in

Nutrient availability drives local seasonal movements of an endangered marine megafauna species


Abstract

Understanding drivers of animal movement is key to predicting species distributions and guiding conservation. Whale sharks (Rhincodon typus) are an endangered species known for broad seasonal migrations across oceans influenced by factors like temperature and prey availability. However, finer-scale local movements within aggregations are less studied. We analysed four years (2016–2019) of sightings data from a year-round aggregation in South Ari Marine Protected Area (SAMPA), the Maldives. Using MODIS-Aqua remote sensing data, we examined seasonal patterns in chlorophyll-a (Chl-a) and sea surface temperature (SST). Generalised additive mixed models (GAMMs) revealed significant seasonal rhythms in SST across SAMPA, and significant seasonal Chl-a variation in the south but not the east of the MPA. In a separate GAMM, we found that seasonal rhythms in shark sightings were significant throughout the MPA but more pronounced in the south than in the east. Chl-a was significantly associated with sightings, with both peaking in the south during the Northeast Monsoon (January–March). SST was not significantly associated with sightings. As Chl-a is tightly linked to the abundance of zooplankton, these findings suggest that whale shark movement within the year-round aggregation is driven by prey availability. The results could be used to inform dynamic management or predict aggregations elsewhere and responses to environmental change.

Data availability

All data generated or analysed during this study are included in this published article and Supplementary_Information_1.zip. Data and code are available at Zenodo.org: https://doi.org/10.5281/zenodo.15755924. MODIS-Aqua Level-2 ocean colour data used in this study are publicly accessible through NASA’s Ocean Biology Processing Group (OBPG) data portal and can be downloaded via NASA’s open data policy (https://science.nasa.gov/researchers/science-information-policy/).

References

  1. Leblond, M., Dussault, C. & Ouellet, J. What drives fine-scale movements of large herbivores? A case study using moose. Ecography 33, 1102–1112 (2010).

    Google Scholar 

  2. Avgar, T., Mosser, A., Brown, G. S. & Fryxell, J. M. Environmental and individual drivers of animal movement patterns across a wide geographical gradient. J. Anim. Ecol. 82, 96–106 (2013).

    Google Scholar 

  3. Allen, A. M. & Singh, N. J. Linking movement ecology with wildlife management and conservation. Front. Ecol. Evol. 3, 155. https://doi.org/10.3389/fevo.2015.00155 (2026).

    Google Scholar 

  4. Sequeira, A., Mellin, C., Rowat, D., Meekan, M. G. & Bradshaw, C. J. A. Ocean-scale prediction of Whale shark distribution. Divers. Distrib. 18, 504–518 (2012).

    Google Scholar 

  5. Sequeira, A. M. M., Mellin, C., Meekan, M. G., Sims, D. W. & Bradshaw, C. J. A. Inferred global connectivity of Whale shark Rhincodon typus populations. J. Fish. Biol. 82, 367–389 (2013).

    Google Scholar 

  6. Alerstam, T. & Bäckman, J. Ecology of animal migration. Curr. Biol. 28, 968–972 (2018).

    Google Scholar 

  7. Rizzo, L. Y. & Schulte, D. A review of humpback whales’ migration patterns worldwide and their consequences to gene flow. J. Mar. Biol. Ass. 89, 995–1002 (2009).

    Google Scholar 

  8. Weimerskirch, H. et al. Lifetime foraging patterns of the wandering albatross: life on the move! J. Exp. Mar. Biol. Ecol. 450, 68–78 (2014).

    Google Scholar 

  9. Gilmour, M. E. et al. Evaluation of MPA designs that protect highly mobile megafauna now and under climate change scenarios. Global Ecol. Conserv. 35, e02070. https://doi.org/10.1016/j.gecco.2022.e02070 (2022).

    Google Scholar 

  10. Cooke, S. J. et al. Animal migration in the anthropocene: threats and mitigation options. Biol. Rev. 99, 1242–1260 (2024).

    Google Scholar 

  11. Valsecchi, S. et al. Analysis of the Temporal and Spatial variability of Whale shark (Rhincodon typus) aggregation in the South Ari marine protected Area, Maldives, Indian ocean. Eur. Zool. J. 88, 684–697 (2021).

