Social behaviour in sharks and rays: Analysis, patterns and implications for conservation

Fish and Fisheries

Volumn 13, Issue 4, 2012, Pages 399-417

  Jacoby, David M P ^a,b   Croft, Darren P ^b   Sims, David W ^a,c  

^a THE LABORATORY   (United Kingdom)

^b UNIVERSITY OF EXETER   (United Kingdom)

^c UNIVERSITY OF PLYMOUTH   (United Kingdom)

Author keywords

Aggregation; Fisheries impact; Sharks; Social behaviour; Social networks; Social organization

Indexed keywords

ABUNDANCE; AGGREGATION BEHAVIOR; ANTHROPOGENIC EFFECT; HABITAT CONSERVATION; HABITAT USE; LONGEVITY; MORTALITY; PHILOPATRY; SHARK; SOCIAL BEHAVIOR; SOCIAL NETWORK; SOCIAL ORGANIZATION; VULNERABILITY;

CHONDRICHTHYES; ELASMOBRANCHII;

EID: 84869496014     PISSN: 14672960     EISSN: 14672979     Source Type: Journal    
DOI: 10.1111/j.1467-2979.2011.00436.x     Document Type: Article

References (121)

1. Allee, W.C. and Dickinson, J.C.J. (1954) Dominance and subordination in the smooth dogfish mustelus canis (Mitchill). Physiological Zoology27, 356-364.
2. Barber, I. and Wright, H.A. (2001) How strong are familiarity preferences in shoaling fish?Animal Behaviour61, 975-979.
3. Baum, J.K., Myers, R.A., Kehler, D.G., Worm, B., Harley, S.J. and Doherty, P.A. (2003) Collapse and conservation of shark populations in the Northwest Atlantic. Science299, 389-392.
4. Caraco, T. (1979) Time budgeting and group size: a test of theory. Ecology60, 618-627.
5. Carey, F.G., Scharold, J.V. and Kalmijn, A.J. (1990) Movements of blue sharks (Prionace glauca) in depth and course. Marine Biology106, 329-342.
6. Carrier, J.C., Pratt, H.L. Jr and Martin, L.K. (1994) Group reproductive behaviors in free-living nurse sharks, Ginglymostoma cirratum. Copeia1994, 646-656.
7. Chapman, D.D., Babcock, E.A., Gruber, S.H. et al. (2009) Long-term natal site-fidelity by immature lemon sharks (Negaprion brevirostris) at a subtropical island. Molecular Ecology18, 3500-3507.
8. Clarke, T.A. (1971) The ecology of the scalloped hammerhead shark, Sphyrna lewini, in Hawaii. Pacific Science25, 133-144.
9. Connor, R.C., Heithaus, M.R. and Barre, L.M. (2001b) Complex social structure, alliance stability and mating access in a bottlenose dolphin 'super-alliance'. Proceedings of the Royal Society of London. Series B: Biological Sciences268, 263-267.
10. Côté, I.M. and Poulin, R.B. (1995) Parasitism and group size in social animals: a meta-analysis. Behavioral Ecology6, 159-165.
11. Couzin, I.D., Krause, J., James, R., Ruxton, G.D. and Franks, N.R. (2002) Collective memory and spatial sorting in animal groups. Journal of Theoretical Biology218, 1-11.
12. Couzin, I.D., Krause, J., Franks, N.R. and Levin, S.A. (2005) Effective leadership and decision-making in animal groups on the move. Nature 433, 513-516.
13. Croft, D.P., Arrowsmith, B.J., Bielby, J. et al. (2003) Mechanisms underlying shoal composition in the Trinidadian guppy, Poecilia reticulata. Oikos100, 429-438.
14. Croft, D.P., Krause, J. and James, R. (2004) Social networks in the guppy (Poecilia reticulata). Proceedings of the Royal Society B: Biological Sciences271, 516-519.
15. Croft, D.P., James, R., Ward, A.J.W., Botham, M.S., Mawdsley, D. and Krause, J. (2005) Assortative interactions and social networks in fish. Oecologia143, 211-219.
16. Croft, D.P., Morrell, L.J., Wade, A.S. et al. (2006a) Predation risk as a driving force for sexual segregation: a cross-population comparison. American Naturalist167, 867-878.
17. Croft, D.P., James, R., Thomas, P. et al. (2006b) Social structure and co-operative interactions in a wild population of guppies (Poecilia reticulata). Behavioral Ecology and Sociobiology59, 644-650.
18. Croft, D.P., James, R. and Krause, J. (2008) Exploring Animal Social Networks, Vol 1. Princeton University Press, Princeton, New Jersey.
