Skip to main content
SpringerLink
Log in

Male ruff colour as a rank signal in a monomorphic-horned mammal: behavioural correlates

  • Original Paper
  • Published:
The Science of Nature Aims and scope Submit manuscript

Abstract

Coexistence of individuals within a social group is possible through the establishment of a hierarchy. Social dominance is achieved through aggressive interactions, and, in wild sheep and goats, it is related mainly to age, body size and weapon size as rank signals. Adult male Himalayan tahr are much larger than females and subadult males. They have a prominent neck ruff, ranging in colour from yellow (5.5–9.5 years old, i.e. young adults, golden males) to brown (7.5–14.5 years old, i.e. older individuals, pale and dark brown males), with golden males being the most dominant. We investigated the social behaviour of male tahr and analysed the relationships between ruff colour, courtship and agonistic behaviour patterns during the rut. Colour classes varied in their use of several behaviour patterns (male dominance: approach, stare, horning vegetation; courtship: low stretch, naso-genital contact, rush). Golden-ruffed males used more threats than darker ones. Pale brown and dark brown males addressed threats significantly more often to males of lower or their own colour classes, respectively, whereas golden ones addressed threats to all colour classes, including their own. The courtship of dominant males was characterised by the assertive rush, whereas that of subordinates did not. Ruff colour of male Himalayan tahr may have evolved as a rank signal, homologous to horn size in wild sheep and goats.

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

Access this article

Log in via an institution

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

Instant access to the full article PDF.

Institutional subscriptions

Fig. 1
Fig. 2
Fig. 3
Fig. 4

Similar content being viewed by others

References

  • Alados CL, Escos J (1992) The determinants of social status and the effect of female rank on reproductive success in Dama and Cuvier’s gazelles. Ethol Ecol Evol 4:151–164

    Article  Google Scholar 

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

    Article  CAS  PubMed  Google Scholar 

  • Anderson MJ (2001) A new method for non‐parametric multivariate analysis of variance. Aust Ecol 26:32–46

    Google Scholar 

  • Andersson M, Simmons LW (2006) Sexual selection and mate choice. Trends Ecol Evol 21:296–302

    Article  PubMed  Google Scholar 

  • Barroso FG, Alados CL, Boza J (2000) Social hierarchy in the domestic goat: effect on food habits and production. Appl Anim Behav Sci 69:35–53

    Article  PubMed  Google Scholar 

  • Bartoš L, Fričová B, Bartošová Víchová J, Panama J, Šustr P, Šmídová E (2007) Estimation of the probability of fighting in fallow deer (Dama dama) during the rut. Aggress Behav 33:7–13

    Article  PubMed  Google Scholar 

  • Bergeron P, Grignolio S, Apollonio M, Shipley B, Festa-Bianchet M (2010) Secondary sexual characters signal fighting ability and determine social rank in Alpine ibex (Capra ibex). Behav Ecol Sociobiol 64:1299–1307

    Article  Google Scholar 

  • Bergman TJ, Ho L, Beehner JC (2009) Chest color and social status in male geladas (Theropithecus gelada). Int J Primatol 30:791–806

    Article  Google Scholar 

  • Boesi R (2004) Selezione intrasessuale per colorazione corporea in un grande mammifero: il tahr dell’Himalaya (Bovidae: Caprinae). Unpublished Ph.D. Thesis, University of Siena

  • Brower B (1991) Sherpa of Khumbu: people, livestock and landscape. Oxford University Press, New Dheli

    Google Scholar 

  • Buffa G, Ferrari C, Lovari S (1998) The upper subalpine vegetation of Sagarmatha National Park (Khumbu Himal area, Nepal) and its relationship with Himalayan tahr, musk deer and domestic yak. An outline. In: Baudo R, Tartari G, Munawar M (eds) Top of the world environmental research: Mount Everest Himalayan ecosystem. Backhuys Publisher, Leiden, The Netherlands, pp 167–175

    Google Scholar 

  • Caro TIM (2005) The adaptive significance of coloration in mammals. Bio Science 55:125–136

    Google Scholar 

  • Caro TIM (2013) The colours of extant mammals. Semin Cell Dev Biol 24:542–552

    Article  PubMed  Google Scholar 

  • Clutton-Brock TH (1982) The functions of antlers. Behaviour 79:108–125

    Article  Google Scholar 

  • Clutton-Brock TH, Albon SD (1979) The roaring of red deer and the evolution of honest advertisement. Behaviour 69:145–170

