Individual variation in phenotypic plasticity of the stress axis. Biology Letters 15, 20190260.

51) van Kesteren F, Delehanty B, Westrick SE, Palme R, Boonstra R, Lane JE, Boutin S, McAdam AG, Dantzer B. 2019. Experimental increases in stress hormones alter function of the neuroendocrine stress axis in wild red squirrels without negatively impacting survival and reproduction. Physiological & Biochemical Zoology 92, 445-458.

50) Sehrsweeney M, Wilson DR, Bain M, Boutin S, Lane JE, McAdam AG, Dantzer B. 2019. Is physiological stress state reflected in acoustic structure of vocalizations? An experimental test in wild North American red squirrels. Behavioral Ecology 30, 1030-1040.

49) Fisher DN, Wilson AJ, Boutin S, Dantzer B, Lane JE, Coltman DW, Gorrell JC, McAdam AG. 2019. Indirect effects of territorial neighbours on the timing of spring breeding may counteract changes in selection in North American red squirrels. Journal of Evolutionary Biology Accepted [preprint]

48) McAdam AG, Boutin S, Dantzer B, Lane JE. 2019. Seed masting causes fluctuations in optimum litter size and lag load in a seed predator. The American Naturalist 194, 574-589.

47) Dantzer B, Dubuc C, Braga Goncalves I, Cram DL, Bennett NC, Ganswindt A, Duncan C, Gaynor D, Clutton-Brock TH. 2019. Maternal stress increases cooperative behaviour of offspring in wild meerkats. Philosophical Transactions of the Royal Society B 374, 20180117.

See the press release

46) Siracusa E, Wilson DR, Studd EK, Boutin S, Humphries MM, Dantzer B, Lane JE, McAdam AG. 2019. Red squirrels mitigate costs of territory defence through social plasticity. Animal Behaviour 151, 29-42.

45) Fisher D, Haines J, Boutin S, Dantzer B, Lane JE, Coltman DW, McAdam AG. 2019. Indirect effects on fitness between individuals that have never met via an extended phenotype. Ecology Letters 22, 697-706.

See the press release, Vice, CBC, National Post

44) Sabol AE, Solomon NG, Dantzer B. 2018. How to study socially monogamous behavior in secretive animals? Using social network analyses and automated tracking systems to study the social behavior of prairie voles. Frontiers in Ecology and Evolution 6, 178.

43) Robertson JG, Boutin S, Humphries MM, Dantzer B, Lane JE, McAdam AG. 2018. Individual variation in the dear enemy phenomenon via territorial vocalizations in red squirrels. Behaviour 155, 1073-1096.

42) Wishart A, Williams C, McAdam AG, Boutin S, Dantzer B, Humphries MM, Coltman DW, Lane JE. 2018. Is biasing offspring sex ratio adaptive? A test of Fisher’s principle across multiple generations of a wild mammal in a fluctuating environment. Proceedings of the Royal Society B 285, 20181251.

41) Cram DL, Monaghan P, Gillespie R, Dantzer B, Duncan C, Spence-Jones H, Clutton-Brock T. 2018. Rank-related contrasts in longevity arise from extra-group excursions not delayed senescence in a cooperative mammal. Current Biology 28, 2934-2939.

40) Santicchia F, Dantzer B, van Kesteren F, Palme R, Martinoli A, Ferrari N, Wauters L. 2018. Stress in biological invasions: alien grey squirrels increase physiological stress in native Eurasian red squirrels. Journal of Animal Ecology 87, 1342-1352.

39) Raulo, A, Dantzer, B. 2018. The changing role of the effects of glucocorticoid stress hormones on pro-social behaviour during the evolution of vertebrate social behaviour. Ecology & Evolution 8, 7697-7716.

38) Sheriff MJ, Dantzer B, Love OP, Orrock JL. 2018. Error management theory and the evolution of maternal stress effects. Ecology & Evolution 8, 6473-6482.

37) Haines JL, Coltman DW, Dantzer B, Gorrell JC, Humphries MM, Lane JE, McAdam AG, Boutin S. 2018. Sexually-selected infanticide by male red squirrels in advance of a mast year. Ecology 99, 1242-1244.

36) Siracusa E, Morandini M, Boutin S, Humphries MM, Dantzer B, Lane JE, McAdam AG. 2017. Red squirrel territorial vocalizations deter intrusions by conspecific rivals. Behaviour 154, 1259-1273.