19. Croft, D.P., Madden, J.R., Franks, D.W. and James, R. (2011) Hypothesis testing in animal social networks. Trends in Ecology & Evolution, in press.
20. Cutts, C.J. and Speakman, J.R. (1994) Energy savings in formation flight of pink-footed geese. Journal of Experimental Biology189, 251-261.
21. Darden, S.K. and Croft, D.P. (2008) Male harassment drives females to alter habitat use and leads to segregation of the sexes. Biology Letters4, 449-451.
22. Darden, S.K., James, R., Ramnarine, I.W. and Croft, D.P. (2009) Social implications of the battle of the sexes: sexual harassment disrupts female sociality and social recognition. Proceedings of the Royal Society B: Biological Sciences276, 2651-2656.
23. Dewar, H., Mous, P., Domeier, M., Muljadi, A., Pet, J. and Whitty, J. (2008) Movements and site fidelity of the giant manta ray, Manta birostris, in the Komodo Marine Park, Indonesia. Marine Biology155, 121-133.
24. Domeier, M. and Nasby-Lucas, N. (2007) Annual re-sightings of photographically identified white sharks (Carcharodon carcharias) at an eastern Pacific aggregation site (Guadalupe Island, Mexico). Marine Biology150, 977-984.
25. Dudgeon, C.L., Noad, M.J. and Lanyon, J.M. (2008) Abundance and demography of a seasonal aggregation of zebra sharks Stegostoma fasciatum. Marine Ecology Progress Series368, 269-281.
26. Dunbar, R.I.M. and Shultz, S. (2007) Evolution in the social brain. Science317, 1344-1347.
27. Ebert, D.A. (1991) Observations on the predatory behaviour of the sevengill shark Notorynchus cepedianus. South African Journal of Marine Science11, 455-465.
28. Eckert, S.A. and Stewart, B.S. (2001) Telemetry and satellite tracking of whale sharks, Rhincodon typus, in the Sea of Cortez, Mexico, and the North Pacific Ocean. Environmental Biology of Fishes60, 299-308.
29. Economakis, A. and Lobel, P. (1998) Aggregation behavior of the grey reef shark, Carcharhinus amblyrhynchos, at Johnston Atoll, Central Pacific Ocean. Environmental Biology of Fishes51, 129-139.
30. Feldheim, K.A., Gruber, S.H. and Ashley, M.V. (2002) The breeding biology of lemon sharks at a tropical nursery lagoon. Proceedings of the Royal Society B: Biological Sciences269, 1655-1661.
31. Ford, E. (1921) A contribution to our knowledge of the life-histories of the dogfishes landed at Plymouth. Journal of the Marine Biological Association of the United Kingdom12, 468-505.
32. Frére, C.H., Krützen, M., Mann, J., Connor, R.C., Bejder, L. and Sherwin, W.B. (2010) Social and genetic interactions drive fitness variation in a free-living dolphin population. Proceedings of the National Academy of Sciences107, 19949-19954.
33. Frommen, J., Mehlis, M., Brendler, C. and Bakker, T. (2007) Shoaling decisions in three-spined sticklebacks (Gasterosteus aculeatus) - familiarity, kinship and inbreeding. Behavioral Ecology and Sociobiology61, 533-539.
34. Gilman, E., Clarke, S., Brothers, N. et al. (2008) Shark interactions in pelagic longline fisheries. Marine Policy32, 1-18.
35. Girard, M. and Du Buit, M.-H. (1999) Reproductive biology of two deep-water sharks from the British Isles, Centroscymnus coelolepis and Centrophorus squamosus (Chondrichthyes: Squalidae). Journal of the Marine Biological Association of the United Kingdom79, 923-931.
36. Goodale, E., Beauchamp, G., Magrath, R.D., Nieh, J.C. and Ruxton, G.D. (2010) Interspecific information transfer influences animal community structure. Trends in Ecology & Evolution25, 354-361.
37. Gowans, S., Whitehead, H. and Hooker, S.K. (2001) Social organization in northern bottlenose whales, Hyperoodon ampullatus: not driven by deep-water foraging?Animal Behaviour62, 369-377.
38. Griffiths, S.W. and Magurran, A.E. (1999) Schooling decisions in guppies (Poecilia reticulata) are based on familiarity rather than kin recognition by phenotype matching. Behavioral Ecology and Sociobiology45, 437-443.
