Behavioral Ecology and Sociobiology

, Volume 70, Issue 10, pp 1735–1744 | Cite as

How solitary are white sharks: social interactions or just spatial proximity?

  • R. Findlay
  • E. Gennari
  • M. Cantor
  • D. P. TittensorEmail author
Original Article


White sharks (Carcharodon carcharias) are circumglobally distributed large apex predators. While ecologically important, there is very limited study of their social behaviour. Although evident in other large, apex marine predators (e.g. toothed whales) and smaller elasmobranchs (e.g. blacktip reef sharks), the ability of any large pelagic elasmobranch to demonstrate social preferences, tolerance or grouping behaviour is largely unknown. Here, we test whether white sharks in a near-coastal environment form non-random associations with other conspecifics or simply share the same space at the same time. We photo-identified 323 individuals—74 % juvenile females (175–300 cm)—during chumming events at six different sites in Mossel Bay, South Africa, over a 6-year period (2008–2013), and tested for grouping behaviour. We found evidence for random associations among individuals, though spatio-temporal co-occurrence of white sharks in close proximity was weakly structured according to sex and, potentially, body size. Such biological traits may play a minor part in structuring co-occurrence of individuals at fine spatio-temporal scales, which could reflect ontogenetic preferences in diet and site fidelity, or differing tolerance levels for conspecifics of different sexes and sizes. Our study strengthens the evidence that large pelagic shark species are generally solitary and display limited social behaviour.

Significance statement

Large pelagic shark species are important top predators, but we know little about their social behaviour. We tested the ability of white sharks (C. carcharias) to form groups and display social preferences for other individuals when they congregate at scavenging events in a coastal environment, where social interactions may be more likely. We found that white sharks co-occur at random, displaying no preferred or avoided associations for other individuals. Nevertheless, there was a minor influence of biological traits, with individuals aggregating according to gender and, possibly, body size. While we hypothesise these effects could represent preferences in diet and site fidelity, or more tolerance for similar-sized individuals of the same sex, our study strengthens the evidence that white sharks are mostly solitary foragers.


Carcharodon carcharias Association Social network Group Aggregation Tolerance 



We thank Hal Whitehead for the help and advice on terminology, Lauren Peel for database management, all the volunteers and field specialists at Oceans Research for assistance during fieldwork and the two anonymous reviewers whose thoughtful comments improved our work.


RF was supported by Dalhousie University and Oceans Research; EG by Oceans Research, MC by doctoral scholarships from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq , Brazil) and Killam Trusts (Canada); and DT was supported by UNEP-WCMC.

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed.


