Recent studies of animal personality have focused on its proximate causation and ecological and evolutionary significance in particular, but the question of its development was largely overlooked. The attributes of personality are defined as between-individual differences in behaviour, which are consistent over time (differential consistency) and contexts (contextual generality) and both can be affected by development. We assessed several candidates for personality variables measured in various tests with different contexts over several life-stages (juveniles, older juveniles, subadults and adults) in the Northern common boa. Variables describing foraging/feeding decision and some of the defensive behaviours expressed as individual average values are highly repeatable and consistent. We found two main personality axes-one associated with foraging/feeding and the speed of decision, the other reflecting agonistic behaviour. Intensity of behaviour in the feeding context changes during development, but the level of agonistic behaviour remains the same. The juveniles and adults have a similar personality structure, but there is a period of structural change of behaviour during the second year of life (subadults). These results require a new theoretical model to explain the selection pressures resulting in this developmental pattern of personality. We also studied the proximate factors and their relationship to behavioural characteristics. Physiological parameters (heart and breath rate stress response) measured in adults clustered with variables concerning the agonistic behavioural profile, while no relationship between the juvenile/adult body size and personality concerning feeding/foraging and the agonistic behavioural profile was found. Our study suggests that it is important for studies of personality development to focus on both the structural and differential consistency, because even though behaviour is differentially consistent, the structure can change.

Department of Zoology Faculty of Science Charles University Prague Czech Republic

National Institute of Mental Health Klecany Czech Republic

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Dall SRX, Houston AI, McNamara JM. The behavioural ecology of personality: consistent individual differences from an adaptive perspective. Ecology Letters. 2004;7(8):734–9. 10.1111/j.1461-0248.2004.00618.x DOI

Bell AM, Hankison SJ, Laskowski KL. The repeatability of behaviour: a meta-analysis. Animal Behaviour. 2009;77(4):771–83. 10.1016/j.anbehav.2008.12.022 . PubMed DOI PMC

Stamps J, Groothuis TGG. The development of animal personality: relevance, concepts and perspectives. Biological Reviews. 2010;85(2):301–25. 10.1111/j.1469-185X.2009.00103.x . PubMed DOI

Réale D, Reader SM, Sol D, McDougall PT, Dingemanse NJ. Integrating animal temperament within ecology and evolution. Biological Reviews. 2007;82(2):291–318. 10.1111/j.1469-185X.2007.00010.x . PubMed DOI

Gosling SD. From mice to men: What can we learn about personality from animal research? Psychological Bulletin. 2001;127(1):45–86. 10.1037//0033-2909.127.1.45 . PubMed DOI

Sih A, Bell AM, Johnson JC, Ziemba RE. Behavioral syndromes: An integrative overview. Quarterly Review of Biology. 2004;79(3):241–77. 10.1086/422893 . PubMed DOI

Réale D, Dingemanse N. Animal Personality: eLS.; 2012.

Cote J, Clobert J. Social personalities influence natal dispersal in a lizard. Proceedings of the Royal Society B-Biological Sciences. 2007;274(1608):383–90. 10.1098/rspb.2006.3734 PubMed DOI PMC

Li H, Holleley CE, Elphick M, Georges A, Shine R. The behavioural consequences of sex reversal in dragons. Proceedings of the Royal Society B-Biological Sciences. 2016;283(1832) 10.1098/rspb.2016.0217 DOI

Magnhagen C. Social influence on the correlation between behaviours in young-of-the-year perch. Behavioral Ecology and Sociobiology. 2007;61(4):525–31. 10.1007/s00265-006-0280-3 DOI

Zhao DP, Feng PS. Temperature increase impacts personality traits in aquatic non-native species: Implications for biological invasion under climate change. Current Zoology. 2015;61(6):966–71 PubMed PMC

Snijders L, van Rooij EP, Henskens MFA, van Oers K, Naguib M. Dawn song predicts behaviour during territory conflicts in personality-typed great tits. Animal Behaviour. 2015;109:45–52. 10.1016/j.anbehav.2015.07.037 DOI

