Neotype designation and re-description of Forsskål’s reticulate whipray Himantura uarnak

Philippe Borsa, Collin T. Williams, Ashlie Mcivor, Thierry B. Hoareau, Michael L. Berumen

Research output: Contribution to journalArticlepeer-review

2 Scopus citations


A continuing impediment to the taxonomy of the reticulate whipray Himantura spp. species complex is the absence of a type specimen for H. uarnak (Gmelin [ex Forsskål], 1789). Here, reticulate whipray specimens were sampled from the Jeddah region in the Red Sea, the assumed type locality of H. uarnak, and characterized genetically at the cytochrome-oxidase subunit 1 (CO1) locus. One of these specimens now in the fish collection of the California Academy of Sciences was designated as neotype. The maximum-likelihood phylogeny of all available CO1 gene sequences from the genus Himantura had the following topology: ((H. leoparda, H. uarnak), (H. undulata, (Himantura sp. 2, (H. australis + Himantura sp. 1))), H. tutul), where H. uarnak haplotypes formed a distinct lineage sister to H. leoparda. Based on these CO1 gene sequences, the geographic distribution of H. uarnak includes the eastern Mediterranean, the Red Sea, the East African coast, and the Arabian Sea. At least one lineage in the reticulate whipray species complex remains to be named.
Original languageEnglish (US)
JournalMarine Biodiversity
Issue number2
StatePublished - Mar 17 2021

Bibliographical note

KAUST Repository Item: Exported on 2021-03-23
Acknowledgements: Foremost, thanks are due to M. Tietbohl for donating tissue samples, assisting in field collections, and providing laboratory assistance. Additionally, we are grateful to J. Spaet and members of KAUST’s Reef Ecology Lab for their support at various stages of this project; to D. Catania and L.A. Rocha for allocating a CAS collection number to the neotype of H. uarnak; to X. Chen for sharing reticulate whipray sequences; to M. Bennett-Smith for sharing excellent photographs; to S. Bogorodsky, H. Debelius, and R. Fricke for insightful discussions about the taxonomy and nomenclature of reticulate whiprays; to G. Lim (ICZN) for confirming the validity of the present neotype designation. Useful taxonomic literature was communicated to us by C.-W. Chang, K.-N. Shen, and an anonymous reviewer. Several books including P. Forsskål’s Descriptiones animalium, J.F. Gmelin’s Systema naturae, and Rüppell’s Fische des rothen Meeres were accessed from the Biodiversity Heritage Library website (

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Aquatic Science
  • Oceanography


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