Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Seasonal patterns of adult and subadult white shark presence at coastal aggregation sites in central California

Samantha Andrzejaczek https://orcid.org/0000-0002-9929-7312 A * , Taylor K. Chapple B , Alexandra E. DiGiacomo https://orcid.org/0000-0002-6375-6070 A , Salvador J. Jorgensen C , Scot D. Anderson D , Michael Castleton A , Paul E. Kanive D E , Theodore E. J. Reimer A , Timothy D. White A and Barbara A. Block A
+ Author Affiliations
- Author Affiliations

A Hopkins Marine Station, Stanford University, Pacific Grove, CA, USA.

B Coastal Oregon Marine Experiment Station, Oregon State University, Newport, OR, USA.

C California State University Monterey Bay, Seaside, CA, USA.

D California White Shark Project, Inverness, CA, USA.

E Montana State University, Bozeman, MT, USA.

* Correspondence to: sandrzejaczek@gmail.com

Handling Editor: Chris Lowe

Wildlife Research 52, WR24136 https://doi.org/10.1071/WR24136
Submitted: 24 August 2024  Accepted: 11 April 2025  Published: 6 May 2025

© 2025 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

Gaining insights into seasonal aggregations of marine megafauna and how patterns vary among demographic groups is pivotal for evaluating anthropogenic risk exposure and modeling populations and ecosystem dynamics. In California, adult and subadult white sharks recurrently aggregate on the coast near pinniped colonies in fall and winter months, facilitating comprehensive long-term field studies.

Aims

In this study, we used over 15 years of passive acoustic telemetry data to compare the seasonal dynamics of coastal habitat use for white sharks tagged in central California among four demographic groups (adult females, adult males, subadult females, and subadult males).

Methods

Acoustic tags were deployed on 355 white sharks at coastal aggregation sites and monitored across a coastal array of underwater receivers from 2006 to 2022. The main aggregation sites of the Northeast Pacific (Año Nuevo, the Farallon Islands and Tomales) were continuously monitored, with an expansion of the acoustic network to the south in the latter years of the study.

Key results

White sharks were tracked for an average duration of 594 ± 552 days (mean ± s.d.), with total track durations ranging up to 3235 days. Notably, adult male sharks exhibited the highest residency to central California coastal aggregation sites and demonstrated earlier seasonal peak densities in late October. Adult female presence peaked in early December. Adult sharks displayed distinct seasonal gaps in detection where they have been shown with satellite tags to migrate offshore, with females displaying much longer average detection gaps than for males (averaging 1.5 years vs 0.7 years). In contrast, subadults exhibited higher coastal affinity with more consistent and widespread detections across a higher number of coastal sites throughout the year, often extending beyond the main aggregation areas outside of the peak aggregation season.

Conclusions

We hypothesize that the observed differences between demographic groups are attributed to sex- and size-specific foraging and reproductive strategies. The extended receiver network also showed expansive coastal movements and identified potential undescribed aggregation sites.

Implications

Insights from our extensive acoustic dataset represent a significant advancement in assessing the timing of anthropogenic interactions and modeling both ecosystem and population dynamics.

Keywords: acoustic telemetry, Carcharodon carcharias, elasmobranch, habitat use, movement ecology, sexual segregation, site fidelity, size segregation.

References

Anderson SD, Pyle P (2003) A temporal, sex-specific occurrence pattern among white sharks at the South Farallon Islands, California. California Fish and Game, USA.

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. Marine Biology 158(6), 1233-1237.
| Crossref | Google Scholar | PubMed |

Andrzejaczek S, Meeuwig J, Rowat D, Pierce S, Davies T, Fisher R, Meekan M (2016) The ecological connectivity of whale shark aggregations in the Indian Ocean: a photo-identification approach. Royal Society Open Science 3(11), 160455.
| Crossref | Google Scholar |

Andrzejaczek S, Chapple TK, Jorgensen SJ, Anderson SD, Castleton M, Kanive PE, White TD, Block BA (2022) Multi-decadal high-resolution data reveal the cryptic vertical movement patterns of a large marine predator along the Californian Coast. Frontiers in Marine Science 9, 835576.
| Crossref | Google Scholar |

