Skip to main content
Log in

Free-ranging dogs assess the quantity of opponents in intergroup conflicts

  • Original Paper
  • Published:
Animal Cognition Aims and scope Submit manuscript

Abstract

In conflicts between social groups, the decision of competitors whether to attack/retreat should be based on the assessment of the quantity of individuals in their own and the opposing group. Experimental studies on numerical cognition in animals suggest that they may represent both large and small numbers as noisy mental magnitudes subject to scalar variability, and small numbers (≤4) also as discrete object-files. Consequently, discriminating between large quantities, but not between smaller ones, should become easier as the asymmetry between quantities increases. Here, we tested these hypotheses by recording naturally occurring conflicts in a population of free-ranging dogs, Canis lupus familiaris, living in a suburban environment. The overall probability of at least one pack member approaching opponents aggressively increased with a decreasing ratio of the number of rivals to that of companions. Moreover, the probability that more than half of the pack members withdrew from a conflict increased when this ratio increased. The skill of dogs in correctly assessing relative group size appeared to improve with increasing the asymmetry in size when at least one pack comprised more than four individuals, and appeared affected to a lesser extent by group size asymmetries when dogs had to compare only small numbers. These results provide the first indications that a representation of quantity based on noisy mental magnitudes may be involved in the assessment of opponents in intergroup conflicts and leave open the possibility that an additional, more precise mechanism may operate with small numbers.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3
Fig. 4

Similar content being viewed by others

References

  • Adams ES (1990) Boundary disputes in the territorial ant Azteca trigona: effects of asymmetries in colony size. Anim Behav 39:321–328

    Article  Google Scholar 

  • Agresti A (1996) An introduction to categorical data analysis. Wilely, New York

    Google Scholar 

  • Agrillo C, Dadda M, Serena G, Bisazza A (2008) Do fish count? Spontaneous discrimination of quantity in female mosquitofish. Anim Cogn 11:495–503

    Article  PubMed  Google Scholar 

  • Al Aїn S, Giret N, Grand M, Kreutzer M, Bovet D (2009) The discrimination of discrete and continuous amounts in African grey parrots (Psittacus erithacus). Anim Cogn 12:145–154

    Article  Google Scholar 

  • Altmann J (1974) Observational study of behavior: sampling methods. Behaviour 49:227–267

    Article  CAS  PubMed  Google Scholar 

  • Austad SN (1983) A game theoretical interpretation of male combat in the bowl and doily spider (Frontinella pyramitela). Anim Behav 31:59–73

    Article  Google Scholar 

  • Bekoff M (1979) Scent-marking by free-ranging domestic dogs. Biol behav 4:123–139

    Google Scholar 

  • Beran MJ (2001) Summation and numerousness judgements of sequentially presented sets of items by chimpanzees (Pan troglodytes). J Comp Psychol 115:181–191

    Article  CAS  PubMed  Google Scholar 

  • Beran MJ (2004) Chimpanzees (Pan troglodytes) respond to nonvisible sets after one-by-one addition and removal of items. J Comp Psychol 118:25–36

    Article  PubMed  Google Scholar 

  • Beran MJ (2007) Rhesus monkeys (Macaca mulatta) enumerate large and small sequentially presented sets of items using analog numerical representations. J Exp Psychol 33:42–54

    Google Scholar 

  • Black JM, Owen M (1989) Agonistic behaviour in barnacle goose flocks: assessment, investment and reproductive success. Anim Behav 37:199–209

    Article  Google Scholar 

  • Boitani L, Ciucci P (1995) Comparative social ecology of feral dogs and wolves. Ethol Ecol Evol 7:49–72

    Article  Google Scholar 

  • Boitani L, Francisci F, Ciucci P, Andreoli G (1995) Population biology and ecology of feral dogs in central Italy. In: Serpell J (ed) The domestic dog: its evolution, behaviour and interactions with people. Cambridge university press, Cambridge, pp 217–244

    Google Scholar 

  • Bonanni R, Cafazzo S, Fantini C, Pontier D, Natoli E (2007) Feeding-order in an urban feral domestic cat colony: relationship to dominance rank, sex and age. Anim Behav 74:1369–1379

    Article  Google Scholar 

  • Bonanni R, Valsecchi P, Natoli E (2010a) Pattern of individual participation and cheating in conflicts between groups of free-ranging dogs. Anim Behav 79:957–968

