Register      Login
Pacific Conservation Biology Pacific Conservation Biology Society
A journal dedicated to conservation and wildlife management in the Pacific region.
RESEARCH ARTICLE

Movement patterns of whale sharks in Cenderawasih Bay, Indonesia, revealed through long-term satellite tagging

Megan M. Meyers A F , Malcolm P. Francis B , Mark Erdmann C D , Rochelle Constantine https://orcid.org/0000-0003-3260-539X A E and Abraham Sianipar C
+ Author Affiliations
- Author Affiliations

A Institute of Marine Science, University of Auckland, Private Bag 92019, Auckland, New Zealand.

B National Institute of Water and Atmospheric Research, Greta Point, Wellington, New Zealand.

C Conservation International Indonesia Marine Program, Bali, Indonesia.

D California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118, USA.

E School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand.

F Corresponding author. Email: megan.meyers@research.uwa.edu.au

Pacific Conservation Biology 26(4) 353-364 https://doi.org/10.1071/PC19035
Submitted: 8 September 2019  Accepted: 25 May 2020   Published: 30 June 2020

Abstract

Whale sharks, Rhincodon typus, are found circumglobally in tropical and warm temperate seas, exhibiting a range of residency and movement patterns. To determine spatio-temporal habitat use by juvenile male whale sharks in Cenderawasih Bay, Indonesia, we collected data from June 2015 to November 2016 using 16 fin-mounted satellite tags that provided exceptionally long track durations. Fifteen tags transmitted for 48–534 days (mean = 321 ± 33, s.e.), with 13 tags transmitting for ≥220 days. Four sharks remained within the bay for the duration of the study, while of the 11 sharks that travelled outside the bay, eight left between March and May 2016. They ranged throughout coastal and offshore waters, travelling up to 5144 km away from Cenderawasih Bay, with a mean horizontal speed of only 3.3 km day−1 ± 0.70, s.e. A switching state space model was fitted to satellite fix data to identify behavioural states. It revealed that sharks spent an average of 81% of their time in foraging-related behaviours, mostly in shallow waters (median depth = 35 m), with travelling observed mainly over deeper waters (median depth = 1284 m). The movement patterns reveal variable periods of residency, with individual patterns of horizontal movement most likely in response to different abiotic and biotic factors, including food availability, which may trigger seasonal dispersal.

Additional keywords: telemetry, spatial distribution, West Papua, elasmobranch, Bird’s Head Seascape.


References

Araujo, G., Lucey, A., Labaja, J., So, C. L., Snow, S., and Ponzo, A. (2014). Population structure and residency patterns of whale sharks, Rhincodon typus, at a provisioning site in Cebu, Philippines. PeerJ 2, e543.
Population structure and residency patterns of whale sharks, Rhincodon typus, at a provisioning site in Cebu, Philippines.Crossref | GoogleScholarGoogle Scholar | 25279256PubMed |

Baker, R. (1978). ‘Evolutionary Ecology of Animal Migration.’ (Holmes and Meier Publishers.)

Berumen, M. L., Braun, C. D., Cochran, J. E., Skomal, G. B., and Thorrold, S. R. (2014). Movement patterns of juvenile whale sharks tagged at an aggregation site in the Red Sea. PLoS One 9, e103536.
Movement patterns of juvenile whale sharks tagged at an aggregation site in the Red Sea.Crossref | GoogleScholarGoogle Scholar | 25076407PubMed |

Block, B. A., Dewar, H., Blackwell, S. B., Williams, T. D., Prince, E. D., Farwell, C. J., Boustany, A., Teo, S. L., Seitz, A., Walli, A., and Fudge, D. (2001). Migratory movements, depth preferences, and thermal biology of Atlantic bluefin tuna. Science 293, 1310–1314.
Migratory movements, depth preferences, and thermal biology of Atlantic bluefin tuna.Crossref | GoogleScholarGoogle Scholar | 11509729PubMed |

Block, B. A., Teo, S. L., Walli, A., Boustany, A., Stokesbury, M. J., Farwell, C. J., Weng, K. C., Dewar, H., and Williams, T. D. (2005). Electronic tagging and population structure of Atlantic bluefin tuna. Nature 434, 1121–1127.
Electronic tagging and population structure of Atlantic bluefin tuna.Crossref | GoogleScholarGoogle Scholar | 15858572PubMed |