Appert C, Udyawer V, Simpfendorfer CA, Heupel MR, Scott M, Currey-Randall LM, Harborne AR, Jaine F, Chin A (2023) Use, misuse, and ambiguity of indices of residence in acoustic telemetry studies. Marine Ecology Progress Series 714, 27-44.
| Crossref | Google Scholar |

Arnoldi NS, Carlisle AB, Andrzejaczek S, Castleton MR, Micheli F, Schallert RJ, White TD, Block BA (2024) Salmon shark seasonal site fidelity demonstrates the influence of scale on identifying potential high-use areas and vulnerabilities. Marine Ecology Progress Series 735, 125-140.
| Crossref | Google Scholar |

Becerril-García EE, Martínez-Rincón RO, Galván-Magaña F, Santana-Morales O, Hoyos-Padilla EM (2020) Statistical modelling reveals spatial, temporal, and environmental preferences for white sharks at an oceanic aggregation site. Marine Ecology Progress Series 655, 171-183.
| Crossref | Google Scholar |

Block BA, Jonsen ID, Jorgensen SJ, Winship AJ, Shaffer SA, Bograd SJ, Hazen EL, Foley DG, Breed GA, Harrison A-L, Ganong JE, Swithenbank A, Castleton M, Dewar H, Mate BR, Shillinger GL, Schaefer KM, Benson SR, Weise MJ, Henry RW, Costa DP (2011) Tracking apex marine predator movements in a dynamic ocean. Nature 475(7354), 86-90.
| Crossref | Google Scholar | PubMed |

Boustany AM, Davis SF, Pyle P, Anderson SD, Le Boeuf BJ, Block BA (2002) Expanded niche for white sharks. Nature 415, 35-36.
| Crossref | Google Scholar | PubMed |

Brown JS, Laundré JW, Gurung M (1999) The ecology of fear: optimal foraging, game theory, and trophic interactions. Journal of Mammalogy 80(2), 385-399.
| Crossref | Google Scholar |

Bruce B, Bradford R (2015) Segregation or aggregation? Sex-specific patterns in the seasonal occurrence of white sharks Carcharodon carcharias at the Neptune Islands, South Australia. Journal of Fish Biology 87(6), 1355-1370.
| Crossref | Google Scholar | PubMed |

Butcher PA, Colefax AP, Gorkin RA, Kajiura SM, López NA, Mourier J, Purcell CR, Skomal GB, Tucker JP, Walsh AJ, Williamson JE, Raoult V (2021) The drone revolution of shark science: a review. Drones 5(1), 8.
| Crossref | Google Scholar |

Carlisle AB, Kim SL, Semmens BX, Madigan DJ, Jorgensen SJ, Perle CR, Anderson SD, Chapple TK, Kanive PE, Block BA (2012) Using Stable isotope analysis to understand the migration and trophic ecology of northeastern Pacific white sharks (Carcharodon carcharias). PLoS ONE 7(2), e30492.
| Crossref | Google Scholar |

Chambers JR Jr (1979) Population studies of California sea lions near Diablo Canyon, California. Master of Science in Biological Sciences, California Polytechnic State University, USA.

Chapple TK, Jorgensen SJ, Anderson SD, Kanive PE, Klimley AP, Botsford LW, Block BA (2011) A first estimate of white shark, Carcharodon carcharias, abundance off central California. Biology Letters 7(4), 581-583.
| Crossref | Google Scholar | PubMed |

Chapple TK, Chambert T, Kanive PE, Jorgensen SJ, Rotella JJ, Anderson SD, Carlisle AB, Block BA (2016) A novel application of multi-event modeling to estimate class segregation in a highly migratory oceanic vertebrate. Ecology 97(12), 3494-3502.
| Crossref | Google Scholar | PubMed |

Chapple TK, Cade DE, Goldbogen J, Massett N, Payne N, McInturf AG (2024) Behavioral response of megafauna to boat collision measured via animal-borne camera and IMU. Frontiers in Marine Science 11, 1430961.
| Crossref | Google Scholar |

Christiansen HM, Campana SE, Fisk AT, Cliff G, Wintner SP, Dudley SFJ, Kerr LA, Hussey NE (2016) Using bomb radiocarbon to estimate age and growth of the white shark, Carcharodon carcharias, from the southwestern Indian Ocean. Marine Biology 163(6), 144.
| Crossref | Google Scholar |

