Disconnected Lives: Social Networks and Emotional Regulation in Domestic Dogs
Simple Summary
Abstract
1. Introduction
2. Materials and Methods
2.1. Search Strategy and Scope
2.2. Screening and Inclusion Criteria
2.3. Quantitative Overview and Limitations
3. Theoretical Perspectives on Canine Regulation and Sociality
3.1. The Dog as a Social Regulator
3.2. Attachment and Co-Regulation Across Species
3.3. Social Stress Models
3.4. Summary
4. Global and Demographic Context of Dog Social Life
4.1. Global Dog Populations and Ownership Patterns
4.2. Urbanisation as a Social Shift
4.3. Synthesis: Demography as Destiny
5. Social Life of Free-Ranging Dogs—The Forgotten Model of Canine Society
5.1. Structure and Composition of Free-Ranging Dog Groups
5.2. Cooperation, Affiliation, and Conflict Management
5.3. Comparative Implications for Companion Dogs
6. The Social Reality of Urban Pet Dogs
6.1. Overview
6.2. Patterns of Social Interaction and Affiliative Bonds
6.3. Developmental Roots of Social Dysregulation
6.4. Emotional and Behavioural Consequences
6.5. Synthesis: The Managed and Ecologically Constrained Social World
7. The Fragmented Social World of the Urban Dog—A Theoretical Synthesis
7.1. From Social Flexibility to Social Deficit
7.2. Theoretical Integration: Attachment, Stress Regulation, and the Ecology of Safety
7.3. Implications for Welfare, Behavioural Therapy, and Research
8. Methodological Challenges and Research Gaps
8.1. Underrepresentation and Sampling Bias in Companion-Dog Research
8.2. Existing Methods for Assessing Dog–Dog Relationship Quality
8.3. Comparative Models of Social Co-Regulation in Mammals
8.4. Toward a Multi-Context Ethology of Urban Dogs
8.5. Practical Implications: Rebuilding Micro-Communities
8.6. Ethical and Welfare Reflections
9. Conclusions
Funding
Institutional Review Board Statement
Informed Consent Statement
Data Availability Statement
Conflicts of Interest
References
- Kotrschal, K. Wolf–Dog–Human: Companionship Based on Common Social Tools. Animals 2023, 13, 2729. [Google Scholar] [CrossRef] [PubMed]
- Sipple, N.; Thielke, L.; Smith, A.; Vitale, K.R.; Udell, M.A.R. Intraspecific and Interspecific Attachment between Cohabitant Dogs and Human Caregivers. Integr. Comp. Biol. 2021, 61, 132–139. [Google Scholar] [CrossRef] [PubMed]
- Anderson, K.L.; Casey, R.A.; Cooper, B.; Upjohn, M.M.; Christley, R.M. National Dog Survey: Describing UK Dog and Ownership Demographics. Animals 2023, 13, 1072. [Google Scholar] [CrossRef] [PubMed]
- Carvelli, A.; Scaramozzino, P.; Iacoponi, F.; Condoleo, R.; Della Marta, U. Size, Demography, Ownership Profiles, and Identification Rate of the Owned Dog Population in Central Italy. PLoS ONE 2020, 15, e0240551. [Google Scholar] [CrossRef]
- Naughton, V.; Grzelak, T.; Naughton, P.J. Association Between Household Location (Urban versus Rural) and Fundamental Care Provided to Domestic Dogs (Canis familiaris) in Northern Ireland. In Nutrition and Metabolism of Dogs and Cats; Advances in Experimental Medicine and Biology; Wu, G., Ed.; Springer Nature: Cham, Switzerland, 2024; Volume 1446, pp. 217–236. ISBN 978-3-031-54191-9. [Google Scholar]
- Puurunen, J.; Hakanen, E.; Salonen, M.K.; Mikkola, S.; Sulkama, S.; Araujo, C.; Lohi, H. Inadequate Socialisation, Inactivity, and Urban Living Environment Are Associated with Social Fearfulness in Pet Dogs. Sci. Rep. 2020, 10, 3527. [Google Scholar] [CrossRef]
- Hakanen, E.; Mikkola, S.; Salonen, M.; Puurunen, J.; Sulkama, S.; Araujo, C.; Lohi, H. Active and Social Life Is Associated with Lower Non-Social Fearfulness in Pet Dogs. Sci. Rep. 2020, 10, 13774. [Google Scholar] [CrossRef]
- Udell, M.A.R.; Brubaker, L.; Thielke, L.E.; Wanser, S.S.H.; Rosenlicht, G.; Vitale, K.R. Dog–Human Attachment as an Aspect of Social Cognition: Evaluating the Secure Base Test. In Comparative Cognition; Anderson, J.R., Kuroshima, H., Eds.; Springer: Singapore, 2021; pp. 305–320. ISBN 978-981-16-2027-0. [Google Scholar]
- Nagasawa, M.; Mitsui, S.; En, S.; Ohtani, N.; Ohta, M.; Sakuma, Y.; Onaka, T.; Mogi, K.; Kikusui, T. Oxytocin-Gaze Positive Loop and the Coevolution of Human-Dog Bonds. Science 2015, 348, 333–336. [Google Scholar] [CrossRef]
