Wild giraffes live in extensive groups in the fission fusion system, maintaining long social distances and loose social bonds. Within these groups, resources are widely distributed, agonistic encounters are scarce and the dominance hierarchy was reported in males only, while never deeply analysed. In captivity, the possibility to maintain inter-individual distances is limited and part of the resources is not evenly distributed. Consequently, we suggest that agonistic encounters should be more frequent, leading to the establishment of the dominance hierarchy. Based on the differences in resource-holding potential, we suggested that the rank of an individual would be affected by age and sex. Based on hypotheses of prior ownership, we tested whether rank was positively affected by the time spent in a herd and whether it was stable in adult females, which were present long-term in the same herd. We originally monitored four herds of Rothschild giraffes (Giraffa camelopardalis rothschildii) in Dvůr Králové zoo (n = 8), Liberec zoo (n = 6), and two herds in Prague zoo: Prague 1 (n = 8) and Prague 2 (n = 9). The Prague 1 and Prague 2 herds were then combined and the resulting fifth herd was observed over three consecutive years (2009, 2010, and 2011) (n = 14, 13, and 14, respectively). We revealed a significantly linear hierarchy in Dvůr Králové, Prague 2 and in the combined herd in Prague. Rank was significantly affected by age in all herds; older individuals dominated the younger ones. In females, rank was positively affected by the time spent in the herd and adult females in Prague maintained their rank during three consecutive years. This study represents the first analysis of the dominance hierarchy in the captive giraffe, and discusses the behavioural flexibility of the social structure in response to monopolisable resources in a captive environment.

Department of Animal Science and Food Processing in Tropics and Subtropics Czech University of Life Sciences Prague Prague Czech Republic

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Davies NB, Krebs JR, West SA (2012) An Introduction to Behavioural Ecology: Wiley; 506 p.

Dugatkin LA (1997) Winner and loser effects and the structure of dominance hierarchies. Behavioral Ecology 8: 583–587.

Kaufmann JH (1983) On the definitions and functions of dominance and territoriality. Biological Reviews 58: 1–20.

Boyd R, Silk JB (1983) A method for assigning cardinal dominance ranks. Animal Behaviour 31: 45–58.

De Vries H (1998) Finding a dominance order most consistent with a linear hierarchy: a new procedure and review. Animal Behaviour 55: 827–843. PubMed

Bang A, Deshpande S, Sumana A, Gadagkar R (2010) Choosing an appropriate index to construct dominance hierarchies in animal societies: a comparison of three indices. Animal Behaviour 79: 631–636.

Vervaecke H, De Vries H, Van Elsacker L (1999) An experimental evaluation of the consistency of competitive ability and agonistic dominance in different social contexts in captive bonobos. Behaviour 136: 423–442.

Korona R (1991) On the role of age and body size in risky animal contests. Journal of Theoretical Biology 152: 165–176.

Boogert NJ, Reader SM, Laland KN (2006) The relation between social rank, neophobia and individual learning in starlings. Animal Behaviour 72: 1229–1239.

Hemelrijk CK, Wantia J, Gygax L (2005) The construction of dominance order: comparing performance of five methods using an individual-based model. Behaviour 142: 1037–1058.

Pluháček J, Bartoš L, Čulík L (2006) High-ranking mares of captive plains zebra Equus burchelli have greater reproductive success than low-ranking mares. Applied Animal Behaviour Science 99: 315–329.

Bonabeau E, Theraulaz G, Deneubourg J-L (1999) Dominance orders in animal societies: The self-organization hypothesis revisited. Bulletin of Mathematical Biology 61: 727–757. PubMed

Wey T, Blumstein DT, Shen W, Jordán F (2008) Social network analysis of animal behaviour: a promising tool for the study of sociality. Animal Behaviour 75: 333–344.

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

Favre M, Martin JGA, Festa-Bianchet M (2008) Determinants and life-history consequences of social dominance in bighorn ewes. Animal Behaviour 76: 1373–1380.

Taylor PW, Elwood RW (2003) The mismeasure of animal contests. Animal Behaviour 65: 1195–1202.