39. Gruber, S.H., Nelson, D.R. and Morrissey, J.F. (1988) Patterns of activity and space utilization of lemon sharks, Negaprion brevirostris, in a shallow Bahamian lagoon. Bulletin of Marine Science43, 61-76.
40. Guttal, V. and Couzin, I.D. (2010) Social interactions, information use, and the evolution of collective migration. Proceedings of the National Academy of Sciences107, 16172-16177.
41. Guttridge, T.L., Gruber, S.H., Gledhill, K.S., Croft, D.P., Sims, D.W. and Krause, J. (2009a) Social preferences of juvenile lemon sharks, Negaprion brevirostris. Animal Behaviour78, 543-548.
42. Guttridge, T.L., Myrberg, A.A., Porcher, I.F., Sims, D.W. and Krause, J. (2009b) The role of learning in shark behaviour. Fish and Fisheries10, 450-469.
43. Guttridge, T.L., Gruber, S.H., Krause, J. and Sims, D.W. (2010) Novel acoustic technology for studying free-ranging shark social behaviour by recording individuals' interactions. PLoS ONE5, e9324.
44. Guttridge, T.L., Gruber, S.H., DiBattista, J.D. et al. (2011) Assortative interactions and leadership in a free-ranging population of juvenile lemon shark Negaprion brevirostris. Marine Ecology Progress Series423, 235-245.
45. Hain, T.J.A. and Neff, B.D. (2007) Multiple paternity and kin recognition mechanisms in a guppy population. Molecular Ecology16, 3938-3946.
46. Hamilton, W.D. (1964) The genetical evolution of social behaviour. II. Journal of Theoretical Biology7, 17-52.
47. Hearn, A., Ketchum, J., Klimley, A., Espinoza, E. and Peñaherrera, C. (2010) Hotspots within hotspots? Hammerhead shark movements around Wolf Island, Galapagos Marine Reserve. Marine Biology157, 1899-1915.
48. Heithaus, M.R., Frid, A., Wirsing, A.J. and Worm, B. (2008) Predicting ecological consequences of marine top predator declines. Trends in Ecology & Evolution23, 202-210.
49. Helfman, G.S. and Schultz, E.T. (1984) Social transmission of behavioural traditions in a coral reef fish. Animal Behaviour32, 379-384.
50. Herskin, J. and Steffensen, J.F. (1998) Energy savings in sea bass swimming in a school: measurements of tail beat frequency and oxygen consumption at different swimming speeds. Journal of Fish Biology53, 366-376.
51. Heupel, M. and Simpfendorfer, C. (2005) Quantitative analysis of aggregation behavior in juvenile blacktip sharks. Marine Biology147, 1239-1249.
52. Heupel, M.R., Simpfendorfer, C.A. and Hueter, R.E. (2004) Estimation of shark home ranges using passive monitoring techniques. Environmental Biology of Fishes71, 135-142.
53. Hight, B.V. and Lowe, C.G. (2007) Elevated body temperatures of adult female leopard sharks, Triakis semifasciata, while aggregating in shallow nearshore embayments: evidence for behavioral thermoregulation?Journal of Experimental Marine Biology and Ecology352, 114-128.
54. Hoare, D.J., Krause, J., Peuhkuri, N. and Godin, J.G.J. (2000) Body size and shoaling in fish. Journal of Fish Biology57, 1351-1366.
55. Hoare, D.J., Couzin, I.D., Godin, J.G.J. and Krause, J. (2004) Context-dependent group size choice in fish. Animal Behaviour67, 155-164.
56. Holland, K.N., Meyer, C.G. and Dagorn, L.C. (2009) Inter-animal telemetry: results from first deployment of acoustic 'business card' tags. Endangered Species Research10, 287-293.
57. Hueter, R.E., Heupel, M.R., Heist, E.J. and Keeney, D.B. (2005) Evidence of philopatry in sharks and implications for the management of shark fisheries. Journal of Northwest Atlantic Fishery Science35, 239-247.
58. Hulbert, L.B., Aires-da-Silva, A.M., Gallucci, V.F. and Rice, J.S. (2005) Seasonal foraging movements and migratory patterns of female Lamna ditropis tagged in Prince William Sound, Alaska. Journal of Fish Biology67, 490-509.
59. Jacoby, D.M.P., Busawon, D.S. and Sims, D.W. (2010) Sex and social networking: the influence of male presence on social structure of female shark groups. Behavioral Ecology21, 808-818.