  1. Anderson SD, Chapple TK, Jorgensen SJ, Klimley AP, Block BA (2011) Long-term individual identification and site fidelity of white sharks, Carcharodon carcharias, off California using dorsal fins. Mar Biol 158:1233–1237CrossRefPubMedPubMedCentralGoogle Scholar
  2. Araabi BN, Kehtarnavaz N, Mckinney T, Hillman G, Würsig B (2000) A string matching computer-assisted system for dolphin photoidentification. Ann Biomed Eng 28:1269–1279CrossRefPubMedGoogle Scholar
  3. Bejder L, Fletcher D, Bra S (1998) A method for testing association patterns of social animals. Anim Behav 56:719–725CrossRefPubMedGoogle Scholar
  4. Block BA, Jonsen ID, Jorgensen SJ, et al. (2011) Tracking apex marine predator movements in a dynamic ocean. Nature 475:86–90CrossRefPubMedGoogle Scholar
  5. Bonfil R, Meÿer M, Scholl MC, Johnson R, O’Brien S, Oosthuizen H, Swanson S, Kotze D, Paterson M (2005) Transoceanic migration, spatial dynamics, and population linkages of white sharks. Science 310:100–103CrossRefPubMedGoogle Scholar
  6. Bonfil R, Francis MP, Duffy C, Manning MJ, O’Brien S (2009) Large-scale tropical movements and diving behavior of white sharks Carcharodon carcharias tagged off New Zealand. Aquat Biol 8:115–123CrossRefGoogle Scholar
  7. Bruce B (1992) Preliminary observations on the biology of the white shark, Carcharodon carcharias, in South Australian waters. Mar Freshw Res 43:1–11CrossRefGoogle Scholar
  8. Bruce BD, Bradford R (2012) Habitat use and spatial dynamics of juvenile white sharks, Carcharodon carcharias, in eastern Australia. In: Domeier ML (ed) Global perspectives on the biology and life history of the white shark, 1st edn. CRC Press, Boca Raton, pp. 225–254CrossRefGoogle Scholar
  9. Burgess GH, Bruce BD, Cailliet GM, Goldman KJ, Grubbs RD, Lowe CG, MacNeil MA, Mollet HF, Weng KC, O’Sullivan JB (2014) A re-evaluation of the size of the white shark (Carcharodon carcharias) population off California, USA. PLoS One 9:e98078CrossRefPubMedPubMedCentralGoogle Scholar
  10. Cairns SJ, Schwager SJ (1987) A comparison of association indices. Anim Behav 35:1454–1469CrossRefGoogle Scholar
  11. Castles M, Heinsohn H, Marshall HH, Lee AE, Cowlishaw G, Carter AJ (2014) Social networks created with different techniques are not comparable. Anim Behav 96:59–67CrossRefGoogle Scholar
  12. Cliff G, Van Der Elst RP, Govender A, Witthuhn TK, Bullen EM (1996) First estimates of mortality and population size of white sharks on the South African coast. In: Klimley AP, Ainley DG (eds) Great white sharks: the biology of Carcharodon carcharias. Academic Press, London, pp. 393–400CrossRefGoogle Scholar
  13. Clua E, Buray N, Legendre P, Mourier J, Planes S (2010) Behavioural response of sicklefin lemon sharks Negaprion acutidens to underwater feeding for ecotourism purposes. Mar Ecol-Prog Ser 414:257–266CrossRefGoogle Scholar
  14. Compagno LJV (2002) Sharks of the world: an annotated and illustrated catalogue of shark species known to date, Vol. 2. Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes and Orectolobiformes). Food and Agriculture Organization of the United Nations, RomeGoogle Scholar
  15. Croft DP, James R, Krause J (2008) Exploring animal social networks. Princeton University Press, PrincetonCrossRefGoogle Scholar
  16. Croft DP, Madden JR, Franks DW, James R (2011) Hypothesis testing in animal social networks. Trends Ecol Evol 26:502–507CrossRefPubMedGoogle Scholar
  17. Curtis TH, Kelly JT, Menard KL, Laroche RK, Jones RE, Klimley AP (2006) Observations on the behavior of white sharks scavenging from a whale carcass at Point Reyes, California. Calif Fish Game 92:113–124Google Scholar
  18. Delaney DG, Johnson R, Bester MN, Gennari E (2012) Accuracy of using visual identification of white sharks to estimate residency patterns. PLoS One 7:e34753CrossRefPubMedPubMedCentralGoogle Scholar
  19. Dicken ML (2008) First observations of young of the year and juvenile great white sharks (Carcharodon carcharias) scavenging from a whale carcass. Mar Freshw Res 59:596–602CrossRefGoogle Scholar