    Article  Google Scholar 

  • Clutton-Brock TH, Guinness FE (1982) Red deer: behavior and ecology of two sexes. University of Chicago Press, Chicago and London

    Google Scholar 

  • Clutton-Brock TH, Albon SD, Gibson RM, Guinness FE (1979) The logical stag: adaptive aspects of fighting in red deer Cervus elaphus. Anim Behav 27:211–225

    Article  Google Scholar 

  • Coltman DW, Festa-Bianchet M, Jorgenson JT, Strobeck C (2002) Age-dependent sexual selection in bighorn rams. Proc R Soc Lon B Biol 269:165–172

    Article  CAS  Google Scholar 

  • Corlatti L, Béthaz S, von Hardenberg A, Bassano B, Palme R, Lovari S (2012) Hormones, parasites and male mating tactics in Alpine chamois: identifying the mechanisms of life history trade-offs. Anim Behav 84:1061–1070

    Article  Google Scholar 

  • Corlatti L, Bassano B, Polakova R, Fattorini L, Pagliarella MC, Lovari S (2015) Preliminary analysis of reproductive success in a large mammal with alternative mating tactics, the Northern chamois, Rupicapra rupicapra. Biol J Linn Soc. doi:10.1111/bij.12569

  • Dematteis A, Menzano A, Tizzani P, Karmacharya B, Meneguz PG, Lovari S (2006) Immobilization of Himalayan tahr with a xylazine-ketamine mixture and reversal with atipamezole under field conditions. J Wildl Dis 42:633–639

    Article  CAS  PubMed  Google Scholar 

  • Eccles TR, Shackleton DM (1986) Correlates and consequences of social status in female bighorn sheep. Anim Behav 34:1392–1401

    Article  Google Scholar 

  • Ferretti F, Sforzi A, Lovari S (2011) Behavioural interference between ungulate species: roe are not on velvet with fallow deer. Behav Ecol Sociobiol 65:875–887

    Article  Google Scholar 

  • Forsyth DM (1999) Long‐term harvesting and male migration in a New Zealand population of Himalayan tahr Hemitragus jemlahicus. J Appl Ecol 36:351–362

    Article  Google Scholar 

  • Forsyth DM, Duncan RP, Tustin KG, Gaillard JM (2005) A substantial energetic cost to male reproduction in a sexually dimorphic ungulate. Ecology 86:2154–2163

    Article  Google Scholar 

  • Francisci F, Focardi S, Boitani L (1985) Male and female alpine ibex: phenology of space use and herd size. In: Lovari S (ed) The biology and management of mountain ungulates. Croom Helm, London, pp 124–133

    Google Scholar 

  • Freitas-de-Melo A, Morales-Pineyrua JT, Ungerfeld R (2013) Agonistic male-female and female-female behaviour in pampas deer (Ozotoceros bezoarticus). N West J Zool 9:127–130

    Google Scholar 

  • Frid A (1997) Vigilance by female Dall’s sheep: interactions between predation risk factors. Anim Behav 53:799–808

    Article  Google Scholar 

  • Garel M, Forsyth DM, Loison A, Dubray D, Jullien JM, Tustin KG, Maillard D, Gaillard JM (2011) Age-related male reproductive effort in two mountain ungulates of contrasting sexual size dimorphism. Can J Zool 89:929–937

    Article  Google Scholar 

  • Garratt KJ (1981) Sagarmatha National Park management plan. Department of national parks and wildlife conservation. His Majesty’s Government of Nepal, Kathmandu

    Google Scholar 

  • Gauthreaux SA (1978) The ecological significance of behavioural dominance. In: Bateson PPG, Klopfer PH (eds) Perspectives in ethology, vol 3, Social behavior. Springer US, New York, pp 17–54

    Google Scholar 

  • Geist V (1964) On the rutting behavior of the mountain goat. J Mammal 45:551–568

    Article  Google Scholar 

  • Geist V (1971) Mountain sheep. University of Chicago Press, Chicago and London

    Google Scholar 

  • Geist V (1982) Adaptive behavioural strategies. In: Thomas JW, Toweill DE (eds) Elk of North America. Stackpole Books, Harrisburg, PA, pp 214–277