- Verbeek, P.; Majure, C.A.; Quattrochi, L.; Turner, S.J. The Welfare of Dogs as an Aspect of the Human–Dog Bond: A Scoping Review. Animals 2024, 14, 1985. [Google Scholar] [CrossRef]
- Bonanni, R.; Cafazzo, S.; Abis, A.; Barillari, E.; Valsecchi, P.; Natoli, E. Age-Graded Dominance Hierarchies and Social Tolerance in Packs of Free-Ranging Dogs. Behav. Ecol. 2017, 28, 1004–1020. [Google Scholar] [CrossRef]
- Bhattacharjee, D.; Bhadra, A. Humans Dominate the Social Interaction Networks of Urban Free-Ranging Dogs in India. Front. Psychol. 2020, 11, 2153. [Google Scholar] [CrossRef]
- Mikkola, S.; Salonen, M.; Puurunen, J.; Hakanen, E.; Sulkama, S.; Araujo, C.; Lohi, H. Aggressive Behaviour Is Affected by Demographic, Environmental and Behavioural Factors in Purebred Dogs. Sci. Rep. 2021, 11, 9433. [Google Scholar] [CrossRef] [PubMed]
- Bradshaw, J.W.S.; Pullen, A.J.; Rooney, N.J. Why Do Adult Dogs ‘Play’? Behav. Process. 2015, 110, 82–87. [Google Scholar] [CrossRef] [PubMed]
- Kubinyi, E.; Turcsán, B. From Kin to Canines: Understanding Modern Dog Keeping from Both Biological and Cultural Evolutionary Perspectives. Biol. Futur. 2025, 76, 213–220. [Google Scholar] [CrossRef] [PubMed]
- Porges, S.W. The Polyvagal Theory: Neurophysiological Foundations of Emotions, Attachment, Communication, and Self-Regulation; The Norton Series on Interpersonal Neurobiology; W. W. Norton: New York, NY, USA, 2011; ISBN 978-0-393-70700-7. [Google Scholar]
- Kubinyi, E. Comparative Social Cognition: From Wolf and Dog to Humans. Comp. Cogn. Behav. Rev. 2006, 2, 26–46. [Google Scholar] [CrossRef]
- Aegerter, J.; Fouracre, D.; Smith, G.C. A First Estimate of the Structure and Density of the Populations of Pet Cats and Dogs across Great Britain. PLoS ONE 2017, 12, e0174709. [Google Scholar] [CrossRef]
- Miklosi, A. Dog Behaviour, Evolution, and Cognition; Oxford University Press: Oxford, UK, 2014; ISBN 978-0-19-964666-1. [Google Scholar]
- Cimarelli, G.; Marshall-Pescini, S.; Range, F.; Berghänel, A.; Virányi, Z. Relationship Quality Affects Social Stress Buffering in Dogs and Wolves. Anim. Behav. 2021, 178, 127–140. [Google Scholar] [CrossRef]
- Jean-Joseph, H.; Kortekaas, K.; Range, F.; Kotrschal, K. Context-Specific Arousal During Resting in Wolves and Dogs: Effects of Domestication? Front. Psychol. 2020, 11, 568199. [Google Scholar] [CrossRef]
- Dietz, L.; Arnold, A.-M.K.; Goerlich-Jansson, V.C.; Vinke, C.M. The Importance of Early Life Experiences for the Development of Behavioural Disorders in Domestic Dogs. Behaviour 2018, 155, 83–114. [Google Scholar] [CrossRef]
- Stahl, A.; Barnard, S.; Diana, A.; Udell, M.A.R.; Croney, C. Attachment Style and Social Behavior in Dogs from Commercial Breeding Kennels. Appl. Anim. Behav. Sci. 2024, 273, 106238. [Google Scholar] [CrossRef]
- Capellà Miternique, H.; Gaunet, F. Coexistence of Diversified Dog Socialities and Territorialities in the City of Concepción, Chile. Animals 2020, 10, 298. [Google Scholar] [CrossRef]
- Heurlin, J.; Barabás, G.; Roth, L.S.V. Behavioural Synchronisation between Different Groups of Dogs and Wolves and Their Owners/Handlers: Exploring the Effect of Breed and Human Interaction. PLoS ONE 2024, 19, e0302833. [Google Scholar] [CrossRef] [PubMed]
- Richards, E.M.; Silver, Z.A.; Santos, L.R. Impact of the Dog–Human Bond on Canine Social Evaluation: Attachment Predicts Preference toward Prosocial Actors. Animals 2023, 13, 2480. [Google Scholar] [CrossRef] [PubMed]
- Wirobski, G.; Range, F.; Schaebs, F.S.; Palme, R.; Deschner, T.; Marshall-Pescini, S. Life Experience Rather than Domestication Accounts for Dogs’ Increased Oxytocin Release during Social Contact with Humans. Sci. Rep. 2021, 11, 14423. [Google Scholar] [CrossRef] [PubMed]
- Caldwell, H.K. Oxytocin and Vasopressin: Powerful Regulators of Social Behavior. Neuroscientist 2017, 23, 517–528. [Google Scholar] [CrossRef]
- Menon, R.; Neumann, I.D. Detection, Processing and Reinforcement of Social Cues: Regulation by the Oxytocin System. Nat. Rev. Neurosci. 2023, 24, 761–777. [Google Scholar] [CrossRef]
- Menon, R.; Süß, T.; Oliveira, V.E.D.M.; Neumann, I.D.; Bludau, A. Neurobiology of the Lateral Septum: Regulation of Social Behavior. Trends Neurosci. 2022, 45, 27–40. [Google Scholar] [CrossRef]