  20. Domeier M, Nasby-Lucas N (2008) Migration patterns of white sharks Carcharodon carcharias tagged at Guadalupe Island, Mexico, and identification of an eastern Pacific shared offshore foraging area. Mar Ecol-Prog Ser 370:221–237CrossRefGoogle Scholar
  21. Domeier ML, Nasby-Lucas N (2013) Two-year migration of adult female white sharks (Carcharodon carcharias) reveals widely separated nursery areas and conservation concerns. Anim Biotelemetry 1:2CrossRefGoogle Scholar
  22. Dudley SFJ, Anderson-reade MD, Thompson GS, McMullen PB (2000) Concurrent scavenging off a whale carcass by great white sharks, Carcharodon carcharias, and tiger sharks, Galeocerdo cuvier. Fish Bull 649:646–649Google Scholar
  23. Dunbar RIM, Shultz S (2007) Evolution in the social brain. Science 317:1344–1347CrossRefPubMedGoogle Scholar
  24. Estrada JA, Rice AN, Natanson LJ, Skomal GB (2014) Use of isotopic analysis of vertebrae in reconstructing ontogenetic feeding ecology in white sharks. Ecol Soc Am 87:829–834Google Scholar
  25. Fallows C, Gallagher AJ, Hammerschlag N (2013) White sharks (Carcharodon carcharias) scavenging on whales and its potential role in further shaping the ecology of an apex predator. PLoS One 8:1–10CrossRefGoogle Scholar
  26. Farine DR (2014) Measuring phenotypic assortment in animal social networks: weighted associations are more robust than binary edges. Anim Behav 89:141–153CrossRefGoogle Scholar
  27. Farine DR (2015) Proximity as a proxy for interactions: issues of scale in social network analysis. Anim Behav 104:1–5CrossRefGoogle Scholar
  28. Ford B, Ellis M, Balcomb C (2000) Killer whales: the natural history and genealogy of Orcinus orca in British Columbia and Washington. University of Washington Press, SeattleGoogle Scholar
  29. Ford E (1921) A contribution to our knowledge of the life-histories of the dogfishes landed at Plymouth. J Mar Biol Assoc UK 12:468–505CrossRefGoogle Scholar
  30. Goldman KJ, Anderson SD (1999) Space utilization and swimming depth of white sharks, Carcharodon carcharias, at the South Farallon Islands, Central California. Environ Biol Fish 56:351–364CrossRefGoogle Scholar
  31. Gubili C, Johnson R, Gennari E, Oosthuizen WH, Kotze D, Meÿer M, Sims DW, Jones CS, Noble LR (2009) Concordance of genetic and fin photo identification in the great white shark, Carcharodon carcharias, off Mossel Bay, South Africa. Mar Biol 156:2199–2207CrossRefGoogle Scholar
  32. Guttridge TL, Gruber SH, DiBattista JD, Feldheim KA, Croft DP, Krause S, Krause J (2011) Assortative interactions and leadership in a free-ranging population of juvenile lemon shark, Negaprion brevirostris. Mar Ecol-Prog Ser 423:235–245CrossRefGoogle Scholar
  33. Guttridge TL, Gruber SH, Gledhill KS, Croft DP, Sims DW, Krause J (2009) Social preferences of juvenile lemon sharks, Negaprion brevirostris. Anim Behav 78:543–548CrossRefGoogle Scholar
  34. Hamady LL (2014) Age, movement, and feeding ecology of Northwest Atlantic white sharks estimated from ecogeochemical profiles in vertebrae. Dissertation, Massachusetts Institute of Technology and Woods Hole Oceanographic InstitutionGoogle Scholar
  35. Heithaus MR, Frid A, Wirsing AJ, Worm B (2008) Predicting ecological consequences of marine top predator declines. Trends Ecol Evol 23:202–210CrossRefPubMedGoogle Scholar
  36. Hewitt AM (2014) Demographics of a seasonal aggregation of white sharks at Seal Island, False Bay, South Africa. Dissertation, University of Cape TownGoogle Scholar
  37. Hight BV, Lowe CG (2007) Elevated body temperatures of adult female leopard sharks, Triakis semifasciata, while aggregating in shallow nearshore embayments: evidence for behavioral thermoregulation? J Exp Mar Biol Ecol 352:114–128CrossRefGoogle Scholar
  38. Hinde R (1976) Interactions, relationships and social structure. Man 11:1–17CrossRefGoogle Scholar
  39. Jacoby DMP, Busawon DS, Sims DW (2010) Sex and social networking: the influence of male presence on social structure of female shark groups. Behav Ecol 21:808–818CrossRefGoogle Scholar