    Google Scholar 

  • Geist V (1986) New evidence of high frequency of antler wounding in cervids. Can J Zool 64:380–384

    Article  Google Scholar 

  • Gerald MS (2001) Primate colour predicts social status and aggressive outcome. Anim Behav 61:559–566

    Article  Google Scholar 

  • Gosling LM (1986) The evolution of mating strategies in male antelopes. In: Rubestein DI, Wrangham RW (eds) Ecological aspects of social evolution. Princeton University Press, Princeton, pp 244–281

    Google Scholar 

  • Hess R (2002) The ecological niche of Markhor Capra Falconeri between wild goat Capra aegagrus and Asiatic ibex Capra ibex. Unpublished Ph.D. Thesis, University of Zürich

  • Hillman JC (1987) Group size and association patterns of the common eland (Tragelaphus oryx). J Zool 213:641–663

    Article  Google Scholar 

  • Hinde RA (1970) Animal behaviour. McGraw-Hill, New York

    Google Scholar 

  • Hogg JT (1984) Mating in bighorn sheep: multiple creative male strategies. Science 225:526–529

    Article  CAS  PubMed  Google Scholar 

  • Hogg JT, Forbes SH (1997) Mating in bighorn sheep: frequent male reproduction via a high-risk “unconventional” tactic. Behav Ecol Sociobiol 41:33–48

    Article  Google Scholar 

  • Jarman P (1974) The social organisation of antelope in relation to their ecology. Behaviour 48:215–267

    Article  Google Scholar 

  • Locati M, Lovari S (1988) Social behaviour of the Apennine chamois: a comparison between sexes and management suggestions. Atti 1° convegno nazionale dei biologi della selvaggina, Bologna: 561–567

  • Lovari S (1985) Behavioural repertoire of the Abruzzo chamois, Rupicapra pyrenaica ornata Neumann, 1899 (Artiodactyla: Bovidae). Säugetierk Mitt 32:113–116

    Google Scholar 

  • Lovari S (1992) Observations on the Himalayan tahr and other ungulates of the Sagarmatha National Park, Khumbu Himal, Nepal. Oecol Mont 1:51–52

    Google Scholar 

  • Lovari S, Ale SB (2001) Are there multiple mating strategies in blue sheep? Behav Process 53:131–135

    Article  Google Scholar 

  • Lovari S, Ale SB, Boesi R (2005) Notes on the large mammal community of Sagarmatha National Park (Nepal). In: Sham FA (ed) Proceedings of the First International Karakorum Conference. Pakistan Acad. of Geological Sciences & Ev-K2-CNR, PanGraphics (Pvd) Ltd., Islamabad, pp 225–230

  • Lovari S, Pellizzi B, Boesi R, Fusani L (2009) Mating dominance amongst male Himalayan tahr: blonds do better. Behav Process 81:20–25

    Article  CAS  Google Scholar 

  • Machlis L, Dodd PWD, Fentress JC (1985) The pooling fallacy: problems arising when individuals contribute more than one observation to the data set. Z Tierpsychol 68:201–214

    Article  Google Scholar 

  • Mainguy J, Côté SD, Cardinal E, Houle M (2008) Mating tactics and mate choice in relation to age and social rank in male mountain goats. J Mammal 89:626–635

    Article  Google Scholar 

  • Martin P, Bateson P (2007) Measuring behaviour: an introductory guide. Cambridge University Press, Cambridge

    Book  Google Scholar 

  • Martin AM, Presseault-Gauvin H, Festa-Bianchet M, Pelletier F (2013) Male mating competitiveness and age-dependent relationship between testosterone and social rank in bighorn sheep. Behav Ecol Sociobiol 67:919–928

    Article  Google Scholar 

  • McElligott AG, Gammell MP, Harty HC, Paini DR, Murphy DT, Walsh JT, Hayden TJ (2001) Sexual size dimorphism in fallow deer (Dama dama): do larger, heavier males gain greater mating success? Behav Ecol Sociobiol 49:266–272

    Article  Google Scholar 

  • McFarland D (1981) The Oxford companion to animal behaviour. Oxford University Press, Oxford

    Google Scholar 

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

    Google Scholar 

  • Mundry R, Sommer C (2007) Discriminant function analysis with nonindependent data: consequences and an alternative. Anim Behav 74:965–976