- Kappeler, P.M.; Silk, J.S.; Burkart, J.M.; Van Schaik, C.P. Primate Behavior and Human Universals: Exploring the Gap. In Mind the Gap; Kappeler, P.M., Silk, J., Eds.; Springer: Berlin/Heidelberg, Germany, 2010; pp. 3–15. ISBN 978-3-642-02724-6. [Google Scholar]
- Plotnik, J.M.; De Waal, F.B.M. Asian Elephants (Elephas maximus) Reassure Others in Distress. PeerJ 2014, 2, e278. [Google Scholar] [CrossRef]
- Kikusui, T. Neuroendocrinology of Social Buffering in Group Living Animals. Jpn. J. Anim. Psychol. 2018, 68, 67–75. [Google Scholar] [CrossRef][Green Version]
- Kikusui, T.; Winslow, J.T.; Mori, Y. Social Buffering: Relief from Stress and Anxiety. Philos. Trans. R. Soc. B Biol. Sci. 2006, 361, 2215–2228. [Google Scholar] [CrossRef]
- Atzil, S.; Gao, W.; Fradkin, I.; Barrett, L.F. Growing a Social Brain. Nat. Hum. Behav. 2018, 2, 624–636. [Google Scholar] [CrossRef]
- Clutton-Brock, T. Social Evolution in Mammals. Science 2021, 373, eabc9699. [Google Scholar] [CrossRef]
- Sundman, A.-S.; Van Poucke, E.; Svensson Holm, A.-C.; Faresjö, Å.; Theodorsson, E.; Jensen, P.; Roth, L.S.V. Long-Term Stress Levels Are Synchronized in Dogs and Their Owners. Sci. Rep. 2019, 9, 7391. [Google Scholar] [CrossRef]
- Schöberl, I.; Wedl, M.; Beetz, A.; Kotrschal, K. Psychobiological Factors Affecting Cortisol Variability in Human-Dog Dyads. PLoS ONE 2017, 12, e0170707. [Google Scholar] [CrossRef]
- Buttner, A.P.; Awalt, S.L.; Strasser, R. Early Life Adversity in Dogs Produces Altered Physiological and Behavioral Responses during a Social Stress-buffering Paradigm. J. Exp. Anal. Behav. 2023, 120, 6–20. [Google Scholar] [CrossRef] [PubMed]
- Ryan, M.G.; Storey, A.E.; Anderson, R.E.; Walsh, C.J. Physiological Indicators of Attachment in Domestic Dogs (Canis familiaris) and Their Owners in the Strange Situation Test. Front. Behav. Neurosci. 2019, 13, 162. [Google Scholar] [CrossRef] [PubMed]
- Pinelli, C.; Scandurra, A.; Mastellone, V.; D’Aniello, B.; Pacifico, E.; Di Lucrezia, A.; Amato, R.; Pero, M.E.; Lombardi, P. Unfamiliar Human-Induced Social Buffering Effect in Kennel-Residing Australian Cattle Dogs. Appl. Anim. Behav. Sci. 2023, 267, 106039. [Google Scholar] [CrossRef]
- Hansen Wheat, C.; Larsson, L.; Berner, P.; Temrin, H. Human-directed Attachment Behavior in Wolves Suggests Standing Ancestral Variation for Human–Dog Attachment Bonds. Ecol. Evol. 2022, 12, e9299. [Google Scholar] [CrossRef]
- Lenkei, R.; Újváry, D.; Bakos, V.; Faragó, T. Adult, Intensively Socialized Wolves Show Features of Attachment Behaviour to Their Handler. Sci. Rep. 2020, 10, 17296. [Google Scholar] [CrossRef]
- Bonanni, R.; Natoli, E.; Cafazzo, S.; Valsecchi, P. Free-Ranging Dogs Assess the Quantity of Opponents in Intergroup Conflicts. Anim. Cogn. 2011, 14, 103–115. [Google Scholar] [CrossRef]
- Handlin, L.; Hydbring-Sandberg, E.; Nilsson, A.; Ejdebäck, M.; Jansson, A.; Uvnäs-Moberg, K. Short-Term Interaction between Dogs and Their Owners: Effects on Oxytocin, Cortisol, Insulin and Heart Rate—An Exploratory Study. Anthrozoös 2011, 24, 301–315. [Google Scholar] [CrossRef]
- Engel, G.L. The Need for a New Medical Model: A Challenge for Biomedicine. Science 1977, 196, 129–136. [Google Scholar] [CrossRef]
- Quervel-Chaumette, M.; Faerber, V.; Faragó, T.; Marshall-Pescini, S.; Range, F. Investigating Empathy-Like Responding to Conspecifics’ Distress in Pet Dogs. PLoS ONE 2016, 11, e0152920. [Google Scholar] [CrossRef] [PubMed]
- Beerda, B.; Schilder, M.B.H.; Van Hooff, J.A.R.A.M.; De Vries, H.W.; Mol, J.A. Chronic Stress in Dogs Subjected to Social and Spatial Restriction. I. Behavioral Responses. Physiol. Behav. 1999, 66, 233–242. [Google Scholar] [CrossRef] [PubMed]
- Bryce, C.M. Dogs as Pets and Pests: Global Patterns of Canine Abundance, Activity, and Health. Integr. Comp. Biol. 2021, 61, 154–165. [Google Scholar] [CrossRef] [PubMed]
- Sadia Authoy, A.; Chanda, A.; Datta, A.; Al Faruk, M.S.; Kamal, T. Prevalence of Ectoparasites in Pet Dog and Cat in Dhaka, Bangladesh. Res. J. Vet. Pract. 2023, 11, 42–59. [Google Scholar] [CrossRef]