Ferretti F, Jorgensen S, Chapple TK, De Leo G, Micheli F (2015) Reconciling predator conservation with public safety. Frontiers in Ecology and the Environment 13(8), 412-417.
| Crossref | Google Scholar |

Francis MP (1996) Observations on a pregnant white shark with a review of reproductive biology. In ‘Great white sharks: the biology of Carcharodon carcharias’. (Eds AP Klimley, DG Ainley) pp. 157–172. (Academic Press: San Diego, CA, USA)

Francis MP, Duffy C, Lyon W (2015) Spatial and temporal habitat use by white sharks (Carcharodon carcharias) at an aggregation site in southern New Zealand. Marine and Freshwater Research 66(10), 900-918.
| Crossref | Google Scholar |

Goldman KJ, Anderson SD (1999) Space utilization and swimming depth of white sharks, Carcharodon carcharias, at the South Farallon Islands, Central California. Environmental Biology of Fishes 56(4), 351-364.
| Crossref | Google Scholar |

Goldman KJ, McCosker JE, Anderson SD, Klimley AP (1996) Temperature, swimming depth, and movements of a white shark at the South Farallon Islands, California. In ‘Great white sharks’. (Eds AP Klimley, AD Ainley) pp. 111–120. (Elsevier)

Harcourt R, Sequeira AMM, Zhang X, Roquet F, Komatsu K, Heupel M, McMahon C, Whoriskey F, Meekan M, Carroll G, Brodie S, Simpfendorfer C, Hindell M, Jonsen I, Costa DP, Block B, Muelbert M, Woodward B, Weise M, Aarestrup K, Biuw M, Boehme L, Bograd SJ, Cazau D, Charrassin J-B, Cooke SJ, Cowley P, de Bruyn PJN, Jeanniard du Dot T, Duarte C, Eguíluz VM, Ferreira LC, Fernández-Gracia J, Goetz K, Goto Y, Guinet C, Hammill M, Hays GC, Hazen EL, Hückstädt LA, Huveneers C, Iverson S, Jaaman SA, Kittiwattanawong K, Kovacs KM, Lydersen C, Moltmann T, Naruoka M, Phillips L, Picard B, Queiroz N, Reverdin G, Sato K, Sims DW, Thorstad EB, Thums M, Treasure AM, Trites AW, Williams GD, Yonehara Y, Fedak MA (2019) Animal-borne telemetry: an integral component of the ocean observing toolkit. Frontiers in Marine Science 6, 326.
| Crossref | Google Scholar |

Hazen EL, Abrahms B, Brodie S, Carroll G, Jacox MG, Savoca MS, Scales KL, Sydeman WJ, Bograd SJ (2019) Marine top predators as climate and ecosystem sentinels. Frontiers in Ecology and the Environment 17(10), 565-574.
| Crossref | Google Scholar |

Hazen EL, Savoca MS, Clark-Wolf TJ, Czapanskiy M, Rabinowitz PM, Abrahms B (2024) Ecosystem sentinels as early-warning indicators in the anthropocene. Annual Review of Environment and Resources 49, 573-598.
| Crossref | Google Scholar |

Hoenner X, Huveneers C, Steckenreuter A, Simpfendorfer C, Tattersall K, Jaine F, Atkins N, Babcock R, Brodie S, Burgess J, Campbell H, Heupel M, Pasquer B, Proctor R, Taylor MD, Udyawer V, Harcourt R (2018) Australia’s continental-scale acoustic tracking database and its automated quality control process. Scientific Data 5(1), 170206.
| Crossref | Google Scholar |

Hoyos-Padilla EM, Klimley AP, Galván-Magaña F, Antoniou A (2016) Contrasts in the movements and habitat use of juvenile and adult white sharks (Carcharodon carcharias) at Guadalupe Island, Mexico. Animal Biotelemetry 4(1), 14.
| Crossref | Google Scholar |

Hutchinson M, Scott M, Bauer R, Anderson J, Coffey DM, Holland K, Meyer C, O’Sullivan J, Royer M (2023) Habitat use and movement patterns of adult male and juvenile scalloped hammerhead sharks Sphyrna lewini throughout the Hawaiian archipelago. Endangered Species Research 52, 41-64.
| Crossref | Google Scholar |

