Baited-Associated Aggregation of Spinner Sharks in Hulhumale, Maldives: Preliminary Observations and Photo-Identification Tools
Abstract
1. Introduction
2. Materials and Methods
2.1. Study Site
2.2. Spinner Shark Observations
2.3. Spinner Shark Photo-Identification Protocols
2.3.1. Standard Protocol
2.3.2. Preliminary Protocol
2.4. Statistical Analysis
3. Results
3.1. Photo-Identification
3.1.1. Standard Protocol
3.1.2. Preliminary Protocol
3.2. Laser Photogrammetry
4. Discussion
5. Conclusions
Author Contributions
Funding
Institutional Review Board Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
- Compagno, L.J.V. FAO Species Catalogue. Vol. 4. Sharks of the World. An Annotated and Illustrated Catalogue of Shark Species Known to Date. Part 2; Fisheries Synopsis No. 125; Food and Agriculture Organization of the United Nations: Rome, Italy, 1984. [Google Scholar]
- Burgess, G.H. Carcharhinus brevipinna. IUCN Red List. Threat. Species 2009, e.T39368A10182758. Available online: https://doi.org/10.2305/IUCN.UK.2009-2.RLTS.T39368A10182758.en (accessed on 7 January 2026).
- De Maddalena, A.; Baensch, H.; Heim, W. Sharks of the Mediterranean. An Illustrated Study of All Species; McFarland & Co.: Jefferson, NC, USA, 2015; p. 204. [Google Scholar]
- Castro, J.I. The Sharks of North America; Oxford University Press: New York, NY, USA, 2011; p. 613. [Google Scholar]
- Branstetter, S. Age and growth estimates for blacktip, Carcharhinus limbatus, and spinner, C. brevipinna, sharks from the northwestern Gulf of Mexico. Copeia 1987, 4, 964–974. [Google Scholar] [CrossRef]
- Allen, B.R.; Wintner, S.P. Age and growth of the spinner shark (Carcharhinus brevipinna) off the KwaZulu-Natal coast, South Africa. S. Afr. J. Mar. Sci. 2002, 24, 1–8. [Google Scholar] [CrossRef]
- Capapé, C.; Hemida, F.; Seck, A.A.; Diatta, Y.; Guélorget, O.; Zaouali, J. Distribution and reproductive biology of the spinner shark, Carcharhinus brevipinna (Müller and Henle, 1841) (Chondrichthyes: Carcharhinidae). Isr. J. Ecol. Evol. 2003, 49, 311–322. [Google Scholar] [CrossRef]
- Capapè, C.; Seck, A.A.; Diatta, Y.; Reynaud, C.; Hemida, F.; Zaouali, J. Reproductive biology of the blacktip shark, Carcharhinus limbatus (Chondrichthyes: Carcharhinidae) off West and North African coasts. Cybium 2004, 28, 275–284. [Google Scholar]
- Joung, S.J.; Liao, Y.Y.; Liu, K.M.; Chen, C.T.; Leu, L.C. Age, growth, and reproduction of the spinner shark, Carcharhinus brevipinna, in the northeastern waters of Taiwan. Zool. Stud. 2005, 44, 102–110. [Google Scholar]
- Carlson, J.K.; Baremore, I.E. Growth dynamics of the spinner shark (Carcharhinus brevipinna) off the United States southeast and Gulf of Mexico coasts: A comparison of methods. Fish. Bull. 2005, 103, 280–291. [Google Scholar]
- Aubrey, C.; Snelson, F.F., Jr. Early life history of the spinner shark (Carcharhinus brevipinna) in a Florida nursery. Fish. Bull. 2007, 105, 527–535. [Google Scholar]
- White, W.T. Catch composition and reproductive biology of whaler sharks (Carcharhiniformes: Carcharhinidae) caught by fisheries in Indonesia. J. Fish Biol. 2007, 71, 1512–1543. [Google Scholar] [CrossRef]