Miller R, Laskowski KL, Schiestl M, Bugnyar T, Schwab C. Socially Driven Consistent Behavioural Differences during Development in Common Ravens and Carrion Crows. Plos One. 2016;11(2) 10.1371/journal.pone.0148822 PubMed DOI PMC

Sommer-Trembo C, Zimmer C, Jourdan J, Bierbach D, Plath M. Predator experience homogenizes consistent individual differences in predator avoidance. Journal of Ethology. 2016;34(2):155–65. 10.1007/s10164-016-0460-1 DOI

Vetter SG, Brandstatter C, Macheiner M, Suchentrunk F, Gerritsmann H, Bieber C. Shy is sometimes better: personality and juvenile body mass affect adult reproductive success in wild boars, Sus scrofa. Animal Behaviour. 2016;115:193–205. 10.1016/j.anbehav.2016.03.026 DOI

Sinn DL, Moltschaniwskyj NA. Personality traits in dumpling squid (Euprymna tasmanica): Context-specific traits and their correlation with biological characteristics. Journal of Comparative Psychology. 2005;119(1):99–110. 10.1037/0735-7036.119.1.99 . PubMed DOI

Kralj-Fišer S, Schuett W. Studying personality variation in invertebrates: why bother? Animal Behaviour. 2014;91:41–52. 10.1016/j.anbehav.2014.02.016 DOI

Carere C, Drent PJ, Privitera L, Koolhaas JM, Groothuis TGG. Personalities in great tits, Parus major: stability and consistency. Animal Behaviour. 2005;70:795–805. 10.1016/j.anbehav.2005.01.003 DOI

Aplin LM, Farine DR, Mann RP, Sheldon BC. Individual-level personality influences social foraging and collective behaviour in wild birds. Proceedings of the Royal Society B-Biological Sciences. 2014;281(1789) 10.1098/rspb.2014.1016 PubMed DOI PMC

Bousquet CAH, Petit O, Arrive M, Robin JP, Sueur C. Personality tests predict responses to a spatial-learning task in mallards, Anas platyrhynchos. Animal Behaviour. 2015;110:145–54. 10.1016/j.anbehav.2015.09.024 DOI

McCowan LSC, Mainwaring MC, Prior NH, Griffith SC. Personality in the wild zebra finch: exploration, sociality, and reproduction. Behavioral Ecology. 2015;26(3):735–46. 10.1093/beheco/aru239 DOI

Class B, Brommer JE. Senescence of personality in a wild bird. Behavioral Ecology and Sociobiology. 2016;70(5):733–44. 10.1007/s00265-016-2096-0 DOI

Réale D, Martin J, Coltman DW, Poissant J, Festa-Bianchet M. Male personality, life-history strategies and reproductive success in a promiscuous mammal. Journal of Evolutionary Biology. 2009;22(8):1599–607. 10.1111/j.1420-9101.2009.01781.x . PubMed DOI

Careau V, Garland T. Performance, Personality, and Energetics: Correlation, Causation, and Mechanism. Physiological and Biochemical Zoology. 2012;85(6):543–71. 10.1086/666970 . PubMed DOI

Montiglio PO, Garant D, Pelletier F, Réale D. Personality differences are related to long-term stress reactivity in a population of wild eastern chipmunks, Tamias striatus. Animal Behaviour. 2012;84(4):1071–9. 10.1016/j.anbehav.2012.08.010 DOI

Briard L, Dorn C, Petit O. Personality and Affinities Play a Key Role in the Organisation of Collective Movements in a Group of Domestic Horses. Ethology. 2015;121(9):888–902. 10.1111/eth.12402 DOI

Sih A, Kats LB, Maurer EF. Behavioural correlations across situations and the evolution of antipredator behaviour in a sunfish-salamander system. Animal Behaviour. 2003;65:29–44. 10.1006/anbe.2002.2025 DOI

Wilson ADM, Krause J. Personality and metamorphosis: is behavioral variation consistent across ontogenetic niche shifts? Behavioral Ecology. 2012;23(6):1316–23. 10.1093/beheco/ars123 DOI