- Smith, L.M.; Hartmann, S.; Munteanu, A.M.; Dalla Villa, P.; Quinnell, R.J.; Collins, L.M. The Effectiveness of Dog Population Management: A Systematic Review. Animals 2019, 9, 1020. [Google Scholar] [CrossRef]
- Boitani, L.; Francisci, F.; Ciucci, P.; Andreoli, G. Population Biology and Ecology of Feral Dogs in Central Italy. In The Domestic Dog: Its Evolution, Behaviour and Interactions with People; Cambridge University Press: Cambridge, UK, 1995; pp. 217–244. ISBN 978-0-521-41529-3. [Google Scholar]
- Baquero, O.S.; Queiroz, M.R. Size, Spatial and Household Distribution, and Rabies Vaccination Coverage of the Brazilian Owned-dog Population. Transbound. Emerg. Dis. 2019, 66, 1693–1700. [Google Scholar] [CrossRef]
- Kubinyi, E. The Link Between Companion Dogs, Human Fertility Rates, and Social Networks. Curr. Dir. Psychol. Sci. 2025, 34, 232–239. [Google Scholar] [CrossRef]
- Andersen, S.S.; Meyer, I.; Forkman, B.; Nielsen, S.S.; Sandøe, P. Regulating Companion Dog Welfare: A Comparative Study of Legal Frameworks in Western Countries. Animals 2021, 11, 1660. [Google Scholar] [CrossRef]
- Rowan, A.; Kartal, T. Dog Population & Dog Sheltering Trends in the United States of America. Animals 2018, 8, 68. [Google Scholar] [CrossRef]
- McMillan, K.M.; Harrison, X.A.; Upjohn, M.M.; Christley, R.M.; Casey, R.A. Estimation of the Size, Density, and Demographic Distribution of the UK Pet Dog Population in 2019. Sci. Rep. 2024, 14, 31746. [Google Scholar] [CrossRef] [PubMed]
- FEDIAF Facts & Figures 2022 Published with Annual Report 2024. Available online: https://europeanpetfood.org/wp-content/uploads/2024/06/FEDIAF-Facts-Figures-2022_Online100.pdf (accessed on 26 November 2025).
- Murray, J.K.; Gruffydd-Jones, T.J.; Roberts, M.A.; Browne, W.J. Assessing Changes in the UK Pet Cat and Dog Populations: Numbers and Household Ownership. Vet. Rec. 2015, 177, 259. [Google Scholar] [CrossRef] [PubMed]
- Westgarth, C.; Pinchbeck, G.L.; Bradshaw, J.W.; Dawson, S.; Gaskell, R.M.; Christley, R.M. Factors Associated with Dog Ownership and Contact with Dogs in a UK Community. BMC Vet. Res. 2007, 3, 5. [Google Scholar] [CrossRef] [PubMed]
- Bhalla, S.J.; Kemmers, R.; Vasques, A.; Vanak, A.T. ‘Stray Appetites’: A Socio-Ecological Analysis of Free-Ranging Dogs Living alongside Human Communities in Bangalore, India. Urban Ecosyst. 2021, 24, 1245–1258. [Google Scholar] [CrossRef]
- Menchetti, L.; Calipari, S.; Guelfi, G.; Catanzaro, A.; Diverio, S. My Dog Is Not My Cat: Owner Perception of the Personalities of Dogs and Cats Living in the Same Household. Animals 2018, 8, 80. [Google Scholar] [CrossRef]
- Mellor, D. Updating Animal Welfare Thinking: Moving beyond the “Five Freedoms” towards “A Life Worth Living”. Animals 2016, 6, 21. [Google Scholar] [CrossRef]
- Pedretti, G.; Canori, C.; Costantini, E.; Palme, R.; Valsecchi, P.; Marshall-Pescini, S. Intra and Interspecific Audience Effect on Domestic Dogs’ Behavioural Displays and Facial Expressions. Sci. Rep. 2024, 14, 9546. [Google Scholar] [CrossRef]
- Huber, A.; Barber, A.L.A.; Faragó, T.; Müller, C.A.; Huber, L. Investigating Emotional Contagion in Dogs (Canis familiaris) to Emotional Sounds of Humans and Conspecifics. Anim. Cogn. 2017, 20, 703–715. [Google Scholar] [CrossRef]
- Corsetti, S.; Natoli, E.; Palme, R.; Viggiano, E. Intraspecific Interactions Decrease Stress Affecting Welfare in Shelter Dogs: A Comparison of Four Different Housing Conditions. Animals 2023, 13, 1828. [Google Scholar] [CrossRef]
- Forrest, R.; Awawdeh, L.; Esam, F.; Pearson, M.; Waran, N. Potential Owner-Related Risk Factors That May Contribute to Obesity in Companion Dogs in Aotearoa New Zealand. Animals 2022, 12, 267. [Google Scholar] [CrossRef]
- Hoffmann, L. Impact of Urbanization on Canine Health in Germany. J. Anim. Health 2024, 4, 41–53. [Google Scholar] [CrossRef]
- Shyan, M.R.; Fortune, K.A.; King, C. “Bark Parks”-A Study on Interdog Aggression in a Limited-Control Environment. J. Appl. Anim. Welf. Sci. 2003, 6, 25–32. [Google Scholar] [CrossRef] [PubMed]
- Ward, C. Greeting Behavior between Dogs in a Dog Park. Pet Behav. Sci. 2020, 10, 1–14. [Google Scholar] [CrossRef]