Huveneers C, Apps K, Becerril-García EE, Bruce B, Butcher PA, Carlisle AB, Chapple TK, Christiansen HM, Cliff G, Curtis TH, Daly-Engel TS, Dewar H, Dicken ML, Domeier ML, Duffy CAJ, Ford R, Francis MP, French GCA, Galván-Magaña F, García-Rodríguez E, Gennari E, Graham B, Hayden B, Hoyos-Padilla EM, Hussey NE, Jewell OJD, Jorgensen SJ, Kock AA, Lowe CG, Lyons K, Meyer L, Oelofse G, Oñate-González EC, Oosthuizen H, O’Sullivan JB, Ramm K, Skomal G, Sloan S, Smale MJ, Sosa-Nishizaki O, Sperone E, Tamburin E, Towner AV, Wcisel MA, Weng KC, Werry JM (2018) Future research directions on the ‘Elusive’ white shark. Frontiers in Marine Science 5, 455.
| Crossref | Google Scholar |

Hyde CA, Notarbartolo di Sciara G, Sorrentino L, Boyd C, Finucci B, Fowler SL, Kyne PM, Leurs G, Simpfendorfer CA, Tetley MJ, Womersley F, Jabado RW (2022) Putting sharks on the map: a global standard for improving shark area-based conservation. Frontiers in Marine Science 9, 968853.
| Crossref | Google Scholar |

Jorgensen SJ, Reeb CA, Chapple TK, Anderson S, Perle C, Van Sommeran SR, Fritz-Cope C, Brown AC, Klimley AP, Block BA (2009) Philopatry and migration of Pacific white sharks. Proceedings of the Royal Society B: Biological Sciences 277, 679-688.
| Crossref | Google Scholar |

Jorgensen SJ, Arnoldi NS, Estess EE, Chapple TK, Rückert M, Anderson SD, Block BA (2012) Eating or meeting? Cluster analysis reveals intricacies of white shark (Carcharodon carcharias) migration and offshore behavior. PLoS ONE 7(10), e47819.
| Crossref | Google Scholar |

Jorgensen SJ, Anderson S, Ferretti F, Tietz JR, Chapple T, Kanive P, Bradley RW, Moxley JH, Block BA (2019) Killer whales redistribute white shark foraging pressure on seals. Scientific Reports 9(1), 6153.
| Crossref | Google Scholar |

Kanive PE, Rotella JJ, Jorgensen SJ, Chapple TK, Anderson SD, Klimley AP, Block BA (2015) Estimating apparent survival of sub-adult and adult white sharks (Carcharodon carcharias) in central California using mark–recapture methods. Frontiers in Marine Science 2, 19.
| Crossref | Google Scholar |

Kanive PE, Rotella JJ, Jorgensen SJ, Chapple TK, Hines JE, Anderson SD, Block BA (2019) Size-specific apparent survival rate estimates of white sharks using mark–recapture models. Canadian Journal of Fisheries and Aquatic Sciences 76(11), 2027-2034.
| Crossref | Google Scholar |

Kanive PE, Rotella JJ, Chapple TK, Anderson SD, White TD, Block BA, Jorgensen SJ (2021) Estimates of regional annual abundance and population growth rates of white sharks off central California. Biological Conservation 257, 109104.
| Crossref | Google Scholar |

Kanive PE, Rotella JJ, Chapple TK, Anderson SD, Hoyos-Padilla M, Klimley AP, Galván-Magaña F, Andrzejaczek S, Block BA, Jorgensen SJ (2023) Connectivity between white shark populations off Central California, USA and Guadalupe Island, Mexico. Frontiers in Marine Science 10, 1210969.
| Crossref | Google Scholar |

Klimley AP (1985) The areal distribution and autoecology of the white shark, Carcharodon carcharias, off the west coast of North America. Memoirs of the Southern California Academy of Sciences 9, 15-40.
| Google Scholar |

Klimley AP (1994) The predatory behavior of the white shark. American Scientist 82(2), 122-133.
| Google Scholar |

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(1), e55048.
| Crossref | Google Scholar |

Lohmann KJ, Lohmann CMF, Endres CS (2008) The sensory ecology of ocean navigation. Journal of Experimental Biology 211(11), 1719.
| Crossref | Google Scholar |

Long DJ, Hanni KD, Pyle P, Roletto J, Jones RE, Bandar R (1996) White shark predation on four pinniped species in central California waters: geographic and temporal patterns inferred from wounded carcasses. In ‘Great white sharks: the biology of Carcharodon carcharias’. (Eds AP Klimley, DG Ainley) pp. 263–274. (Academic Press: San Diego, CA, USA)