  40. Jacoby DMP, Croft DP, Sims DW (2012) Social behaviour in sharks and rays: analysis, patterns and implications for conservation. Fish Fish 13:399–417CrossRefGoogle Scholar
  41. Johnson R, Bester MN, Dudley SF, Oosthuizen WH, Meÿer M, Hancke L, Gennari E (2009) Coastal swimming patterns of white sharks (Carcharodon carcharias) at Mossel Bay, South Africa. Environ Biol Fish 85:189–200CrossRefGoogle Scholar
  42. Johnson R, Kock A (2006) South Africa’s white shark cage-diving industry—is there cause for concern? In: Nel DC, Peschak TP (eds). Finding a balance: white shark conservation and recreational safety in the inshore waters of Cape Town, South Africa; proceedings of a specialist workshop. WWF South Africa Report Series, South Africa, pp 40–59Google Scholar
  43. Jorgensen SJ, Reeb CA, Chapple TK, Anderson S, Perle C, Van Sommeran SR, Fritz-Cope C, Brown AC, Klimley AP, Block BA (2010) Philopatry and migration of Pacific white sharks. Proc R Soc Lond B 277:679–688CrossRefGoogle Scholar
  44. Kime DE, Hews EA (1982) The effect of temperature on steroid biosynthesis by testes of the dogfish, Scyliorhinus caniculus. Comparative biochemistry and physiology. Biochem Mol Biol 4:675–679Google Scholar
  45. Kirkman SP, Oosthuizen WH, Meÿer MA, Kotze PG, Roux JP, Underhill LG (2007) Making sense of censuses and dealing with missing data: trends in pup counts of Cape fur seal, Arctocephalus pusillus pusillus, for the period 1972–2004. Afr J Mar Sci 29:161–176CrossRefGoogle Scholar
  46. Klimley AP, Nelson DR (1984) Diel movement patterns of the scalloped hammerhead shark (Sphyrna lewini) in relation to el Bajo Espiritu Santo: a refuging central-position social system. Behav Ecol Sociobiol 15:45–54CrossRefGoogle Scholar
  47. Kock A, O’Riain MJ, Mauff K, Meÿer M, Kotze D, Griffiths C (2013) Residency, habitat use and sexual segregation of white sharks, Carcharodon carcharias in False Bay, South Africa. PLoS One 8:e55048CrossRefPubMedPubMedCentralGoogle Scholar
  48. Krause J, Ruxton G (2002) Living in groups. Oxford University Press, OxfordGoogle Scholar
  49. Laroche RK, Kock AA, Dill LM, Oosthuizen WH (2007) Effects of provisioning ecotourism activity on the behaviour of white sharks, Carcharodon carcharias. Mar Ecol-Prog Ser 338:199–209CrossRefGoogle Scholar
  50. Lee KA, Huveneers C, Gimenez O, Peddemors V, Harcourt RG (2014) To catch or to sight? A comparison of demographic parameter estimates obtained from mark-recapture and mark-resight models. Biodivers Conserv 23:2781–2800Google Scholar
  51. Lessa R, Santana FM, Paglerani R (1999) Age, growth and stock structure of the oceanic whitetip shark, Carcharhinus longimanus, from the southwestern equatorial Atlantic. Fish Res 42:21–30CrossRefGoogle Scholar
  52. Mourier J, Vercelloni J, Planes S (2012) Evidence of social communities in a spatially structured network of a free-ranging shark species. Anim Behav 83:389–401CrossRefGoogle Scholar
  53. Mull CG, Yopak KE, Dulvy NK (2011) Does more maternal investment mean a larger brain? Evolutionary relationships between reproductive mode and brain size in chondrichthyans. Mar Freshw Res 62:567–575CrossRefGoogle Scholar
  54. Northcutt G (1977) Elasmobranch central nervous system organization and its possible evolutionary significance. Am Zool 429:411–429CrossRefGoogle Scholar
  55. Orams MB (2002) Feeding wildlife as a tourism attraction: a review of issues and impacts. Tour Manag 23:281–293CrossRefGoogle Scholar
  56. Powter D, Gladstone W (2009) Habitat-mediated use of space by juvenile and mating adult Port Jackson sharks, Heterodontus portusjacksoni, in Eastern Australia. Pac Sci 63:1–14CrossRefGoogle Scholar
  57. Psorakis I, Voelkl B, Garroway CJ, et al. (2015) Inferring social structure from temporal data. Behav Ecol Sociobiol 5:857–866CrossRefGoogle Scholar
  58. Robbins RL (2007) Environmental variables affecting the sexual segregation of great white sharks (Carcharodon carcharias) at the Neptune Islands, South Australia. J Fish Biol 70:1350–1364CrossRefGoogle Scholar