60. Johnson, D.D.P., Kays, R., Blackwell, P.G. and Macdonald, D.W. (2002) Does the resource dispersion hypothesis explain group living?Trends in Ecology & Evolution17, 563-570.
61. Jorgensen, S.J., Reeb, C.A., Chapple, T.K. et al. (2009) Philopatry and migration of Pacific white sharks. Proceedings of the Royal Society B: Biological Sciences277, 679-688.
62. Klimley, A.P. (1985) Schooling in sphyrna-lewini, a species with low-risk of predation - a non-egalitarian state. Zeitschrift Fur Tierpsychologie-Journal of Comparative Ethology70, 297-319.
63. Klimley, A.P. (1987) The determinants of sexual segregation in the scalloped hammerhead shark, Sphyrna lewini. Environmental Biology of Fishes18, 27-40.
64. Klimley, A.P. and Nelson, D.R. (1984) Diel movement patterns of the scalloped hammerhead shark (Sphyrna lewini) in relation to El Bajo Espiritu Santo: a refuging central-position social system. Behavioral Ecology and Sociobiology15, 45-54.
65. Knip, D.M., Heupal, M.R. and Simpfendorfer, C. (2010) Sharks in nearshore environments: models, importance and consequences. Marine Ecology Progress Series402, 1-11.
66. Krause, J. and Ruxton, G.D. (2002) Living in Groups, Vol 1. Oxford University Press, Oxford.
67. Krause, J., Butlin, R.K., Peuhkuri, N. and Pritchard, V.L. (2000) The social organization of fish shoals: a test of the predictive power of laboratory experiments for the field. Biological Reviews75, 477-501.
68. Krause, J., Lusseau, D. and James, R. (2009) Animal social networks: an introduction. Behavioral Ecology and Sociobiology63, 967-973.
69. Krause, J., Wilson, A.D.M. and Croft, D.P. (2011) New technology facilitates the study of social networks. Trends in Ecology & Evolution26, 5-6.
70. Lachlan, R.F., Crooks, L. and Laland, K.N. (1998) Who follows whom? Shoaling preferences and social learning of foraging information in guppies. Animal Behaviour56, 181-190.
71. Litvinov, F. (2006) On the role of dense aggregations of males and juveniles in the functional structure of the range of the blue shark Prionace glauca. Journal of Ichthyology46, 613-624.
72. Lusseau, D. (2003) The emergent properties of a dolphin social network. Proceedings of the Royal Society of London. Series B: Biological Sciences270, S186-S188.
73. Lusseau, D., Whitehead, H. and Gero, S. (2008) Incorporating uncertainty into the study of animal social networks. Animal Behaviour75, 1809-1815.
74. Magurran, A.E. (1990) The adaptive significance of schooling as an anti-predator defence in fish. Annales Zoologici Fennici27, 51-66.
75. Magurran, A.E., Seghers, B.H., Shaw, P.W. and Carvalho, G.R. (1994) Schooling preferences for familiar fish in the guppy, Poecilia reticulata. Journal of Fish Biology45, 401-406.
76. Martin, R.A., Rossmo, D.K. and Hammerschlag, N. (2009) Hunting patterns and geographic profiling of white shark predation. Journal of Zoology279, 111-118.
77. McKibben, J.N. and Nelson, D.R. (1986) Patterns of movement and grouping of gray reef sharks, Carcharhinus amblyrhynchos, at Enewetak, Marshall Islands. Bulletin of Marine Science38, 89-110.
78. McLaughlin, R.H. and O'Gower, A.K. (1971) Life history and underwater studies of a Heterodont shark.. Ecological Monographs 41, 271-289.
79. Morrell, L.J., Croft, D.P., Dyer, J.R.G. et al. (2008) Association patterns and foraging behaviour in natural and artificial guppy shoals. Animal Behaviour76, 855-864.
80. Morrissey, J.F. and Gruber, S.H. (1993) Habitat selection by juvenile lemon sharks, Negaprion brevirostris. Environmental Biology of Fishes38, 311-319.
81. Mucientes, G.R., Queiroz, N., Sousa, L.L., Tarroso, P. and Sims, D.W. (2009) Sexual segregation of pelagic sharks and the potential threat from fisheries. Biology Letters5, 156-159.
82. Myers, R.A. and Worm, B. (2003) Rapid worldwide depletion of predatory fish communities. Nature423, 280-283.