Martin RA, Rossmo DK, Hammerschlag N (2009) Hunting patterns and geographic profiling of white shark predation. Journal of Zoology 279(2), 111-118.
| Crossref | Google Scholar |

McInturf AG, Bowman J, Schulte JM, Newton KC, Vigil B, Honig M, Pelletier S, Cox N, Lester O, Cantor M, Chapple TK (2023) A unified paradigm for defining elasmobranch aggregations. ICES Journal of Marine Science 80, 1551-1566.
| Crossref | Google Scholar |

Moxley JH, Nicholson TE, Van Houtan KS, Jorgensen SJ (2019) Non-trophic impacts from white sharks complicate population recovery for sea otters. Ecology and Evolution 9(11), 6378-6388.
| Crossref | Google Scholar | PubMed |

Mull CG, Andrzejaczek S, Udyawer V, Dwyer RG (2022) Advances in methods, understanding, and applications of elasmobranch movement ecology. In ‘Biology of sharks and their relatives’. 3rd edn. (Eds JC Carrier, CA Simpfendorfer, MR Heithaus, KE Yopak) pp. 357–399. (CRC Press)

Natanson LJ, Skomal GB (2015) Age and growth of the white shark, Carcharodon carcharias, in the western North Atlantic Ocean. Marine and Freshwater Research 66(5), 387-398.
| Crossref | Google Scholar |

National Marine Sanctuaries (2024) Greater Farallones National Marine Sanctuary condition report: 2010–2022. US Department of Commerce, National Oceanic and Atmospheric Administration, National Ocean Service.

Niella Y, Meyer L, Clarke TM, Dennis JD, Pederson H, Huveneers C (2024) Effects of wildlife tourism on white shark associative behaviour. Animal Behaviour 215, 227-239.
| Crossref | Google Scholar |

Nosal AP, Cartamil DP, Ammann AJ, Bellquist LF, Ben-Aderet NJ, Blincow KM, Burns ES, Chapman ED, Freedman RM, Klimley AP, Logan RK, Lowe CG, Semmens BX, White CF, Hastings PA (2021) Triennial migration and philopatry in the critically endangered soupfin shark Galeorhinus galeus. Journal of Applied Ecology 58(8), 1570-1582.
| Crossref | Google Scholar |

Palacios MD, Stewart JD, Croll DA, Cronin MR, Trejo-Ramírez A, Stevens GMW, Lezama-Ochoa N, Zilliacus KM, González−Armas R, Notarbartolo di Sciara G, Galván−Magaña F (2023) Manta and devil ray aggregations: conservation challenges and developments in the field. Frontiers in Marine Science 10, 1148234.
| Crossref | Google Scholar |

Pratt HL (1996) Reproduction in the male white shark. In ‘Great white sharks: the biology of Carcharodon carcharias’. (Eds AP Klimley, DG Ainley) pp. 131–138. (Academic Press: San Diego, CA, USA)

Pratt HL, Jr., Pratt TC, Knotek RJ, Carrier JC, Whitney NM (2022) Long-term use of a shark breeding ground: three decades of mating site fidelity in the nurse shark, Ginglymostoma cirratum. PLoS ONE 17(10), e0275323.
| Crossref | Google Scholar |

R Core Team (2020) R: a language and environment for statistical computing. Vienna, Austria. Available at http://www.R-project.org/

Schilds A, Mourier J, Huveneers C, Nazimi L, Fox A, Leu ST (2019) Evidence for non-random co-occurrences in a white shark aggregation. Behavioral Ecology and Sociobiology 73(10), 138.
| Crossref | Google Scholar |

Sims DW (2005) Differences in habitat selection and reproductive strategies of male and female sharks. In ‘Sexual segregation in vertebrates: ecology of the two sexes’. (Eds KE Ruckstuhl, P Neuhaus) pp. 127–147. (Cambridge University Press: Cambridge)

Spaet JLY, Manica A, Brand CP, Gallen C, Butcher PA (2020) Environmental conditions are poor predictors of immature white shark Carcharodon carcharias occurrences on coastal beaches of eastern Australia. Marine Ecology Progress Series 653, 167-179.
| Crossref | Google Scholar |

