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Carlgren’s hesitation allayed: redescription and systematics of Heteranthus verruculatus Klunzinger, 1877 (Cnidaria, Actiniaria), with a redefinition of Heteranthidae Carlgren, 1900
Yap, N.W.L.; Quek, Z.B.R.; Tan, R.; Nugroho, D.A.; Lee, J.N.; Berumen, M.L.; Tan, K.S.; Huang, D. (2021). Carlgren’s hesitation allayed: redescription and systematics of Heteranthus verruculatus Klunzinger, 1877 (Cnidaria, Actiniaria), with a redefinition of Heteranthidae Carlgren, 1900. Contributions to Zoology 90(2): 155-182. https://dx.doi.org/10.1163/18759866-bja10015
In: Contributions to Zoology. SPB Academic Publishing: Amsterdam. ISSN 1383-4517; e-ISSN 1875-9866, more
Peer reviewed article  

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Keywords
    Actiniaria [WoRMS]; Actinioidea Rafinesque, 1815 [WoRMS]; Heteranthidae Carlgren, 1900 [WoRMS]; Heteranthus verruculatus Klunzinger, 1877 [WoRMS]; Phymanthus Milne Edwards & Haime, 1851 [WoRMS]
    Marine/Coastal
Author keywords
    Actinioidea; Indo-West Pacific; longitudinal fission; Phymanthus; taxonomy; zooxanthellate

Authors  Top 
  • Yap, N.W.L.
  • Quek, Z.B.R.
  • Tan, R.
  • Nugroho, D.A.
  • Lee, J.N.
  • Berumen, M.L.
  • Tan, K.S.
  • Huang, D.

Abstract
    Species boundaries delineating tropical sea anemones (Cnidaria, Actiniaria) of the zooxanthellate genus, Heteranthus , are unclear. There are currently two valid Heteranthus species: type species Heteranthus verruculatus , first reported from Koseir, Egypt, and H. insignis , from Poulo Condore, Vietnam. In describing the latter from a single, poorly preserved specimen, zoologist Oskar Carlgren expressed apprehension with traits he had used to establish this species. Carlgren’s doubts persisted later in writing when he found a similar-looking sea anemone from the Great Barrier Reef. Crucial details to positively identify either species have since remained limited. Here, we re-diagnosed Heteranthus and re-described its type species based on observations of specimens we have obtained from Singapore and Pulau Ambon (Indonesia), and of museum material collected elsewhere across the Indo-West Pacific region (n > 180). Supported by molecular phylogenetic evidence, the family Heteranthidae was reinstated and re-diagnosed. Heteranthus verruculatus is encountered in the lower intertidal region amongst seagrass, in rocky crevices, or coral rubble. It occurs as solitary individuals or in clonal clusters, well-camouflaged against the substratum. Individuals were observed to frequently propagate by longitudinal fission, resulting in a varied appearance. Type material of H. verruculatus and H. insignis were re-examined and as we found no differences between them, the two were synonymised. We inferred that Carlgren probably misinterpreted cnidae and histological data in defining H. insignis as a distinct species. This revision clarifies the taxonomy and geographic range of H. verruculatus, an Indo-West Pacific species that is found from the Red Sea to subtropical Australia and Hawaii.

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