Parker GA (1974) Assessment strategy and the evolution of fighting behaviour. Journal of Theoretical Biology 47: 223–243. PubMed

Hammerstein P, Parker GA (1982) The asymmetric war of attrition. Journal of Theoretical Biology 96: 647–682.

Kemp DJ, Wiklund C (2004) Residency effects in animal contests. Proceedings of the Royal Society B-Biological Sciences 271: 1707–1711. PubMed PMC

Cote SD (2000) Dominance hierarchies in female mountain goats: Stability, aggressiveness and determinants of rank. Behaviour 137: 1541–1566.

Wrangham RW (1980) An Ecological Model of Female-Bonded Primate Groups. Behaviour 75: 262–300.

Thompson KV (1993) Aggressive-behavior and dominance hierarchies in female sable antelope, Hippotragus niger—implications for captive management. Zoo Biology 12: 189–202.

Barroso FG, Alados CL, Boza J (2000) Social hierarchy in the domestic goat: effect on food habits and production. Applied Animal Behaviour Science 69: 35–53. PubMed

Veiberg V, Loe LE, Mysterud A, Langvatn R, Stenseth NC (2004) Social rank, feeding and winter weight loss in red deer: any evidence of interference competition? Oecologia 138: 135–142. PubMed

Taillon J, Côté SD (2007) Social rank and winter forage quality affect aggressiveness in white-tailed deer fawns. Animal Behaviour 74: 265–275.

Jørgensen GHM, Andersen IL, Bøe KE (2007) Feed intake and social interactions in dairy goats—The effects of feeding space and type of roughage. Applied Animal Behaviour Science 107: 239–251.

Brand R (2007) Evolutionary ecology of giraffes (Giraffa camelopardalis) in Etosha National Park, Namibia.

Dutoit JT (1990) Feeding-height stratification among African browsing ruminants. African Journal of Ecology 28: 55–61.

Jepsen JU, Topping CJ (2004) Modelling roe deer (Capreolus capreolus) in a gradient of forest fragmentation: behavioural plasticity and choice of cover. Canadian Journal of Zoology-Revue Canadienne De Zoologie 82: 1528–1541.

Sol D, Timmermans S, Lefebvre L (2002) Behavioural flexibility and invasion success in birds. Animal Behaviour 63: 495–502.

Boydston EE, Kapheim KM, Watts HE, Szykman M, Holekamp KE (2003) Altered behaviour in spotted hyenas associated with increased human activity. Animal Conservation 6: 207–219.

Estes RD (1991) The Behavior Guide to African Mammals: Including Hoofed Mammals, Carnivores, Primates: University of California Press; 611 p.

Kingdon J (2003) The Kingdon Field Guide to African Mammals: Helm; 476 p.

Coe MJ (1967) Necking behaviour in giraffe. Journal of Zoology 151: 313-&.

Bercovitch FB, Berry PSM (2013) Age proximity influences herd composition in wild giraffe. Journal of Zoology 290: 281–286.

Bercovitch FB, Berry PSM (2013) Herd composition, kinship and fission—fusion social dynamics among wild giraffe. African Journal of Ecology 51: 206–216.

Carter KD, Brand R, Carter JK, Shorrocks B, Goldizen AW (2013) Social networks, long-term associations and age-related sociability of wild giraffes. Animal Behaviour 86: 901–910.

Carter KD, Seddon JM, Frere CH, Carter JK, Goldizen AW (2013) Fission-fusion dynamics in wild giraffes may be driven by kinship, spatial overlap and individual social preferences. Animal Behaviour 85: 385–394.

Dagg AI (1970) Tactile Encounters in a Herd of Captive Giraffes. Journal of Mammalogy 51: 279–287.

van der Jeugd HP, Prins HHT (2000) Movements and group structure of giraffe (Giraffa camelopardalis) in Lake Manyara National Park, Tanzania. Journal of Zoology 251: 15–21.

Le Pendu Y, Ciofolo I, Gosser A (2000) The social organization of giraffes in Niger. African Journal of Ecology 38: 78–85.

Bercovitch FB, Berry PSM (2010) Ecological determinants of herd size in the Thornicroft’s giraffe of Zambia. African Journal of Ecology 48: 962–971.

