1.
McCauley, D. J. et al. Marine defaunation: Animal loss in the global ocean. Science 347, 1255641 (2015).
PubMed Article CAS Google Scholar
2.
Dulvy, N. K. et al. Extinction risk and conservation of the world’s sharks and rays. eLife 3, 590 (2014).
Article Google Scholar
3.
Hutchings, J. A., Myers, R. A., García, V. B., Lucifora, L. O. & Kuparinen, A. Life-history correlates of extinction risk and recovery potential. Ecol. Appl. 22, 1061–1067 (2012).
PubMed Article Google Scholar
4.
Frisk, M. & Miller, T. J. Life histories and vulnerability to exploitation of elasmobranchs: Inferences from elasticity, perturbation and phylogenetic analyses. Artic. J. Northwest Atl. Fish. Sci. https://doi.org/10.2960/J.v35.m514 (2005).
Article Google Scholar
5.
Carr, L. A. et al. Illegal shark fishing in the Galápagos Marine Reserve. Mar. Policy 39, 317–321 (2013).
Article Google Scholar
6.
Dharmadi, F. & Satria, F. African Journal of Marine Science Fisheries management and conservation of sharks in Indonesia. Afr. J. Mar. Sci. 37, 249–258 (2015).
Article Google Scholar
7.
Heupel, M., Carlson, J. & Simpfendorfer, C. Shark nursery areas: Concepts, definition, characterization and assumptions. Mar. Ecol. Prog. Ser. 337, 287–297 (2007).
ADS Article Google Scholar
8.
Meylan, P. A., Meylan, A. B. & Gray, J. A. The ecology and migrations of sea turtles 8. Tests of the developmental habitat hypothesis. Bull. Am. Museum Nat. Hist. 357, 1–70 (2011).
Article Google Scholar
9.
Jennings, D. E., Gruber, S. H., Franks, B. R., Kessel, S. T. & Robertson, A. L. Effects of large-scale anthropogenic development on juvenile lemon shark (Negaprion brevirostris) populations of Bimini, Bahamas. Environ. Biol. Fishes 83, 369–377 (2008).
Article Google Scholar
10.
Kinney, M. J. & Simpfendorfer, C. A. Reassessing the value of nursery areas to shark conservation and management. Conserv. Lett. 2, 53–60 (2009).
Article Google Scholar
11.
Healy, T. J., Hill, N. J., Chin, A. & Barnett, A. A global review of elasmobranch tourism activities, management and risk. Mar. Policy 118, 103964 (2020).
Article Google Scholar
12.
White, T. D. et al. Assessing the effectiveness of a large marine protected area for reef shark conservation. Biol. Conserv. 207, 64–71 (2017).
Article Google Scholar
13.
Claudet, J., Loiseau, C., Sostres, M. & Correspondence, M. Z. Underprotected marine protected areas in a global biodiversity hotspot. One Earth https://doi.org/10.1016/j.oneear.2020.03.008 (2020).
Article Google Scholar
14.
Pierce, S. & Norman, B. Rhincodon typus. IUCN Red List Threat. Species e-T19488A2, (2016).
15.
CITES. Convention on international trade in endangered species of wild fauna and flora. Amendments to Appendices I and II of CITES. (2000).
16.
Convention on Migratory Species. Proposal for the inclusion of the whale shark (Rhincodon typus) on Appendix I of the convention CMS convention on migratory species. (2017).
17.
Simpfendorfer, C. A. & Dulvy, N. K. Bright spots of sustainable shark fishing. Curr. Biol. 27, R97–R98 (2017).
CAS PubMed Article Google Scholar
18.
Reeve-Arnold, K. E., Kinni, J., Newbigging, R., Pierce, S. J. & Roques, K. Sustaining whale shark tourism in a diminishing population. In (Hamad bin Khalifa University Press, HBKU Press, 2016). https://doi.org/10.5339/qproc.2016.iwsc4.49.
19.
Pravin, P. Whale Shark in the Indian Coast—Need for conservation. Curr. Sci. 79, 310–315 (2000).