    Article  Google Scholar 

  • Parkes JP, Tustin KG (1989) The trophy potential of Himalayan tahr (Hemitragus jemlahicus) in New Zealand. N Z J Ecol 12:121–124

    Google Scholar 

  • Pelletier F, Festa-Bianchet M (2006) Sexual selection and social rank in bighorn rams. Anim Behav 71:649–655

    Article  Google Scholar 

  • Pelletier F, Bauman J, Festa-Bianchet M (2003) Fecal testosterone in bighorn sheep (Ovis canadensis): behavioural and endocrine correlates. Can J Zool 81:1678–1684

    Article  CAS  Google Scholar 

  • Preston BT, Stevenson IR, Pemberton JM, Wilson K (2001) Dominant rams lose out by sperm depletion. Nature 409:681–682

    Article  CAS  PubMed  Google Scholar 

  • Preston BT, Stevenson IR, Pemberton JM, Coltman DW, Wilson K (2003) Overt and covert competition in a promiscuous mammal: the importance of weaponry and testes size to male reproductive success. Proc R Soc Lon B Biol 270:633–640

    Article  CAS  Google Scholar 

  • Rachlow JL, Berkeley EV, Berger J (1998) Correlates of male mating strategies in white rhinos (Ceratotherium simum). J Mammal 79:1317–1324

    Article  Google Scholar 

  • Rencher AC (1995) Methods of multivariate analysis. Wiley, New York

    Google Scholar 

  • Rice CG (1987) Dermal shields of Himalayan tahr (Hemitragus jemlahicus). J Bombay Nat Hist Soc 84:671–673

    Google Scholar 

  • Rowell TE (1974) The concept of social dominance. Behav Biol 11:131–154

    Article  CAS  PubMed  Google Scholar 

  • Sankar K, Johnsingh AJT, Acharya B (2004) Blue bull or Nilgai Boselaphus tragocamelus Pallas 1766. In: Sankar K, Goyal SP (eds) Ungulates of India. Envis Bulletin: Wildlife and Protected Areas, Vol 07, No 1. Wildlife Institute of India, Dehradun, India, pp 29–40

  • Schaller GB (1963) The mountain gorilla: ecology and behavior. University Chicago Press, Chicago and London

    Google Scholar 

  • Schaller GB (1972) The Serengeti lion: a study of predator—prey relations. University of Chicago Press, Chicago and London

  • Schaller GB (1973) Observations on Himalayan tahr (Hemitragus jemlahicus). J Bombay Nat Hist Soc 70:1–24

    Google Scholar 

  • Schaller GB (1977) Mountain monarchs: wild sheep and goats of the Himalaya. University of Chicago Press, Chicago and London

    Google Scholar 

  • Setchell JM, Smith T, Wickings EJ, Knapp LA (2008) Social correlates of testosterone and ornamentation in male mandrills. Horm Behav 54:365–372

    Article  CAS  PubMed  Google Scholar 

  • Syme GJ (1974) Competitive orders as measures of social dominance. Anim Behav 22:931–940

    Article  Google Scholar 

  • Taborsky M, Oliveira RF, Brockmann HJ (2008) The evolution of alternative reproductive tactics: concepts and questions. In: Oliveira RF, Taborsky M, Brockmann HJ (eds) Alternative reproductive tactics: an integrative approach. Cambridge University Press, Cambridge, pp 1–22

    Chapter  Google Scholar 

  • Tear TH, Mosley JC, Ables ED (1997) Landscape-scale foraging decisions by reintroduced Arabian oryx. J Wildl Manag 61:1142–1154

    Article  Google Scholar 

  • Thirgood S, Langbein J, Putman RJ (1999) Intraspecific variation in ungulate mating strategies: the case of the flexible fallow deer. Adv Stud Behav 28:333–361

    Article  Google Scholar 

  • Trivers R (1985) Social evolution. The Benjamin/Cummings Publishing Company, Menlo Park

    Google Scholar 

  • Tustin KG, Parkes JP (1988) Daily movement and activity of female and juvenile Himalayan thar (Hemitragus jemlahicus) in the eastern Southern Alps, New Zealand. N Z J Ecol 11:51–59