35) Ren T, Boutin S, Humphries MM, Dantzer B, Gorrell JC, Coltman DW, McAdam AG, Wu M. 2017. Seasonal, spatial and maternal effects on gut microbiome in wild red squirrels. Microbiome 5, 163.

34) Cooper EB, Taylor RW, Kelley AD, Martinig AR, Boutin S, Humphries MM, Dantzer B, Lane JE, McAdam AG. 2017. Personality is correlated with natal dispersal in North American red squirrels (Tamiasciurus hudsonicus). Behaviour 154, 939-961.

33) Siracusa E, Boutin S, Humphries MM, Gorrell JC, Coltman DW, Dantzer B, Lane JE, McAdam AG. 2017. Familiarity with neighbours affects intrusion risk in territorial red squirrels (Tamiasciurus hudsonicus). Animal Behaviour 133, 11-20

32) Dantzer B, Braga Goncalves I, Spence-Jones HC, Bennett NC, Heistermann M, Ganswindt A, Gaynor D, Dubuc C, Gaynor D, Manser MB, Clutton-Brock T. 2017. The influence of stress hormones and aggression on cooperative behavior in wild meerkats. Proceedings of the Royal Society Series B 284, 20171248.

See the press release

31) Hämäläinen A, McAdam AG, Dantzer B, Lane JE, Haines JA, Humphries MM, Boutin S. 2017. Fitness consequences of peak reproductive effort in a resource pulse system. Scientific Reports 7, 9335.

30) Fisher DN, Boutin S, Dantzer B, Humphries MM, Lane JE, McAdam AG. 2017. Multilevel and sex-specific selection on competitive traits in North American red squirrels. Evolution 71, 1841-1854.

29) Sheriff MJ, Bell A, Boonstra R, Dantzer B, Lavergne S, McGhee KE, MacLeod KJ, Winandy L, Zimmer C, Love OP. Integrating ecological and evolutionary context in the study of maternal stress. Integrative and Comparative Biology 57, 437-449.

28) Dantzer B, Rubenstein DR. 2017. Introduction to symposium: the developmental and proximate mechanisms causing individual variation in cooperative behavior. Integrative and Comparative Biology 57, 560-565.

27) Dantzer B, Swanson EM. 2017. Does hormonal pleiotropy constrain the independent evolution of performance and life history traits? Integrative and Comparative Biology 57, 372-384.

26) Dantzer B, Bennett NC, Clutton-Brock T. 2017. Social conflict and costs of cooperation in meerkats are reflected in measures of stress hormones. Behavioral Ecology 28, 1131-1141.

25) Dantzer B, Garratt M. 2017. Sex differences in telomeres and lifespan in Soay sheep: From the beginning to the end. Molecular Ecology 26, 3090-3092 (invited commentary).

24) Dubuc C, English S, Thavarajah N, Dantzer B, Sharp S, Spence-Jones H, Gaynor D, Clutton-Brock T. 2017. Food availability increases eviction in a cooperative mammal. Biology Letters 20160961.

23) Dantzer B, Santicchia F, van Kesteren F, Palme R, Martinoli A, Wauters LA. 2016. Measurement of fecal glucocorticoid metabolite levels in Eurasian red squirrels (Sciurus vulgaris): effects of temporary capture, sex, reproductive condition, and season. Journal of Mammalogy 97, 1385-1398.

22) Dantzer, B, Westrick, SE, van Kesteren, F. 2016. Relationships between endocrine traits and life histories in wild animals: insights, problems, and potential pitfalls. Integrative & Comparative Biology 56, 185-197.

21) Goncalves, IB, Heistermann, M, Santema, P, Dantzer, B, Mausbach, J, Ganswindt, A, Manser, MB. 2016. Validation of a fecal glucocorticoid assay to assess adrenocortical activity in meerkats using physiological and biological stimuli. PLoS One 11, e0153161.

20) Benson-Amram, S, Dantzer, B, Stricker, G, Swanson, EM, Holekamp, KE. 2016. Brain size predicts problem-solving ability in mammalian carnivores. PNAS 113, 2532-2537.

See the videopress release, The New York TimesNational Science FoundationScientific AmericanBrainDecoderPopular ScienceNew ScientistARS Technica.

19) Speakman, JR, Blount, JD, Bronikowski, AM, Buffenstein, R, Isaksson, C, Kirkwood, TBL, Monaghan, P, Ozanne, SE, Beulieu, M, Briga, M, Carr, SK, Christensen, LL, Cocheme, HM, Cram, DL, Dantzer, B, Harper, JM, Jurk, D, King, A, Noguera, JC, Salin, K, Sild, E, Simons, MJP, Smith, S, Stier, A, Tobler, M, Vitikainen, E, Peaker, M, Selman, C. 2015. Oxidative stress and life histories: a consensus view on several unresolved issues. Ecology & Evolution 5, 5745-5757.

