Acquiring knowledge about the environment is crucial for survival. Animals, often driven by their exploratory tendencies, gather valuable information regarding food resources, shelter, mating partners, etc. However, neophobia, or avoiding novel environmental stimuli, can constrain their exploratory behaviour. While neophobia can reduce potential predation risks, decreased exploratory behaviour resulting from it may limit the ability to discover highly rewarding resources. Dogs (Canis familiaris) living in semi-urban and urban environments as free-ranging populations, although subject to various selection forces, typically have negligible predation pressure. These dogs are scavengers in human-dominated environments; thus, selection against object-neophobia can provide benefits when searching for novel food resources. Although captive pack-living dogs are known to be less neophobic than their closest living ancestors, wolves (Canis lupus), little is known about free-ranging dogs' behavioural responses to novel objects, particularly in foraging contexts. Using an object choice experiment, we tested 259 free-ranging dogs from two age classes, adult and juvenile, to investigate their object-neophobia in a scavenging context. We employed a between-subject study design, providing dogs with a familiar and a potentially novel object, both baited with equal, hidden food items. Adult and juvenile dogs significantly inspected the novel object first compared to the familiar one, even when the hidden food item was partially visible. To validate these findings, we compared novel objects with different strengths of olfactory cues (baited vs. false-baited) and found that they were inspected comparably by adults and juveniles. No significant differences were found in the latencies to inspect the objects, suggesting that free-ranging dogs may still be cautious when exploring their environments. These results indicate that free-ranging dogs, evidently from an early ontogenetic phase, do not show object-neophobia, as demonstrated by their preference for novel over familiar food sources. We conclude that little to no constraint of neophobia on exploratory behaviour in semi-urban and urban-dwelling animals can guide foraging decision-making processes, providing adaptive benefits.

Centre for Animal Health and Welfare Jockey Club College of Veterinary Medicine and Life Sciences City University of Hong Kong Hong Kong SAR Hong Kong

Department of Botany and Zoology Masaryk University Brno Czech Republic

Department of Endangered Species Management Wildlife Institute of India Dehradun Uttarakhand India

Department of Infectious Diseases and Public Health Jockey Club College of Veterinary Medicine and Life Sciences City University of Hong Kong Hong Kong SAR Hong Kong

The Dog Lab Behaviour and Ecology Lab Department of Biological Sciences Indian Institute of Science Education and Research Kolkata Mohanpur West Bengal India

Zobrazit více v PubMed

Alley TR. Food Neophobia. Philadelphia: Woodhead Publishing; 2018. Conceptualization and measurement of human food neophobia; pp. 169–192. DOI

Banerjee A, Bhadra A. The more the merrier:dogs can assess quantities in food-choice tasks. Current Science. 2019;117(6):1095–1100. doi: 10.18520/cs/v117/i6/1095-1100. DOI

Bates D, Mächler M, Bolker BM, Walker SC. Fitting linear mixed-effects models using lme4. Journal of Statistical Software. 2015;67(1):1–48. doi: 10.18637/jss.v067.i01. DOI

Beckman AK, Richey BMS, Rosenthal GG. Behavioral responses of wild animals to anthropogenic change: insights from domestication. Behavioral Ecology and Sociobiology. 2022;76(7):105. doi: 10.1007/s00265-022-03205-6. DOI

Bhadra A, Bhattacharjee D, Paul M, Bhadra A. The meat of the matter: a thumb rule for scavenging dogs? Ethology Ecology & Evolution. 2015;28(4):427–440. doi: 10.1080/03949370.2015.1076526. DOI

Bhattacharjee D, Bhadra A. Humans dominate the social interaction networks of urban free-ranging dogs in India. Frontiers in Psychology. 2020;11:2153. doi: 10.3389/fpsyg.2020.02153. PubMed DOI PMC

Bhattacharjee D, Bhadra A. Response to short-lived human overcrowding by free-ranging dogs. Behavioral Ecology and Sociobiology. 2021;75:11. doi: 10.1007/s00265-021-03052-x. DOI

Bhattacharjee D, Bhadra A. Adjustment in the point-following behaviour of free-ranging dogs—roles of social petting and informative-deceptive nature of cues. Animal Cognition. 2022;25(3):571–579. doi: 10.1007/s10071-021-01573-6. PubMed DOI

Bhattacharjee D, Dasgupta S, Biswas A, Deheria J, Gupta S, Nikhil Dev N, Udell M, Bhadra A. Practice makes perfect: familiarity of task determines success in solvable tasks for free-ranging dogs (Canis lupus familiaris) Animal Cognition. 2017a;20:771–776. doi: 10.1007/s10071-017-1097-3. PubMed DOI