- Trisko, R.K.; Sandel, A.A.; Smuts, B. Affiliation, Dominance and Friendship among Companion Dogs. Behaviour 2016, 153, 693–725. [Google Scholar] [CrossRef]
- Howse, M.S.; Anderson, R.E.; Walsh, C.J. Social Behaviour of Domestic Dogs (Canis familiaris) in a Public off-Leash Dog Park. Behav. Process. 2018, 157, 691–701. [Google Scholar] [CrossRef]
- Baranyiová, E.; Holub, A.; Tyrlík, M.; Janáčková, B.; Ernstová, M. The Influence of Urbanization on the Behaviour of Dogs in the Czech Republic. Acta Vet. Brno 2005, 74, 401–409. [Google Scholar] [CrossRef]
- Berghänel, A.; Lazzaroni, M.; Ferenc, M.; Pilot, M.; El Berbri, I.; Marshall-Pescini, S.; Range, F. Cofeeding at Rich Clumped Food Patches in Free-Ranging Dogs: Social Tolerance or Scramble Competition? Behav. Ecol. Sociobiol. 2025, 79, 51. [Google Scholar] [CrossRef]
- Corrieri, L.; Adda, M.; Miklósi, Á.; Kubinyi, E. Companion and Free-Ranging Bali Dogs: Environmental Links with Personality Traits in an Endemic Dog Population of South East Asia. PLoS ONE 2018, 13, e0197354. [Google Scholar] [CrossRef]
- Melo, S.N.D.; Da Silva, E.S.; Barbosa, D.S.; Teixeira-Neto, R.G.; Lacorte, G.A.; Horta, M.A.P.; Cardoso, D.T.; Werneck, G.L.; Struchiner, C.J.; Belo, V.S. Effects of Gender, Sterilization, and Environment on the Spatial Distribution of Free-Roaming Dogs: An Intervention Study in an Urban Setting. Front. Vet. Sci. 2020, 7, 289. [Google Scholar] [CrossRef]
- Kolkmeyer, C.A.; Baum, J.; Warlich-Zach, N.; Gansloßer, U. From “Husky” to “Bulldog”–Behavioural Correlates between Castration and Breed Groups in the Domestic Dog (Canis lupus familiaris). BMC Vet. Res. 2024, 20, 238. [Google Scholar] [CrossRef]
- Arroube, A.; Pereira, A.F. Dog Neuter, Yes or No? A Summary of the Motivations, Benefits, and Harms, with Special Emphasis on the Behavioral Aspect. Animals 2025, 15, 1063. [Google Scholar] [CrossRef]
- Kolkmeyer, C.A.; Zambrano Cardona, A.M.; Gansloßer, U. Personality Unleashed: Surveying Correlation of Neuter Status and Social Behaviour in Mixed-Breed Male Dogs across Weight Classes. Animals 2024, 14, 2445. [Google Scholar] [CrossRef] [PubMed]
- Pal, S.K. Parental Care in Free-Ranging Dogs, Canis familiaris. Appl. Anim. Behav. Sci. 2005, 90, 31–47. [Google Scholar] [CrossRef]
- Paul, M.; Bhadra, A. Do Dogs Live in Joint Families? Understanding Allo-Parental Care in Free-Ranging Dogs. arXiv 2016, arXiv:1607.01138. [Google Scholar] [CrossRef]
- Silk, M.J.; Cant, M.A.; Cafazzo, S.; Natoli, E.; McDonald, R.A. Elevated Aggression Is Associated with Uncertainty in a Network of Dog Dominance Interactions. Proc. R. Soc. B Biol. Sci. 2019, 286, 20190536. [Google Scholar] [CrossRef]
- Bhattacharjee, D.; Sau, S.; Bhadra, A. ‘Bolder’ Together—Response to Human Social Cues in Groups of Free-Ranging Dogs. Behaviour 2020, 157, 363–384. [Google Scholar] [CrossRef]
- Bonanni, R.; Cafazzo, S. The Social Organisation of a Population of Free-Ranging Dogs in a Suburban Area of Rome. In The Social Dog; Elsevier: Amsterdam, The Netherlands, 2014; pp. 65–104. ISBN 978-0-12-407818-5. [Google Scholar]
- Cools, A.K.A.; Van Hout, A.J.-M.; Nelissen, M.H.J. Canine Reconciliation and Third-Party-Initiated Postconflict Affiliation: Do Peacemaking Social Mechanisms in Dogs Rival Those of Higher Primates? Ethology 2008, 114, 53–63. [Google Scholar] [CrossRef]
- Hansen Wheat, C.; Wynne, C.D.L. The Unfulfilled Potential of Dogs in Studying Behavioural Ecology and Evolution during the Anthropocene. Anim. Behav. 2025, 219, 123020. [Google Scholar] [CrossRef]
- Bonanni, R.; Cafazzo, S.; Valsecchi, P.; Natoli, E. Effect of Affiliative and Agonistic Relationships on Leadership Behaviour in Free-Ranging Dogs. Anim. Behav. 2010, 79, 981–991. [Google Scholar] [CrossRef]
- Caeiro, C.; Guo, K.; Mills, D. Dogs and Humans Respond to Emotionally Competent Stimuli by Producing Different Facial Actions. Sci. Rep. 2017, 7, 15525. [Google Scholar] [CrossRef]
- Gfrerer, N.; Taborsky, M.; Würbel, H. Benefits of Intraspecific Social Exposure in Adult Swiss Military Dogs. Appl. Anim. Behav. Sci. 2018, 201, 54–60. [Google Scholar] [CrossRef]
- Iotchev, I.B.; Egerer, A.; Grafe, S.; Adorján, A.; Kubinyi, E. Encounters between Pairs of Unfamiliar Dogs in a Dog Park. Biol. Future 2019, 70, 156–165. [Google Scholar] [CrossRef]