Borrell, A., Cardona, L., Kumarran, R. P., and Aguilar, A. (2011). Trophic ecology of elasmobranchs caught off Gujarat, India, as inferred from stable isotopes. ICES Journal of Marine Science 68, 547–554.
Trophic ecology of elasmobranchs caught off Gujarat, India, as inferred from stable isotopes.Crossref | GoogleScholarGoogle Scholar |

Cade, D. E., Levenson, J. J., Cooper, R., de la Parra, R., Webb, D. H., and Dove, A. D. M. (2020). Whale sharks increase swimming effort while filter feeding, but appear to maintain high foraging efficiencies. The Journal of Experimental Biology , .
Whale sharks increase swimming effort while filter feeding, but appear to maintain high foraging efficiencies.Crossref | GoogleScholarGoogle Scholar | 32366692PubMed |

Cagua, E. F., Cochran, J. E., Rohner, C. A., Prebble, C. E., Sinclair-Taylor, T. H., Pierce, S. J., and Berumen, M. L. (2015). Acoustic telemetry reveals cryptic residency of whale sharks. Biology Letters 11, 20150092.
Acoustic telemetry reveals cryptic residency of whale sharks.Crossref | GoogleScholarGoogle Scholar | 25832816PubMed |

Castro, A., Stewart, B., Wilson, S., Hueter, R., Meekan, M., Motta, P., Bowen, B., and Karl, S. (2007). Population genetic structure of earth’s largest fish, the whale shark (Rhincodon typus). Molecular Ecology 16, 5183–5192.
Population genetic structure of earth’s largest fish, the whale shark (Rhincodon typus).Crossref | GoogleScholarGoogle Scholar | 18092992PubMed |

Colman, J. (1997). A review of the biology and ecology of the whale shark. Journal of Fish Biology 51, 1219–1234.
A review of the biology and ecology of the whale shark.Crossref | GoogleScholarGoogle Scholar | 29991171PubMed |

Compagno, L. J. (2001). ‘Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date.’ (Food and Agriculture Organization: Rome.)

Diamant, S., Rohner, C. A., Kiszka, J. J., Guillemain d’Echon, A., Guillemain d’Echon, T., Sourisseau, E., and Pierce, S. J. (2018). Movements and habitat use of satellite-tagged whale sharks off western Madagascar. Endangered Species Research 36, 49–58.
Movements and habitat use of satellite-tagged whale sharks off western Madagascar.Crossref | GoogleScholarGoogle Scholar |

Eckert, S. A., Dolar, L. L., Kooyman, G. L., Perrin, W., and Rahman, R. A. (2002). Movements of whale sharks (Rhincodon typus) in South-east Asian waters as determined by satellite telemetry. Journal of Zoology 257, 111–115.
Movements of whale sharks (Rhincodon typus) in South-east Asian waters as determined by satellite telemetry.Crossref | GoogleScholarGoogle Scholar |

Escalle, L., Murua, H., Amande, J. M., Arregui, I., Chavance, P., Delgado de Molina, A., Gaertner, D., Fraile, I., Filmalter, J. D., and Santiago, J. (2016). Post‐capture survival of whale sharks encircled in tuna purse‐seine nets: tagging and safe release methods. Aquatic Conservation 26, 782–789.
Post‐capture survival of whale sharks encircled in tuna purse‐seine nets: tagging and safe release methods.Crossref | GoogleScholarGoogle Scholar |

Fox, S., Foisy, I., De La Parra Venegas, R., Galván Pastoriza, B. E., Graham, R. T., Hoffmayer, E. R., Holmberg, J., and Pierce, S. J. (2013). Population structure and residency of whale sharks Rhincodon typus at Utila, Bay Islands, Honduras. Journal of Fish Biology 83, 574–587.
Population structure and residency of whale sharks Rhincodon typus at Utila, Bay Islands, Honduras.Crossref | GoogleScholarGoogle Scholar | 23991875PubMed |