Urszan TJ, Torok J, Hettyey A, Garamszegi LZ, Herczeg G. Behavioural consistency and life history of Rana dalmatina tadpoles. Oecologia. 2015;178(1):129–40. 10.1007/s00442-014-3207-0 . PubMed DOI

Carazo P, Noble DWA, Chandrasoma D, Whiting MJ. Sex and boldness explain individual differences in spatial learning in a lizard. Proceedings of the Royal Society B-Biological Sciences. 2014;281(1782) 10.1098/rspb.2013.3275 PubMed DOI PMC

McEvoy J, While GM, Sinn DL, Carver S, Wapstra E. Behavioural syndromes and structural and temporal consistency of behavioural traits in a social lizard. Journal of Zoology. 2015;296(1):58–66. 10.1111/jzo.12217 DOI

Mell H, Josserand R, Decenciere B, Artacho P, Meylan S, Le Galliard JF. Do personalities co-vary with metabolic expenditure and glucocorticoid stress response in adult lizards? Behavioral Ecology and Sociobiology. 2016;70(6):951–61. 10.1007/s00265-016-2117-z DOI

Bergmüller R, Taborsky M. Animal personality due to social niche specialisation. Trends in Ecology & Evolution. 2010;25(9):504–11. 10.1016/j.tree.2010.06.012 PubMed DOI

Stamps JA, Biro PA. Personality and individual differences in plasticity. Current Opinion in Behavioral Sciences. 2016;12:18–23. 10.1016/j.cobeha.2016.08.008 DOI

Müller T, Müller C. Behavioural phenotypes over the lifetime of a holometabolous insect. Frontiers in Zoology. 2015;12 10.1186/1742-9994-12-s1-s8 PubMed DOI PMC

Guenther A, Trillmich F. Photoperiod influences the behavioral and physiological phenotype during ontogeny. Behavioral Ecology. 2013;24(2):402–11. 10.1093/beheco/ars177 DOI

Groothuis TGG, Trillmich F. Unfolding Personalities: The Importance of Studying Ontogeny. Developmental Psychobiology. 2011;53(6):641–55. 10.1002/dev.20574 . PubMed DOI

Lessells CM, Boag PT. Unrepeatable repeatabilities—A Common mistake. Auk. 1987;104(1):116–21

Nakagawa S, Schielzeth H. Repeatability for Gaussian and non-Gaussian data: a practical guide for biologists. Biological Reviews. 2010;85(4):935–56. 10.1111/j.1469-185X.2010.00141.x . PubMed DOI

Bell AM, Stamps JA. Development of behavioural differences between individuals and populations of sticklebacks, Gasterosteus aculeatus. Animal Behaviour. 2004;68:1339–48. 10.1016/j.anbehav.2004.05.007 DOI

Sinn DL, Gosling SD, Moltschaniwskyj NA. Development of shy/bold behaviour in squid: context-specific phenotypes associated with developmental plasticity. Animal Behaviour. 2008;75:433–42. 10.1016/j.anbehav.2007.05.008 DOI

Dingemanse NJ, Both C, Drent PJ, Van Oers K, Van Noordwijk AJ. Repeatability and heritability of exploratory behaviour in great tits from the wild. Animal Behaviour. 2002;64:929–38. 10.1006/anbe.2002.2006 DOI

Weiss A, King JE, Hopkins WD. A cross-setting study of chimpanzee (Pan troglodytes) personality structure and development: Zoological parks and Yerkes National Primate Research Center. American Journal of Primatology. 2007;69(11):1264–77. 10.1002/ajp.20428 . PubMed DOI PMC

Wuerz Y, Krueger O. Personality over ontogeny in zebra finches: long-term repeatable traits but unstable behavioural syndromes. Frontiers in Zoology. 2015;12 10.1186/1742-9994-12-s1-s9 PubMed DOI PMC

Gyuris E, Fero O, Barta Z. Personality traits across ontogeny in firebugs, Pyrrhocoris apterus. Animal Behaviour. 2012;84(1):103–9. 10.1016/j.anbehav.2012.04.014 DOI

