Behavioral Ecology and Sociobiology

, Volume 57, Issue 1, pp 69–76 | Cite as

Reproductive effort in relation to maternal social rank in reindeer (Rangifer tarandus)

  • Øystein HolandEmail author
  • Robert B. Weladji
  • Hallvard Gjøstein
  • Jouko Kumpula
  • Martin E. Smith
  • Mauri Nieminen
  • Knut H. Røed
Original Article


In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves’ preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves’ body-mass change during their first winter). In the models, we included September females’ body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females’ rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females’ social rank, whereas late preweaning daily growth rate of calves was not influenced by females’ rank. Calves’ preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females’ rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females’ rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.


Social rank Maternal effort Rangifer tarandus 



We wish to thank V. Tervonen of the Finnish Reindeer Herders’ Association and his crew at Kutuharju Reindeer Research Station for valuable assistance. Heikki Törmänen at the Reindeer Research Station in Kaamanen and K.C. Hofman are also thanked for logistic support and valuable assistance in the field. The Norwegian Research Council, the Finnish Game and Fisheries Research Institute and the Norwegian Reindeer Husbandry Research Council supported the work.


  1. Adams LG, Dale BW (1998) Reproductive performance of female Alaskan caribou. J Wildl Manage 62:1184–1195Google Scholar
  2. Akaike H (1973) Information theory as an extension of the maximum likelihood principle. In: Petrov BN, Csaki F (eds) Second international symposium on information theory. Akademiai Kiado, Budapest, pp 267–281Google Scholar
  3. Barrette C, Vandal D (1986) Social rank, dominance, antler size and access to food in snow-bound wild woodland caribou. Behaviour 97:118–146Google Scholar
  4. Birgersson B, Ekvall K (1997) Early growth in male and female fallow deer fawns. Behav Ecol 8:493–499Google Scholar
  5. Boyce MS (1989) The Jackson herd: intensive wildlife management in North America. Cambridge University Press, CambridgeGoogle Scholar
  6. Brouns F, Edwards SA (1994) Social rank and feeding behaviour of group-housed sows fed competitively or ad libitum. Appl Anim Behav Sci 39:225–235CrossRefGoogle Scholar
  7. Bunnell FL (1982) The lambing period of mountain sheep: synthesis, hypothesis, and tests. Can J Zool 60:1–14Google Scholar
  8. Burnham KP, Anderson DR (1998) Model selection and inference: a practical information-theoretic approach. Springer, Berlin Heidelberg New YorkGoogle Scholar
  9. Byers JA (1997) American pronghorn. Social adaptations and the ghosts of predator past. University of Chicago Press, ChicagoGoogle Scholar
  10. Cameron RD, Smith WT, Fancy SG, Gerhart KL, White RG (1993) Calving success of female caribou in relation to body weight. Can J Zool 71:480–486Google Scholar
  11. Clutton-Brock TH, Coulson T (2002) Comparative ungulate dynamics: the devil is in the detail. Philos Trans R Soc Lond B 357:1285–1298CrossRefGoogle Scholar
  12. Clutton-Brock TH, Albon SD, Guinness FE (1981) Parental investment in male and female offspring in polygynous mammals. Nature 289:487–489Google Scholar
  13. Clutton-Brock TH, Albon SD, Guinness FE (1986) Great expectations: dominance, breeding success and offspring sex ratio in red deer. Anim Behav 27:211–225Google Scholar
  14. Clutton-Brock TH, Albon SD, Guinness FE (1987) Interactions between population density and maternal characteristics affecting fecundity and juvenile survival in red deer. J Anim Ecol 56:857–871Google Scholar
  15. Côté SD (2000a) Dominance hierarchies in female mountain goats: stability aggressiveness and determinants of rank. Behaviour 137:1541–1566CrossRefGoogle Scholar
  16. Côté SD (2000b) Determining social rank in ungulates: a comparison of aggressive interactions recorded at a bait site and under natural conditions. Ethology 106:945–955Google Scholar
  17. Côté SD, Festa-Bianchet M (2001) Birthdate, mass and survival in mountain goat kids: effects of maternal characteristics and forage quality. Oecologia 127:230–238CrossRefGoogle Scholar
  18. Eccles TR, Shackleton DM (1986) Correlates and consequences of social status in female bighorn sheep. Anim Behav 34:1392–1401Google Scholar
  19. Eloranta E, Nieminen M (1986) Calving of the experimental reindeer herd in Kaamanen during 1970–1986. Rangifer Spec Issue 1:115–121Google Scholar
