Behavioral Ecology and Sociobiology

, Volume 70, Issue 12, pp 2185–2193 | Cite as

A mixed strategy of counter-singing behavior in male rock hyrax vocal competitions

  • Vlad Demartsev
  • Amiyaal Ilany
  • Adi Barocas
  • Einat Bar Ziv
  • Inbar Schnitzer
  • Lee Koren
  • Eli Geffen
Original Article

Abstract

Maynard-Smith’s game-theoretical framework suggests that conflicts should be resolved through an exchange of agonistic displays which can be as effective as physical attack in securing contested resources. Hence, participants use signaling tactics that are governed by game-theoretical competition rules to maximize their benefit from the interaction. In rock hyraxes, males initiate singing in response to conspecific male songs, possibly to self-advertise and to facilitate comparison of performance by receivers. We hypothesized that hyrax counter-singing strategy can be explained by resident-intruder model and its outcome is determined by the identity and ability of the participants. Natural hyrax singing events and playback-induced events were classified according to the participants’ residence status—resident or bachelor (i.e., non-resident). As predicted by the resident-intruder model, resident listeners tended to reply to conspecific males at high rates (≈40 %), while bachelors showed low reply rates (17 %) to resident initiators. Unexpectedly, the lowest-ranking bachelors showed a tendency to reply to the highest-ranking residents. One possible explanation for this counter-intuitive behavior might be that weaker individuals exploit a situation in which it is unremunerated for a stronger opponent to retaliate (i.e., Napoleon strategy) and seize a self-advertisement opportunity in front of a large audience attending to the high-ranking males’ signal. Overall, we show that male hyraxes employ different counter-singing tactics depending on the residence and rank differences between the participants. This choice of competition tactics can be based on opponents’ individual recognition or/and on direct decoding of opponents’ traits encoded in the song.

Significance statement

In nature, conflicts are usually resolved through signaling rather than by actual fighting. Rock hyrax males often participate in vocal competitions, possibly to reduce the need for physical confrontations. During our study on the dynamics of male hyrax vocal competition, we discovered that hyraxes employ mixed counter-singing strategy dependent on the residence status and social rank differences between the participants. A paradoxical “Napoleon-complex” tactic was observed in interactions between males, when low ranking bachelors were found to interact and challenge the highest-ranking residents. On the other hand, residents were found to interact with opponents regardless of residency and/or rank. On top of the cognitive abilities required to individually recognize an opponent and possibly decode its vocal signals, our results suggest that male hyraxes compare own individual traits to those of the opponent in a mutual signal assessment interaction.

Keywords

Counter-singing Vocal competition Napoleon strategy Resident-intruder model 

Supplementary material

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ESM 2S2-S4: Adult male rock-hyrax songs recorded in Ein-Gedi Natural Reserve, Israel. Sample of recordings used for playback experiments. (WAV 9935 kb)
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ESM 4(WAV 7628 kb)
265_2016_2222_MOESM5_ESM.pdf (684 kb)
ESM 5(PDF 683 kb)