    Article  Google Scholar 

  • Bonanni R, Cafazzo S, Valsecchi P, Natoli E (2010b) Effect of affiliative and agonistic relationships on leadership behaviour in free-ranging dogs. Anim Behav 79:981–991

    Article  Google Scholar 

  • Bradshaw JWS, Nott HMR (1995) Social and communication behaviour of companion dogs. In: Serpell J (ed) The domestic dog: its evolution, behaviour and interactions with people. Cambridge university press, Cambridge, pp 115–130

    Google Scholar 

  • Brannon EM, Terrace HS (1998) Ordering of the numerosities 1 to 9 by monkeys. Science 282:746–749

    Article  CAS  PubMed  Google Scholar 

  • Cafazzo S (2007) Dinamiche sociali in un gruppo di cani domestici (Canis lupus familiaris) liberi in ambiente suburbano. Ph.D. thesis, University of Parma, Parma, Italy

  • Cafazzo S, Valsecchi P, Bonanni R, Natoli E (2010) Dominance in relation to age, sex and competitive contexts in a group of free-ranging domestic dogs. Behav Ecol 21:443–455

    Article  Google Scholar 

  • Call J (2000) Estimating and operating on discrete quantities in orangutans (Pongo pygmaeus). J Comp Psychol 114:136–147

    Article  CAS  PubMed  Google Scholar 

  • Cantlon JF, Brannon EM (2006) Shared system for ordering small and large numbers in monkeys and humans. Psychol Sci 17:401–406

    Article  PubMed  Google Scholar 

  • Cheney DL (1987) Interactions and relationship between groups. In: Smuts BB, Cheney DL, Seyfarth RM, Wrangham RW, Struhsaker TT (eds) Primate societies. University of Chicago Press, Chicago, pp 267–281

    Google Scholar 

  • Clutton-Brock J (1995) Origins of the dog: domestication and early history. In: Serpell J (ed) The domestic dog: its evolution, behaviour and interactions with people. Cambridge university press, Cambridge, pp 7–20

    Google Scholar 

  • Coppinger R, Schneider R (1995) Evolution of working dogs. In: Serpell J (ed) The domestic dog: its evolution, behaviour and interactions with people. Cambridge university press, Cambridge, pp 21–47

    Google Scholar 

  • Cordes S, Brannon EM (2009) Crossing the divide: infants discriminate small from large numerosities. Dev Psychol 45:1583–1594

    Article  PubMed  Google Scholar 

  • Daniels TJ, Bekoff M (1989a) Spatial and temporal resource use by feral and abandoned dogs. Ethology 81:300–312

    Article  Google Scholar 

  • Daniels TJ, Bekoff M (1989b) Population and social biology of free-ranging dogs, Canis familiaris. J Mammal 70:754–762

    Article  Google Scholar 

  • Darlington RB (1990) Regression and linear models. McGraw-Hill, New York

    Google Scholar 

  • Dobson AJ (1990) An introduction to generalized linear models. Chapman & Hall, New York

    Google Scholar 

  • Emmerton J, Renner JC (2009) Local rather than global processing of visual arrays in numerosity discrimination by pigeons (Columbia livia). Anim Cogn 12:511–526

    Article  PubMed  Google Scholar 

  • Enquist M, Leimar O (1987) Evolution of fighting behaviour: the effect of variation in resource value. J Theor Biol 127:187–205

    Article  Google Scholar 

  • Feigenson L, Carey S (2005) On the limits of infants’ quantification of small object arrays. Cognition 97:295–313

    Article  PubMed  Google Scholar 

  • Feigenson L, Carey S, Hauser MD (2002) The representations underlying infants’ choice of more: object files versus analog magnitudes. Psychol Sci 13:150–156

    Article  PubMed  Google Scholar 

  • Feigenson L, Dehaene S, Spelke E (2004) Core systems of number. Trends Cogn Sci 8:307–314

    Article  PubMed  Google Scholar 

  • Flombaum JI, Junge JA, Hauser MD (2005) Rhesus monkeys (Macaca mulatta) spontaneously compute addition operations over large numbers. Cognition 97:315–325

    Article  PubMed  Google Scholar 

  • Font E (1987) Spacing and social organization: urban stray dogs revisited. Appl Anim Behav Sci 17:319–328

    Article  Google Scholar 

  • Gallistel CR, Gelman R (2000) Non-verbal numerical cognition: from reals to integers. Trends Cogn Sci 4:59–65

