Multilevel social organization and space use in reticulated giraffe (Giraffa camelopardalis)

Kimberly L. VanderWaal, Hui Wang, Brenda McCowan, Hsieh Fushing, Lynne A. Isbell

Research output: Contribution to journalReview articlepeer-review

85 Scopus citations


It is increasingly recognized that association patterns of most gregarious animals are nonrandom. However, nonrandom patterns can emerge in any population that exhibits spatial structure, even if individuals associate randomly. In species that lack clearly differentiated social relationships characteristic of socially complex mammals, space use patterns must be considered alongside association patterns in order to establish whether nonrandom association patterns are determined by underlying social structure or are merely an artifact of spatial structure. In this study, we simultaneously consider space use and association patterns for a wild population of reticulated giraffe. We examined whether the giraffe's flexible fission-fusion association patterns were embedded in higher levels of social organization. We identified multilevel social organization in which individuals were members of social cliques. Cliques were embedded in larger subcommunities, which in turn were embedded in communities. The frequency with which 2 individuals were observed together was positively correlated with the extent to which their home range overlapped, implying an underlying role of shared space use in determining association patterns. However, membership in cliques and subcommunities was relatively unrelated to space use patterns for males. For females, space use played a much larger role in determining multitiered social organization, which is consistent with a matrilineal-based society characterized by female philopatry. Although giraffe social interactions are highly fluid in nature, it is apparent that association patterns in giraffe are not the result of random fission-fusion events but are embedded within a structured social network characterized by multiple levels of organization.

Original languageEnglish (US)
Pages (from-to)17-26
Number of pages10
JournalBehavioral Ecology
Issue number1
StatePublished - Jul 2014

Bibliographical note

Funding Information:
National Science Foundation Graduate Research Fellowship (2009068763 to K.L.V.); Doctoral Dissertation Improvement Grant (IOS-1209338), the University of California Wildlife Heath Center, Phoenix Zoo, Explorer’s Club, Oregon Zoo Future for Wildlife Program, American Society of Mammalogists, Cleveland Metroparks Zoo, Cleveland Zoological Society, Sigma Xi (grant number G20101015154371 to K.L.V.); Animal Behavior Society, Northeastern Wisconsin Zoo and a UC Davis Faculty Research Grant (to L.A.I.).


  • Association patterns
  • Community structure
  • Data cloud geometry
  • Network analysis
  • Social structure
  • Spatial overlap


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