Bhattacharjee D, Nikhil Dev N, Gupta S, Sau S, Sarkar R, Biswas A, Banerjee A, Babu D, Mehta D, Bhadra A. Free-ranging dogs show age related plasticity in their ability to follow human pointing. PLOS ONE. 2017b;12(7):e0180643. doi: 10.1371/journal.pone.0180643. PubMed DOI PMC

Bhattacharjee D, Mandal S, Shit P, Varghese George M, Vishnoi A, Bhadra A. Free-ranging dogs are capable of utilising complex human pointing cues. Frontiers in Psychology. 2019;10:2818. doi: 10.3389/fpsyg.2019.02818. PubMed DOI PMC

Bhattacharjee D, Sau S, Bhadra A. Free-ranging dogs understand human intentions and adjust their behavioral responses accordingly. Frontiers in Ecology and Evolution. 2018;6:232. doi: 10.3389/fevo.2018.00232. DOI

Bhattacharjee D, Sau S, Bhadra A. ‘Bolder’ together—response to human social cues in groups of free-ranging dogs. Behaviour. 2020;157(3–4):363–384. doi: 10.1163/1568539X-bja10005. DOI

Bhattacharjee D, Sau S, Das J, Bhadra A. Free-ranging dogs prefer petting over food in repeated interactions with unfamiliar humans. The Journal of Experimental Biology. 2017c;220(24):4654–4660. doi: 10.1242/jeb.166371. PubMed DOI

Biondi LM, Fuentes GM, Córdoba RS, Bó MS, Cavalli M, Paterlini CA, Castano MV, García GO. Variation in boldness and novelty response between rural and urban predatory birds: the Chimango Caracara, Milvago chimango as study case. Behavioural Processes. 2020;173:104064. doi: 10.1016/j.beproc.2020.104064. PubMed DOI

Biswas S, Bhowmik T, Ghosh K, Roy A, Lahiri A, Sarkar S, Bhadra A. Scavengers in the human-dominated landscape: an experimental study. 2023 doi: 10.48550/arXiv.2208.05030.2208.05030 PubMed DOI PMC

Boitani L, Francisci F, Ciucci P, Andreoli G. The domestic dog. Cambridge University Press; Cambridge: 2016. The ecology and behavior of feral dogs: a case study from central Italy; pp. 342–368. DOI

Breck SW, Poessel SA, Mahoney P, Young JK. The intrepid urban coyote: a comparison of bold and exploratory behavior in coyotes from urban and rural environments. Scientific Reports. 2019;9:2104. doi: 10.1038/s41598-019-38543-5. PubMed DOI PMC

Brubaker L, Bhattacharjee D, Ghaste P, Babu D, Shit P, Bhadra A, Udell MAR. The effects of human attentional state on canine gazing behaviour: a comparison of free-ranging, shelter, and pet dogs. Animal Cognition. 2019;22:1129–1139. doi: 10.1007/s10071-019-01305-x. PubMed DOI

Cimarelli G, Juskaite M, Range F, Marshall-Pescini S. Free-ranging dogs match a human’s preference in a foraging task. Current Zoology. 2023:zoad046. doi: 10.1093/cz/zoad046. PubMed DOI PMC

Dall S, Giraldeau L, Olsson O, Mcnamara J, Stephens D. Information and its use by animals in evolutionary ecology. Trends in Ecology & Evolution. 2005;20(4):187–193. doi: 10.1016/j.tree.2005.01.010. PubMed DOI

Dammhahn M, Mazza V, Schirmer A, Göttsche C, Eccard JA. Of city and village mice: behavioural adjustments of striped field mice to urban environments. Scientific Reports. 2020;10:13056. doi: 10.1038/s41598-020-69998-6. PubMed DOI PMC

Day RL, Coe RL, Kendal JR, Laland KN. Neophilia, innovation and social learning: a study of intergeneric differences in callitrichid monkeys. Animal Behaviour. 2003;65:559–571. doi: 10.1006/anbe.2003.2074. DOI

Ditchkoff SS, Saalfeld ST, Gibson CJ. Animal behavior in urban ecosystems: modifications due to human-induced stress. Urban Ecosystems. 2006;9:5–12. doi: 10.1007/s11252-006-3262-3. DOI

Egert-Berg K, Handel M, Goldshtein A, Eitan O, Borissov I, Yovel Y. Fruit bats adjust their foraging strategies to urban environments to diversify their diet. BMC Biology. 2021;19(1):123. doi: 10.1186/s12915-021-01060-x. PubMed DOI PMC