- Geraghty, P.T.; Williamson, J.E.; Macbeth, W.G.; Wintner, S.P.; Harry, A.V.; Ovenden, J.R.; Gillings, M.R. Population expansion and genetic structure in Carcharhinus brevipinna in the southern Indo-Pacific. PLoS ONE 2013, 8, e75169. [Google Scholar] [CrossRef]
- Geraghty, P.T.; Macbeth, W.G.; Williamson, J.E. Aspects of the reproductive biology of dusky, spinner and sandbar sharks (Family carcharhinidae) from the Tasman Sea. Mar. Freshw. Res. 2016, 67, 513–525. [Google Scholar] [CrossRef]
- Santos dos Santos, P.R.; Santoro, P.; de David Chelotti, L. Length–weight relationship of the spinner shark (Carcharhinus brevipinna) on the continental shelf of southern Brazil. Arq. Ciências Mar 2017, 50, 133–135. [Google Scholar]
- Palmrose, K. Reproductive Biology of the Spinner Shark, Carcharhinus brevipinna, off the Southeast U.S. Coast. Master’s Thesis, University of North Florida, Jacksonville, FL, USA, 2021. [Google Scholar]
- Linn, A.K.; Levengood, A.L.; Dudgeon, C.L.; Gustafson, J.A.; Smith, J.L.; Holmes, B.J. Assessment of multiple paternity in spinner shark (Carcharhinus brevipinna) litters from eastern Australian waters. Mar. Freshw. Res. 2025, 76, MF24222. [Google Scholar] [CrossRef]
- Vinyard, E.A.; Portnoy, D.S.; Driggers, W.B., III; Gelsleichter, J.; Hendon, J.M.; Higgs, J.M.; Natanson, L.J.; Frazier, B.S. Age and growth of spinner sharks (Carcharhinus brevipinna) in the western North Atlantic Ocean. Environ. Biol. Fish. 2025, 109, 17. [Google Scholar] [CrossRef]
- Allen, B.R.; Cliff, G. Sharks caught in the protective gill nets off KwaZulu-Natal, South Africa. 9. The spinner shark (Carcharhinus brevipinna) (Müller and Henle). S. Afr. J. Mar. Sci. 2000, 22, 163–179. [Google Scholar] [CrossRef]
- Sivasubramaniam, K. Pelagic sharks in the Indian Ocean. Bay Bengal News 1992, 47, 11–19. [Google Scholar]
- Bass, A.J.; d’Aubrey, J.D.; Kistnasamy, N. Sharks of the East Coast of Southern Africa. The Genus Carcharhinus (Carcharhinidae). I; Oceanographic Research Institute: KwaZulu-Natal, South Africa, 1973; Volume 33, pp. 1–168. [Google Scholar]
- Hussey, N.E.; Wintner, S.P.; Dudley, S.F.J.; Cliff, G.; Cocks, D.T.; MacNeil, M.A. Maternal investment and size-specific reproductive output in carcharhinid sharks. J. Anim. Ecol. 2009, 79, 184–193. [Google Scholar] [CrossRef]
- Rolim, F.A.; Rodrigues, P.F.C.; Gadig, O.B.F.; Motta, F.D.S.; Slerca, M.A.L.; Araujo, N.L.F.; Neves, L.M. Evidences of seasonal aggregations and breaching behaviour of blacktip (Carcharhinus limbatus) and spinner (C. brevipinna) sharks in a southwest Atlantic marine protected area. Aquat. Conserv. Mar. Freshw. Ecosyst. 2025, 35, e70153. [Google Scholar] [CrossRef]
- Sumpton, W.; Lane, B.; Ham, T. Characteristics of the biology and distribution of the spinner shark (Carcharhinus brevipinna) in Queensland, Australia based on data collected from the shark control program. Asian Fish. Sci. 2010, 23, 340–354. [Google Scholar] [CrossRef]
- Russo, F.; De Maddalena, A. On the presence of the spinner shark Carcharhinus brevipinna (Müller & Henle, 1839) (Chondrichthyes: Carcharhinidae) in Maldivian waters. Mar. Biol. J. 2021, 6, 87–91. [Google Scholar] [CrossRef]