    Google Scholar 

  12. Harvey-Carroll, J., Carroll, D., Trivella, C. M. & Connelly, E. Classification of African ground Pangolin behaviour based on accelerometer readouts. Anim. Biotelem. 12, 22. https://doi.org/10.1186/s40317-024-00377-y (2024).

    Google Scholar 

  13. Cashion, T. et al. Shifting seas, shifting boundaries: dynamic marine protected area designs for a changing climate. PLoS One. 15, e0241771. https://doi.org/10.1371/journal.pone.0241771 (2020).

    Google Scholar 

  14. Ovando, D., Liu, O., Molina, R. & Szuwalski, C. Models of marine protected areas must explicitly address Spatial dynamics. Proc. Natl. Acad. Sci. USA. 118 (e2025958118). https://doi.org/10.1073/pnas.2025958118 (2021).

  15. Colman, J. G. A review of the biology and ecology of the Whale shark. J. Fish. Biol. 51, 1219–1234 (1997).

    Google Scholar 

  16. Rohner, C. A. et al. Diet of Whale sharks Rhincodon typus inferred from stomach content and signature fatty acid analyses. Mar. Ecol. Prog Ser. 493, 219–235 (2013).

    Google Scholar 

  17. Montero-Quintana, A. N., Ocampo-Valdez, C. F., Vázquez-Haikin, J. A., Sosa-Nishizaki, O. & Osorio-Beristain, M. Whale shark (Rhincodon typus) predatory flexible feeding behaviors on schooling fish. J. Ethol. 39, 399–410 (2021).

    Google Scholar 

  18. Harvey-Carroll, J. et al. The impact of injury on apparent survival of Whale sharks (Rhincodon typus) in South Ari Atoll marine protected Area, Maldives. Sci. Rep. 11, 937. https://doi.org/10.1038/s41598-020-79101-8 (2021).

    Google Scholar 

  19. 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).

    Google Scholar 

  20. Pierce, S. J. & Norman, B. Rhincodon typus. The IUCN Red List of Threatened Species. e-T19488A2 (2016).

  21. Reynolds, S. D. et al. Regional variation in anthropogenic threats to Indian ocean Whale sharks. Global Ecol. Conserv. 33, e01961. https://doi.org/10.1016/j.gecco.2021.e01961 (2022).

    Google Scholar 

  22. Womersley, F. C. et al. Identifying priority sites for Whale shark ship collision management globally. Sci. Total Environ. 934, 172776. https://doi.org/10.1016/j.scitotenv.2024.172776 (2024).

    Google Scholar 

  23. 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).

    Google Scholar 

  24. Araujo, G. et al. Improving sightings-derived residency Estimation for Whale shark aggregations: A novel metric applied to a global data set. Front. Mar. Sci. 9, 775691. https://doi.org/10.3389/fmars.2022.775691 (2022).

    Google Scholar 

  25. 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. https://doi.org/10.1371/journal.pone.0071883 (2013).

    Google Scholar 

  26. Robinson, D. P. et al. Some like it hot: repeat migration and residency of Whale sharks within an extreme natural environment. PLoS ONE. 12, 9. https://doi.org/10.1371/journal.pone.0185360 (2017).

    Google Scholar 

  27. Taylor, J. Seasonal occurrence, distribution and movements of the Whale shark, Rhincodon typus, at Ningaloo Reef, Western Australia. Mar. Freshw. Res. 47, 637 (1996).

    Google Scholar 

  28. 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).

    Google Scholar 

  29. Graham, R. T. Whale sharks of the Western caribbean: an overview of current research and conservation efforts and future needs for effective management of the species. Gulf Caribb. Res. 19, 149–159 (2007).

    Google Scholar 

  30. Hoffmayer, E. R., Franks, J. S., Driggers, W. B., Oswald, K. J. & Quattro, J. M. Observations of a feeding aggregation of Whale Sharks, rhincodon typus, in the North central Gulf of Mexico. Gulf Caribb. Res. 19, 69–73 (2007).