    Article  PubMed  Google Scholar 

  • Gese EM (2001) Territorial defense by coyotes (Canis latrans) in Yellowstone National Park, Wyoming: who, how, where, when and why. Can J Zool 79:980–987

    Article  Google Scholar 

  • Hammerstein P, Parker GA (1982) The asymmetric war of attrition. J Theor Biol 96:647–682

    Article  Google Scholar 

  • Harrington FH, Mech LD (1979) Wolf howling and its role in territory maintenance. Behaviour 68:207–249

    Article  Google Scholar 

  • Harris S, Cresswell WJ, Forde PG, Trewhella WJ, Wollard T, Wray S (1990) Home-range analysis using radio-telemetry data: a review of problems and techniques particularly as applied to the study of mammals. Mammal Rev 20:97–123

    Article  Google Scholar 

  • Hauser MD, Spelke ES (2004) Evolutionary and developmental foundations of human knowledge: a case study of mathematics. In: Gazzaniga M (ed) The cognitive neurosciences, vol 3. MIT Press, Cambridge

    Google Scholar 

  • Hauser MD, Carey S, Hauser LB (2000) Spontaneous number representation in semi-free-ranging rhesus monkeys. Proc R Soc Lond B 267:829–833

    Article  CAS  Google Scholar 

  • Hauser MD, Tsao F, Garcia P, Spelke ES (2003) Evolutionary foundations of number: spontaneous representations of numerical magnitudes by cotton-top tamarins. Proc R Soc Lond B 270:1441–1446

    Article  Google Scholar 

  • Hofer H, East ML (1993) The commuting system of Serengeti spotted hyenas: how a predator copes with migratory prey. II. Intrusion pressure and commuters’ space use. Anim Behav 46:559–574

    Article  Google Scholar 

  • Johnsson JI, Forser A (2002) Residence duration influences the outcome of territorial conflicts in brown trout (Salmo trutta). Behav Ecol Sociobiol 51:282–286

    Article  Google Scholar 

  • Kilian A, Yaman S, von Fersen L, Gunturkun O (2003) A bottlenose dolphin discriminate visual stimuli differing in numerosity. Learn Behav 31:133–142

    PubMed  Google Scholar 

  • Kitchen DM (2004) Alpha male black howler monkey responses to loud calls: effect of numeric odds, male companion behaviour and reproductive investment. Anim Behav 67:125–139

    Article  Google Scholar 

  • Kitchen DM, Cheney DL, Seyfarth RM (2004) Factors mediating inter-group encounters in savannah baboons (Papio cynocephalus ursinus). Behaviour 141:197–218

    Article  Google Scholar 

  • Krebs JR (1982) Territorial defence in the great tit (Parus major): do residents always win? Behav Ecol Sociobiol 11:185–194

    Article  Google Scholar 

  • Maarschalkerweerd RJ, Endenburg N, Kirpensteijn J, Knol BW (1997) Influence of orchiectomy on canine behaviour. Vet Rec 140:617–619

    Article  CAS  PubMed  Google Scholar 

  • Macdonald DW, Carr GM (1995) Variation in dog society: between resource dispersion and social flux. In: Serpell J (ed) The domestic dog: its evolution, behaviour and interactions with people. Cambridge university press, Cambridge, pp 199–216

    Google Scholar 

  • Maynard Smith J, Parker GA (1976) The logic of asymmetric contests. Anim Behav 24:159–175

    Article  Google Scholar 

  • McComb K, Packer C, Pusey A (1994) Roaring and numerical assessment in contests between groups of female lions, Panthera leo. Anim Behav 47:379–387

    Article  Google Scholar 

  • Mech LD (1970) The wolf: the ecology and behaviour of an endangered species. Natural History Press, Doubleday, New York

    Google Scholar 

  • Meck WH, Church RM (1983) A mode control model of counting and timing processes. J Exp Psychol Anim Behav Process 9:320–334

    Article  CAS  PubMed  Google Scholar 

  • Morrell LJ, Lindstrom J, Ruxton GD (2005) Why are small males aggressive? Proc R Soc Lond B 272:1235–1241

    Article  Google Scholar 

  • Packer C, Scheel D, Pusey AE (1990) Why lions form groups? Food is not enough. Am Nat 136:1–19