Hedrick AV, Kortet R. Sex differences in the repeatability of boldness over metamorphosis. Behavioral Ecology and Sociobiology. 2012;66(3):407–12. 10.1007/s00265-011-1286-z DOI

Petelle MB, McCoy DE, Alejandro V, Martin JGA, Blumstein DT. Development of boldness and docility in yellow-bellied marmots. Animal Behaviour. 2013;86(6):1147–54. 10.1016/j.anbehav.2013.09.016 DOI

Edenbrow M, Croft DP. Environmental and genetic effects shape the development of personality traits in the mangrove killifish Kryptolebias marmoratus. Oikos. 2013;122(5):667–81. 10.1111/j.1600-0706.2012.20556.x DOI

Guenther A, Finkemeier MA, Trillmich F. The ontogeny of personality in the wild guinea pig. Animal Behaviour. 2014;90:131–9. 10.1016/j.anbehav.2014.01.032 DOI

Boissy A. Fear and fearfulness in animals. Quarterly Review of Biology. 1995;70(2):165–91. 10.1086/418981 . PubMed DOI

Carere C, Caramaschi D, Fawcett TW. Covariation between personalities and individual differences in coping with stress: Converging evidence and hypotheses. Current Zoology. 2010;56(6):728–40

de Boer SF, Buwalda B, Koolhaas JM. Untangling the neurobiology of coping styles in rodents: Towards neural mechanisms underlying individual differences in disease susceptibility. Neuroscience and Biobehavioral Reviews. 2017;74:401–22. 10.1016/j.neubiorev.2016.07.008 . PubMed DOI

Stamps JA, Groothuis TGG. Developmental perspectives on personality: implications for ecological and evolutionary studies of individual differences. Philosophical Transactions of the Royal Society B-Biological Sciences. 2010;365(1560):4029–41. 10.1098/rstb.2010.0218 PubMed DOI PMC

Trillmich F, Hudson R. The Emergence of Personality in Animals: The Need for a Developmental Approach. Developmental Psychobiology. 2011;53(6):505–9. 10.1002/dev.20573 . PubMed DOI

Chapman BB, Hegg A, Ljungberg P. Sex and the Syndrome: Individual and Population Consistency in Behaviour in Rock Pool Prawn Palaemon elegans. Plos One. 2013;8(3) 10.1371/journal.pone.0059437 PubMed DOI PMC

Stamps JA, Krishnan VV. Individual differences in the potential and realized developmental plasticity of personality traits. Frontiers in Ecology and Evolution. 2014;2(69) 10.3389/fevo.2014.00069 DOI

Smith BR, Blumstein DT. Structural consistency of behavioural syndromes: does predator training lead to multi-contextual behavioural change? Behaviour. 2012;149(2):187–213. 10.1163/156853912x634133 DOI

Naguib M, Florcke C, van Oers K. Effects of Social Conditions During Early Development on Stress Response and Personality Traits in Great Tits (Parus major). Developmental Psychobiology. 2011;53(6):592–600. 10.1002/dev.20533 . PubMed DOI

Hudson R, Bautista A, Reyes-Meza V, Montor JM, Rodel HG. The Effect of Siblings on Early Development: A Potential Contributor to Personality Differences in Mammals. Developmental Psychobiology. 2011;53(6):564–74. 10.1002/dev.20535 . PubMed DOI

Talarovičová A, Kršková L, Blazeková J. Testosterone enhancement during pregnancy influences the 2D:4D ratio and open field motor activity of rat siblings in adulthood. Hormones and Behavior. 2009;55(1):235–9. 10.1016/j.yhbeh.2008.10.010 . PubMed DOI

Overli O, Sorensen C, Pulman KGT, Pottinger TG, Korzan WJ, Summers CH, et al. Evolutionary background for stress-coping styles: Relationships between physiological, behavioral, and cognitive traits in non-mammalian vertebrates. Neuroscience and Biobehavioral Reviews. 2007;31(3):396–412. 10.1016/j.neubiorev.2006.10.006 . PubMed DOI

