Widest geographic distribution of a shallow and mesophotic antipatharian coral (Anthozoa: Hexacorallia): Antipathes gran

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Widest geographic distribution of a shallow and mesophotic antipatharian coral (Anthozoa: Hexacorallia): Antipathes grandis VERRILL, 1928 – confirmed by morphometric and molecular analyses Erika Gress1,2* , Dennis M. Opresko3, Mercer R. Brugler4,5, Daniel Wagner6, Igor Eeckhaut7,8 and Lucas Terrana7,8

Abstract We provide the first record of the shallow and mesophotic (< 150 m depth) antipatharian coral Antipathes grandis VERRILL, 1928 from the Indian Ocean. First described from Hawaii, A. grandis was recently found on a mesophotic coral reef in SW Madagascar. Its identity was confirmed by means of morphological and molecular analyses (cytochrome c oxidase subunit I). These findings not only expand our knowledge of the potential geographic range of antipatharian corals, but also challenge the belief that only deep-sea (> 200 m depth) benthic taxa have such wide geographic distributions. Keywords: Black coral, Geographic range, Madagascar, Hawaii, Indian Ocean, Antipatharia

Introduction Corals in the order Antipatharia MILNE-EDWARDS & HAIME, 1857 (Anthozoa: Hexacorallia) – commonly known as black corals – inhabit all oceans at depths ranging from 2 m down to 8600 m (Roberts et al. 2009; Wagner et al. 2012). Around 273 antipatharian species, within 45 genera and 7 families, have been described. Despite their importance as habitat providers on reefs at all depths (Love et al. 2007; Tazioli et al. 2007; Suarez et al. 2015; Gress and Andradi-Brown 2018), ecological studies of these corals are still very limited. Moreover, long-term ecological studies are rare. Exceptions include the ecological studies dedicated to Antipathes grandis, which was the first antipatharian species described from Hawaii (Verrill 1928). * Correspondence: [email protected]; [email protected] 1 ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Australia 2 Nekton Foundation, Oxford, England Full list of author information is available at the end of the article

Vast aggregations of A. grandis (VERRILL, 1928) were first documented in the late 1960s on mesophotic reefs off the western side of Maui, Hawaii (Grigg 1964, 1965, 1984, 2001). The arborescent colonies of A. grandis and Antipathes griggi (Opresko 2009) have been harvested for their skeleton to support the black coral jewellery industry since their discovery in Hawaii (Grigg 1984, 2001; Montgomery and France 2006; Wagner et al. 2010). This commercial exploitation of antipatharian corals has been the impetus for ecological studies on these two species for several decades in Hawaii (Grigg 1964, 1965, 1984, 1993, 2001; Kahng and Grigg 2005; Wagner et al. 2010, 2017). The harvesting was presumed to be sustainable (Grigg 1984, 1993, 2001), although the latest findings demonstrate that the populations were declining after several decades of exploitation (Grigg 2004; Montgomery and France 2006). These ecological studies, however, are some of the lengthiest and most detailed ones conducted on any antipatharian species to date.