Francis, M. P., Holdsworth, J. C., and Block, B. A. (2015). Life in the open ocean: seasonal migration and diel diving behaviour of Southern Hemisphere porbeagle sharks (Lamna nasus). Marine Biology 162, 2305–2323.
Life in the open ocean: seasonal migration and diel diving behaviour of Southern Hemisphere porbeagle sharks (Lamna nasus).Crossref | GoogleScholarGoogle Scholar |

Freitas, C., Lydersen, C., Fedak, M. A., and Kovacs, K. M. (2008). A simple new algorithm to filter marine mammal Argos locations. Marine Mammal Science 24, 315–325.
A simple new algorithm to filter marine mammal Argos locations.Crossref | GoogleScholarGoogle Scholar |

Genisa, A. S. (2000). The occurrence of fish species in the Mamberamo estuary, Irian Jaya. In ‘Anonymous Proceedings of the Indo-Tropics Workshop’. (Eds A. G. Ilahude, W. W. Kastoro, and D. P. Praseno.) pp. 135–144. (Research And Development Centre For Oceanology, Indonesian Institute Of Sciences: Jakarta, Indonesia.)

Gleiss, A. C., Wright, S., Liebisch, N., Wilson, R. P., and Norman, B. (2013). Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef. Marine Biology 160, 2981–2992.
Contrasting diel patterns in vertical movement and locomotor activity of whale sharks at Ningaloo Reef.Crossref | GoogleScholarGoogle Scholar |

Grimes, C. B., and Finucane, J. H. (1991). Spatial distribution and abundance of larval and juvenile fish, chlorophyll and macrozooplankton around the Mississippi River discharge plume, and the role of the plume in fish recruitment. Marine Ecology Progress Series 75, 109–119.
Spatial distribution and abundance of larval and juvenile fish, chlorophyll and macrozooplankton around the Mississippi River discharge plume, and the role of the plume in fish recruitment.Crossref | GoogleScholarGoogle Scholar |

Hammerschlag, N., Gallagher, A., and Lazarre, D. (2011). A review of shark satellite tagging studies. Journal of Experimental Marine Biology and Ecology 398, 1–8.
A review of shark satellite tagging studies.Crossref | GoogleScholarGoogle Scholar |

Hays, G. C., Bradshaw, C., James, M., Lovell, P., and Sims, D. (2007). Why do Argos satellite tags deployed on marine animals stop transmitting? Journal of Experimental Marine Biology and Ecology 349, 52–60.
Why do Argos satellite tags deployed on marine animals stop transmitting?Crossref | GoogleScholarGoogle Scholar |

Hearn, A. R., Green, J., Román, M. H., Acuña-Marrero, D., Espinoza, E., and Klimley, A. (2016). Adult female whale sharks make long-distance movements past Darwin Island (Galapagos, Ecuador) in the Eastern Tropical Pacific. Marine Biology 163, 214.
Adult female whale sharks make long-distance movements past Darwin Island (Galapagos, Ecuador) in the Eastern Tropical Pacific.Crossref | GoogleScholarGoogle Scholar |

Heyman, W. D., Graham, R. T., Kjerfve, B., and Johannes, R. E. (2001). Whale sharks Rhincodon typus aggregate to feed on fish spawn in Belize. Marine Ecology Progress Series 215, 275–282.
Whale sharks Rhincodon typus aggregate to feed on fish spawn in Belize.Crossref | GoogleScholarGoogle Scholar |

Himawan, M. R., Tania, C., Noor, B. A., Wijonarno, A., Subhan, B., and Madduppa, H. (2015). Sex and size range composition of whale shark (Rhincodon typus) and their sighting behaviour in relation with fishermen lift-net within Cenderawasih Bay National Park, Indonesia. Aquaculture, Aquarium Conservation and Legislation Bioflux 8, 123–133.

Hoffmayer, E. R., Franks, J. S., and Shelley, J. P. (2005). Recent observations of the whale shark (Rhincodon typus) in the northcentral Gulf of Mexico. Gulf and Caribbean Research 17, 117–120.
Recent observations of the whale shark (Rhincodon typus) in the northcentral Gulf of Mexico.Crossref | GoogleScholarGoogle Scholar |

Hsu, H., Joung, S., Liao, Y., and Liu, K. (2007). Satellite tracking of juvenile whale sharks, Rhincodon typus, in the northwestern Pacific. Fisheries Research 84, 25–31.
Satellite tracking of juvenile whale sharks, Rhincodon typus, in the northwestern Pacific.Crossref | GoogleScholarGoogle Scholar |