Mayer M, Shine R, Brown GP. Bigger babies are bolder: effects of body size on personality of hatchling snakes. Behaviour. 2016;153(3):313–23. 10.1163/1568539x-00003343 DOI

Besson AA, Lagisz M, Senior AM, Hector KL, Nakagawa S. Effect of maternal diet on offspring coping styles in rodents: a systematic review and meta-analysis. Biological Reviews. 2016;91(4):1065–80. 10.1111/brv.12210 . PubMed DOI

Nyqvist MJ, Gozlan RE, Cucherousset J, Britton JR. Behavioural Syndrome in a Solitary Predator Is Independent of Body Size and Growth Rate. Plos One. 2012;7(2) 10.1371/journal.pone.0031619 PubMed DOI PMC

Trník M, Albrechtová J, Kratochvíl L. Persistent Effect of Incubation Temperature on Stress-Induced Behavior in the Yucatan Banded Gecko (Coleonyx elegans). Journal of Comparative Psychology. 2011;125(1):22–30. 10.1037/a0021186 . PubMed DOI

Siviter H, Deeming DC, Rosenberger J, Burman OHP, Moszuti SA, Wilkinson A. The impact of egg incubation temperature on the personality of oviparous reptiles. Animal Cognition. 2017;20(1):109–16. 10.1007/s10071-016-1030-1 . PubMed DOI PMC

Carter AW, Paitz RT, McGhee KE, Bowden RM. Turtle hatchlings show behavioral types that are robust to developmental manipulations. Physiology & Behavior. 2016;155:46–55. 10.1016/j.physbeh.2015.11.034 PubMed DOI

Hawlena D, Boochnik R, Abramsky Z, Bouskila A. Blue tail and striped body: why do lizards change their infant costume when growing up? Behavioral Ecology. 2006;17(6):889–96. 10.1093/beheco/arl023 DOI

Wilson D, Heinsohn R, Endler JA. The adaptive significance of ontogenetic colour change in a tropical python. Biology Letters. 2007;3(1):40–3. 10.1098/rsbl.2006.0574 . PubMed DOI PMC

Landová E, Jančúchová-Lásková J, Musilová V, Kadochová S, Frynta D. Ontogenetic switch between alternative antipredatory strategies in the leopard gecko (Eublepharis macularius): defensive threat versus escape. Behavioral Ecology and Sociobiology. 2013;67(7):1113–22. 10.1007/s00265-013-1536-3 DOI

Wolf M, van Doorn GS, Leimar O, Weissing FJ. Life-history trade-offs favour the evolution of animal personalities. Nature. 2007;447(7144):581–4. 10.1038/nature05835 . PubMed DOI

Clark CW. Antipredator behavior and the asset-protection principle. Behavioral Ecology. 1994;5(2):159–70. 10.1093/beheco/5.2.159 DOI

Frynta D, Vejvodová T, Šimková O. Sex allocation and secondary sex ratio in Cuban boa (Chilabothrus angulifer): mother's body size affects the ratio between sons and daughters. Science of Nature. 2016;103(5–6) 10.1007/s00114-016-1369-9 PubMed DOI

Le Galliard JF, Paquet M, Cisel M, Montes-Poloni L. Personality and the pace-of-life syndrome: variation and selection on exploration, metabolism and locomotor performances. Functional Ecology. 2013;27(1):136–44. 10.1111/1365-2435.12017 DOI

Herzog HA, Burghardt GM. Development of antipredator responses in snakes. III. Stability of individual and litter differences over the first year of life. Ethology. 1988;77(3):250–8

Herzog HA, Burghardt GM. Development of antipredator responses in snakes. I. Defensive and open-field behaviors in newborns and adults of three species of garter snakes (Thamnophis melanogaster, T. sirtalis, T. butleri). Journal of Comparative Psychology. 1986;100(4):372–9. 10.1037/0735-7036.100.4.372 DOI

Caspi A, Roberts BW, Shiner RL. Personality development: Stability and change. Annual Review of Psychology. 2005;56:453–84. 10.1146/annurev.psych.55.090902.141913 . PubMed DOI

