First records of the genus Sphenopus from temperate waters and lower mesophotic depths

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First records of the genus Sphenopus from temperate waters and lower mesophotic depths Hiroki Kise 1

&

Takuo Higashiji 2 & Masanori Nonaka 3 & Daisuke Uyeno 4 & James Davis Reimer 1,5

Received: 15 November 2019 / Revised: 6 January 2020 / Accepted: 7 January 2020 # Senckenberg Gesellschaft für Naturforschung 2020

Abstract Azooxanthellate brachycnemic zoantharians have so far only been reported from shallow depths (< 35 m) in tropical and subtropical regions. The present study reports for the first time on an azooxanthellate brachycnemic zoantharian, Sphenopus cf. exilis, from warm temperate waters. The record of this free-living, sand-dwelling species is from the mouth of Kagoshima Bay, Kagoshima, Japan. In addition, its congener S. marsupialis was observed on a sandy bottom at 114 m depth at Okinawa, Japan, which represents the first record of Sphenopus from lower mesophotic depths. These records from temperate to tropical sandy seafloors imply that free-living Sphenopus species may be much more widely and commonly distributed across the Indo-Pacific Ocean than is currently thought. Keywords Azooxanthellate . Distribution . Northernmost record . Solitary . Japan

Introduction Studies on the taxonomy and biogeography of zooxanthellate scleractinian corals and other anthozoans have been extensively conducted over the past three decades as global climate change, and marine pollution have been predicted to have impacts on these symbiotic organisms (e.g., Fukami et al. 2004; Reimer et al. 2014; Hall et al. 2015; Madin et al. 2016). However, studies on azooxanthellate anthozoans have been generally limited to certain taxa such as scleractinian corals, octocorals, and antipatharians (e.g., Cairns 2007; Pérez et al. 2016; Yesson et al. 2017). Communicated by B.W. Hoeksema * Hiroki Kise [email protected] 1

Graduate School of Engineering and Science, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan

2

Okinawa Churaumi Aquarium, Okinawa Churashima Foundation, Motobu, Okinawa, Japan

3

Okinawa Churashima Research Center, Okinawa Churashima Foundation, Motobu, Okinawa, Japan

4

Graduate School of Science and Engineering, Kagoshima University, 1-21-35 Korimoto, Kagoshima 890-0065, Japan

5

Tropical Biosphere Research Center, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan

Zoantharians of the suborder Brachycnemina Haddon & Shackleton, 1891 usually have symbiotic relationships with Symbiodiniaceae with the exception of three species within the genus Palythoa Lamouroux, 1816 and all species of the genus Sphenopus Steenstrup, 1856 (Irei et al. 2015). Due to their photosymbiotic nature, zooxanthellate brachycnemic zoantharians are generally restricted to shallow tropical and subtropical waters (Reimer et al. 2012). On the other hand, most species of the suborder Macrocnemina Haddon & Shackleton, 1891 are azooxanthellate and epizoic and are often limited to deeper waters (Swain 2010; Sinniger et al. 2013; Reimer et al. 2019). Regarding azooxanthellate