  20. Espmark Y (1964) Studies in dominance-subordination relationship in a group of semi-domesticated reindeer (Rangifer tarandus L.). Anim Behav 12:420–426Google Scholar
  21. Espmark Y (1980) Effect of maternal pre-partum undernutrition on early mother-calf relationships in reindeer. Proc 2nd Int Reindeer/Caribou Symp. Røros, NorwayGoogle Scholar
  22. Fairbanks WS (1993) Birth rate, birth weight, and survival in pronghorn fawns. J Mammal 74:129–135Google Scholar
  23. Fancy SG, White RG (1985) Energy expenditures by caribou while cratering in snow. J Wildl Manage 49:987–993Google Scholar
  24. Festa-Bianchet M (1988) Birthdate and survival in bighorn lambs (Ovis canadensis). J Zool 214:653–661Google Scholar
  25. Festa-Bianchet M (1991) The social system of bighorn sheep; grouping patterns, kinship and female dominace rank. Anim Behav 42:71–82Google Scholar
  26. Festa-Bianchet M, Jorgenson JT (1998) Selfish mothers: reproductive expenditure and resource availability in bighorn ewes. Behav Ecol 9:144–150Google Scholar
  27. Festa-Bianchet M, Jorgenson JT, Bérubé CH, Portier C, Wishart WD (1997) Body mass and survival in bighorn sheep. Can J Zool 75:1372–1379Google Scholar
  28. Flydal K, Reimers E (2002) Relationship between calving time and physical condition in three wild reindeer Rangifer tarandus populations in southern Norway. Wildl Biol 8:145–151Google Scholar
  29. Fournier F, Festa-Bianchet M (1995) Social dominance in adult female mountain goats. Anim Behav 49:1449–1459CrossRefGoogle Scholar
  30. Gaillard J-M, Sempéré AJ, Boutin JM, Van Laere G, Boisaubert B (1992) Effects of age and body weight on the proportion of females breeding in a population of roe deer (Capreolus capreolus). Can J Zool 70:1541–1545Google Scholar
  31. Gaillard J-M, Festa-Bianchet M, Yoccoz NG, Loison A, Toigo C (2000) Temporal variation in fitness components and population dynamics of large herbivores. Annu Rev Ecol Syst 31:367–393CrossRefGoogle Scholar
  32. Geist V (1999) Deer of the world. Their evolution, behaviour and ecology. San-Hill, LondonGoogle Scholar
  33. Guilhem C, Gerard JF, Bideau E (2002) Rank acquisition through birth order in Mouflon sheep (Ovis gmelini) ewes. Ethology 108:63–73CrossRefGoogle Scholar
  34. Guinness FE, Clutton-Brock TH, Albon SD (1978) Factors affecting calf mortality in red deer. J Anim Ecol 47:812–832Google Scholar
  35. Hall MJ (1983) Social organization in an enclosed group of red deer (Cervus elaphus L.) on Rhum. Z Tierpsychol 61:250–262Google Scholar
  36. Hamilton WJ, Blaxter KL (1980) Reproduction in farmed red deer. 1. Hind and stag fertility. J Agric Sci 95:261–273Google Scholar
  37. Hewison AJM, Gaillard J-M (1999) Successful sons or advantaged daughters? The Trivers-Willard model on sex-biased maternal investment in ungulates. TREE 14:229–234CrossRefPubMedGoogle Scholar
  38. Hirotani A (1990) Social organization of reindeer (Rangifer tarandus), with special reference to relationships among females. Can J Zool 68:743–749Google Scholar
  39. Holand Ø, Røed KH, Mysterud A, Kumpula J, Nieminen M, Smith ME (2003) The effect of sex ratio and male age structure on reindeer calving. J Wildl Manage 67:25–33Google Scholar
  40. Holand Ø, Gjøstein H, Losvar A, KumpulaJ, Smith ME, Røed KH, Nieminen M, Weladji RB (2004) Social rank in female reindeer (Rangifer tarandus)—effects of body mass, antler size and age. J Zool Lond (in press)Google Scholar
  41. Jönsson KI (1997) Capital and income breeding as alternative tactics of resource use in reproduction. Oikos 78:57–66Google Scholar
  42. Keech MA, Bowyer RT, Ver Hoef JM, Boertje RD, Dale BW, Stephenson TR (2000) Life-history consequences of maternal condition in Alaska moose. J Wildl Manage 64:175–180Google Scholar
  43. Klein DR (1996) Arctic ungulates at the northern edge of terrestrial life. Rangifer 16:51–56Google Scholar
  44. Kojola I (1989) Mother’s dominance status and differential investment in reindeer calves. Anim Behav 38:177–185Google Scholar
  45. Kojola I (1997) Behavioural correlates of female social status and birth mass of male and female calves in reindeer. Ethology 103:809–814Google Scholar
  46. Kruuk LEB, Clutton-Brock TH, Rose KE, Guinness FE (1999) Early determinants of lifetime reproducive success differ between the sexes in red deer. Proc R Soc Lond B Biol Sci 266:1655–1661CrossRefPubMedGoogle Scholar
  47. Kumpula J (2001) Productivity of the semi-domesticated reindeer (Rangifer T. Tarandus L.) stock and carrying capacity of pastures in Finland during the 1960–1990’s. PhD Thesis, University of OuluGoogle Scholar