    Google Scholar 

  31. De La Venegas, P. An unprecedented aggregation of Whale sharks, Rhincodon typus, in Mexican coastal waters of the Caribbean sea. PLoS ONE. 6, e18994. https://doi.org/10.1371/journal.pone.0018994 (2011).

    Google Scholar 

  32. Macena, B. C. L. & Hazin, F. H. V. Whale shark (Rhincodon typus) seasonal occurrence, abundance and demographic structure in the Mid-Equatorial Atlantic ocean. PLoS ONE. 11, e0164440. https://doi.org/10.1371/journal.pone.0164440 (2016).

    Google Scholar 

  33. Sequeira, A. M. M., Mellin, C., Fordham, D. A., Meekan, M. G. & Bradshaw, C. J. A. Predicting current and future global distributions of Whale sharks. Global Change Biol. 20, 778–789 (2014).

    Google Scholar 

  34. Báez, J. C., Barbosa, A. M., Pascual, P., Ramos, M. L. & Abascal, F. Ensemble modeling of the potential distribution of the Whale shark in the Atlantic. Ocean. Ecol. Evol. 10, 175–184 (2020).

    Google Scholar 

  35. Jaramillo-Gil, S., Pardo, M. A., Vázquez‐Haikin, A. & Bolaños‐Jiménez, J. Sosa‐Nishizaki O. Whale shark abundance forecast: the interannual hotspot effect. J. Appl. Ecol. 60, 954–966 (2023).

    Google Scholar 

  36. Carroll, D. & Harvey-Carroll, J. The influence of light on elasmobranch behavior and physiology: a review. Front. Mar. Sci. 10, 1225067. https://doi.org/10.3389/fmars.2023.1225067 (2023).

    Google Scholar 

  37. Robinson, D. P. et al. Whale sharks, Rhincodon typus, aggregate around offshore platforms in Qatari waters of the Arabian Gulf to feed on fish spawn. PLoS ONE. 8, e58255. https://doi.org/10.1371/journal.pone.0058255 (2013).

    Google Scholar 

  38. Afonso, P., McGinty, N. & Machete, M. Dynamics of Whale shark occurrence at their fringe oceanic habitat. PLoS ONE. 9, e102060. https://doi.org/10.1371/journal.pone.0102060 (2014).

    Google Scholar 

  39. Cárdenas-Palomo, N., Herrera‐Silveira, J., Velázquez‐Abunader, I., Reyes, O. & Ordoñez, U. Distribution and feeding habitat characterization of Whale sharks Rhincodon typus in a protected area in the North Caribbean sea. J. Fish. Biol. 86, 668–686 (2015).

    Google Scholar 

  40. Herdianti, T. P. et al. Habitat assessment of Whale sharks (Rhincodon typus) in Saleh Bay, indonesia: linking chlorophyll-a and sea surface temperature using aqua modis data. Geogr. Tech. 22, 52–64 (2025).

    Google Scholar 

  41. Anderson, R. C. & Ahmed, H. The shark fisheries of the Maldives. Ministry of Fisheries and Agriculture, Republic of Maldives and Food and Agriculture Organization of the United Nations. (1993).

  42. 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. https://doi.org/10.7717/peerj.515 (2014).

    Google Scholar 

  43. Zimmerhackel, J. S., Kragt, M. E., Rogers, A. A., Ali, K. & Meekan, M. G. Evidence of increased economic benefits from shark-diving tourism in the Maldives. Mar. Policy. 100, 21–26 (2019).

    Google Scholar 

  44. Riley, M., Hale, M., Harman, A. & Rees, R. Analysis of Whale shark Rhincodon typus aggregations near South Ari Atoll, Maldives Archipelago. Aquat. Biol. 8, 145–150 (2010).

    Google Scholar 

  45. Cagua, E. F. et al. Acoustic telemetry reveals cryptic residency of Whale sharks. Biol. Lett. 11, 20150092. https://doi.org/10.1098/rsbl.2015.0092 (2015).

    Google Scholar 

  46. 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, 1487–1495 (2018).