Simmons RE, Scheepers L (1996) Winning by a neck: Sexual selection in the evolution of giraffe. American Naturalist 148: 771–786.

Mitchell G, Van Sittert SJ, Skinner JD (2009) Sexual selection is not the origin of long necks in giraffes. Journal of Zoology 278: 281–286.

Wilkinson DM, Ruxton GD (2012) Understanding selection for long necks in different taxa. Biological Reviews 87: 616–630. 10.1111/j.1469-185X.2011.00212.x PubMed DOI

Fennessy J, Brenneman R (2010) Giraffa camelopardalis ssp. rothschildi. IUCN 2013. IUCN Red List of Threatened Species. Version 2013.1. http://www.iucnredlist.org/details/full/174469/0: IUCN.

Bashaw MJ, Bloomsmith MA, Maple TL, Bercovitch FB (2007) The structure of social relationships among captive female giraffe (Giraffa camelopardalis). Journal of Comparative Psychology 121: 46–53. PubMed

Langman VA (1977) Cow-calf Relationships in Giraffe (Giraffa camelopardalis giraffa). Zeitschrift für Tierpsychologie 43: 264–286. 10.1107/S0108767309007235 PubMed DOI

Tarou LR, Bashaw MJ, Maple TL (2000) Social attachment in giraffe: Response to social separation. Zoo Biology 19: 41–51. PubMed

Bashaw MJ (2011) Consistency of captive giraffe behavior under two different management regimes. Zoo Biology 30: 371–378. 10.1002/zoo.20338 PubMed DOI

Malyjurková L, Hejzlarová M, Jůnková Vymyslická P, Brandlová K (2014) Social Preferences of Translocated Giraffes (Giraffa Camelopardalis Giraffa) in Senegal: Evidence for Friendship Among Females? Agricultura Tropica et Subtropica 47: 8.

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

Seeber P, Ciofolo I, Ganswindt A (2012) Behavioural inventory of the giraffe (Giraffa camelopardalis). BMC Research Notes 5: 650 10.1186/1756-0500-5-650 PubMed DOI PMC

Bashaw M, Bloomsmith M, Maple T, Bercovitch F (2007) The structure of social relationships among captive female giraffe (Giraffa camelopardalis). J Comp Psychol 121: 46–53. PubMed

Bashaw M (2010) Consistency of captive giraffe behavior under two different management regimes. Zoo Biol 29: 1–8. 10.1002/zoo.20239 PubMed DOI

De Vries H, Appleby MC (2000) Finding an appropriate order for a hierarchy: a comparison of the I&SI and the BBS methods. Animal Behaviour 59: 239–245. PubMed

Appleby MC (1983) The probability of linearity in hierarchies. Animal Behaviour 31: 600–608.

Archie EA, Morrison TA, Foley CAH, Moss CJ, Alberts SC (2006) Dominance rank relationships among wild female African elephants, Loxodonta africana. Animal Behaviour 71: 117–127.

Cafazzo S, Valsecchi P, Bonanni R, Natoli E (2010) Dominance in relation to age, sex, and competitive contexts in a group of free-ranging domestic dogs. Behavioral Ecology 21: 443–455.

Clutton-Brock TH, Albon SD, Guinness FE (1984) Maternal dominance, breeding success and birth sex ratios in red deer. Nature 308: 358–360.

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. Animal Behaviour 66: 601–605.

Clutton-Brock TH, Albon SD, Gibson RM, Guinness FE (1979) Logical stag—adaptive aspects of fighting in red deer (Cervus-elaphus L.). Animal Behaviour 27: 211–225.

Martin P, Bateson P (2007) Measuring Behaviour: An Introductory Guide: Cambridge University Press; 222 p.

Dagg AI, Foster JB (1976) The giraffe: its biology, behavior, and ecology: Van Nostrand Reinhold Co; 10.1016/0376-6357(79)90040-8 DOI

Shorrocks B, Croft DP (2009) Necks and networks: a preliminary study of population structure in the reticulated giraffe (Giraffa camelopardalis reticulata de Winston). African Journal of Ecology 47: 374–381.