Sulikowski JA, Williams LJ, Domeier ML (2012) The use of a nonlethal technique to assess the reproductive biology of the white shark, Carcharodon carcharias. In ‘Global perspectives on the biology and life history of the white shark’. (Ed. ML Domeier) pp. 467–476. (CRC Press)

Tanaka KR, Van Houtan KS, Mailander E, Dias BS, Galginaitis C, O’Sullivan J, Lowe CG, Jorgensen SJ (2021) North Pacific warming shifts the juvenile range of a marine apex predator. Scientific Reports 11(1), 3373.
| Crossref | Google Scholar |

Udyawer V, Dwyer RG, Hoenner X, Babcock RC, Brodie S, Campbell HA, Harcourt RG, Huveneers C, Jaine FRA, Simpfendorfer CA, Taylor MD, Heupel MR (2018) A standardised framework for analysing animal detections from automated tracking arrays. Animal Biotelemetry 6(1), 17.
| Crossref | Google Scholar |

Wearmouth VJ, Sims DW (2008) Sexual segregation in marine fish, reptiles, birds and mammals: behaviour patterns, mechanisms and conservation implications. Advances in Marine Biology 54, 107-170.
| Crossref | Google Scholar | PubMed |

Weng KC, Boustany AM, Pyle P, Anderson SD, Brown A, Block BA (2007a) Migration and habitat of white sharks (Carcharodon carcharias) in the eastern Pacific Ocean. Marine Biology 152(4), 877-894.
| Crossref | Google Scholar |

Weng KC, O’Sullivan JB, Lowe CG, Winkler CE, Dewar H, Block BA (2007b) Movements, behavior and habitat preferences of juvenile white sharks Carcharodon carcharias in the eastern Pacific. Marine Ecology Progress Series 338, 211-224.
| Crossref | Google Scholar |

White CF, Lyons K, Jorgensen SJ, O’Sullivan J, Winkler C, Weng KC, Lowe CG (2019) Quantifying habitat selection and variability in habitat suitability for juvenile white sharks. PLoS ONE 14(5), e0214642.
| Crossref | Google Scholar |

Wilson SG, Jonsen ID, Schallert RJ, Ganong JE, Castleton MR, Spares AD, Boustany AM, Stokesbury MJW, Block BA (2015) Tracking the fidelity of Atlantic bluefin tuna released in Canadian waters to the Gulf of Mexico spawning grounds. Canadian Journal of Fisheries and Aquatic Sciences 72(11), 1700-1717.
| Crossref | Google Scholar |

Winton MV, Fay G, Skomal GB (2023) An open spatial capture–recapture framework for estimating the abundance and seasonal dynamics of white sharks at aggregation sites. Marine Ecology Progress Series 715, 1-25.
| Crossref | Google Scholar |

Womersley FC, Humphries NE, Queiroz N, Vedor M, da Costa I, Furtado M, Tyminski JP, Abrantes K, Araujo G, Bach SS, Barnett A, Berumen ML, Bessudo Lion S, Braun CD, Clingham E, Cochran JEM, de la Parra R, Diamant S, Dove ADM, Dudgeon CL, Erdmann MV, Espinoza E, Fitzpatrick R, Cano JG, Green JR, Guzman HM, Hardenstine R, Hasan A, Hazin FHV, Hearn AR, Hueter RE, Jaidah MY, Labaja J, Ladino F, Macena BCL, Morris JJ, Norman BM, Peñaherrera-Palma C, Pierce SJ, Quintero LM, Ramírez-Macías D, Reynolds SD, Richardson AJ, Robinson DP, Rohner CA, Rowat DRL, Sheaves M, Shivji MS, Sianipar AB, Skomal GB, Soler G, Syakurachman I, Thorrold SR, Webb DH, Wetherbee BM, White TD, Clavelle T, Kroodsma DA, Thums M, Ferreira LC, Meekan MG, Arrowsmith LM, Lester EK, Meyers MM, Peel LR, Sequeira AMM, Eguíluz VM, Duarte CM, Sims DW (2022) Global collision-risk hotspots of marine traffic and the world’s largest fish, the whale shark. Proceedings of the National Academy of Sciences 119(20), e2117440119.
| Crossref | Google Scholar |

Zar JH (1998) ‘Biostatistical analysis.’ 4th edn. (Prentice-Hall: NJ, USA)