83. Myers, R.A., Baum, J.K., Shepherd, T.D., Powers, S.P. and Peterson, C.H. (2007) Cascading effects of the loss of apex predatory sharks from a coastal ocean. Science315, 1846-1850.
84. Myrberg, A.A. Jr and Gruber, S.H. (1974) The behavior of the bonnethead shark, Sphyrna tiburo. Copeia1974, 358-374.
85. Northcutt, R.G. (1977) Elasmobranch central nervous system organization and its possible evolutionary significance. American Zoologist17, 411-429.
86. Pitcher, T.J. (1983) Heuristic definitions of fish shoaling behaviour. Animal Behaviour31, 611-613.
87. Powter, D.M. and Gladstone, W. (2009) Habitat-mediated use of space by juvenile and mating adult port jackson sharks, Heterodontus portusjacksoni, in Eastern Australia. Pacific Science63, 1-14.
88. Riley, M.J., Hale, M.S., Harman, A. and Rees, R.G. (2010) Analysis of whale shark Rhincodon typus aggregations near South Ari Atoll, Maldives Archipelago. Aquatic Biology8, 145-150.
89. Robbins, R.L. (2007) Environmental variables affecting the sexual segregation of great white sharks Carcharodon carcharias at the Neptune Islands South Australia. Journal of Fish Biology70, 1350-1364.
90. Robbins, W.D., Hisano, M., Connolly, S.R. and Choat, J.H. (2006) Ongoing collapse of coral-reef shark populations. Current Biology16, 2314-2319.
91. Rowat, D., Meekan, M., Engelhardt, U., Pardigon, B. and Vely, M. (2007) Aggregations of juvenile whale sharks (Rhincodon typus) in the Gulf of Tadjoura, Djibouti. Environmental Biology of Fishes80, 465-472.
92. Scott, G.W., Gibbs, L. and Holding, J. (1997) Group 'resting' behaviour in a population of captive bull huss (Scyliorhinus stellaris). Aquarium Sciences and Conservation1, 251-254.
93. Semeniuk, C.A.D. and Dill, L.M. (2006) Anti-predator benefits of mixed-species groups of cowtail stingrays (Pastinachus sephen) and whiprays (Himantura uarnak) at rest. Ethology112, 33-43.
94. Semeniuk, C.A.D. and Rothley, K.D. (2008) Costs of group-living for a normally solitary forager: effects of provisioning tourism on southern stingrays Dasyatis americana.. Marine Ecology Progress Series357, 271-282.
95. Silliman, W.R. and Gruber, S.H. (1999) Behavioral biology of the spotted eagle ray, Aetobatus narinari (Euphrasen, 1790), in Bimini, Bahamas; an interim report. Bahamas Journal of Science7, 13-20.
96. Simpfendorfer, C.A. and Milward, N.E. (1993) Utilisation of a tropical bay as a nursery area by sharks of the families Carcharhinidae and Sphyrnidae. Environmental Biology of Fishes37, 337-345.
97. Sims, D.W. (2003) Tractable models for testing theories about natural strategies: foraging behaviour and habitat selection of free-ranging sharks. Journal of Fish Biology63, 53-73.
98. Sims, D.W. (2005) Differences in habitat selection and reproductive strategies of male and female sharks. In: Sexual Segregation in Vertebrates: Ecology of the Two Sexes (eds K. Ruckstuhl and P. Neuhaus). Cambridge University Press, Cambridge, pp. 127-147.
99. Sims, D.W. (2008) Sieving a living: a review of the biology, ecology and conservation status of the plankton-feeding basking shark Cetorhinus maximus, Chapter 3. In: Advances in Marine Biology, (ed. D.W. Sims) Vol. 54. Academic Press, London, pp. 171-220.
100. Sims, D.W. (2010) Tracking and analysis techniques for understanding free-ranging shark movements and behavior. In: Sharks and Their Relatives II. Biodiversity, Adaptive Physiology, and Conservation (eds J.C. Carrier, J.A. Musick and M.R. Heithaus). CRC Press Taylor & Francis Group, Boca Raton, Florida, pp. 351-392.
101. Sims, D.W., Southall, E.J., Quayle, V.A. and Fox, A.M. (2000) Annual social behaviour of basking sharks associated with coastal front areas. Proceedings of the Royal Society B Biological Sciences267, 1897-1904.
