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In conclusion, because of the small sample sizes inherently available in studies comparing behavior in wolves and dogs, it is critical that continued, standardized studies on wolf dog comparisons are encouraged to further uncover the resolution in behavioral variation during domestication. CH and HT designed the study.

CH conducted the experiments and prepared data for analyses. All authors reviewed the manuscript prior to submission. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. This manuscript has been released as a preprint on BioRxiv Hansen Wheat et al. Video S2. Bates, D. Fitting linear mixed-effects models using lme4. Belyaev, D. Domestication in the silver fox Vulpes Fulvus Desm : changes in physiological boundaries of the sensitive period of primary socialization.

Regular handling early in the nursing period eliminates fear responses toward human beings in wild and domestic rabbits. Boissy, A.

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Fear and fearfulness in animals. Boogert, N. The relation between social rank, neophobia and individual learning in starlings. Bray, E. Effects of maternal investment, temperament, and cognition on guide dog success.

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Bremner-Harrison, S. Behavioural trait assessment as a release criterion: boldness predicts early death in a reintroduction programme of captive-bred swift fox Vulpes Velox. Burnham, K. Methods Res. Christensen, J. Do horses generalise between objects during habituation? Clark, M. Environmental effects on the ontogey of exploratory and escape behaviors of mongolian gerbils.

Coppinger, R. Google Scholar. Even minimal human contact linked to nursing reduces fear responses toward humans in rabbits. Dobney, K. Genetics and animal domestication: new windows on an elusive process. Fear-related behaviour of dogs in veterinary practice. Driscoll, C. From wild animals to domestic pets, an evolutionary view of domestication. Feddersen-Petersen, D.

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The ontogeny of social play and agonistic behaviour in selected canid species. Fentress, J. Observations on the behavioural developement of a hand-reared male timber wolf. Fox, M.

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A comparative study of the development of facial expressions in canids; wolf, coyote and foxes. Behaviour 36, 49— Socio-ecological implications of individual differences in wolf litters: a developmental and evolutionary perspective. Behaviour 41, — Frank, H. Comparison of problem-solving performance in six-week-old wolves and dogs. Comparative manipulation-test performance in ten-week-old wolves Canis lupus and Alaskan malamutes Canis familiaris : a piagetian interpretation. Freedman, D.

Critical period in the social development of dogs. Science 31, — Friard, O. Methods Ecol. Species-specific differences and similarities in the behavior of hand-raised dog and wolf pups in social situations with humans. Selective breeding for differential aggression in mice provides evidence for heterochrony in social behaviours. Goddard, M.


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A factor analysis of fearfulness in potential guide dogs. Goodwin, D.

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Paedomorphosis affects agonistic visual signals of domestic dogs. Gray, J. The Psychology of Fear and Stress. Griffin, A. Social learning about predators: a review and prospectus. Hansen Wheat, C. Wolf Canis lupus hybrids highlight the importance of human-directed play behavior during domestication of dogs Canis familiaris. Dogs, but not wolves, lose their sensitivity towards novelty with age. Hemsworth, P.


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Behavioural responses of domestic pigs and cattle to humans and novel stimuli. Jones, R. Modification of fear in domestic chicks, Gallus gallus domesticus , via regular handling and early environmental enrichment. Animal Behaviour, Short Communications 43, — King, T. Fear of novel and startling stimuli in domestic dogs. Klinghammer, E.

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Frank Dordrecht: Dr. Junk Publishers , 31— Kruska, D. Mamalian domestication and its effect on brain structure and behaviour. Kuznetsova, A. Leiner, L. Behavioural fear and heart rate responses of horses after exposure to novel objects: effects of habituation. Lenth, R.


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R package version 1. Ley, J. Assessing fear of novel and startling stimuli in domestic dogs. Lindblad-Toh, K. Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature , — Lord, K. A comparison of the sensory development of wolves Canis lupus lupus and dogs Canis lupus familiaris. Ethology , — Mainwaring, M.