Hueter, R. E., Tyminski, J. P., and de la Parra, R. (2013). Horizontal movements, migration patterns, and population structure of whale sharks in the Gulf of Mexico and northwestern Caribbean Sea. PLoS One 8, e71883.
Horizontal movements, migration patterns, and population structure of whale sharks in the Gulf of Mexico and northwestern Caribbean Sea.Crossref | GoogleScholarGoogle Scholar | 23991000PubMed |

Jonsen, I. (2016). Joint estimation over multiple individuals improves behavioural state inference from animal movement data. Scientific Reports 6, 20625.
Joint estimation over multiple individuals improves behavioural state inference from animal movement data.Crossref | GoogleScholarGoogle Scholar | 26853261PubMed |

Jonsen, I. D., Flemming, J. M., and Myers, R. A. (2005). Robust state–space modeling of animal movement data. Ecology 86, 2874–2880.
Robust state–space modeling of animal movement data.Crossref | GoogleScholarGoogle Scholar |

Jonsen, I. D., Myers, R. A., and James, M. C. (2007). Identifying leatherback turtle foraging behaviour from satellite telemetry using a switching state-space model. Marine Ecology Progress Series 337, 255–264.
Identifying leatherback turtle foraging behaviour from satellite telemetry using a switching state-space model.Crossref | GoogleScholarGoogle Scholar |

Le Fevre, J. (1987). Aspects of the biology of frontal systems. Advances in Marine Biology 23, 163–299.
Aspects of the biology of frontal systems.Crossref | GoogleScholarGoogle Scholar |

Maguire, J. (2006). The state of world highly migratory, straddling and other high seas fishery resources and associated species. In ‘The State of World Highly Migratory, Straddling and Other High Seas Fishery Resources and Associated Species’. (Food and Agriculture Organization: Rome.)

Mangubhai, S., Erdmann, M. V., Wilson, J. R., Huffard, C. L., Ballamu, F., Hidayat, N. I., Hitipeuw, C., Lazuardi, M. E., Pada, D., and Purba, G. (2012). Papuan Bird’s Head Seascape: emerging threats and challenges in the global center of marine biodiversity. Marine Pollution Bulletin 64, 2279–2295.
Papuan Bird’s Head Seascape: emerging threats and challenges in the global center of marine biodiversity.Crossref | GoogleScholarGoogle Scholar | 22863353PubMed |

McCoy, E., Burce, R., David, D., Aca, E. Q., Hardy, J., Labaja, J., Snow, S. J., Ponzo, A., and Araujo, G. (2018). Long-term photo-identification reveals the population dynamics and strong site fidelity of adult whale sharks to the coastal waters of Donsol, Philippines. Frontiers in Marine Science 5, 271.
Long-term photo-identification reveals the population dynamics and strong site fidelity of adult whale sharks to the coastal waters of Donsol, Philippines.Crossref | GoogleScholarGoogle Scholar |

McKinney, J. A., Hoffmayer, E. R., Holmberg, J., Graham, R. T., Driggers, W. B., de la Parra-Venegas, R., Galván-Pastoriza, B. E., Fox, S., Pierce, S. J., and Dove, A. D. (2017). Long-term assessment of whale shark population demography and connectivity using photo-identification in the Western Atlantic Ocean. PLoS One 12, e0180495.
Long-term assessment of whale shark population demography and connectivity using photo-identification in the Western Atlantic Ocean.Crossref | GoogleScholarGoogle Scholar | 28817569PubMed |

Meyers, M. M. (2017). Demography and movement ecology of whale sharks in Cenderawasih Bay, Indonesia. M.Sc. Thesis, University of Auckland.