Smith J, Memenis S, Holekamp K (2007) Rank-related partner choice in the fission—fusion society of the spotted hyena (Crocuta crocuta). Behavioral Ecology and Sociobiology 61: 753–765.

Dugatkin LA, Druen M (2004) The social implications of winner and loser effects. Proceedings of the Royal Society B-Biological Sciences 271: S488–S489. PubMed PMC

Wirtu G, Pope CE, Vaccaro J, Sarrat E, Cole A, Godke RA, et al. (2004) Dominance hierarchy in a herd of female eland antelope (Taurotragus oryx) in captivity. Zoo Biology 23: 323–333.

Ceacero F, García AJ, Landete-Castillejos T, Bartošová J, Bartoš L, Gallego L, et al. (2012) Benefits for Dominant Red Deer Hinds under a Competitive Feeding System: Food Access Behavior, Diet and Nutrient Selection. Plos One 7: e32780 10.1371/journal.pone.0032780 PubMed DOI PMC

Hass CC (1991) Social status in female bighorn sheep (Ovis canadensis): expression, development and reproductive correlates. Journal of Zoology 225: 509–523.

Alados CL, Escós JM (1992) The determinants of social status and the effect of female rank on reproductive success in Dama and Cuvier's gazelles. Ethology Ecology & Evolution 4: 151–164.

Correa LA, Zapata B, Samaniego H, Soto-Gamboa M (2013) Social structure in a family group of Guanaco (Lama guanicoe, Ungulate): Is female hierarchy based on ‘prior attributes’ or ‘social dynamics’? Behavioural Processes 98: 92–97. 10.1016/j.beproc.2013.05.003 PubMed DOI

Holand O, Gjostein H, Losvar A, Kumpula J, Smith ME, Roed KH, et al. (2004) Social rank in female reindeer (Rangifer tarandus): effects of body mass, antler size and age. Journal of Zoology 263: 365–372.

Roden C, Vervaecke H, Van Elsacker L (2005) Dominance, age and weight in American bison males (Bison bison) during non-rut in semi-natural conditions. Applied Animal Behaviour Science 92: 169–177.

Cransac N, Aulagnier S (1996) Factors influencing hierarchy in a captive herd of eland Taurotragus oryx. Aggressive Behavior 22: 209–213.

Sarova R, Spinka M, Stehulova I, Ceacero F, Simeckova M, Kotrba R, et al. (2013) Pay respect to the elders: age, more than body mass, determines dominance in female beef cattle. Animal Behaviour 86: 1315–1323.

Kingdon J (1988) East African Mammals: An Atlas of Evolution in Africa, Volume 3, Part B: Large Mammals: University of Chicago Press; 450 p.

Leuthold BM, Leuthold W (1978) Daytime activity patterns of gerenuk and giraffe in Tsavo National Park, Kenya. East African Wildlife Journal 16: 12.

Ciofolo I, Le Pendu Y (2002) The feeding behaviour of giraffe in Niger. Mammalia 66: 183–194.

Coe M (1998) Some aspects of the interaction between mammalian herbivores and Acacia eriobola E. Mey. Transactions of the Royal Society of South Africa 53: 141–147.

Foster JB (1966) The giraffe of Nairobi national park: home range, sex ratios, the herd, and food. African Journal of Ecology 4: 139–148.

Hejcmanová P, Homolka M, Antonínová M, Hejcman M, Podhájecká V (2010) Diet composition of western Derby eland (Taurotragus derbianus derbianus) in the dry season in a natural and a managed habitat in Senegal using faecal analyses. South African Journal of Wildlife Research 40: 27–34.

Napper CJ, Sharp SP, McGowan A, Simeoni M, Hatchwell BJ (2013) Dominance, not kinship, determines individual position within the communal roosts of a cooperatively breeding bird. Behavioral Ecology and Sociobiology 67: 2029–2039.

Nepusz T, Vicsek T (2013) Hierarchical Self-Organization of Non-Cooperating Individuals. Plos One 8: e8.1449 PubMed PMC

Further behavioural parameters support reciprocity and milk theft as explanations for giraffe allonursing