18) Dantzer, B, Fletcher, QE. 2015. Telomeres shorten more slowly in slow-aging wild animals than in fast-aging ones. Experimental Gerontology 71, 38-47. *Shared first authorship

17) Fletcher, QE*, Dantzer, B*, Boonstra, R. 2015. The impact of reproduction on the hippocampal-hypothalamic-pituitary-adrenal stress axis of free-living male northern red backed voles (Myodes rutilus).  General and Comparative Endocrinology 224, 136-147*Shared first authorship

16) Holekamp, KE, Dantzer B, Stricker G, Yoshida KCS, Benson-Amram, S. 2015. Brains, brawn and sociality: a hyaena’s tail. Animal Behaviour 103, 237-248.

15) Dantzer, B. 2015. Mothers shape ecological communities. Science 347, 822-823 (invited perspective).

14) Dantzer, B, Fletcher, QE, Boonstra, R, Sheriff, MJ. 2014. Stress physiology – a transparent or opaque window into the status, management, and conservation of a species? Conservation Physiology 2, 1-18

13) Swanson, EM, Dantzer, B*. 2014. Insulin-like growth factor-1 is associated with life history variation across Mammalia. Proceedings of the Royal Society Series B 281, 20132458. *Shared first authorship

12) Boonstra, R*, Dantzer, B*, Delehanty, B*, Fletcher, QE*, Sheriff, MJ*. 2014. Equipped for life in the boreal forest: the role of the stress axis in mammals. Arctic 67, 82-97. *All authors contributed equally

11) Dantzer, B, Newman, AEM, Boonstra, R, Palme, R, Boutin, S, Humphries, MM, McAdam, AG. 2013. Density triggers maternal hormones that increase adaptive offspring growth in a wild mammal. Science 340, 1215-1217

Highlighted by Ed Yong blog,, Nature, CBC coverage, Quirks and Quarks Radio, newspapers

10) Creel, SR*, Dantzer, B*, Goymann, W*, Rubenstein, D*. 2013. The ecology of stress: effects of the social environment. Functional Ecology (invited submission) 27, 66-80. *All authors contributed equally

9) Dantzer, B, Boutin, S, Humphries, MM, McAdam, AG. 2012. Behavioral responses of territorial red squirrels to natural and experimental variation in population density. Behavioral Ecology and Sociobiology 66:865-878

8) Taylor, RW, Boon, AK, Dantzer, B, Reale, D, Humphries, MM, Boutin, S, Gorrell, JC, Coltman, DW, Boutin, S, Humphries, MM, McAdam, AG. 2012. Low heritabilities, but genetic and maternal correlations between red squirrel behaviours. Journal of Evolutionary Biology 25:614-624.

7) Dantzer, B*, Swanson, EM.* 2012. Mediation of vertebrate life-histories via insulin-like growth factor-1. Biological Reviews 87:414-429. *Shared first authorship

6) Dantzer, B, McAdam, AG, Palme, R, Boutin, S, Boonstra, R. 2011. How does diet affect fecal steroid hormone metabolite concentrations? An experimental test in red squirrels. General and Comparative Endocrinology 174:124-131.

5) Sheriff, MJ, Dantzer, B, Delehanty, B, Palme, R, Boonstra, R. 2011. Measuring stress in wildlife: techniques for measuring glucocorticoids. Oecologia 166:869-897.

4) Dantzer, B, McAdam, AG, Palme, R, Boutin, S, Humphries, MM, Boonstra, R. 2011. Maternal androgens and behaviour in free-ranging North American red squirrels. Animal Behaviour 81:469-479.

3) Dantzer, B, McAdam, AG, Palme, R, Fletcher, QE, Boutin, S, Humphries, MM, Boonstra, R. 2010. Fecal cortisol metabolites in free-ranging North American red squirrels: assay validation and the effects of reproductive condition. General and Comparative Endocrinology 167: 279-286.

2) Dantzer, BJ & Jaeger, RG. 2007. Detection of the sexual identity of conspecifics through volatile chemical signals in a territorial salamander. Ethology 113:214-222.

1) Dantzer, BJ & Jaeger, RG. 2007. Male red-backed salamanders can determine the reproductive status of females through volatile chemical signals. Herpetologica 63:176-183.