- Chen, S.; Wu, Z.; Sleipness, O.R.; Wang, H. Benefits and Conflicts: A Systematic Review of Dog Park Design and Management Strategies. Animals 2022, 12, 2251. [Google Scholar] [CrossRef]
- Trisko, R.K.; Smuts, B.B. Dominance Relationships in a Group of Domestic Dogs (Canis lupus familiaris). Behaviour 2015, 152, 677–704. [Google Scholar] [CrossRef]
- Vékony, K.; Pongrácz, P. Many Faces of Dominance: The Manifestation of Cohabiting Companion Dogs’ Rank in Competitive and Non-Competitive Scenarios. Anim. Cogn. 2024, 27, 12. [Google Scholar] [CrossRef]
- Lyons, D. Developmental Cascades Linking Stress Inoculation, Arousal Regulation, and Resilience. Front. Behav. Neurosci. 2009, 3, 32. [Google Scholar] [CrossRef] [PubMed]
- Guardini, G.; Bowen, J.; Mariti, C.; Fatjó, J.; Sighieri, C.; Gazzano, A. Influence of Maternal Care on Behavioural Development of Domestic Dogs (Canis familiaris) Living in a Home Environment. Animals 2017, 7, 93. [Google Scholar] [CrossRef] [PubMed]
- Pongrácz, P.; Sztruhala, S.S. Forgotten, But Not Lost—Alloparental Behavior and Pup–Adult Interactions in Companion Dogs. Animals 2019, 9, 1011. [Google Scholar] [CrossRef] [PubMed]
- Řezáč, P.; Viziová, P.; Dobešová, M.; Havlíček, Z.; Pospíšilová, D. Factors Affecting Dog–Dog Interactions on Walks with Their Owners. Appl. Anim. Behav. Sci. 2011, 134, 170–176. [Google Scholar] [CrossRef]
- Westgarth, C.; Christley, R.M.; Pinchbeck, G.L.; Gaskell, R.M.; Dawson, S.; Bradshaw, J.W.S. Dog Behaviour on Walks and the Effect of Use of the Leash. Appl. Anim. Behav. Sci. 2010, 125, 38–46. [Google Scholar] [CrossRef]
- Mehrkam, L.R.; Wynne, C.D.L. Owner Attention Facilitates Social Play in Dog–Dog Dyads (Canis lupus familiaris): Evidence for an Interspecific Audience Effect. Anim. Cogn. 2021, 24, 341–352. [Google Scholar] [CrossRef] [PubMed]
- Foyer, P.; Wilsson, E.; Wright, D.; Jensen, P. Early Experiences Modulate Stress Coping in a Population of German Shepherd Dogs. Appl. Anim. Behav. Sci. 2013, 146, 79–87. [Google Scholar] [CrossRef]
- Cafazzo, S.; Marshall-Pescini, S.; Lazzaroni, M.; Virányi, Z.; Range, F. The Effect of Domestication on Post-Conflict Management: Wolves Reconcile While Dogs Avoid Each Other. R. Soc. Open Sci. 2018, 5, 171553. [Google Scholar] [CrossRef] [PubMed]
- Range, F.; Ritter, C.; Virányi, Z. Testing the Myth: Tolerant Dogs and Aggressive Wolves. Proc. R. Soc. B Biol. Sci. 2015, 282, 20150220. [Google Scholar] [CrossRef]
- Awalt, S.L.; Boghean, L.; Klinkebiel, D.; Strasser, R. A Dog’s Life: Early Life Histories Influence Methylation of Glucocorticoid (NR3C1) and Oxytocin (OXTR) Receptor Genes, Cortisol Levels, and Attachment Styles. Dev. Psychobiol. 2024, 66, e22482. [Google Scholar] [CrossRef]
- Hennessy, M.B.; Willen, R.M.; Schiml, P.A. Psychological Stress, Its Reduction, and Long-Term Consequences: What Studies with Laboratory Animals Might Teach Us about Life in the Dog Shelter. Animals 2020, 10, 2061. [Google Scholar] [CrossRef]
- Majecka, K.; Pąsiek, M.; Pietraszewski, D.; Smith, C. Behavioural Outcomes of Housing for Domestic Dog Puppies (Canis lupus familiaris). Appl. Anim. Behav. Sci. 2020, 222, 104899. [Google Scholar] [CrossRef]
- Bray, E.E.; MacLean, E.L.; Hare, B.A. Context Specificity of Inhibitory Control in Dogs. Anim. Cogn. 2014, 17, 15–31. [Google Scholar] [CrossRef]
- Yoo, Y.; Son, Y.-H.; Zoh, K.-J. Analysis of Dog-Related Outdoor Public Space Conflicts Using Complaint Data. J. Korean Inst. Landsc. Archit. 2024, 52, 34–45. [Google Scholar] [CrossRef]
- Hubrecht, R.C. A Comparison of Social and Environmental Enrichment Methods for Laboratory Housed Dogs. Appl. Anim. Behav. Sci. 1993, 37, 345–361. [Google Scholar] [CrossRef]
- Wells, D.L. A Review of Environmental Enrichment for Kennelled Dogs, Canis familiaris. Appl. Anim. Behav. Sci. 2004, 85, 307–317. [Google Scholar] [CrossRef]
- Lopes, J.V.S.R.; Daud, N.M.; Young, R.J.; De Azevedo, C.S. To Pet or to Enrich? Increasing Dogs’ Welfare in Veterinary Clinics/Shelters: A Pilot Study. J. Vet. Behav. 2022, 52–53, 31–36. [Google Scholar] [CrossRef]