Card DC, Schield DR, Adams RH, Corbin AB, Perry BW, Andrew AL, et al. Phylogeographic and population genetic analyses reveal multiple species of Boa and independent origins of insular dwarfism. Molecular Phylogenetics and Evolution. 2016;102:104–16. doi: 10.1016/j.ympev.2016.05.034 PubMed DOI PMC

Hynková I, Starostová Z, Frynta D. Mitochondrial DNA Variation Reveals Recent Evolutionary History of Main Boa constrictor Clades. Zoological Science. 2009;26(9):623–31. 10.2108/zsj.26.623 . PubMed DOI

Reed R, Rodda G. Giant Constrictors: Biological and Management Profiles and an Establishment Risk Assessment for Nine Large Species of Pythons, Anacondas, and the Boa Constrictor: U.S. Geological Survey Open-File Report; 2009. 302 p.

Gardner-Santana LC, Beaupre SJ. Timber Rattlesnakes (Crotalus horridus) Exhibit Elevated and Less Variable Body Temperatures during Pregnancy. Copeia. 2009;(2):363–8. 10.1643/cp-07-271 DOI

Schuett GW, Hardy DL, Earley RL, Greene HW. Does prey size induce head skeleton phenotypic plasticity during early ontogeny in the snake Boa constrictor? Journal of Zoology. 2005;267:363–9. 10.1017/s0952836905007624 DOI

Lind AJ, Welsh HH. Ontogenetic changes in foraging behavior and habitat use by the Oregon garter snake, Thamnophis atratus hydrophilus. Animal Behaviour. 1994;48(6):1261–73. 10.1006/anbe.1994.1362 DOI

Arnold SJ, Bennett AF. Behavioral variation in natural populations. III. Antipredator displays in the garter snake Thamnophis radix. Animal Behaviour. 1984;32(NOV):1108–18. 10.1016/s0003-3472(84)80227-4 DOI

Montgomery GG, Rand AS. Movements, body temperature and hunting strategy of a Boa constrictor. Copeia. 1978;(3):532–3

Webb JK, Brook BW, Shine R. Does foraging mode influence life history traits? A comparative study of growth, maturation and survival of two species of sympatric snakes from south-eastern Australia. Austral Ecology. 2003;28(6):601–10. 10.1046/j.1442-9993.2003.t01-1-01316.x DOI

Herzog HA, Bailey BD. Development of antipredator responses in snakes. II. Effects of recent feeding on defensive behaviors of juvenile garter snakes (Thamnophis sirtalis). Journal of Comparative Psychology. 1987;101(4):387–9. 10.1037/0735-7036.101.4.387 DOI

Secor SM. Regulation of digestive performance: a proposed adaptive response. Comparative Biochemistry and Physiology a-Molecular and Integrative Physiology. 2001;128(3):565–77 PubMed

Ott BD, Secor SM. Adaptive regulation of digestive performance in the genus Python. Journal of Experimental Biology. 2007;210(2):340–56. 10.1242/jeb.02626 PubMed DOI

Herzog H, Burghardt G. Prey movement and predatory behavior of juvenile western yellow-bellied racers, Coluber constrictor mormon. Herpetologica. 1974:285–9.

Candland DK, Nagy ZM. Open field—Some comparative data. Annals of the New York Academy of Sciences. 1969;159(A3):831–&. 10.1111/j.1749-6632.1969.tb12982.x PubMed DOI

Archer J. Tests for emotionality in rats and mice—review. Animal Behaviour. 1973;21(MAY):205–35. 10.1016/s0003-3472(73)80065-x PubMed DOI

Ossenkopp KP, Sorenson L, Mazmanian DS. Factor-analysis of Open-field behavior in the rat (Rattus norvegicus)–Application of the 3-way Parafac model to a longitudinal data set. Behavioural Processes. 1994;31(2–3):129–44. 10.1016/0376-6357(94)90001-9 . PubMed DOI

Chiszar D, Carter T, Knight L, Simonsen L, Taylor S. Investigatory behavior in Plains garter snake (Thamnophis radix) and several additional species. Animal Learning & Behavior. 1976;4(3):273–8. 10.3758/bf03214049 DOI