- Anderson, R.C.; Ahmed, H. The Shark Fisheries of the Maldives; Ministry of Fisheries and Agriculture, Republic of Maldives: Malé, Maldives; Food and Agriculture Organization of the United Nations: Malé, Maldives, 1993; p. 73. [Google Scholar]
- Adam, M.S.; Merrett, N.R.; Anderson, R.C. Additions to the fish fauna of the Maldive Islands. Part I: An annotated checklist of the deep demersal fishes of the Maldive Islands. Ichthyol. Bull. JLB Smith Inst. Ichthyol. 1998, 67, 1–19. [Google Scholar]
- Anderson, R.; Randall, J.; Kuiter, R. New records of fishes from the Maldive Islands with notes on other species. Ichthyol. Bull. 1998, 67, 20–36. [Google Scholar]
- De Maddalena, A.; Galli, P. Sharks of the Maldives; Editoriale Magenes: Milano, Italy, 2017; p. 216. [Google Scholar]
- Ali, K.; Sinan, H. National Plan of Action for the Conservation and Management of Sharks in the Maldives; Ministry of Fisheries and Agriculture, Republic of Maldives: Malé, Maldives, 2015; p. 44. [Google Scholar]
- Branstetter, S. Problems associated with the identification and separation of the spinner shark, Carcharhinus brevipinna, and the blacktip shark, Carcharhinus limbatus. Copeia 1982, 2, 461–465. [Google Scholar] [CrossRef]
- Kajiura, S.M.; Tellman, S.L. Quantification of massive seasonal aggregations of blacktip sharks (Carcharhinus limbatus) in southeast Florida. PLoS ONE 2016, 11, e0150911. [Google Scholar] [CrossRef] [PubMed]
- Bethea, D.M.; Buckel, J.A.; Carlson, J.K. Foraging ecology of the early life stages of four sympatric shark species. Mar. Ecol. Prog. Ser. 2004, 268, 245–264. [Google Scholar] [CrossRef]
- Martín, G.; Espinoza, M.; Heupel, M.; Simpfendorfer, C.A. Estimating marine protected area network benefits for reef sharks. J. Appl. Ecol. 2020, 57, 1969–1980. [Google Scholar] [CrossRef]
- Micarelli, P.; Pireddu, M.; Persia, D.; Sanna, M.; Vicariotto, C.; Pacifico, A.; Storelli, P.; Mahrer, M.; Venanzi, E.E.; Reinero, F.R. Observations on an aggregation of grey reef sharks (Carcharhinus amblyrhynchos) in the Mozambique Channel off the coast of Nosy Be (Madagascar) and tools for photo-identification—A new aggregation nursery site? Biology 2024, 13, 661. [Google Scholar] [CrossRef]
- Marshall, A.D.; Pierce, S.J. Use and abuse of photographic identification in sharks and rays. J. Fish Biol. 2012, 80, 1361–1379. [Google Scholar] [CrossRef]
- Sims, D.W.; Speedy, C.D.; Fox, A.M. Movements and growth of a female basking shark re-sighted after a three-year period. J. Mar. Biol. Assoc. UK 2000, 80, 1141–1142. [Google Scholar] [CrossRef]
- Carraro, R.; Gladstone, W. Habitat preferences and site fidelity of the ornate wobbegong shark (Orectolobus ornatus) on rocky reefs of New South Wales. Pac. Sci. 2006, 60, 207–223. [Google Scholar] [CrossRef]
- Dudgeon, C.L.; Noad, M.J.; Lanyon, J.M. Abundance and demography of a seasonal aggregation of zebra sharks (Stegostoma fasciatum). Mar. Ecol. Prog. Ser. 2008, 368, 269–281. [Google Scholar] [CrossRef]
- Buray, N.; Mourier, J.; Planes, S.; Clua, E. Underwater photo-identification of sicklefin lemon sharks, Negaprion acutidens, at Moorea (French Polynesia). Cybium 2009, 33, 21–27. [Google Scholar]