    Google Scholar 

  47. Ministry of Climate Change, Environment, and Energy. SAMPA Management Plan ((IUL)438-ENV/438/2025/54). Presidents Office – Gazzette. (2025). Available from: https://www.gazette.gov.mv/iulaan/326014

  48. Cariño, M. C. M. R. et al. Planktonic foraminifera fluxes and their response to the Asian monsoon: insights from the Maldives, Indian ocean. Front. Earth Sci. 11, 1141263. https://doi.org/10.3389/feart.2023.1141263 (2023).

    Google Scholar 

  49. Dryden, C. et al. An ecological assessment of coral reefs in the South Ari Marine Protected Area. Malé, Maldives: IUCN and Government of Maldives.46pp (2020).

  50. Government of Maldives. Regulation No. 2024/R-96 on Whale Shark Conservation [Amendment To the Protected Species Regulation 2021/R-25] (Malé, 2024).

  51. UNEP-WCMC and IUCN. Protected Planet: [The World Database on Protected Areas (WDPA)]. Cambridge, UK: UNEP-WCMC and IUCN. (2025). Available from: https://www.protectedplanet.net/en

  52. OCHA Regional Office for Asia and the Pacific (ROAP). Maldives – Subnational Administrative Boundaries (COD-AB). Humanitarian Data Exchange. (2024). Available from: https://data.humdata.org/dataset/cod-ab-mdv

  53. QGIS Association. QGIS Geographic Information System. (2022). Available from: https://qgis.org/

  54. Huot, Y. et al. Does chlorophyll a provide the best index of phytoplankton biomass for primary productivity studies? Biogeosciences Discuss. 4, 707–745 (2007).

    Google Scholar 

  55. Druon, J. N. et al. Satellite-based indicator of zooplankton distribution for global monitoring. Sci. Rep. 9 https://doi.org/10.1038/s41598-019-41212-2 (2019).

  56. NASA Ocean Biology Processing Group. Aqua MODIS Level 2 Ocean Color Data, Version R2022.0. NASA Ocean Biology Distributed Active Archive Center. (2022). Available from: https://oceancolor.gsfc.nasa.gov/data/10.5067/AQUA/MODIS/L2/OC/2022

  57. R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2024).

  58. Wickham, H. et al. Welcome to the tidyverse. JOSS 4, 1686. https://doi.org/10.21105/joss.01686 (2019).

    Google Scholar 

  59. Wood, S. N. Generalized Additive Models: an Introduction with R 2nd Edition (Chapman and Hall/CRC, 2017).

  60. Pedersen, E. J., Miller, D. L. & Simpson, G. L. Ross N. Hierarchical generalized additive models in ecology: an introduction with Mgcv. PeerJ 7, e6876. https://doi.org/10.7717/peerj.6876 (2019).

    Google Scholar 

  61. Carroll, D. et al. Declining harbour seal abundance in a previously recovering meta-population. PLoS One. 20, e0326933. https://doi.org/10.1371/journal.pone.0326933 (2025).

    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. https://doi.org/10.7717/peerj.543 (2014).

    Google Scholar 

  63. Anderson, R. C., Adam, M. S. & Goes, J. I. From monsoons to mantas: seasonal distribution of Manta alfredi in the Maldives. Fish. Oceanogr. 20, 104–113 (2011).

    Google Scholar 

  64. Castelblanco-Martínez, D. N. et al. Detecting, counting and following the giants of the sea: a review of monitoring methods for aquatic megavertebrates in the Caribbean. Wildl. Res. 46, 545–556 (2019).

    Google Scholar 

  65. Cubaynes, H. C., Fretwell, P. T., Bamford, C., Gerrish, L. & Jackson, J. A. Whales from space: four Mysticete species described using new VHR satellite imagery. Mar. Mammal Sci. 35, 466–491 (2019).

    Google Scholar 

  66. Hancock, J. W. et al. The Big Fish Network: Using new technology to incentivise citizen science engagement in the Maldives. The 4th International Whale Shark Conference, Doha, Qatar. Hamad bin Khalifa University Press. (2016). Available from: https://www.qscience.com/content/papers/10.5339/qproc.iwsc4.21 (2016).