  59. Ryklief R, Pistorius P, Johnson R (2014) Spatial and seasonal patterns in sighting rate and life-history composition of the white shark (Carcharodon carcharias) at Mossel Bay, South Africa. Afr J Mar Sci 36:449–453CrossRefGoogle Scholar
  60. Simpfendorfer CA, Milward NE (1993) Utilisation of a tropical bay as a nursery area by sharks of the families Carcharhinidae and Sphyrnidae. Environ Biol Fish 37:337–345CrossRefGoogle Scholar
  61. Sims D (2003) Tractable models for testing theories about natural strategies : foraging behaviour and habitat selection of free-ranging sharks. J Fish Biol 63:53–73CrossRefGoogle Scholar
  62. Sims D, Nash J, Morritt D (2001) Movements and activity of male and female dogfish in a tidal sea lough: alternative behavioural strategies and apparent sexual segregation. Mar Biol 139:1165–1175CrossRefGoogle Scholar
  63. Sims DW, Southall EJ, Wearmouth VJ, Hutchinson N, Budd GC, Morritt D (2005) Refuging behaviour in the nursehound Scyliorhinus stellaris (Chondrichthyes: Elasmobranchii): preliminary evidence from acoustic telemetry. J Mar Biol Assoc UK 85:2–5CrossRefGoogle Scholar
  64. Sperone E, Micarelli P, Andreotti S, Spinetti S, Andreani A, Serena F, Brunelli E, Tripepi S (2010) Social interactions among bait-attracted white sharks at Dyer Island (South Africa). Mar Biol Res 6:408–414CrossRefGoogle Scholar
  65. Stehfest KM, Patterson TA, Dagorn L, Holland KN, Itano D, Semmens JM (2013) Network analysis of acoustic tracking data reveals the structure and stability of fish aggregations in the ocean. Anim Behav 85:839–848CrossRefGoogle Scholar
  66. Towner AV, Wcisel MA, Reisinger RR, Edwards D, Jewell OJD (2013) Gauging the threat: the first population estimate for white sharks in South Africa using photo identification and automated software. PLoS One 8:e66035CrossRefPubMedPubMedCentralGoogle Scholar
  67. Wearmouth V, Sims D (2008) Sexual segregation in marine fish, reptiles, birds and mammals: behaviour patterns, mechanisms and conservation implications. Adv Mar Biol 54:107–170CrossRefPubMedGoogle Scholar
  68. Weng KC, Boustany AM, Pyle P, Anderson SD, Brown A, Block BA (2007) Migration and habitat of white sharks (Carcharodon carcharias) in the eastern Pacific Ocean. Mar Biol 152:877–894CrossRefGoogle Scholar
  69. Whitehead H (1999) Testing association patterns of social animals. Anim Behav 57:26–29CrossRefGoogle Scholar
  70. Whitehead H (2003) Sperm whale societies: social evolution in the ocean. University of Chicago Press, ChicagoGoogle Scholar
  71. Whitehead H (2008a) Analyzing animal societies: quantitative methods for vertebrate social analysis. University of Chicago Press, ChicagoCrossRefGoogle Scholar
  72. Whitehead H (2008b) Precision and power in the analysis of social structure using associations. Anim Behav 75:1093–1099CrossRefGoogle Scholar
  73. Whitehead H (2009) SOCPROG programs: analysing animal social structures. Behav Ecol Sociobiol 63:765–78Google Scholar
  74. Whitehead H, Bejder L, Andrea Ottensmeyer C (2005) Testing association patterns: issues arising and extensions. Anim Behav 69:e1CrossRefGoogle Scholar
  75. Whitehead H, Dufault S (1999) Techniques for analyzing vertebrate social structure using identified individuals: review and recommendations. Adv Study Behav 28:33–74CrossRefGoogle Scholar
  76. Yopak KE, Lisney TJ, Collin SP, Montgomery JC (2007) Variation in brain organization and cerebellar foliation in chondrichthyans: sharks and holocephalans. Brain Behav Evol 69:280–300CrossRefPubMedGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2016

Authors and Affiliations

  1. 1.Department of Biology, LSC Room 4089Dalhousie UniversityHalifaxCanada
  2. 2.Oceans ResearchMossel BaySouth Africa
  3. 3.South African Institute for Aquatic BiodiversityGrahamstownSouth Africa
  4. 4.United Nations Environment Programme World Conservation Monitoring CentreCambridgeUK

Personalised recommendations