    Google Scholar 

  • Ungerfeld R, Freitas-de-Melo A (2014) Pampas deer (Ozotoceros bezoarticus) male-female agonistic behavior toward high and low social ranked females. Acta Ethol 17:167–171

    Article  Google Scholar 

  • Valdez R, Cardenas M, Sanchez J (1991) Disruptive mating behavior by subadult Armenian wild sheep in Iran. Appl Anim Behav Sci 29:165–171

    Article  Google Scholar 

  • Walther FR (1984) Communication and expression in hoofed mammals. Indiana University Press, Bloomington

    Google Scholar 

  • West PM, Packer C (2002) Sexual selection, temperature, and the lion’s mane. Science 297:1339–1343

    Article  CAS  PubMed  Google Scholar 

  • West JP, Packer C (2003) Melanin, nutrition, and the lion’s mane - Response. Science 299:660

  • Wilkinson PF, Shank CC (1976) Rutting-fight mortality among musk oxen on Banks Island, Northwest Territories, Canada. Anim Behav 24:756–758

    Article  Google Scholar 

  • Willisch CS, Neuhaus P (2009) Alternative mating tactics and their impact on survival in adult male Alpine ibex (Capra ibex ibex). J Mammal 90:1421–1430

    Article  Google Scholar 

  • Zahavi A (1975) Mate selection—a selection for a handicap. J Theor Biol 53:205–214

    Article  CAS  PubMed  Google Scholar 

  • Zahavi A (1987) The theory of signal selection and some of its implications. In: Delfino VP (ed) International symposium of biological evolution. Adriatica Editrice, Bari, pp 305–327

    Google Scholar 

Download references

Acknowledgments

We are grateful to NAST and to DNPWC (especially to its late Director General, Narayan Paudel) for backing our project and for granting permission to work in Sagarmatha National Park. Special thanks are due to L. Fattorini for guidance in the statistical treatment of our data, as well as to A. Dematteis and P. Tizzani for kindly providing horn measurements of tahr. We are also indebted to M. Festa-Bianchet, D. Jennings, L. Corlatti, F. Ferretti and three anonymous reviewers for greatly improving previous drafts of this paper; to M. Apollonio, A. Falchetti and S. Severgnini for help during field observations; to L. Sonam Sherpa, his family and his staff for lodging and friendship. Without the backing of A. Da Polenza, Hari K. Shresta and the personnel of Ev-K2-CNR, this work could not have been carried out. Our work was financially supported by Ev-K2-CNR and by the Italian Ministry of Foreign Affairs. S. L. had the idea of this research, collected field data (1991–1992), supervised part of data analyses and participated in writing up all drafts of this paper; N. F. analysed data, conducted statistical tests and participated in writing up all drafts of this paper; R. B. collected field data (2002–2003) and participated in data analyses; A. B. participated in data analyses and in writing up parts of this paper.

Author information

Author notes

    Authors and Affiliations

    1. Department of Life Sciences, Research Unit of Ecology, Ethology and Wildlife Management, University of Siena, via P.A. Mattioli 4, 53100, Siena, Italy

      S. Lovari, N. Fattorini, R. Boesi & A. Bocci

    2. Ev-K2-CNR Agency, Bergamo, Italy

      S. Lovari

    Authors
    1. S. Lovari
      View author publications

      You can also search for this author in PubMed Google Scholar

    2. N. Fattorini
      View author publications

      You can also search for this author in PubMed Google Scholar

    3. R. Boesi
      View author publications

      You can also search for this author in PubMed Google Scholar

    4. A. Bocci
      View author publications

      You can also search for this author in PubMed Google Scholar

    Corresponding author

    Correspondence to S. Lovari.

    Additional information

    Communicated by: Sven Thatje

    S. Lovari and N. Fattorini contributed equally to this work.

    Rights and permissions

    Reprints and permissions

    About this article

    Check for updates. Verify currency and authenticity via CrossMark

    Cite this article

    Lovari, S., Fattorini, N., Boesi, R. et al. Male ruff colour as a rank signal in a monomorphic-horned mammal: behavioural correlates. Sci Nat 102, 39 (2015). https://doi.org/10.1007/s00114-015-1290-7

    Download citation

    • Received:

    • Revised:

    • Accepted:

    • Published:

    • DOI: https://doi.org/10.1007/s00114-015-1290-7

    Keywords

    Search

    Navigation