- Anderson, S.D.; Chapple, T.K.; Jorgensen, S.J.; Klimley, A.P.; Block, B.A. Long-term individual identification and site fidelity of white sharks, Carcharodon carcharias, off California using dorsal fins. Mar. Biol. 2011, 158, 1233–1237. [Google Scholar] [CrossRef] [PubMed]
- Boube, T.; Azam, C.-S.; Guilbert, A.; Trujillo, J.E.; Femmami, N.; Bersani, F.; Laurent, E.; Mourier, J.; Kunovsky, A.; Bousseyroux, A.; et al. Population characteristics and seasonal occurrence of the great hammerhead shark, Sphyrna mokarran (Rüppell, 1837) in the Western Tuamotu Archipelago, French Polynesia. Front. Mar. Sci. 2023, 10, 1234059. [Google Scholar] [CrossRef]
- Micarelli, P.; Bonsignori, D.; Compagno, L.J.V.; Pacifico, A.; Romano, C.; Reinero, F.R. Analysis of sightings of white sharks in Gansbaai (South Africa). Eur. Zool. J. 2021, 88, 363–374. [Google Scholar] [CrossRef]
- Reinero, F.R.; Marsella, A.; Pacifico, A.; Vicariotto, C.; Maule, L.; Mahrer, M.; Micarelli, P. Influence of environmental factors on the surface feeding behaviour of immature male whale sharks in the Gulf of Tadjoura (Djibouti). Conservation 2024, 4, 792–811. [Google Scholar] [CrossRef]
- Reinero, F.R.; Bocchi, F.; Perisic, N.; Crouch, J.; Pacifico, A.; Asshauer, L.; Vicariotto, C.; Micarelli, P. First insights into social behavioral patterns between pairs of bait-attracted mature female tiger sharks from Fuvahmulah Island, Maldives. Behav. Process 2025, 229, 105216. [Google Scholar] [CrossRef]
- Reinero, F.R.; Marsella, A.; Vitale, G.; Pacifico, A.; Mahrer, M.; Micarelli, P. Environmental drivers of immature whale shark surface sightings in the Gulf of Tadjoura, Djibouti. Conservation 2025, 5, 68. [Google Scholar] [CrossRef]
- Rohner, C.A.; Richardson, A.J.; Marshall, A.D.; Weeks, S.J.; Pierce, S.J. How large is the world’s largest fish? Measuring whale sharks Rhincodon typus with laser photogrammetry. J. Fish Biol. 2011, 78, 378–385. [Google Scholar] [CrossRef]
- Klimley, A.P.; Anderson, S.D. Residency patterns of white sharks at the South Farallon Islands, California. In Great White Sharks: The Biology of Carcharodon carcharias; Klimley, A.P., Ainley, D.G., Eds.; Academic Press: Cambridge, MA, USA, 1996; pp. 365–373. [Google Scholar]
- Mourier, J.; Vercelloni, J.; Planes, S. Evidence of social communities in a spatially structured network of a free-ranging shark species. Anim. Behav. 2012, 83, 389–401. [Google Scholar] [CrossRef]
- Chin, A.; Mourier, J.; Rummer, J.L. Blacktip reef sharks (Carcharhinus melanopterus) show high capacity for wound healing and recovery following injury. Conserv. Physiol. 2015, 3, cov062. [Google Scholar] [CrossRef]
- Andreotti, S.; Rutzen, M.; van der Walt, S.; Von der Heyden, S.; Henriques, R.; Meÿer, M.; Matthee, C.A. An integrated mark-recapture and genetic approach to estimate the population size of white sharks in South Africa. Mar. Ecol. Prog. Ser. 2016, 552, 241–253. [Google Scholar] [CrossRef]