- Biswas, S.; Ghosh, K.; Ghosh, S.; Biswas, A.; Bhadra, A. What Is in a Scent? Understanding the Role of Scent Marking in Social Dynamics and Territoriality of Free-Ranging Dogs. Behav. Ecol. Sociobiol. 2025, 79, 3. [Google Scholar] [CrossRef]
- Bräuer, J.; Blasi, D. Dogs Display Owner-Specific Expectations Based on Olfaction. Sci. Rep. 2021, 11, 3291. [Google Scholar] [CrossRef] [PubMed]
- Berns, G.S.; Brooks, A.M.; Spivak, M. Scent of the Familiar: An fMRI Study of Canine Brain Responses to Familiar and Unfamiliar Human and Dog Odors. Behav. Process. 2015, 110, 37–46. [Google Scholar] [CrossRef] [PubMed]
- Van Der Borg, J.A.M.; Schilder, M.B.H.; Vinke, C.M.; De Vries, H. Dominance in Domestic Dogs: A Quantitative Analysis of Its Behavioural Measures. PLoS ONE 2015, 10, e0133978. [Google Scholar] [CrossRef]
- Hinde, R.A. Interactions, Relationships and Social Structure. Man 1976, 11, 1–17. [Google Scholar] [CrossRef]
- Stephan, G.; Leidhold, J.; Hammerschmidt, K. Pet Dogs Home Alone: A Video-Based Study. Appl. Anim. Behav. Sci. 2021, 244, 105463. [Google Scholar] [CrossRef]
- Powell, L.; Stefanovski, D.; Dreschel, N.A.; Serpell, J. The Impacts of Household Factors and Proxies of Human Social Determinants of Health on Dog Behavior. Prev. Vet. Med. 2025, 239, 106520. [Google Scholar] [CrossRef]
- Flannigan, G.; Dodman, N.H. Risk Factors and Behaviors Associated with Separation Anxiety in Dogs. J. Am. Vet. Med. Assoc. 2001, 219, 460–466. [Google Scholar] [CrossRef]
- Bakos, V.; Vékony, K.; Pongrácz, P. Hierarchy-Dependent Behaviour of Dogs in the Strange Situation Test: High-Ranking Dogs Show Less Stress and Behave Less Friendly with a Stranger in the Presence of Their Owner. Animals 2025, 15, 1916. [Google Scholar] [CrossRef] [PubMed]
- Mariti, C.; Carlone, B.; Sighieri, C.; Campera, M.; Gazzano, A. Dog Behavior in the Ainsworth Strange Situation Test during Separation from the Owner and from the Cohabitant Dog. Dog Behav. 2018, 4, 1–8. [Google Scholar] [CrossRef]
- Crockford, C.; Deschner, T.; Wittig, R.M. The Role of Oxytocin in Social Buffering: What Do Primate Studies Add? In Behavioral Pharmacology of Neuropeptides: Oxytocin; Current Topics in Behavioral Neurosciences; Hurlemann, R., Grinevich, V., Eds.; Springer International Publishing: Cham, Switzerland, 2017; Volume 35, pp. 155–173. ISBN 978-3-319-63738-9. [Google Scholar]
- Palagi, E.; Cordoni, G. Intraspecific Motor and Emotional Alignment in Dogs and Wolves: The Basic Building Blocks of Dog–Human Affective Connectedness. Animals 2020, 10, 241. [Google Scholar] [CrossRef]
- Yu, D.; Bao, L.; Yin, B. Emotional Contagion in Rodents: A Comprehensive Exploration of Mechanisms and Multimodal Perspectives. Behav. Process. 2024, 216, 105008. [Google Scholar] [CrossRef]
- Witczak, L.R.; Simmons, T.C.; Bales, K.L. Social Bond Paradoxes. In The Oxford Handbook of Evolutionary Psychology and Behavioral Endocrinology; Welling, L.L.M., Shackelford, T.K., Eds.; Oxford University Press: Oxford, UK, 2019; pp. 334–350. ISBN 978-0-19-064973-9. [Google Scholar]
- Deri, S.; Rappaz, J.; Aiello, L.M.; Quercia, D. Coloring in the Links: Capturing Social Ties as They Are Perceived. Proc. ACM Hum.-Comput. Interact. 2018, 2, 1–18. [Google Scholar] [CrossRef]
- Fincham, F.D.; Rogge, R. Understanding Relationship Quality: Theoretical Challenges and New Tools for Assessment. J. Fam. Theory Rev. 2010, 2, 227–242. [Google Scholar] [CrossRef]
- Gottfredson, R.K.; Wright, S.L.; Heaphy, E.D. A Critical Review of Relationship Quality Measures: Is a Fresh Start Needed? An Agenda to Move Forward. J. Occup. Organ. Psychol. 2022, 95, 624–659. [Google Scholar] [CrossRef]

| Region/Country | Dominant Dog Population Type | % Free-Ranging | % Single-Dog Households | Key Urban Constraints |
|---|---|---|---|---|
| India/Brazil/Africa [12,53] | Community and semi-owned dogs integrated into human environments | 70–90% | N/A | High population density, open access to streets, shared resources |
| Western Europe/US [3,54,55] | Fully owned, confined companion dogs | <5% | 70–80% | Strict leash laws, limited off-leash space, apartment living |
| Global Trend [56,57] | Shift from free-ranging to owned populations | – | Increasing single-dog ownership | Urban density, legal control, risk-averse culture |