Google Scholar
20.
Li, W., Wang, Y. & Norman, B. A preliminary survey of whale shark Rhincodon typus catch and trade in China: An emerging crisis. J. Fish Biol. 80, 1608–1618 (2012).
CAS PubMed Article PubMed Central Google Scholar
21.
Hearn, A. R. et al. Adult female whale sharks make long-distance movements past Darwin Island (Galapagos, Ecuador) in the Eastern Tropical Pacific. Mar. Biol. 163, 1–12 (2016).
Article Google Scholar
22.
Wilson, S. G., Polovina, J. J., Stewart, B. S. & Meekan, M. G. Movements of whale sharks (Rhincodon typus) tagged at Ningaloo Reef, Western Australia. Mar. Biol. 148, 1157–1166 (2006).
Article Google Scholar
23.
Hueter, R. E., Tyminski, J. P. & de la Parra, R. Horizontal movements, migration patterns, and population structure of whale sharks in the Gulf of Mexico and northwestern Caribbean Sea. PLoS ONE 8, e71883 (2013).
ADS CAS PubMed PubMed Central Article Google Scholar
24.
Robinson, D. P. et al. Some like it hot: Repeat migration and residency of whale sharks within an extreme natural environment. PLoS ONE 12, e0185360 (2017).
PubMed PubMed Central Article CAS Google Scholar
25.
Araujo, G. et al. Photo-ID and telemetry highlight a global whale shark hotspot in Palawan, Philippines. Sci. Rep. 9, 1–12 (2019).
Article CAS Google Scholar
26.
Bradshaw, C. J. A., Fitzpatrick, B. M., Steinberg, C. C., Brook, B. W. & Meekan, M. G. Decline in whale shark size and abundance at Ningaloo Reef over the past decade: The world’s largest fish is getting smaller. Biol. Conserv. 141, 1894–1905 (2008).
Article Google Scholar
27.
Speed, C. W. et al. Scarring patterns and relative mortality rates of Indian Ocean whale sharks. J. Fish Biol. 72, 1488–1503 (2008).
Article Google Scholar
28.
Lester, E. et al. Multi-year patterns in scarring, survival and residency of whale sharks in Ningaloo Marine Park, Western Australia. Mar. Ecol. Prog. Ser. 634, 115–125 (2020).
ADS Article Google Scholar
29.
Rowat, D. & Brooks, K. S. A review of the biology, fisheries and conservation of the whale shark Rhincodon typus. J. Fish Biol. 80, 1019–1056 (2012).
CAS PubMed Article PubMed Central Google Scholar
30.
Cochran, J. E. M. et al. Multi-method assessment of whale shark (Rhincodon typus) residency, distribution, and dispersal behavior at an aggregation site in the Red Sea. PLoS ONE 14, e0222285 (2019).
CAS PubMed PubMed Central Article Google Scholar
31.
Copping, J. P., Stewart, B. D., McClean, C. J., Hancock, J. & Rees, R. Does bathymetry drive coastal whale shark (Rhincodon typus) aggregations?. PeerJ 6, e4904 (2018).
PubMed PubMed Central Article Google Scholar
32.
Norman, B. M. et al. Undersea constellations: The global biology of an endangered marine megavertebrate further informed through citizen science. Bioscience 67, 1029–1043 (2017).
Article Google Scholar
33.
Donati, G. et al. New insights into the South Ari atoll whale shark, Rhincodon typus, aggregation. In (Hamad bin Khalifa University Press, HBKU Press, 2016). https://doi.org/10.5339/qproc.2016.iwsc4.16.
34.
Riley, M. J., Hale, M. S., Harman, A. & Rees, R. G. Analysis of whale shark Rhincodon typus aggregations near South Ari Atoll, Maldives Archipelago. Aquat. Biol. 8, 145–150 (2010).
Article Google Scholar
35.
Rowat, D., Meekan, M. G., Engelhardt, U., Pardigon, B. & Vely, M. Aggregations of juvenile whale sharks (Rhincodon typus) in the Gulf of Tadjoura, Djibouti. Environ. Biol. Fishes 80, 465–472 (2007).