- Maroen, M.; Rowlinson, M.; Matthee, C.; Andreotti, S. The effectiveness of the semi-automated Identifin software to accurately fast-track dorsal fin photographic identifications of sharks. Aquat. Conserv. Mar. Freshw. Ecosyst. 2024, 34, e4159. [Google Scholar] [CrossRef]
- Andreotti, S.; Holtzhausen, P.; Rutzen, M.; Meÿer, M.; Van der Walt, S.; Herbst, B.; Matthee, C.A. Semi-automated software for dorsal fin photographic identification of marine species: Application to Carcharodon carcharias. Mar. Biodivers. 2018, 48, 1655–1660. [Google Scholar] [CrossRef]
- Domeier, M.L.; Nasby-Lucas, N. Annual re-sightings of photographically identified white sharks (Carcharodon carcharias) at an eastern Pacific aggregation site (Guadalupe Island, Mexico). Mar. Biol. 2006, 150, 977–984. [Google Scholar] [CrossRef]
- Meekan, M.G.; Bradshaw, C.J.A.; Press, M.; McLean, C.; Richards, A.; Quasnichka, S.; Taylor, J.G. Population size and structure of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia. Mar. Ecol. Prog. Ser. 2006, 319, 275–285. [Google Scholar] [CrossRef]
- Marshall, A.D.; Dudgeon, C.L.; Bennett, M.B. Size and structure of a photographically identified population of manta rays Manta alfredi in southern Mozambique. Mar. Biol. 2011, 158, 1111–1124. [Google Scholar] [CrossRef]
- Pratt, H.L.; Carrier, J.C. A review of elasmobranch reproductive behavior with a case study on the nurse shark, Ginglymostoma cirratum. Environ. Biol. Fish. 2001, 60, 157–188. [Google Scholar] [CrossRef]
- Gubili, C.; Johnson, R.; Gennari, E.; Oosthuizen, W.H.; Kotze, D.; Meyer, M.; Sims, D.W.; Jones, C.S.; Noble, L.R. Concordance of genetic and fin photo-identification in the great white shark (Carcharodon carcharias) off Mossel Bay, South Africa. Mar. Biol. 2009, 156, 2199–2207. [Google Scholar] [CrossRef]
- Clua, E.; Buray, N.; Legendre, P.; Mourier, J.; Planes, S. Behavioural response of sicklefin lemon sharks (Negaprion acutidens) to underwater feeding for ecotourism purposes. Mar. Ecol. Prog. Ser. 2010, 414, 257–266. [Google Scholar] [CrossRef]
- Maljković, A.; Côté, I.M. Effects of tourism-related provisioning on the trophic signatures and movement patterns of an apex predator, the Caribbean reef shark (Carcharhinus perezi). Biol. Conserv. 2011, 144, 859–865. [Google Scholar] [CrossRef]
- Hammerschlag, N.; Gallagher, A.J.; Wester, J.; Luo, J.; Ault, J.S. Don’t bite the hand that feeds: Assessing ecological impacts of provisioning ecotourism on an apex marine predator. Funct. Ecol. 2012, 26, 567–576. [Google Scholar] [CrossRef]
- Brunnschweiler, J.M.; Barnett, A. Opportunistic visitors: Long-term behavioural response of bull sharks (Carcharhinus leucas) to food provisioning in Fiji. PLoS ONE 2013, 8, e58522. [Google Scholar] [CrossRef] [PubMed]
- Séguigne, C.; Bègue, M.; Meyer, C.; Mourier, J.; Clua, É. Provisioning ecotourism does not increase tiger shark (Galeocerdo cuvier) site fidelity. Sci. Rep. 2023, 13, 7785. [Google Scholar] [CrossRef]
- Micarelli, P.; Reinero, F.R.; Pacifico, A.; Giglio, G.; Mahrer, M.; Sperone, E. Observations on the frequency of sightings of white sharks in the population of Gansbaai, South Africa, in the presence of cage diving. Biology 2025, 14, 762. [Google Scholar] [CrossRef]
