Carlgren’s hesitation allayed: redescription and systematics of Heteranthus verruculatus Klunzinger, 1877 (Cnidaria, Actiniaria), with a redefinition of Heteranthidae Carlgren, 1900

Nicholas Wei Liang Yap, Zheng Bin Randolph Quek, Ria Tan, Dharma Arif Nugroho, Jen Nie Lee, Michael L. Berumen, Koh Siang Tan, Danwei Huang

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Species boundaries delineating tropical sea anemones (Cnidaria, Actiniaria) of the zooxanthellate genus, Heteranthus Klunzinger, 1877, are unclear. There are currently two valid Heteranthus species: type species Heteranthus verruculatus Klunzinger, 1877, first reported from Koseir, Egypt, and H. insignis Carlgren, 1943, 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 Carlgren, 1900 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.
Original languageEnglish (US)
Pages (from-to)1-28
Number of pages28
JournalContributions to Zoology
StatePublished - Nov 27 2020

Bibliographical note

KAUST Repository Item: Exported on 2020-12-02
Acknowledgements: We thank the ongoing efforts of the ‘Anemone Army’ over the many years; without their keen eyesight and persistent enthusiasm, many of the tiny specimens featured in this study would not have been found or collected. This study also benefited from the collections of sea anemones made during the Comprehensive Marine Biodiversity Survey (CMBS, 2010–2015) and associated workshops led by the National Parks Board and National University of Singapore (NUS). The CMBS workshops were supported by generous contributions from Asia Pacific Breweries Singapore, Care-for-Nature Trust Fund, Keppel Care Foundation, Shell Companies in Singapore and The Air Liquide Group. NWL Yap also thanks Ms Sudhanshi Jain and Mr Chim Chee Kong for their assistance in overseas fieldwork. For permit arrangements granted to conduct fieldwork overseas, we thank: Dr Roberto Arrigoni (Saudi Arabia); Drs Yasser Salama and Mostafa Ahmed (Egypt); Profs Bella Galil and Menachem Goren, Mr Golan and Mr Ziv Neder (Israel); Mr Lutfi Afiq-Rosli (Malaysia); Dr Saleh Yahya, Ms Kalma, and Mr Ahmad (Tanzania, Zanzibar). We also thank the personnel from the following institutions that granted permission to examine type material and voucher specimens, all of which were crucial in helping reconstruct the identity of Heteranthus verruculatus. For this, NWL Yap gratefully acknowledges the following staff from the respective institutes: Ms Miranda Lowe and Mr Andrew Cabrinovic (nhm); Dr Dwi Listyo Rahayu (lipi); Dr Magalie Castelin (mnhn); Dr Robyn Cumming (mtq); Dr Lena Gustavsson, Dr Mattias Forshage, Ms Emma Wahlberg, Ms Emily Dock Åkerman (nrs); Dr Gustav Paulay and Ms Amanda M Bemis (ufm); Dr Jane Fromont and Mr Oliver Gomez (wam); Drs Lüter Carsten and Birger Neuhaus (zmb); and Ms Iffah Iesa, Mr Kelvin Lim Kok Peng and Mr Chua Keng Soon (zrc). We also thank Ms Royale Hardenstine from the King Abdullah University of Science and Technology for sharing the specimen of Phymanthus loligo collected from Thuwal, Saudi Arabia. Dr Andrea Crowther generously shared information on a sample of Heteranthus collected from Australia. On logistical support and help with molecular work, we thank the staff of Tropical Marine Science Institute and the Reef Ecology Laboratory, NUS: Ms Gan Bin Qi, Ms Helen Wong, Ms Serina Lee, Mr Ahmed Aliyar, Mr Jackson Chan, Mr Marc Chan and Aden Ip. We are grateful to Mr Lok Kok Sheng for permission to use his photograph in this publication. Prof Peter Ng and Dr Darren Yeo for their constructive inputs to this study, as with the three anonymous reviewers and Prof Bert Hoeksema whose comments greatly improved this manuscript. Fieldwork in Malaysia was conducted with the following permit: jtlm 630-7Jld.9(9). Computational work for this article was partially performed on resources of the National Supercomputing Centre, Singapore ( The St John’s Island National Marine Laboratory provided the facilities necessary for conducting this study – the Laboratory is a National Research Infrastructure under the National Research Foundation (nrf), Prime Minister’s Office, Singapore. This research was also supported by the nrf, under its Marine Science R&D Programme (MSRDP-P03 and MSRDP-P38).


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