Article Google Scholar
36.
Cagua, E. F. et al. Acoustic telemetry reveals cryptic residency of whale sharks. Biol. Lett. 11, 20150092 (2015).
PubMed PubMed Central Article Google Scholar
37.
Thomson, J. A. et al. Feeding the world’s largest fish: Highly variable whale shark residency patterns at a provisioning site in the Philippines. R. Soc. Open Sci. 4, 170394 (2017).
ADS PubMed PubMed Central Article Google Scholar
38.
Perry, C. T. et al. Comparing length-measurement methods and estimating growth parameters of free-swimming whale sharks (Rhincodon typus) near the South Ari Atoll, Maldives. Mar. Freshw. Res. 69, 1487 (2018).
Article Google Scholar
39.
Riley, M. J., Harman, A. & Rees, R. G. Evidence of continued hunting of whale sharks Rhincodon typus in the Maldives. Environ. Biol. Fishes 86, 371–374 (2009).
Article Google Scholar
40.
Cagua, E. F., Collins, N., Hancock, J. & Rees, R. Whale shark economics: A valuation of wildlife tourism in South Ari Atoll. Maldives. PeerJ 2, e515 (2014).
PubMed Article Google Scholar
41.
Arzoumanian, Z., Holmberg, J. & Norman, B. An astronomical pattern-matching algorithm for computer-aided identification of whale sharks Rhincodon typus. J. Appl. Ecol. 42, 999–1011 (2005).
Article Google Scholar
42.
Bradshaw, C. J. A., Mollett, H. F. & Meekan, M. G. Inferring population trends for the world’s largest fish from mark recapture estimates of survival. J. Anim. Ecol. 76, 480–489 (2007).
PubMed Article Google Scholar
43.
Holmberg, J., Norman, B. & Arzoumanian, Z. Estimating population size, structure, and residency time for whale sharks Rhincodon typus through collaborative photo-identification. Endanger. Species Res. 7, 39–53 (2009).
Article Google Scholar
44.
Rowat, D., Gore, M., Meekan, M. G., Lawler, I. R. & Bradshaw, C. J. A. Aerial survey as a tool to estimate whale shark abundance trends. J. Exp. Mar. Biol. Ecol. 368, 1–8 (2009).
Article Google Scholar
45.
Acuña-Marrero, D. et al. Whale shark (Rhincodon typus) seasonal presence, residence time and habitat use at darwin island, galapagos marine reserve. PLoS ONE 9, e115946 (2014).
ADS PubMed PubMed Central Article CAS Google Scholar
46.
Schwarz, C. J., Bailey, R. E., Irvine, J. R. & Dalziel, F. C. Estimating salmon spawning escapement using capture-recapture methods. Can. J. Fish. Aquat. Sci. 50, 1181–1197 (1993).
Article Google Scholar
47.
Schwarz, C. J. & Arnason, A. N. A general methodology for the analysis of capture-recapture experiments in open populations. Biometrics 52, 860 (1996).
MathSciNet MATH Article Google Scholar
48.
Whitehead, H. Analysis of animal movement using opportunistic individual identifications: Application to sperm whales. Ecology 82, 1417–1432 (2001).
Article Google Scholar
49.
QGIS.org. QGIS Geographic Information System. Open Source Geospatial Foundation Project. (2020).
50.
Van Tienhoven, A. M., Den Hartog, J. E., Reijns, R. A. & Peddemors, V. M. A computer-aided program for pattern-matching of natural marks on the spotted raggedtooth shark Carcharias taurus. J. Appl. Ecol. 44, 273–280 (2007).
Article Google Scholar
51.
RStudio Team. RStudio: Integrated Development for R. (2015).
52.
Laake, J. L. RMark: An R interface for analysis of capture-recapture data with MARK. AFSC Processed Rep. 2013-01 Alaska Fish. Sci. Cent., NOAA, Natl. Mar. Fish. Serv., (2013). https://doi.org/10.1017/CBO9781107415324.004.