- Werry, J.M.; Clua, E. Sex-based spatial segregation of adult bull sharks (Carcharhinus leucas) in the New Caledonian great lagoon. Aquat. Living Resour. 2013, 26, 281–288. [Google Scholar] [CrossRef]
- Bruce, B.D.; Bradford, R.W. Segregation or aggregation? Sex-specific patterns in the seasonal occurrence of white sharks (Carcharodon carcharias) at the Neptune Islands, South Australia. J. Fish Biol. 2015, 87, 1355–1370. [Google Scholar] [CrossRef]
- Spaet, J.L.Y.; Patterson, T.A.; Bradford, R.W.; Butcher, P.A. Spatiotemporal distribution patterns of immature Australasian white sharks (Carcharodon carcharias). Sci. Rep. 2020, 10, 10169. [Google Scholar] [CrossRef]
- Pillans, R.D.; Rochester, W.; Babcock, R.C.; Thomson, D.P.; Haywood, M.D.E.; Vanderklift, M.A. Long-term acoustic monitoring reveals site fidelity, reproductive migrations, and sex-specific differences in habitat use and migratory timing in a large coastal shark (Negaprion acutidens). Front. Mar. Sci. 2021, 8, 616633. [Google Scholar] [CrossRef]









| N. Photo-ID | Date Photo-ID in 2024 | Date Photo-ID 2025 |
|---|---|---|
| N.7-R1 | 1 November 2024 | 11 November 2024 |
| N.13-R2 | 1 November 2024 | 11 and 12 November 2024 |
| N.14-R3 | 1 November 2024 | 11, 12 and 13 November 2024 |
| N.16-R4 | 3 November 2024 | 11, 12 and 13 November 2024 |
| N.18-R5 | 3 November 2024 | 10 and 11 November 2024 |
| N.19-R6 | 3 November 2024 | 11 and 13 November 2024 |
| Year | Minimum TL (cm) | Maximum TL (cm) | Mean TL (cm) | SD (cm) |
|---|---|---|---|---|
| 2024 | 178 | 243 | 208.6 | 17.1 |
| 2025 | 114 | 246 | 207.8 | 32.3 |
| Females | ||||
| 2024 | 178 | 243 | 209.8 | 18.3 |
| 2025 | 202 | 220 | 211 | 12.7 |
| Males | ||||
| 2024 | 201 | 202 | 201.5 | 0.7 |
| 2025 | 190 | 246 | 213.3 | 18.9 |
| Unsexed | ||||
| 2025 | 114 | 240 | 177 | 89.1 |
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content. |
© 2026 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license.
Share and Cite
Reinero, F.R.; Pireddu, M.; Ridella, G.; Valenti, L.; Pacifico, A.; Ellero, F.; Micarelli, P. Baited-Associated Aggregation of Spinner Sharks in Hulhumale, Maldives: Preliminary Observations and Photo-Identification Tools. Oceans 2026, 7, 22. https://doi.org/10.3390/oceans7020022
Reinero FR, Pireddu M, Ridella G, Valenti L, Pacifico A, Ellero F, Micarelli P. Baited-Associated Aggregation of Spinner Sharks in Hulhumale, Maldives: Preliminary Observations and Photo-Identification Tools. Oceans. 2026; 7(2):22. https://doi.org/10.3390/oceans7020022
Chicago/Turabian StyleReinero, Francesca Romana, Marco Pireddu, Giacomo Ridella, Lorenzo Valenti, Antonio Pacifico, Francesca Ellero, and Primo Micarelli. 2026. "Baited-Associated Aggregation of Spinner Sharks in Hulhumale, Maldives: Preliminary Observations and Photo-Identification Tools" Oceans 7, no. 2: 22. https://doi.org/10.3390/oceans7020022
APA StyleReinero, F. R., Pireddu, M., Ridella, G., Valenti, L., Pacifico, A., Ellero, F., & Micarelli, P. (2026). Baited-Associated Aggregation of Spinner Sharks in Hulhumale, Maldives: Preliminary Observations and Photo-Identification Tools. Oceans, 7(2), 22. https://doi.org/10.3390/oceans7020022