Eötvös CB, Magura T, Lövei GL. A meta-analysis indicates reduced predation pressure with increasing urbanization. Landscape and Urban Planning. 2018;180:54–59. doi: 10.1016/j.landurbplan.2018.08.010. DOI

Fischer JD, Cleeton SH, Lyons TP, Miller JR. Urbanization and the predation paradox: the role of trophic dynamics in structuring vertebrate communities. BioScience. 2012;62(9):809–818. doi: 10.1525/bio.2012.62.9.6. DOI

Gordon DM. The fusion of behavioral ecology and ecology. Behavioral Ecology. 2011;22(2):225–230. doi: 10.1093/beheco/arq172. DOI

Greenberg R. Animal innovation. Oxford University Press; Oxford: 2003. The role of neophobia and neophilia in the development of innovative behaviour of birds; pp. 175–196. DOI

Greenberg R, Mettke-Hofmann C. Current ornithology. Volume 16. Boston: Springer US; 2001. Ecological aspects of neophobia and neophilia in birds; pp. 119–178. DOI

Greggor AL, Clayton NS, Fulford AJC, Thornton A. Street smart: faster approach towards litter in urban areas by highly neophobic corvids and less fearful birds. Animal Behaviour. 2016;117:123–133. doi: 10.1016/j.anbehav.2016.03.029. PubMed DOI PMC

Greggor AL, Thornton A, Clayton NS. Neophobia is not only avoidance: improving neophobia tests by combining cognition and ecology. Current Opinion in Behavioral Sciences. 2015;6:82–89. doi: 10.1016/j.cobeha.2015.10.007. DOI

Griffin AS, Netto K, Peneaux C. Neophilia, innovation and learning in an urbanized world: a critical evaluation of mixed findings. Current Opinion in Behavioral Sciences. 2017;16:15–22. doi: 10.1016/j.cobeha.2017.01.004. DOI

Grunst AS, Grunst ML, Pinxten R, Eens M. Personality and plasticity in neophobia levels vary with anthropogenic disturbance but not toxic metal exposure in urban great tits. Science of the Total Environment. 2019;656:997–1009. doi: 10.1016/j.scitotenv.2018.11.383. PubMed DOI

Hartig F. DHARMa: residual diagnostics for hierarchical regression models. R package version 0.4.6http://florianhartig.github.io/DHARMa/ 2020

Hothorn T, Zeileis A. Diagnostic checking in regression relationships. R News. 2011;2(3):7–10.

Jarjour C, Evans JC, Routh M, Morand-Ferron J. Does city life reduce neophobia? A study on wild black-capped chickadees. Behavioral Ecology. 2019;31(1):123–131. doi: 10.1093/beheco/arz167. DOI

Kaulfuß P, Mills DS. Neophilia in domestic dogs (Canis familiaris) and its implication for studies of dog cognition. Animal Cognition. 2008;11:553–556. doi: 10.1007/s10071-007-0128-x. PubMed DOI

Kimball MG, Lattin CR. The seven deadly sins of neophobia experimental design. SSRN Electronic Journal. 2023 doi: 10.2139/ssrn.4511040. PubMed DOI

Klump BC, Major RE, Farine DR, Martin JM, Aplin LM. Is bin-opening in cockatoos leading to an innovation arms race with humans? Current Biology. 2022;32(17):R910–R911. doi: 10.1016/j.cub.2022.08.008. PubMed DOI

Lazzaroni M, Schär J, Baxter E, Gratalon J, Range F, Marshall-Pescini S, Dale R. Village dogs match pet dogs in reading human facial expressions. PeerJ. 2023;11:e15601. doi: 10.7717/peerj.15601. PubMed DOI PMC

Magle SB, Hunt VM, Vernon M, Crooks KR. Urban wildlife research: past, present, and future. Biological Conservation. 2012;155:23–32. doi: 10.1016/j.biocon.2012.06.018. DOI

Mangalam M, Singh M. Differential foraging strategies: motivation, perception and implementation in urban free-ranging dogs, Canis familiaris. Animal Behaviour. 2013;85:763–770. doi: 10.1016/j.anbehav.2013.01.019. DOI

Markus N, Hall L. Foraging behaviour of the black flying-fox (Pteropus alecto) in the urban landscape of Brisbane, Queensland. Wildlife Research. 2004;31(3):345–355. doi: 10.1071/WR01117. DOI