53.
Schwarz, C., Arnason, A., Cooch, E. & White, G. Jolly-Seber models in MARK. Progr. MARK–a gentle Introd. 18th Ed. (2018).
54.
Cooch, E. & White, G. Program MARK: A gentle introduction (13th ed.). available online with MARK Program. (2006).
55.
Whitehead, H. SOCPROG programs: Analysing animal social structures. Behav. Ecol. Sociobiol. 63, 765–778 (2009).
Article Google Scholar
56.
Whitehead, H. Selection of models of lagged identification rates and lagged association rates using AIC and QAIC. Commun. Stat. Simul. Comput. 36, 1233–1246 (2007).
MathSciNet MATH Article Google Scholar
57.
Buckland, S. T. & Garthwaite, P. H. Quantifying precision of mark-recapture estimates using the bootstrap and related methods. Biometrics 47, 255 (1991).
Article Google Scholar
58.
Rohner, C. A. et al. No place like home? High residency and predictable seasonal movement of whale sharks off Tanzania. Front. Mar. Sci. 7, 423 (2020).
Article Google Scholar
59.
Norman, B. M., Whitty, J. M., Beatty, S. J., Reynolds, S. D. & Morgan, D. L. Do they stay or do they go? Acoustic monitoring of whale sharks at Ningaloo Marine Park, Western Australia. J. Fish Biol. 91, 1713–1720 (2017).
CAS PubMed Article Google Scholar
60.
Araujo, G. et al. Population structure and residency patterns of whale sharks, Rhincodon typus, at a provisioning site in Cebu, Philippines. PeerJ 2014, e543 (2014).
Article Google Scholar
61.
Prebble, C. et al. Limited latitudinal ranging of juvenile whale sharks in the Western Indian Ocean suggests the existence of regional management units. Mar. Ecol. Prog. Ser. 601, 167–183 (2018).
ADS Article Google Scholar
62.
Araujo, G. et al. Population structure and residency patterns of whale sharks, Rhincodon typus, at a provisioning site in Cebu, Philippines. PeerJ 2, e543 (2014).
PubMed PubMed Central Article Google Scholar
63.
Akhilesh, K. V. et al. Landings of whale sharks Rhincodon typus Smith, 1828 in Indian waters since protection in 2001 through the Indian Wildlife (Protection) Act, 1972. Environ. Biol. Fishes 96, 713–722 (2013).
Article Google Scholar
64.
Heyman, W., Graham, R., Kjerfve, B. & Johannes, R. Whale sharks Rhincodon typus aggregate to feed on fish spawn in Belize. Mar. Ecol. Prog. Ser. 215, 275–282 (2001).
ADS Article Google Scholar
65.
Meekan, M. et al. Population size and structure of whale sharks Rhincodon typus at Ningaloo Reef, Western Australia. Mar. Ecol. Prog. Ser. 319, 275–285 (2006).
ADS Article Google Scholar
66.
Cochran, J. E. M. et al. Population structure of a whale shark Rhincodon typus aggregation in the Red Sea. J. Fish Biol. 89, 1570–1582 (2016).
CAS PubMed Article PubMed Central Google Scholar
67.
Araujo, G. et al. Population structure, residency patterns and movements of whale sharks in Southern Leyte, Philippines: Results from dedicated photo-ID and citizen science. Aquat. Conserv. Mar. Freshw. Ecosyst. 27, 237–252 (2017).
Article Google Scholar
68.
Robinson, D. P. et al. Population structure, abundance and movement of whale sharks in the Arabian Gulf and the Gulf of Oman. PLoS ONE 11, e0158593 (2016).
PubMed PubMed Central Article CAS Google Scholar
69.
McCoy, E. et al. Long-term photo-identification reveals the population dynamics and strong site fidelity of adult whale sharks to the coastal waters of Donsol, Philippines. Front. Mar. Sci. 5, 271 (2018).
Article Google Scholar
70.
Araujo, G. et al. In-water methods reveal population dynamics of a green turtle Chelonia mydas foraging aggregation in the Philippines. Endanger. Species Res. 40, 207–218 (2019).
