The adaptive value of morphological, behavioural and life-history traits in reproductive female wolves

Daniel R. Stahler, Daniel R. Macnulty, Robert K. Wayne, Bridgett Marie vonHoldt, Douglas W. Smith

Research output: Contribution to journalArticlepeer-review

87 Scopus citations


Reproduction in social organisms is shaped by numerous morphological, behavioural and life-history traits such as body size, cooperative breeding and age of reproduction, respectively. Little is known, however, about the relative influence of these different types of traits on reproduction, particularly in the context of environmental conditions that determine their adaptive value. Here, we use 14 years of data from a long-term study of wolves (Canis lupus) in Yellowstone National Park, USA, to evaluate the relative effects of different traits and ecological factors on the reproductive performance (litter size and survival) of breeding females. At the individual level, litter size and survival improved with body mass and declined with age (c. 4-5 years). Grey-coloured females had more surviving pups than black females, which likely contributed to the maintenance of coat colour polymorphism in this system. The effect of pack size on reproductive performance was nonlinear as litter size peaked at eight wolves and then declined, and litter survival increased rapidly up to three wolves, beyond which it increased more gradually. At the population level, litter size and survival decreased with increasing wolf population size and canine distemper outbreaks. The relative influence of these different-level factors on wolf reproductive success followed individual > group > population. Body mass was the primary determinant of litter size, followed by pack size and population size. Body mass was also the main driver of litter survival, followed by pack size and disease. Reproductive gains because of larger body size and cooperative breeding may mitigate reproductive losses because of negative density dependence and disease. These findings highlight the adaptive value of large body size and sociality in promoting individual fitness in stochastic and competitive environments.

Original languageEnglish (US)
Pages (from-to)222-234
Number of pages13
JournalJournal of Animal Ecology
Issue number1
StatePublished - Jan 2013

All Science Journal Classification (ASJC) codes

  • Ecology, Evolution, Behavior and Systematics
  • Animal Science and Zoology


  • Body mass
  • Canine distemper virus
  • Coat colour
  • Cooperative breeding
  • Density dependence
  • Disease
  • Group size
  • Large carnivores
  • Senescence
  • Sociality


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