Marshall-Pescini S, Cafazzo S, Virányi Z, Range F. Integrating social ecology in explanations of wolf–dog behavioral differences. Current Opinion in Behavioral Sciences. 2017b;16:80–86. doi: 10.1016/j.cobeha.2017.05.002. DOI

Marshall-Pescini S, Virányi Z, Kubinyi E, Range F. Motivational factors underlying problem solving: comparing wolf and dog puppies’ explorative and neophobic behaviors at 5 6, and 8 weeks of age. Frontiers in Psychology. 2017a;8:180. doi: 10.3389/fpsyg.2017.00180. PubMed DOI PMC

Mazza V, Czyperreck I, Eccard JA, Dammhahn M. Cross-context responses to novelty in rural and urban small mammals. Frontiers in Ecology and Evolution. 2021;9:661971. doi: 10.3389/fevo.2021.661971. DOI

Meddock TD, Osborn DR. Neophobia in wild and laboratory mice. Psychonomic Science. 1968;12(5):223–223. doi: 10.3758/BF03331280. DOI

Mettke-Hofmann C, Rowe KC, Hayden TJ, Canoine V. Effects of experience and object complexity on exploration in garden warblers (Sylvia borin) Journal of Zoology. 2006;268(4):405–413. doi: 10.1111/j.1469-7998.2005.00037.x. DOI

Mettke-Hofmann C, Winkler H, Leisler B. The significance of ecological factors for exploration and neophobia in parrots. Ethology. 2002;108(3):249–272. doi: 10.1046/j.1439-0310.2002.00773.x. DOI

Miller R, Lambert ML, Frohnwieser A, Brecht KF, Bugnyar T, Crampton I, Garcia-Pelegrin E, Gould K, Greggor AL, Izawa E-I, Kelly DM, Li Z, Luo Y, Luong LB, Massen JJM, Nieder A, Reber SA, Schiestl M, Seguchi A, Sepehri P, Stevens JR, Taylor AH, Wang L, Wolff LM, Zhang Y, Clayton NS. Socio-ecological correlates of neophobia in corvids. Current Biology. 2022;32(1):74–85. doi: 10.1016/j.cub.2021.10.045. PubMed DOI

Moretti L, Hentrup M, Kotrschal K, Range F. The influence of relationships on neophobia and exploration in wolves and dogs. Animal Behaviour. 2015;107:159–173. doi: 10.1016/j.anbehav.2015.06.008. PubMed DOI PMC

Murray MH, St. Clair CC. Predictable features attract urban coyotes to residential yards. The Journal of Wildlife Management. 2017;81(4):593–600. doi: 10.1002/jwmg.21223. DOI

Oro D, Genovart M, Tavecchia G, Fowler MS, Martínez-Abraín A. Ecological and evolutionary implications of food subsidies from humans. Ecology Letters. 2013;16(12):1501–1514. doi: 10.1111/ele.12187. PubMed DOI

Paul M, Sen Majumder S, Sau S, Nandi AK, Bhadra A. High early life mortality in free-ranging dogs is largely influenced by humans. Scientific Reports. 2016;6:19641. doi: 10.1038/srep19641. PubMed DOI PMC

Quinn JL, Patrick SC, Bouwhuis S, Wilkin TA, Sheldon BC. Heterogeneous selection on a heritable temperament trait in a variable environment. Journal of Animal Ecology. 2009;78(6):1203–1215. doi: 10.1111/j.1365-2656.2009.01585.x. PubMed DOI

R Core Team . R Foundation for Statistical Computing; Vienna, Austria: 2020.

Ramírez F, Afán I, Bouten W, Carrasco JL, Forero MG, Navarro J. Humans shape the year-round distribution and habitat use of an opportunistic scavenger. Ecology and Evolution. 2020;10(11):4716–4725. doi: 10.1002/ece3.6226. PubMed DOI PMC

Range F, Marshall-Pescini S. Comparing wolves and dogs: current status and implications for human ‘self-domestication’. Trends in Cognitive Sciences. 2022;26:337–349. doi: 10.1016/j.tics.2022.01.003. PubMed DOI

Roy A, Lahiri A, Nandi S, Manchalwar A, Bhadra A. Ready, set, yellow! color preference of Indian free-ranging dogs. bioRxiv. 2024. DOI

Sarkar R, Bhadra A. How do animals navigate the urban jungle? A review of cognition in urban-adapted animals. Current Opinion in Behavioral Sciences. 2022;46:101177. doi: 10.1016/j.cobeha.2022.101177. DOI