Moore, C., Mills, M., Arrigo, K., Berman-Frank, I., Bopp, L., Boyd, P., Galbraith, E., Geider, R. J., Guieu, C., and Jaccard, S. (2013). Processes and patterns of oceanic nutrient limitation. Nature Geoscience 6, 701–710.
Processes and patterns of oceanic nutrient limitation.Crossref | GoogleScholarGoogle Scholar |

Morales, J. M., Haydon, D. T., Frair, J., Holsinger, K. E., and Fryxell, J. M. (2004). Extracting more out of relocation data: building movement models as mixtures of random walks. Ecology 85, 2436–2445.
Extracting more out of relocation data: building movement models as mixtures of random walks.Crossref | GoogleScholarGoogle Scholar |

Motta, P. J., Maslanka, M., Hueter, R. E., Davis, R. L., De la Parra, R., Mulvany, S. L., Habegger, M. L., Strother, J. A., Mara, K. R., and Gardiner, J. M. (2010). Feeding anatomy, filter-feeding rate, and diet of whale sharks Rhincodon typus during surface ram filter feeding off the Yucatan Peninsula, Mexico. Zoology 113, 199–212.
Feeding anatomy, filter-feeding rate, and diet of whale sharks Rhincodon typus during surface ram filter feeding off the Yucatan Peninsula, Mexico.Crossref | GoogleScholarGoogle Scholar | 20817493PubMed |

Muchtar, M. (2004). IndoTROPICS studies on the plume of the Mamberamo River into the Bismarck Sea, West Papua, Indonesia. Continental Shelf Research 24, 2521–2533.
IndoTROPICS studies on the plume of the Mamberamo River into the Bismarck Sea, West Papua, Indonesia.Crossref | GoogleScholarGoogle Scholar |

Norman, B. M. (1999). Aspects of the biology and ecotourism industry of the whale shark Rhincodon typus in north-western Australia. M.Res. Thesis, Murdoch University, Perth.

Norman, B. M., and Stevens, J. D. (2007). Size and maturity status of the whale shark (Rhincodon typus) at Ningaloo Reef in Western Australia. Fisheries Research 84, 81–86.
Size and maturity status of the whale shark (Rhincodon typus) at Ningaloo Reef in Western Australia.Crossref | GoogleScholarGoogle Scholar |

Norman, B. M., Holmberg, J. A., Arzoumanian, Z., Reynolds, S. D., Wilson, R. P., Rob, D., Pierce, S. J., Gleiss, A. C., de la Parra, R., and Galvan, B. (2017). Undersea constellations: the global biology of an endangered marine megavertebrate further informed through citizen science. Bioscience 67, 1029–1043.
Undersea constellations: the global biology of an endangered marine megavertebrate further informed through citizen science.Crossref | GoogleScholarGoogle Scholar |

Olson, D., Hitchcock, G., Mariano, A., Ashjian, C., Peng, G., Nero, R., and Podesta, G. (1994). Life on the edge: marine life and fronts. Oceanography 7, 52–60.
Life on the edge: marine life and fronts.Crossref | GoogleScholarGoogle Scholar |

Prebble, C. E., Rohner, C. A., Pierce, S. J., Robinson, D. P., Jaidah, M. Y., Bach, S. S., and Trueman, C. N. (2018). Limited latitudinal ranging of juvenile whale sharks in the Western Indian Ocean suggests the existence of regional management units. Marine Ecology Progress Series 601, 167–183.
Limited latitudinal ranging of juvenile whale sharks in the Western Indian Ocean suggests the existence of regional management units.Crossref | GoogleScholarGoogle Scholar |

Prentice, M. L., and Hope, G. S. (2007). Climate of Papua. The Ecology of Papua 1, 479–494.

R Core Development (2016). R: A language and environment for statistical computing. R foundation for statistical computing, Vienna, Austria. Available at: https://www.r-project.org/ (accessed 18 June 2020)

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 Biology 8, 145–150.
Analysis of whale shark Rhincodon typus aggregations near South Ari Atoll, Maldives archipelago.Crossref | GoogleScholarGoogle Scholar |

Rohner, C., Pierce, S., Marshall, A., Weeks, S., Bennett, M., and Richardson, A. (2013a). Trends in sightings and environmental influences on a coastal aggregation of manta rays and whale sharks. Marine Ecology Progress Series 482, 153–168.
Trends in sightings and environmental influences on a coastal aggregation of manta rays and whale sharks.Crossref | GoogleScholarGoogle Scholar |

Rohner, C. A., Couturier, L. I., Richardson, A. J., Pierce, S. J., Prebble, C. E., Gibbons, M. J., and Nichols, P. D. (2013b). Diet of whale sharks Rhincodon typus inferred from stomach content and signature fatty acid analyses. Marine Ecology Progress Series 493, 219–235.
Diet of whale sharks Rhincodon typus inferred from stomach content and signature fatty acid analyses.Crossref | GoogleScholarGoogle Scholar |