    Article  Google Scholar 

  • Pal SK, Ghosh B, Roy S (1998) Agonistic behaviour of free-ranging dogs (Canis familiaris) in relation to season, sex and age. Appl Anim Behav Sci 59:331–348

    Article  Google Scholar 

  • Parker GA (1974) Assessment strategy and the evolution of fighting behaviour. J Theor Biol 47:223–243

    Article  CAS  PubMed  Google Scholar 

  • Parker GA, Rubenstein DI (1981) Role assessment, reserve strategy, and acquisition of information in asymmetric animal conflicts. Anim Behav 29:221–240

    Article  Google Scholar 

  • Pepperberg IM (2006) Grey parrot numerical competence: a review. Anim Cogn 9:377–391

    Article  PubMed  Google Scholar 

  • Peters RP, Mech LD (1975) Scent-marking in wolves. Am Sci 63:628–637

    CAS  PubMed  Google Scholar 

  • Pisa EP, Agrillo C (2009) Quantity discrimination in felines: a preliminary investigation of the domestic cat (Felis silvestris catus). J Ethol 27:289–293

    Article  Google Scholar 

  • Platt JR, Johnson DM (1971) Localization of position within an homogeneous behaviour chain: effects of error contingencies. Learn Motiv 2:386–414

    Article  Google Scholar 

  • Price EO (1984) Behavioral aspects of animal domestication. Q Rev Biol 59:1–32

    Article  Google Scholar 

  • Revkin SK, Piazza M, Izard V, Cohen L, Dehaene S (2008) Does subitizing reflect numerical estimation? Psychol Sci 19:607–614

    Article  PubMed  Google Scholar 

  • Sillero-Zubiri C, Macdonald DW (1998) Scent-marking and territorial behaviour of ethiopian wolves Canis simensis. J Zool Lond 245:351–361

    Article  Google Scholar 

  • Tanner CJ (2006) Numerical assessment affects aggression and competitive ability: a team-fighting strategy for the ant Formica xerophila. Proc R Soc London B Biol Sci 273:2737–2742

    Article  Google Scholar 

  • Tobias J (1997) Asymmetric territorial contests in the European robin: the role of settlement costs. Anim Behav 54:9–21

    Article  PubMed  Google Scholar 

  • Uller C, Jaeger R, Guidry G, Martin C (2003) Salamanders (Plethodon cinereus) go for more: rudiments of number in a species of basal vertebrate. Anim Cogn 6:105–112

    PubMed  Google Scholar 

  • Vilà C, Savolainen P, Maldonado JE, Amorin IR, Rice JE, Honeycutt RL, Crendall KA, Lundeberg J, Wayne RK (1997) Multiple and ancient origins of the domestic dog. Science 276:1687–1689

    Article  PubMed  Google Scholar 

  • Ward C, Smuts BB (2007) Quantity-based judgments in the domestic dog (Canis lupus familiaris). Anim Cogn 10:71–80

    Article  PubMed  Google Scholar 

  • West RE, Young RJ (2002) Do domestic dogs show any evidence of being able to count? Anim Cogn 5:183–186

    PubMed  Google Scholar 

  • Wilson ML, Hauser MD, Wrangham RW (2001) Does participation in intergroup conflict depend on numerical assessment, range location or rank for wild chimpanzee? Anim Behav 61:1203–1216

    Article  Google Scholar 

Download references

Acknowledgments

We are deeply indebted to Christian Agrillo who provided fundamental suggestions and relevant literature. We also would like to thank Annamaria Andreozzi and Mirella De Paolis for helping with the dog census; Mario Di Traglia, Alessandro Giuliani and Orazio Rossi for statistical advices; Oliver P. Hoener for providing long stimulating discussions about territoriality and the methods for assessing it; Gina Raihani and Marion L. East for language revision; Rolf O. Peterson and Manuela Piazza for providing useful literature; the three anonymous referee whose suggestions greatly improved the manuscript. A special thank goes also to Luis Nieder who provided support and facilities. Finally, this research was partially funded by University of Parma with FIL 2005 and FIL 2006 to Paola Valsecchi, and it complies with all laws of the country (Italy) in which it was performed.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Roberto Bonanni.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Bonanni, R., Natoli, E., Cafazzo, S. et al. Free-ranging dogs assess the quantity of opponents in intergroup conflicts. Anim Cogn 14, 103–115 (2011). https://doi.org/10.1007/s10071-010-0348-3

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10071-010-0348-3

Keywords

Navigation