  48. Lenvik D, Bø E, Fjellheim A (1988) Utvalgsstrategi i reinflokken. III. Reinkalvens høstvekt relatert til mødrenes vekt og alder. Nor Landbruksforsk 2:65–69Google Scholar
  49. Loison A, Langvatn R (1998) Short and long-term effects of winter and spring weather on growth and survival of red deer in Norway. Oecologia 116:489–500CrossRefGoogle Scholar
  50. Lovari S, Rosto G (1985) Feeding rate and social stress of female chamois foraging in groups. In: Lovari S (ed) The biology and management of mountain ungulates. Croom Helm, London, pp 102–105Google Scholar
  51. Masteller MA, Bailey JA (1988) Agonistic behaviour among mountain goats foraging in winter. Can J Zool 66:2585–2588Google Scholar
  52. Mysterud A, Holand Ø, Røed KH, Gjøstein H, Kumpula J, Nieminen M (2003) Effects of age, density and sex ratio on reproductive effort in male reindeer. J Zool 261:341–344CrossRefGoogle Scholar
  53. Noldus Information Technology (1998) Matman, reference manual, version 1.0 Windows. WageningenGoogle Scholar
  54. Oftedal OT (1985) Pregnancy and lacation. In: Hudson RJ, White RG (eds) Bioenergetics of wild herbivores. CRC, Boca Raton, pp 215–238Google Scholar
  55. Rachlow JL, Bowyer RT (1991) Interannual variation in timing and synchrony of parturition in Dall’s sheep. J Mammal 72:487–492Google Scholar
  56. Rutberg AT (1986) Dominance and fitness consequences in American bison cows. Behaviour 96:62–91Google Scholar
  57. Røed KH, Holand Ø, Smith ME, Gjøstein H, Kumpula J, Nieminen M (2002) Reproductive success in reindeer males in herds with varying male composition. Mol Ecol 11:1239–1243CrossRefPubMedGoogle Scholar
  58. Sadleir PMFS (1984) Ecological consequences of lactation. Acta Zool Fenn 171:179–182Google Scholar
  59. SAS (1999) SAS/STAT guide for personal computers. Version 8. SAS Institute, Cary, NCGoogle Scholar
  60. Skogland T (1984) The effect of food and maternal conditions on fetal growth and size in wild reindeer. Rangifer 4:39–46Google Scholar
  61. Skogland T (1989) Comparative social organisation of wild reindeer in relation to food, mates and predator avoidance. Adv Ethol 29:1–74Google Scholar
  62. Thouless CR (1990) Feeding competition between grazing red deer hinds. Anim Behav 40:105–111Google Scholar
  63. Trivers RL, Willard DE (1973) Natural selection of parental ability to vary the sex ratio in offspring. Science 179:90–92PubMedGoogle Scholar
  64. Unsworth JW, Pac DF, White GC, Bartmann RM (1999) Mule deer survival in Colorado, Idaho, and Montana. J Wildl Manage 63:315–326Google Scholar
  65. Veiberg V, Loe LE, Mysterud A, Langvatn R, Stenseth NC (2004) Social rank, feeding time and winter weight loss in red deer: any evidence of interference competition? Oecologia 138:135–142CrossRefPubMedGoogle Scholar
  66. Vries H de (1995) An improved test of linearity in dominance hierarchies containing unknown or tied relationships. Anim Behav 50:1375–1389CrossRefGoogle Scholar
  67. Vries H de (1998) Finding a dominance order most consistent with a linear hierarchy: a new procedure and review. Anim Behav 55:827–843CrossRefPubMedGoogle Scholar
  68. Weladji RB, Holand Ø (2003) Global climate change and reindeer: effects of winter weather on the autumn weight and growth of calves. Oecologia 136:317–323CrossRefPubMedGoogle Scholar
  69. Weladji RB, Holand Ø, Steinheim G, Lenvik D (2003) Sex-specific preweaning maternal care in reindeer (Rangifer tarandus t.). Behav Ecol Sociol 53:308–311Google Scholar
  70. White RG (1992) Nutrition in relation to season, lactation and growth of northern temperate deer. In: Brown RD (ed) The biology of deer. Springer, Berlin Heidelberg New York, pp 407–417Google Scholar
  71. Wierenga HK (1990) Social dominance in dairy cattle and the influences of housing and management. Appl Anim Behav Sci 27:201–229CrossRefGoogle Scholar
  72. Wolf JB, Brodie ED, Cheverud JM, Moore AJ, Wade MJ (1998) Evolutionary consequences of indirect genetic effects. TREE 13:64–69CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2004

Authors and Affiliations

  • Øystein Holand
    • 1
    Email author
  • Robert B. Weladji
    • 1
  • Hallvard Gjøstein
    • 1
  • Jouko Kumpula
    • 2
  • Martin E. Smith
    • 1
  • Mauri Nieminen
    • 2
  • Knut H. Røed
    • 3
  1. 1.Department of Animal SciencesAgricultural University of NorwayÅsNorway
  2. 2.Finnish Game and Fisheries Research InstituteReindeer Research StationKaamanenFinland
  3. 3.Department of Morphology, Genetics and Aquatic BiologyNorwegian School of Veterinary MedicineOsloNorway

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