Rohner, C. A., Armstrong, A. J., Pierce, S. J., Prebble, C. E., Cagua, E. F., Cochran, J. E., Berumen, M. L., and Richardson, A. J. (2015). Whale sharks target dense prey patches of sergestid shrimp off Tanzania. Journal of Plankton Research 37, 352–362.
Whale sharks target dense prey patches of sergestid shrimp off Tanzania.Crossref | GoogleScholarGoogle Scholar | 25814777PubMed |

Rohner, C. A., Richardson, A. J., Jaine, F. R., Bennett, M. B., Weeks, S. J., Cliff, G., Robinson, D. P., Reeve-Arnold, K. E., and Pierce, S. J. (2018). Satellite tagging highlights the importance of productive Mozambican coastal waters to the ecology and conservation of whale sharks. PeerJ 6, e4161.
Satellite tagging highlights the importance of productive Mozambican coastal waters to the ecology and conservation of whale sharks.Crossref | GoogleScholarGoogle Scholar | 30065862PubMed |

Rowat, D., and Brooks, K. (2012). A review of the biology, fisheries and conservation of the whale shark Rhincodon typus. Journal of Fish Biology 80, 1019–1056.
A review of the biology, fisheries and conservation of the whale shark Rhincodon typus.Crossref | GoogleScholarGoogle Scholar | 22497372PubMed |

Rowat, D., and Gore, M. (2007). Regional scale horizontal and local scale vertical movements of whale sharks in the Indian Ocean off Seychelles. Fisheries Research 84, 32–40.
Regional scale horizontal and local scale vertical movements of whale sharks in the Indian Ocean off Seychelles.Crossref | GoogleScholarGoogle Scholar |

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 Fishes 80, 465–472.
Aggregations of juvenile whale sharks (Rhincodon typus) in the Gulf of Tadjoura, Djibouti.Crossref | GoogleScholarGoogle Scholar |

Sale, A., Luschi, P., Mencacci, R., Lambardi, P., Hughes, G. R., Hays, G. C., Benvenuti, S., and Papi, F. (2006). Long-term monitoring of leatherback turtle diving behaviour during oceanic movements. Journal of Experimental Marine Biology and Ecology 328, 197–210.
Long-term monitoring of leatherback turtle diving behaviour during oceanic movements.Crossref | GoogleScholarGoogle Scholar |

Schmidt, J., Chien-Chi, C., Sheikh, S., Meekan, M., Norman, B., and Joung, S. (2010). Paternity analysis in a litter of whale shark embryos. Endangered Species Research 12, 117–124.
Paternity analysis in a litter of whale shark embryos.Crossref | GoogleScholarGoogle Scholar |

Sequeira, A. M. M., Mellin, C., Rowat, D., Meekan, M. G., and Bradshaw, C. J. (2012). Ocean-scale prediction of whale shark distribution. Diversity and Distributions 18, 504–518.
Ocean-scale prediction of whale shark distribution.Crossref | GoogleScholarGoogle Scholar |

Sequeira, A. M. M., Mellin, C., Meekan, M., Sims, D., and Bradshaw, C. (2013). Inferred global connectivity of whale shark Rhincodon typus populations. Journal of Fish Biology 82, 367–389.
Inferred global connectivity of whale shark Rhincodon typus populations.Crossref | GoogleScholarGoogle Scholar |

Sims, D. W. (2008). Sieving a living: a review of the biology, ecology and conservation status of the plankton‐feeding basking shark Cetorhinus maximus. Advances in Marine Biology 54, 171–220.
Sieving a living: a review of the biology, ecology and conservation status of the plankton‐feeding basking shark Cetorhinus maximus.Crossref | GoogleScholarGoogle Scholar | 18929065PubMed |

Sims, D. W., and Quayle, V. A. (1998). Selective foraging behaviour of basking sharks on zooplankton in a small-scale front. Nature 393, 460–464.
Selective foraging behaviour of basking sharks on zooplankton in a small-scale front.Crossref | GoogleScholarGoogle Scholar |

