Abstract
The thorny seahorse, Hippocampus histrix Kaup, 1856, has long been treated as a single, widely distributed species ranging from the Red Sea and western Indian Ocean to the western Pacific. Morphological and molecular evidence now demonstrate that this broad concept encompasses at least three diagnosable species. Examination of Indian Ocean specimens combined with mitochondrial COI sequence analysis reveals a new lineage herein described as Hippocampus amandavincentae sp. nov., based on material from southeastern India and corroborated by published sequences from Madagascar, Mozambique, the Seychelles, and the northern Red Sea. The new species is diagnosed by a shorter snout, five coronet spines, and spines present on all principal trunk and tail ridges. A published COI sequence from the northern Red Sea places H. amandavincentae in apparent sympatry with Hippocampus jayakari Boulenger, 1900, in that region, indicating that the distributional boundary between these two species requires targeted field investigation. Hippocampus histrix sensu stricto is redescribed and its distribution restricted to the Pacific Ocean, including Japan, Indonesia, and the Philippines, based on direct examination of type material. Hippocampus jayakari, described from the Gulf of Oman, is confirmed as a distinct species of the northwestern Indian Ocean, readily distinguished by four cleithral spines with the ventral-most bifurcate, a shorter and deeper snout, and spines confined to alternating tail rings. Together, these three species represent geographically coherent lineages whose delineation resolves over a century of misidentification in museum collections and online databases, stabilises the application of long-used names, and provides a practical framework for regional conservation assessment and future systematic research on spiny seahorses across the Indo-Pacific.
References
- Boehm, J.T., Woodall, L., Teske, P.R., Lourie, S.A., Baldwin, C., Waldman, J. & Hickerson, M. (2013) Marine dispersal and barriers drive Atlantic seahorse diversification. Journal of Biogeography, 40 (10), 1839–1849. https://doi.org/10.1111/jbi.12127
- Borsa, P., Sembiring, A., Fauvelot, C. & Chen, W.-J. (2014) Resurrection of Indian Ocean humbug damselfish, Dascyllus abudafur (Forsskål) from synonymy with its Pacific Ocean sibling, Dascyllus aruanus (L.). Comptes Rendus Biologies, 337 (12), 709–716. https://doi.org/10.1016/j.crvi.2014.09.001
- Boulenger, G.A. (1900) Description of a new sea-horse (Hippocampus) from Muscat. The Annals and Magazine of Natural History, Series 7, 6 (31), 51–52. https://doi.org/10.1080/00222930008678338
- Chang, C.-H., Shao, K.-T., Lin, H.-Y., Chiu, Y.-C., Lee, M.-Y., Liu, S.-H. & Lin, P.-L. (2017) DNA barcodes of the native ray-finned fishes in Taiwan. Molecular Ecology Resources, 17 (4), 796–805. https://doi.org/10.1111/1755-0998.12601
- Cheng, R., Fang, Y., Ge, Y., Liu, Q. & Zhang, G. (2017) Complete mitochondrial genome sequence of the Jayakar’s seahorse Hippocampus jayakari Boulenger, 1900 (Gasterosteiformes: Syngnathidae). Mitochondrial DNA Part B, 2 (2), 593–594. https://doi.org/10.1080/23802359.2017.1372704
- Chernomor, O., von Haeseler, A. & Minh, B.Q. (2016) Terrace aware data structure for phylogenomic inference from supermatrices. Systematic Biology, 65 (6), 997–1008. https://doi.org/10.1093/sysbio/syw037
- Coleman, R.R., Eble, J.A., DiBattista, J.D., Rocha, L.A., Randall, J.E., Berumen, M.L. & Bowen, B.W. (2016) Regal phylogeography: range-wide survey of the marine angelfish Pygoplites diacanthus reveals evolutionary partitions between the Red Sea, Indian Ocean, and Pacific Ocean. Molecular Phylogenetics and Evolution, 100, 243–253. https://doi.org/10.1016/j.ympev.2016.04.005
- Dawson, C.E. (1985) Indo-Pacific Pipefishes (Red Sea to the Americas). Gulf Coast Research Laboratory, Ocean Springs, Mississippi, 230 pp.
- Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32 (5), 1792–1797. https://doi.org/10.1093/nar/gkh340
- GBIF.org (2026) Global Biodiversity Information Facility. Available from: https://www.gbif.org (accessed 26 May 2026)
- Hamilton, H., Saarman, N., Short, G., Sellas, A.B., Moore, B., Hoang, T., Grace, C.L., Gomon, M., Crow, K. & Simison, W.B. (2017) Molecular phylogeny and patterns of diversification in syngnathid fishes. Molecular Phylogenetics and Evolution, 107, 388–403. https://doi.org/10.1016/j.ympev.2016.10.003
- Han, S.-Y., Kim, J.-K., Kai, Y. & Senou, H. (2017) Seahorses of the Hippocampus coronatus complex: taxonomic revision, and description of Hippocampus haema, a new species from Korea and Japan (Teleostei, Syngnathidae). ZooKeys, 712, 113–139. https://doi.org/10.3897/zookeys.712.14955
- Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- Jaonalison, H., Durand, J.-D., Mahafina, J., Valade, P., Collet, A., Cerqueira, F. & Ponton, D. (2022) Application of DNA barcoding for monitoring Madagascar fish biodiversity in coastal areas. Diversity, 14 (5), 377. https://doi.org/10.3390/d14050377
- Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14 (6), 587–589. https://doi.org/10.1038/nmeth.4285
- Kaup, J.J. (1856) Catalogue of Lophobranchiate Fish in the Collection of the British Museum. Trustees of the British Museum, London, 76 pp. https://doi.org/10.5962/bhl.title.21150
- Kimmerling, N., Zuqert, O., Amitai, G., Gurevich, T., Armoza-Zvuloni, R., Kolesnikov, I., Berenshtein, I., Melamed, S., Gilad, S., Benjamin, S., Rivlin, A., Ohavia, M., Paris, C.B., Holzman, R., Kiflawi, M. & Sorek, R. (2018) Quantitative species-level ecology of reef fish larvae via metabarcoding. Nature Ecology & Evolution, 2 (2), 306–316. https://doi.org/10.1038/s41559-017-0413-2
- Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35 (6), 1547–1549. https://doi.org/10.1093/molbev/msy096
- Lourie, S.A., Foster, S.J., Cooper, E.W.T. & Vincent, A.C.J. (2004) A Guide to the Identification of Seahorses. Project Seahorse and TRAFFIC North America, Washington D.C., 114 pp.
- Lourie, S.A., Pollom, R.A. & Foster, S.J. (2016) A global revision of the seahorses Hippocampus Rafinesque 1810 (Actinopterygii: Syngnathiformes): taxonomy and biogeography with recommendations for further research. Zootaxa, 4146 (1), 1–66. https://doi.org/10.11646/zootaxa.4146.1.1
- Lourie, S.A., Vincent, A.C.J. & Hall, H.J. (1999) Seahorses: An Identification Guide to the World’s Species and Their Conservation. Project Seahorse, London, 214 pp.
- Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32 (1), 268–274. https://doi.org/10.1093/molbev/msu300
- Nickel, J. & Cursons, R. (2012) Genetic diversity and population structure of the pot-belly seahorse Hippocampus abdominalis in New Zealand. New Zealand Journal of Marine and Freshwater Research, 46 (2), 207–218. https://doi.org/10.1080/00288330.2011.632014
- Rambaut, A. (2018) FigTree v1.4.4. Available from: https://tree.bio.ed.ac.uk/software/figtree/ (accessed 1 January 2024)
- Ratnasingham, S. & Hebert, P.D.N. (2007) BOLD: the Barcode of Life Data System (http://www.barcodinglife.org). Molecular Ecology Notes, 7 (3), 355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x
- Sabaj, M.H. (2020) Codes for natural history collections in ichthyology and herpetology. Copeia, 108 (3), 593–669. https://doi.org/10.1643/ASIHCODONS2020
- Shalu, K., Dahanukar, N., Raghavan, R. & Ranjeet, K. (2021) By-catch-associated demographics of two threatened seahorses from the south-east coast of India. Marine and Freshwater Research, 73 (3), 343–350. https://doi.org/10.1071/MF21127
- Short, G., Harasti, D. & Hamilton, H. (2019) Hippocampus whitei Bleeker, 1855, a senior synonym of the southern Queensland seahorse H. procerus Kuiter, 2001: molecular and morphological evidence (Teleostei, Syngnathidae). ZooKeys, 824, 109–133. https://doi.org/10.3897/zookeys.824.30921
- Short, G., Smith, R., Harasti, D. & Claassens, L. (2024) A new record and species of pygmy pipehorse of the genus Cylix (Teleostei, Syngnathidae) from South Africa and the African continent. Ichthyology & Herpetology, 112 (3), 315–327. https://doi.org/10.1643/i2023053
- Silveira, R.B., Siccha-Ramirez, R., Silva, J.R.S. & Oliveira, C. (2014) Morphological and molecular evidence for the occurrence of three Hippocampus species (Teleostei: Syngnathidae) in Brazil. Zootaxa, 3861 (4), 317–332. https://doi.org/10.11646/zootaxa.3861.4.2
- Singh, K.V., Lakra, W.S., Gopalakrishnan, A., Modayil, M.J., Malakar, A.K. & Sobti, R.C. (2011) Molecular identification and phylogenetic relationship of seahorse, Hippocampus kuda (Bleeker 1852) using mitochondrial 16S rRNA and COI gene sequences from east and west coasts of India. Indian Journal of Animal Sciences, 81 (1), 97–101.
- Song, H. & Mabuchi, K. (2014) Complete mitochondrial genome sequence of the thorny seahorse Hippocampus histrix (Gasterosteiformes: Syngnathidae). Mitochondrial DNA, 25 (1), 7–8. https://doi.org/10.3109/19401736.2013.775263
- Teske, P.R. & Beheregaray, L.B. (2009) Evolution of seahorses’ upright posture was linked to Oligocene expansion of seagrass habitats. Biology Letters, 5 (4), 521–523. https://doi.org/10.1098/rsbl.2009.0152
- Wang, X., Zhong, H., Guo, J. & Hou, F. (2020) Morphology and molecular identification of the zoological origin of medicinal seahorses in Chinese herbal markets. Mitochondrial DNA Part A, 31 (8), 335–345. https://doi.org/10.1080/24701394.2020.1815719
- Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R. & Hebert, P.D.N. (2005) DNA barcoding Australia’s fish species. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 360 (1462), 1847–1857. https://doi.org/10.1098/rstb.2005.1716
- Weigt, L.A., Baldwin, C.C., Driskell, A., Smith, D.G., Ormos, A. & Reyier, E.A. (2012) Using DNA barcoding to assess Caribbean reef fish biodiversity: expanding taxonomic and geographic coverage. PLoS ONE, 7 (7), e41059. https://doi.org/10.1371/journal.pone.0041059
- Whitley, G.P. (1964) Presidential address: a survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89 (1), 11–127.
- Wilson, A.B., Vincent, A., Ahnesjö, I. & Meyer, A. (2001) Male pregnancy in seahorses and pipefishes (family Syngnathidae): rapid diversification of paternal brood pouch morphology inferred from a molecular phylogeny. Journal of Heredity, 92 (2), 159–166. https://doi.org/10.1093/jhered/92.2.159
- Zhang, Y.-H., Qin, G., Zhang, H.-X., Wang, X. & Lin, Q. (2017) DNA barcoding reflects the diversity and variety of brooding traits of fish species in the family Syngnathidae along China’s coast. Fisheries Research, 185, 137–144. https://doi.org/10.1016/j.fishres.2016.09.015