Sarkar R, Bhowmick A, Dasgupta D, Banerjee R, Chakraborty P, Nayek A, Sreelekshmi R, Roy A, Sonowal R, Mondal AB, Bhadra A. Eating smart: free-ranging dogs follow an optimal foraging strategy while scavenging in groups. Frontiers in Ecology and Evolution. 2023;11:212. doi: 10.3389/fevo.2023.1099543. DOI

Sarkar R, Sau S, Bhadra A. Scavengers can be choosers: a study on food preference in free-ranging dogs. Applied Animal Behaviour Science. 2019;216:38–44. doi: 10.1016/j.applanim.2019.04.012. DOI

Sen Majumder S, Bhadra A, Ghosh A, Mitra S, Bhattacharjee D, Chatterjee J, Nandi AK, Bhadra A. To be or not to be social: foraging associations of free-ranging dogs in an urban ecosystem. Acta Ethologica. 2014;17(1):1–8. doi: 10.1007/s10211-013-0158-0. DOI

Sih A, Bell A, Johnson JC. Behavioral syndromes: an ecological and evolutionary overview. Trends in Ecology & Evolution. 2004;19:372–378. doi: 10.1016/j.tree.2004.04.009. PubMed DOI

Sen Majumder S, Chatterjee A, Bhadra A. A dog’s day with humans-time activity budget of free-ranging dogs in India. Current Science. 2014;106:874–878.

Sih A, Stamps J, Yang LH, McElreath R, Ramenofsky M. Behavior as a key component of integrative biology in a human-altered world. Integrative and Comparative Biology. 2010;50(6):934–944. doi: 10.1093/icb/icq148. PubMed DOI

Siniscalchi M, d’Ingeo S, Fornelli S, Quaranta A. Are dogs red–green colour blind? Royal Society Open Science. 2017;4(11):170869. doi: 10.1098/rsos.170869. PubMed DOI PMC

Sloan Wilson D, Clark AB, Coleman K, Dearstyne T. Shyness and boldness in humans and other animals. Trends in Ecology & Evolution. 1994;9:442–446. doi: 10.1016/0169-5347(94)90134-1. PubMed DOI

Stöwe M, Bugnyar T, Heinrich B, Kotrschal K. Effects of Group Size on Approach to Novel Objects in Ravens (Corvus corax) Ethology. 2006a;112(11):1079–1088. doi: 10.1111/j.1439-0310.2006.01273.x. DOI

Stöwe M, Bugnyar T, Loretto M-C, Schloegl C, Range F, Kotrschal K. Novel object exploration in ravens (Corvus corax): effects of social relationships. Behavioural Processes. 2006b;73(1):68–75. doi: 10.1016/j.beproc.2006.03.015. PubMed DOI

Takola E, Krause ET, Müller C, Schielzeth H. Novelty at second glance: a critical appraisal of the novel object paradigm based on meta-analysis. Animal Behaviour. 2021;180:123–142. doi: 10.1016/j.anbehav.2021.07.018. DOI

Thompson MJ, Evans JC, Parsons S, Morand-Ferron J. Urbanization and individual differences in exploration and plasticity. Behavioral Ecology. 2018;29(6):1415–1425. doi: 10.1093/beheco/ary103. DOI

Tryjanowski P, Møller AP, Morelli F, Biaduń W, Brauze T, Ciach M, Czechowski P, Czyz S, Dulisz B, Goławski A, Hetmański T, Indykiewicz P, Mitrus C, Myczko Ł, Nowakowski JJ, Polakowski M, Takacs V, Wysocki D, Zduniak P. Urbanization affects neophilia and risk-taking at bird-feeders. Scientific Reports. 2016;6:28575. doi: 10.1038/srep28575. PubMed DOI PMC

Vincze E, Papp S, Preiszner B, Seress G, Bókony V, Liker A. Habituation to human disturbance is faster in urban than rural house sparrows. Behavioral Ecology. 2016;27(5):1304–1313. doi: 10.1093/beheco/arw047. DOI

Wagner DL, Grames EM, Forister ML, Berenbaum MR, Stopak D. Insect decline in the Anthropocene: death by a thousand cuts. Proceedings of the National Academy of Sciences of the United States of America. 2021;118(2):e2023989118. doi: 10.1073/pnas.2023989118. PubMed DOI PMC

Wat KKY, Banks PB, McArthur C. Linking animal personality to problem-solving performance in urban common brushtail possums. Animal Behaviour. 2020;162:35–45. doi: 10.1016/j.anbehav.2020.01.013. DOI