Sims, D. W., Southall, E. J., Richardson, A. J., Reid, P. C., and Metcalfe, J. D. (2003). Seasonal movements and behaviour of basking sharks from archival tagging: no evidence of winter hibernation. Marine Ecology Progress Series 248, 187–196.
Seasonal movements and behaviour of basking sharks from archival tagging: no evidence of winter hibernation.Crossref | GoogleScholarGoogle Scholar |

Sleeman, J. C., Meekan, M. G., Wilson, S. G., Polovina, J. J., Stevens, J. D., Boggs, G. S., and Bradshaw, C. J. (2010). To go or not to go with the flow: environmental influences on whale shark movement patterns. Journal of Experimental Marine Biology and Ecology 390, 84–98.
To go or not to go with the flow: environmental influences on whale shark movement patterns.Crossref | GoogleScholarGoogle Scholar |

Stevens, J. (2007). Whale shark (Rhincodon typus) biology and ecology: a review of the primary literature. Fisheries Research 84, 4–9.
Whale shark (Rhincodon typus) biology and ecology: a review of the primary literature.Crossref | GoogleScholarGoogle Scholar |

Stewart, B. S. (2011). Workshop and Monitoring Training for Whale Sharks in Cendrawasih Bay National Park, West Papua 2–7 May 2011, Nabire, Papua. Hubbs-SeaWorld Research Institute Technical Report 2011-375: 1–27.

Stewart, B. S., and Wilson, S. G. (2005). Threatened fishes of the world: Rhincodon typus (Smith 1828) (Rhincodontidae). Environmental Biology of Fishes 74, 184–185.
Threatened fishes of the world: Rhincodon typus (Smith 1828) (Rhincodontidae).Crossref | GoogleScholarGoogle Scholar |

Tania, C., Blæsbjerg, M., Himawan, M. R., Noor, B. A., Suruan, S. S., Pranata, B., and Stewart, B. S. (2016). Characteristics of whale sharks (Rhincodon typus) in Teluk Cenderawasih National Park, Indonesia. Qscience Proceedings 62, .
Characteristics of whale sharks (Rhincodon typus) in Teluk Cenderawasih National Park, Indonesia.Crossref | GoogleScholarGoogle Scholar |

Taylor, G. (1994). ‘Whale Sharks: The Giants of Ningaloo Reef.’ (Angus and Robertson: Sydney.)

Taylor, J. (1996). Seasonal occurrence, distribution and movements of the whale shark, Rhincodon typus, at Ningaloo Reef, Western Australia. Marine and Freshwater Research 47, 637–642.
Seasonal occurrence, distribution and movements of the whale shark, Rhincodon typus, at Ningaloo Reef, Western Australia.Crossref | GoogleScholarGoogle Scholar |

Taylor, J., and Pearce, A. (1999). Ningaloo Reef currents: implications for coral spawn dispersal, zooplankton and whale shark abundance. Journal of the Royal Society of Western Australia 82, 57–65.

Thomson, J. A., Araujo, G., Labaja, J., McCoy, E., Murray, R., and Ponzo, A. (2017). Feeding the world’s largest fish: highly variable whale shark residency patterns at a provisioning site in the Philippines. Royal Society Open Science 4, 170394.
Feeding the world’s largest fish: highly variable whale shark residency patterns at a provisioning site in the Philippines.Crossref | GoogleScholarGoogle Scholar | 28989750PubMed |

Turchin, P. (1991). Translating foraging movements in heterogeneous environments into the spatial distribution of foragers. Ecology 72, 1253–1266.
Translating foraging movements in heterogeneous environments into the spatial distribution of foragers.Crossref | GoogleScholarGoogle Scholar |

Wijeratne, S., Pattiaratchi, C., and Proctor, R. (2018). Estimates of surface and subsurface boundary current transport around Australia. Journal of Geophysical Research 123, 3444–3466.

Wilson, S., Polovina, J., Stewart, B., and Meekan, M. (2006). Movements of whale sharks (Rhincodon typus) tagged at Ningaloo Reef, Western Australia. Marine Biology 148, 1157–1166.
Movements of whale sharks (Rhincodon typus) tagged at Ningaloo Reef, Western Australia.Crossref | GoogleScholarGoogle Scholar |