102. Sims, D., Nash, J. and Morritt, D. (2001) Movements and activity of male and female dogfish in a tidal sea lough: alternative behavioural strategies and apparent sexual segregation. Marine Biology139, 1165-1175.
103. Sims, D.W., Southall, E.J., Wearmouth, V.J., Hutchinson, N., Budd, G.C. and Morritt, D. (2005) Refuging behaviour in the nursehound Scyliorhinus stellaris (Chondrichthyes: Elasmobranchii): preliminary evidence from acoustic telemetry. Journal of the Marine Biological Association of the UK85, 1137-1140.
104. Sims, D.W., Wearmouth, V.J., Southall, E.J. et al. (2006) Hunt warm, rest cool: bioenergetic strategy underlying diel vertical migration of a benthic shark. Journal of Animal Ecology75, 176-190.
105. Sperone, E., Micarelli, P., Andreotti, S. et al. (2010) Social interactions among bait-attracted white sharks at Dyer Island (South Africa). Marine Biology Research6, 408-414.
106. Springer, S. (1967) Social organization of shark populations. In: Sharks, Skates and Rays (eds P.W. Gilbert, R.F. Mathewson and D.P. Rall). John Hopkins Press, Baltimore, pp. 149-174.
107. Sumpter, D.J.T. (2006) The principles of collective animal behaviour. Philosophical Transactions of the Royal Society B: Biological Sciences361, 5-22.
108. Theberge, M.M. and Dearden, P. (2006) Detecting a decline in whale shark Rhincodon typus sightings in the Andaman Sea, Thailand, using ecotourist operator-collected data. Oryx40, 337-342.
109. Tibbetts, E.A. and Dale, J. (2007) Individual recognition: it is good to be different. Trends in Ecology & Evolution22, 529-537.
110. Topelko, K.N. and Dearden, P. (2005) The shark watching industry and its potential contribution to shark conservation. Journal of Ecotourism4, 108-128.
111. Ward, A.J.W., Botham, M.S., Hoare, D.J. et al. (2002) Association patterns and shoal fidelity in the three-spined stickleback. Proceedings of the Royal Society of London Series B: Biological Sciences269, 2451-2455.
112. Ward, A.J.W., Webster, M.M. and Hart, P.J.B. (2007) Social recognition in wild fish populations. Proceedings of the Royal Society B: Biological Sciences274, 1071-1077.
113. Ward, A.J.W., Sumpter, D.J.T., Couzin, I.D., Hart, P.J.B. and Krause, J. (2008) Quorum decision-making facilitates information transfer in fish shoals. Proceedings of the National Academy of Sciences105, 6948-6953.
114. Wearmouth, V.J. and Sims, D.W. (2008) Sexual segregation in marine fish, reptiles, birds and mammals: behaviour patterns, mechanisms and conservation implications. Advances in Marine Biology54, 107-170.
115. Weng, K., Boustany, A., Pyle, P., Anderson, S., Brown, A. and Block, B. (2007) Migration and habitat of white sharks (Carcharodon carcharias) in the eastern Pacific Ocean. Marine Biology152, 877-894.
116. Wetherbee, B.M., Gruber, S.H. and Rosa, R.S. (2007) Movement patterns of juvenile lemon sharks Negaprion brevirostris within Atol das Rocas, Brazil: a nursery characterized by tidal extremes. Marine Ecology Progress Series343, 283-293.
117. Wey, T., Blumstein, D.T., Shen, W. and Jordán, F. (2008) Social network analysis of animal behaviour: a promising tool for the study of sociality. Animal Behaviour75, 333-344.
118. Whitney, N.M., Pratt, J.H.L. and Carrier, J.C. (2004) Group courtship, mating behaviour and siphon sac function in the whitetip reef shark, Triaenodon obesus. Animal Behaviour68, 1435-1442.
119. Wilson, S.G. (2004) Basking sharks (Cetorhinus maximus) schooling in the southern Gulf of Maine. Fisheries Oceanography13, 283-286.
120. Wolf, J.B.W., Mawdsley, D., Trillmich, F. and James, R. (2007) Social structure in a colonial mammal: unravelling hidden structural layers and their foundations by network analysis. Animal Behaviour74, 1293-1302.
121. Worm, B., Barbier, E.B., Beaumont, N. et al. (2006) Impacts of biodiversity loss on ocean ecosystem services. Science314, 787-790.