References

  1. Adkins-Regan E (2005) Hormones and animal social behavior. Princeton University Press, PrincetonGoogle Scholar
  2. Aragon P, Lopez P, Martin J (2006) Roles of male residence and relative size in the social behavior of Iberian rock lizards, Lacerta monticola. Behav Ecol Sociobiol 59:762–769CrossRefGoogle Scholar
  3. Bar Ziv E, Ilany A, Demartsev V, Barocas A, Geffen E, Koren L (2016) Individual, social, and sexual niche traits affect copulation success in a polygynandrous mating system. Behav Ecol Sociobiol 70:901–912CrossRefGoogle Scholar
  4. Barlow G, Rogers W, Fraley N (1986) Do Midas cichlids win through prowess or daring? It depends. Behav Ecol Sociobiol 19:1–8CrossRefGoogle Scholar
  5. Barocas A, Ilany A, Koren L, Kam M, Geffen E (2011) Variance in centrality within rock hyrax social networks predicts adult longevity. PLoS One 6:e22375CrossRefPubMedPubMedCentralGoogle Scholar
  6. Behr O, Knörnschild M, von Helversen O (2009) Territorial counter-singing in male sac-winged bats (Saccopteryx bilineata): low-frequency songs trigger a stronger response. Behav Ecol Sociobiol 63:433–442CrossRefGoogle Scholar
  7. Bergman M, Olofsson M, Wiklund C (2010) Contest outcome in a territorial butterfly: the role of motivation. Proc R Soc Lond B 277:3027–3033CrossRefGoogle Scholar
  8. Blas J, Cabezas S, Figuerola J, Lopez L, Tanferna A, Hiraldo F (2013) Carotenoids and skin coloration in a social raptor. J Raptor Res 47:174–184CrossRefGoogle Scholar
  9. Bolyard KJ, Rowland WJ (2000) The effects of spatial context and social experience on the territorial aggression of male threespine stickleback. Behaviour 137:845–864CrossRefGoogle Scholar
  10. Demartsev V, Kershenbaum A, Ilany A, Barocas A, Bar Ziv E, Koren L, Geffen E (2014) Male hyraxes increase song complexity and duration in the presence of alert individuals. Behav Ecol 25:1451–1458CrossRefGoogle Scholar
  11. Dirienzo N, Hedrick AV (2014) Animal personalities and their implications for complex signaling. Curr Zool 60:381–386CrossRefGoogle Scholar
  12. Elias DO, Botero CA, Andrade MCB, Mason AC, Kasumovic MM (2010) High resource valuation fuels “desperado” fighting tactics in female jumping spiders. Behav Ecol 21:868–875CrossRefGoogle Scholar
  13. Fischer J, Kitchen D, Seyfarth R, Cheney D (2004) Baboon loud calls advertise male quality: acoustic features and their relation to rank, age, and exhaustion. Behav Ecol Sociobiol 56:140–148CrossRefGoogle Scholar
  14. Fourie PB (1977) Acoustic communication in the Rock Hyrax (Procavia capensis. Z Tierpsychol 44:194–219Google Scholar
  15. Frommolt K, Goltsman M, MacDonald D (2003) Barking foxes, Alopex lagopus: field experiments in individual recognition in a territorial mammal. Anim Behav 65:509–518CrossRefGoogle Scholar
  16. Gammell MP, De Vries H, Jennings DJ, Carlin CM, Hayden TJ (2003) David's score: a more appropriate dominance ranking method than Clutton-Brock et al.'s index. Anim Behav 66:601–605CrossRefGoogle Scholar
  17. Getty T (1987) Dear enemies and the prisoners-dilemma—why should territorial neighbors form defensive coalitions. Am Zool 27:327–336CrossRefGoogle Scholar
  18. Getty T (1989) Are dear enemies in a war of attrition? Anim Behav 37:337–339CrossRefGoogle Scholar
  19. Grafen A (1987) The logic of divisively asymmetric contests—respect for ownership and the desperado effect. Anim Behav 35:462–467CrossRefGoogle Scholar
  20. Hurd PL (2006) Resource holding potential, subjective resource value, and game theoretical models of aggressiveness signalling. J Theor Biol 241:639–648CrossRefPubMedGoogle Scholar
  21. Ilany A, Barocas A, Koren L, Kam M, Geffen E (2011) Do singing rock hyraxes exploit conspecific calls to gain attention? PLoS One 6:e28612CrossRefPubMedPubMedCentralGoogle Scholar
  22. Ilany A, Barocas A, Kam M, Ilany T, Geffen E (2013) The energy cost of singing in wild rock hyrax males: evidence for an index signal. Anim Behav 85:995–1001CrossRefGoogle Scholar
  23. Innocent TM, West SA, Sanderson JL, Hyrkkanen N, Reece SE (2011) Lethal combat over limited resources: testing the importance of competitors and kin. Behav Ecol 22:923–931CrossRefPubMedPubMedCentralGoogle Scholar
  24. Jennings DJ, Elwood RW, Carlin CM, Hayden TJ, Gammell MP (2012) Vocal rate as an assessment process during fallow deer contests. Behav Process 91:152–158CrossRefGoogle Scholar
  25. Jenssen TA, Decourcy KR, Congdon JD (2005) Assessment in contests of male lizards (Anolis carolinensis): how should smaller males respond when size matters? Anim Behav 69:1325–1336CrossRefGoogle Scholar
  26. Just W, Morris MR (2003) The Napoleon Complex: why smaller males pick fights. Evol Ecol 17:509–522CrossRefGoogle Scholar
  27. Kitchen DM, Seyfarth RM, Fischer J, Cheney DL (2003) Loud calls as indicators of dominance in male baboons (Papio cynocephalus ursinus. Behav Ecol Sociobiol 53:374–384Google Scholar