| Existing Method | Typical Scope | Populations | What It Captures | What It Misses |
|---|---|---|---|---|
| Behavioural ethograms | Long-term, narrow scope | Multi-dog households, day-cares, shelters | Dominance and affiliation patterns | Broader social context, network structure |
| Owner or guardian surveys | Short term, broad scope | Urban pet dogs | Focused on human–dog relationship. Dyadic interaction, perceived behavioural issues | Broader social context, network structure |
| Dog-park observations | Short term, broad scope | Urban pet dogs | High-density play and conflict behaviour | Stable relationships, network structure |
| Network analysis | Long term, broad scope | Free ranging dogs | Structure and frequency of contact | Emotional valence, reciprocity, stability |
| Functional Dimension | Theoretical and Interpretive Meaning | Example Behavioural/Relational Indicators | Possible Data Sources or Methods |
|---|---|---|---|
| Tension (arousal/constraint) | Expresses the degree of physiological and behavioural arousal within relational patterns; high tension indicates low perceived safety or regulatory overload | Body stiffness, scanning, speed of movement, micro-pauses (“freezes”), displacement behaviours | Observation, video coding, heart rate variability or cortisol |
| Relational valence | Reflects the overall affective tone and balance of participation within interactions; indicates whether exchanges are cooperative, neutral, or conflictual | Frequency of positive vs. negative behaviours, reciprocity, ease of repair after conflict | Field observation, video coding, guardian reports |
| Bond stability | Describes the range and heterogeneity of dogs encountered within the individual’s social environment; includes both brief and recurrent contacts | Number of distinct partners encountered over a defined period, variety of contexts (home, park, walks), proportion of novel vs. familiar dogs | Structured observation, short guardian logs of daily encounters, annotated video samples |
| Social integration | Represents the individual’s embeddedness within a broader social network; linked to access to social support and group cohesion | Number and diversity of partners, network centrality, isolation, bridging roles | Social network analysis, observational mapping |
| Contact Zone | Description | Typical Behaviours or Signals |
|---|---|---|
| Distal | Interaction initiated at a distance; communication occurs through posture, orientation, and gaze (usually beyond 5–10 m, depending on terrain and visibility) | Sustained gaze, curved approach, intentional stop, body angling, tension release through movement, scent marking |
| Mid-range | Mutual awareness and approach within visible and olfactory range, without physical contact (typically 1–5 m) | Parallel orientation, slow approach, posture signalling intent, relaxed stance, micro-pauses, scent marking and overmarking |
| Proximal | Close proximity allowing brief or partial physical contact (within roughly 1 m or one body length) | Sniffing, nudging, shoulder or side contact, short affiliative pauses, shared resting area |
| Tactile | Full physical contact; overlap of personal space or sustained touch (direct contact, 0 m) | Play, grooming, leaning, lying together, affiliative body pressure |
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Grynkiewicz, A.; Reinholz, A.; Imbir, K. Disconnected Lives: Social Networks and Emotional Regulation in Domestic Dogs. Animals 2026, 16, 398. https://doi.org/10.3390/ani16030398
Grynkiewicz A, Reinholz A, Imbir K. Disconnected Lives: Social Networks and Emotional Regulation in Domestic Dogs. Animals. 2026; 16(3):398. https://doi.org/10.3390/ani16030398
Chicago/Turabian StyleGrynkiewicz, Agnieszka, Anna Reinholz, and Kamil Imbir. 2026. "Disconnected Lives: Social Networks and Emotional Regulation in Domestic Dogs" Animals 16, no. 3: 398. https://doi.org/10.3390/ani16030398
APA StyleGrynkiewicz, A., Reinholz, A., & Imbir, K. (2026). Disconnected Lives: Social Networks and Emotional Regulation in Domestic Dogs. Animals, 16(3), 398. https://doi.org/10.3390/ani16030398