Article Google Scholar
71.
Sleeman, J. C. et al. To go or not to go with the flow: Environmental influences on whale shark movement patterns. J. Exp. Mar. Biol. Ecol. 390, 84–98 (2010).
Article Google Scholar
72.
Calambokidis, J., Laake, J. L. & Klimek, A. Abundance and population structure of seasonal gray whales in the Pacific Northwest, 1998–2008. Sc/62/Brg32, Vol. 2008 (2010).
73.
Branstetter, S. Early Life-History Implications of Selected Carcharhinoid and Lamnoid Sharks of the Northwest Atlantic. Elasmobranchs as Living Resour. Adv. Biol. Ecol. Syst. Status Fish. (1990).
74.
Parker, J. H. & Gischler, E. Modern foraminiferal distribution and diversity in two atolls from the Maldives, Indian Ocean. Mar. Micropaleontol. 78, 30–49 (2011).
ADS Article Google Scholar
75.
Halvorsen, M. B., Casper, B. M., Woodley, C. M., Carlson, T. J. & Popper, A. N. Threshold for onset of injury in Chinook salmon from exposure to impulsive pile driving sounds. PLoS ONE 7, e38968 (2012).
ADS CAS PubMed PubMed Central Article Google Scholar
76.
Haskell, P. J. et al. Monitoring the effects of tourism on whale shark Rhincodon typus behaviour in Mozambique. ORYX 49, 492–499 (2015).
Article Google Scholar
77.
Quiros, A. L. Tourist compliance to a Code of Conduct and the resulting effects on whale shark (Rhincodon typus) behavior in Donsol, Philippines. Fish. Res. 84, 102–108 (2007).
Article Google Scholar
78.
Araujo, G., Vivier, F., Labaja, J. J., Hartley, D. & Ponzo, A. Assessing the impacts of tourism on the world’s largest fish Rhincodon typus at Panaon Island, Southern Leyte, Philippines. Aquat. Conserv. Mar. Freshw. Ecosyst. 27, 986–994 (2017).
Article Google Scholar
79.
Finger, J. S. et al. Rate of movement of juvenile lemon sharks in a novel open field, are we measuring activity or reaction to novelty?. Anim. Behav. 116, 75–82 (2016).
Article Google Scholar
80.
Cade, D. E. et al. Whale sharks increase swimming effort while filter feeding, but appear to maintain high foraging efficiencies. J. Exp. Biol. 223, jeb.224402 (2020).
Article Google Scholar
81.
Archie, E. A. Wound healing in the wild: stress, sociality, and energetic costs affect wound healing in natural populations. Parasite Immunol. 35, n/a-n/a (2013).
82.
Baker, M. R., Swanson, P. & Young, G. Injuries from non-retention in gillnet fisheries suppress reproductive maturation in escaped fish. PLoS ONE 8, e69615 (2013).
ADS CAS PubMed PubMed Central Article Google Scholar
83.
Neat, F. C., Taylor, A. C. & Huntingford, F. A. Proximate costs of fighting in male cichlid fish: The role of injuries and energy metabolism. Anim. Behav. 55, 875–882 (1998).
CAS PubMed Article Google Scholar
84.
Meekan, M. G., Fuiman, L. A., Davis, R., Berger, Y. & Thums, M. Swimming strategy and body plan of the world’s largest fish: Implications for foraging efficiency and thermoregulation. Front. Mar. Sci. 2, 64 (2015).
Article Google Scholar
85.
Chin, A., Mourier, J. & Rummer, J. L. Blacktip reef sharks (Carcharhinus melanopterus) show high capacity for wound healing and recovery following injury. Conserv. Physiol. 3(1) (2015).
86.
Tierney, K. B. & Farrell, A. P. The relationships between fish health, metabolic rate, swimming performance and recovery in return-run sockeye salmon, Oncorhynchus nerka (Walbaum). J. Fish Dis. 27, 663–671 (2004).
CAS PubMed Article Google Scholar
87.