  67. Ali, K. & Sinan, H. National plan of action for the conservation and management of sharks in the Maldives. Ministry of Fisheries and Agriculture. (2015). Available from: https://faolex.fao.org/docs/pdf/mdv163507.pdf

  68. The President’s Office, Republic of Maldives. Whale sharks declared protected species under the Environmental Protection and Preservation Act. The President’s Office. (2023). Available from: https://presidency.gov.mv/Press/Article/28993

  69. Sequeira, A. M. M. et al. Overhauling ocean Spatial planning to improve marine megafauna conservation. Front. Mar. Sci. 6, 639. https://doi.org/10.3389/fmars.2019.00639 (2019).

    Google Scholar 

  70. Carroll, D., Infantes, E. & Pagan, E. V. Approaching a population-level assessment of body size in pinnipeds using drones, an early warning of environmental degradation. Remote Sens. Ecol. Conserv. 11, 156–171 (2024).

    Google Scholar 

  71. Infantes, E. et al. An automated work-flow for pinniped surveys: A new tool for monitoring population dynamics. Front. Ecol. Evol. 11, 905309. https://doi.org/10.3389/fevo.2022.905309 (2022).

    Google Scholar 

  72. Gayford, J. H., Pearse, W. D., De La Parra Venegas, R. & Whitehead, D. A. Quantifying the behavioural consequences of shark ecotourism. Sci. Rep. 13, 12938. https://doi.org/10.1038/s41598-023-39560-1 (2023).

    Google Scholar 

  73. Li, H., Li, X., Song, D., Nie, J. & Liang, S. Prediction on daily Spatial distribution of chlorophyll-a in coastal seas using a synthetic method of remote sensing, machine learning and numerical modeling. Sci. Total Environ. 910, 168642. https://doi.org/10.1016/j.scitotenv.2023.168642 (2024).

    Google Scholar 

  74. Motta, P. J. et al. 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 (2010).

    Google Scholar 

  75. Ketchum, J. T., Galván-Magaña, F. & Klimley, A. P. Segregation and foraging ecology of Whale sharks, Rhincodon typus, in the Southwestern Gulf of California. Environ. Biol. Fishes. 96, 779–795 (2013).

    Google Scholar 

  76. 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. https://doi.org/10.3389/fmars.2015.00064 (2015).

    Google Scholar 

  77. Whitehead, H. Analysis of animal movement using opportunistic individual identifications: application to sperm whales. Ecology 82, 1417–1432 (2001).

    Google Scholar 

Download references

Acknowledgements

The authors thank all MWSR (formally MWSRP) staff and volunteers for their role in collecting and compiling whale shark data. Whale shark sightings data were collected under the MWSRP Protected species research permit (A149508) from the Environmental Protection Agency and Ministry of Fisheries and Research permit (OTHR)30-D/PRIV/2019/1172 awarded by the Ministry of Fisheries, Marine Resources and Agriculture, Malé, Maldives.

Funding

Open access funding provided by University of Gothenburg. DC was funded by the Wild Animal Initiative Fellowship, grant number F-2023-00005.

Author information

Authors and Affiliations

Authors

Contributions

Conceptualisation: DC and JHC. Sightings data collection: IHZ and CCP. Data compilation and analysis: DC. Writing of original manuscript: DC, IHZ, and JHC. Editing of manuscript: All authors.

Corresponding author

Correspondence to
Daire Carroll.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary Information

Below is the link to the electronic supplementary material.

Supplementary Material 1

Rights and permissions

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

Reprints and permissions

About this article

Cite this article

Carroll, D., Zareer, I.H., Pérez, C.C. et al. Nutrient availability drives local seasonal movements of an endangered marine megafauna species.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-38138-x

Download citation

  • Received:

  • Accepted:

  • Published:

  • DOI: https://doi.org/10.1038/s41598-026-38138-x

Keywords

  • Seasonal rhythms
  • Remote sensing
  • Marine megafauna
  • Animal behaviour
  • Conservation
  • Marine protected areas


Source: Ecology - nature.com

Environmental and societal costs of maize production decrease by addressing the uncertainty in nitrogen rate recommendations

High-resolution forecasting of soil thermal regimes using different deep learning frameworks under climate change

Back to Top