Suchomelová P, Jančúchová-Lásková J, Landová E, Frynta D. Experimental assessment of social interactions in two species of the genus Teratoscincus (Gekkota). Behavioural Processes. 2015;120:14–24. 10.1016/j.beproc.2015.08.005 . PubMed DOI

Le Galliard JF, Paquet M, Mugabo M. An experimental test of density-dependent selection on temperament traits of activity, boldness and sociability. Journal of Evolutionary Biology. 2015;28(5):1144–55. 10.1111/jeb.12641 . PubMed DOI

Lopez P, Hawlena D, Polo V, Amo L, Martin J. Sources of individual shy-bold variations in antipredator behaviour of male Iberian rock lizards. Animal Behaviour. 2005;69:1–9. 10.1016/j.anbehav.2004.05.010 DOI

Brodie ED. Consistency of individual differences in antipredator behavior and color pattern in the garter snake, Thamnophis ordinoides. Animal Behaviour. 1993;45(5):851–61. 10.1006/anbe.1993.1106 DOI

Herzog HA, Bowers BB, Burghardt GM. Development of antipredator responses in snakes. V. Species differences in ontogenetic trajectories. Developmental Psychobiology. 1992;25(3):199–211. 10.1002/dev.420250305 . PubMed DOI

Herzog HA, Bowers BB, Burghardt GM. Development of antipredator responses in snakes. IV. Interspecific and intraspecific differences in habituation of defensive behavior. Developmental Psychobiology. 1989;22(5):489–508. 10.1002/dev.420220507 . PubMed DOI

Aloisi AM, Ceccarelli I, Lupo C. Behavioural and hormonal effects of restraint stress and formalin test in male and female rats. Brain Research Bulletin. 1998;47(1):57–62. 10.1016/s0361-9230(98)00063-x . PubMed DOI

Lillywhite HB, Zippel KC, Farrell AP. Resting and maximal heart rates in ectothermic vertebrates. Comparative Biochemistry and Physiology a-Molecular and Integrative Physiology. 1999;124(4):369–82. 10.1016/s1095-6433(99)00129-4 PubMed DOI

Carrasco JL, Jover L. Concordance correlation coefficient applied to discrete data. Statistics in Medicine. 2005;24(24):4021–34. 10.1002/sim.2397 . PubMed DOI

Team RDC. R: a language and environment for statistical computing. In: Computing RFfS, editor. Vienna, Austria: 2012.

Statsoft S. Statistica. 6.0 ed2001.

Budaev SV. Using Principal Components and Factor Analysis in Animal Behaviour Research: Caveats and Guidelines. Ethology. 2010;116(5):472–80. 10.1111/j.1439-0310.2010.01758.x DOI

Flores D, Tousignant A, Crews D. Incubation temperature affects the behavior of adult leopard geckos (Eublepharis macularius). Physiology & Behavior. 1994;55(6):1067–72. 10.1016/0031-9384(94)90389-1 PubMed DOI

Luttbeg B, Sih A. Risk, resources and state- dependent adaptive behavioural syndromes. Philosophical Transactions of the Royal Society B-Biological Sciences. 2010;365(1560):3977–90. 10.1098/rstb.2010.0207 PubMed DOI PMC

Dingemanse NJ, Wolf M. Recent models for adaptive personality differences: a review. Philosophical Transactions of the Royal Society B-Biological Sciences. 2010;365(1560):3947–58. 10.1098/rstb.2010.0221 PubMed DOI PMC

Herczeg G, Ab Ghani NI, Merila J. Evolution of stickleback feeding behaviour: genetics of population divergence at different ontogenetic stages. Journal of Evolutionary Biology. 2013;26(5):955–62. 10.1111/jeb.12103 . PubMed DOI

Verbeek MEM, Drent PJ, Wiepkema PR. Consistent individual differences in early exploratory behavior of male Great tits. Animal Behaviour. 1994;48(5):1113–21. 10.1006/anbe.1994.1344 DOI