McGregor, F., Richardson, A. J., Armstrong, A. J., Armstrong, A. O. & Dudgeon, C. L. Rapid wound healing in a reef manta ray masks the extent of vessel strike. PLoS ONE 14, e0225681 (2019).
CAS PubMed PubMed Central Article Google Scholar
88.
Tort, L. Stress and immune modulation in fish. Dev. Comp. Immunol. 35, 1366–1375 (2011).
CAS PubMed Article Google Scholar
89.
Mateus, A. P., Anjos, L., Cardoso, J. R. & Power, D. M. Chronic stress impairs the local immune response during cutaneous repair in gilthead sea bream (Sparus aurata L.). Mol. Immunol. 87, 267–283 (2017).
CAS PubMed Article Google Scholar
90.
Environmental Protection Agency. Maldivian Whale Shark Tourist Encounter Guidelines. (2009).
91.
Leston, F. A. L. Monitoring Tourist Pressure on Whale Shark (Rhincodon typus) Behaviour in South Ari MPA, Maldive) Behaviour in South Ari MPA, Maldive (The University of Edinburgh, Edinburgh, 2016).
Google Scholar
92.
Kallsen, H. Regulation of Whale Shark Tourism: A Data Driven Approach for the South Ari Marine Protected Area (Syddansk Universitet, Odense, 2018).
Google Scholar
93.
Montero-Quintana, A. N., Vázquez-Haikin, J. A., Merkling, T., Blanchard, P. & Osorio-Beristain, M. Ecotourism impacts on the behaviour of whale sharks: An experimental approach. ORYX 54, 270–275 (2020).
Article Google Scholar
94.
Bouyoucos, I. A., Simpfendorfer, C. A. & Rummer, J. L. Estimating oxygen uptake rates to understand stress in sharks and rays. Rev. Fish Biol. Fish. 29, 297–311 (2019).
Article Google Scholar
95.
Semeniuk, C. A. D., Bourgeon, S., Smith, S. L. & Rothley, K. D. Hematological differences between stingrays at tourist and non-visited sites suggest physiological costs of wildlife tourism. Biol. Conserv. 142, 1818–1829 (2009).
Article Google Scholar
96.
Van Rijn, J. A. & Reina, R. D. Distribution of leukocytes as indicators of stress in the Australian swellshark, Cephaloscyllium laticeps. Fish Shellfish Immunol. 29, 534–538 (2010).
PubMed Article CAS PubMed Central Google Scholar
97.
Barnett, A., Payne, N. L., Semmens, J. M. & Fitzpatrick, R. Ecotourism increases the field metabolic rate of whitetip reef sharks. Biol. Conserv. 199, 132–136 (2016).
Article Google Scholar
98.
Mau, R. Managing for conservation and recreation: The Ningaloo whale shark experience. J. Ecotourism 7, 213–225 (2008).
Article Google Scholar
99.
Martin, R. A. A review of behavioural ecology of whale sharks (Rhincodon typus). Fish. Res. 84, 10–16 (2007).
ADS Article Google Scholar
100.
Skomal, G. B. & Mandelman, J. W. The physiological response to anthropogenic stressors in marine elasmobranch fishes: A review with a focus on the secondary response. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 126, 146–155 (2012).
Article CAS Google Scholar
101.
Pankhurst, N. W. The endocrinology of stress in fish: An environmental perspective. Gen. Comp. Endocrinol. 170, 265–275 (2011).
CAS PubMed Article PubMed Central Google Scholar
102.
Renshaw, G. M. C., Kutek, A. K., Grant, G. D. & Anoopkumar-Dukie, S. Forecasting elasmobranch survival following exposure to severe stressors. Comp. Biochem. Physiol. Part A Mol. Integr. Physiol. 126, 101–112 (2012).
Article CAS Google Scholar
103.
Lester, E. et al. Using an electronic monitoring system and photo identification to understand effects of tourism encounters on whale sharks in Ningaloo Marine Park. Tour. Mar. Environ. 14, 121–131 (2019).
Article Google Scholar More