  28. Kokko H, Rintamaki PT, Alatalo RV, Hoglund J, Karvonen E, Lundberg A (1999) Female choice selects for lifetime lekking performance in black grouse males. Proc R Soc Lond B 266:2109–2115CrossRefGoogle Scholar
  29. Kokko H, Lopez-Sepulcre A, Morrell LJ (2006) From hawks and doves to self-consistent games of territorial behavior. Am Nat 167:901–912CrossRefPubMedGoogle Scholar
  30. Koren L, Geffen E (2009) Complex call in male rock hyrax (Procavia capensis): a multi-information distributing channel. Behav Ecol Sociobiol 63:581–590CrossRefGoogle Scholar
  31. Koren L, Geffen E (2011) Individual identity is communicated through multiple pathways in male rock hyrax (Procavia capensis) songs. Behav Ecol Sociobiol 65:675–684CrossRefGoogle Scholar
  32. Koren L, Mokady O, Geffen E (2008) Social status and cortisol levels in singing rock hyraxes. Horm Behav 54:212–216CrossRefPubMedGoogle Scholar
  33. Lindstrom K (2001) Effects of resource distribution on sexual selection and the cost of reproduction in sandgobies. Am Nat 158:64–74CrossRefPubMedGoogle Scholar
  34. Lotem A (1998) Higher levels of begging behavior by small nestlings: a case of a negatively correlated handicap. Isr J Zool 44:29–45Google Scholar
  35. Mateos C, Carranza J (1997) The role of bright plumage in male-male interactions in the ring-necked pheasant. Anim Behav 54:1205–1214CrossRefPubMedGoogle Scholar
  36. Maynard-Smith J, Parker GA (1976) Logic of asymmetric contests. Anim Behav 24:159–175CrossRefGoogle Scholar
  37. Michod RE, Hasson O (1990) On the evolution of reliable indicators of fitness. Am Nat 135:788–808CrossRefGoogle Scholar
  38. Miyai CA, Carretero Sanches FH, Costa TM, Colpo KD, Volpato GL, Barreto RE (2011) The correlation between subordinate fish eye colour and received attacks: a negative social feedback mechanism for the reduction of aggression during the formation of dominance hierarchies. Zoology 114:335–339CrossRefPubMedGoogle Scholar
  39. Morrell LJ, Lindstrom J, Ruxton GD (2005) Why are small males aggressive? Proc R Soc Lond B 272:1235–1241CrossRefGoogle Scholar
  40. Olsson M, Shine R (2000) Ownership influences the outcome of male-male contests in the scincid lizard, Niveoscincus microlepidotus. Behav Ecol 11:587–590CrossRefGoogle Scholar
  41. Reby D, McComb K, Cargnelutti B, Darwin C, Fitch WT, Clutton-Brock T (2005) Red deer stags use formants as assessment cues during intrasexual agonistic interactions. Proc R Soc Lond B 272:941–947CrossRefGoogle Scholar
  42. Reddon AR, Balk D, Balshine S (2013) Probing aggressive motivation during territorial contests in a group-living cichlid fish. Behav Process 92:47–51CrossRefGoogle Scholar
  43. Russ BE, Comins JA, Smith R, Hauser MD (2010) Recognizing and respecting claims over resources in free-ranging rhesus monkeys, Macaca mulatta. Anim Behav 80:563–569CrossRefGoogle Scholar
  44. Schradin C, Schneider C, Lindholm AK (2010) The nasty neighbour in the striped mouse (Rhabdomys pumilio) steals paternity and elicits aggression. Front Zool 7:19CrossRefPubMedPubMedCentralGoogle Scholar
  45. Searcy WA, Beecher MD (2009) Song as an aggressive signal in songbirds. Anim Behav 78:1281–1292CrossRefGoogle Scholar
  46. Shuster SM, Wade MJ (2003) Mating systems and strategies. Princeton University Press, PrincetonGoogle Scholar
  47. Sih A, Hanser SF, McHugh KA (2009) Social network theory: new insights and issues for behavioral ecologists. Behav Ecol Sociobiol 63:975–988CrossRefGoogle Scholar
  48. Smith IP, Huntingford FA, Atkinson RJA, Taylor AC (1994) Strategic decisions during agonistic behaviour in the velvet swimming crab (Necora puber L. Anim Behav 47:885–894CrossRefGoogle Scholar
  49. Svensson PA, Lehtonen TK, Wong BBM (2012) A high aggression strategy for smaller males. PLoS One 7:e43121CrossRefPubMedPubMedCentralGoogle Scholar
  50. Temeles EJ (1994) The role of neighbors in territorial systems—when are they dear enemies. Anim Behav 47:339–350CrossRefGoogle Scholar
  51. Umbers KDL, Osborne L, Keogh JS (2012) The effects of residency and body size on contest initiation and outcome in the territorial dragon, Ctenophorus decresii. PLoS One 7:e47143CrossRefPubMedPubMedCentralGoogle Scholar
  52. Yack JE, Gill S, Drummond-Main C, Sherratt TN (2014) Residency duration and shelter quality influence vibratory signalling displays in a territorial caterpillar. Ethology 120:354–364CrossRefGoogle Scholar
  53. Ydenberg RC, Giraldeau LA, Falls JB (1988) Neighbours, strangers, and the asymmetric war of attrition. Anim Behav 36:343–347CrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  1. 1.Department of ZoologyTel Aviv UniversityTel AvivIsrael
  2. 2.The Mina and Everard Goodman Faculty of Life SciencesBar-Ilan UniversityRamat-GanIsrael
  3. 3.Department of Zoology and Physiology and Program in EcologyUniversity of WyomingLaramieUSA

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