Stowe M, Kotrschal K. Behavioural phenotypes may determine whether social context facilitates or delays novel object exploration in ravens (Corvus corax). Journal of Ornithology. 2007;148:S179–S84. 10.1007/s10336-007-0145-1 DOI

Herborn KA, Coffey J, Larcombe SD, Alexander L, Arnold KE. Oxidative profile varies with personality in European greenfinches. Journal of Experimental Biology. 2011;214(10):1732–9. 10.1242/jeb.051383 PubMed DOI

Montiglio PO, Garant D, Bergeron P, Messier GD, Réale D. Pulsed resources and the coupling between life-history strategies and exploration patterns in eastern chipmunks (Tamias striatus). Journal of Animal Ecology. 2014;83(3):720–8. 10.1111/1365-2656.12174 . PubMed DOI

Jones KA, Godin JGJ. Are fast explorers slow reactors? Linking personality type and anti-predator behaviour. Proceedings of the Royal Society B-Biological Sciences. 2010;277(1681):625–32. 10.1098/rspb.2009.1607 PubMed DOI PMC

Verbeek MEM, Boon A, Drent PJ. Exploration, aggressive behavior and dominance in pair-wise confrontations of juvenile male great tits patent 11–12. 1996. September.

van Oers K, Drent PJ, de Goede P, van Noordwijk AJ. Realized heritability and repeatability of risk-taking behaviour in relation to avian personalities. Proceedings of the Royal Society B-Biological Sciences. 2004;271(1534):65–73. 10.1098/rspb.2003.2518 PubMed DOI PMC

Koolhaas JM, Korte SM, De Boer SF, Van Der Vegt BJ, Van Reenen CG, Hopster H, et al. Coping styles in animals: current status in behavior and stress-physiology. Neuroscience and Biobehavioral Reviews. 1999;23(7):925–35. 10.1016/s0149-7634(99)00026-3 . PubMed DOI

Koolhaas JM, de Boer SF, Coppens CM, Buwalda B. Neuroendocrinology of coping styles: Towards understanding the biology of individual variation. Frontiers in Neuroendocrinology. 2010;31(3):307–21. 10.1016/j.yfrne.2010.04.001 . PubMed DOI

DuRant SE, Romero LM, Talent LG, Hopkins WA. Effect of exogenous corticosterone on respiration in a reptile. General and Comparative Endocrinology. 2008;156(1):126–33. 10.1016/j.ygcen.2007.12.004 . PubMed DOI

Langkilde T, Shine R. How much stress do researchers inflict on their study animals? A case study using a scincid lizard, Eulamprus heatwolei. Journal of Experimental Biology. 2006;209(6):1035–43. 10.1242/jeb.02112 PubMed DOI

Carere C, van Oers K. Shy and bold great tits (Parus major): body temperature and breath rate in response to handling stress. Physiology & Behavior. 2004;82(5):905–12. 10.1016/j.physbeh.2004.07.009 PubMed DOI

Stinner JN, Ely DL Blood pressure during routine activity, stress, and feeding in Black racer snakes (Coluber constrictor). American Journal of Physiology. 1993;264(1):R79–R84 PubMed

Secor SM, Hicks JW, Bennett AF. Ventilatory and cardiovascular responses of a python (Python molurus) to exercise and digestion. Journal of Experimental Biology. 2000;203(16):2447–54 PubMed

Stamps JA, Krishnan VV. Combining Information from Ancestors and Personal Experiences to Predict Individual Differences in Developmental Trajectories. American Naturalist. 2014;184(5):647–57. 10.1086/678116 . PubMed DOI

del Barrio V, Moreno-Rosset C, Lopez-Martinez R, Olmedo M. Anxiety, depression and personality structure. Personality and Individual Differences. 1997;23(2):327–35. 10.1016/s0191-8869(97)00030-5 DOI

Gullone E, Moore S. Adolescent risk-taking and the five-factor model of personality. Journal of Adolescence. 2000;23(4):393–407. 10.1006/jado.2000.0327 . PubMed DOI

Inter-individual differences in laboratory rats as revealed by three behavioural tasks