Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae) from Wakayama, Japan
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Species Diversity 20: 141–151 25 November 2015 DOI: 10.12782/sd.20.2.141 A New Species of Thyone (Echinodermata: Holothuroidea: Dendrochirotida: Phyllophoridae) from Wakayama, Japan Yusuke Yamana1,3, Atsushi Hirai2, and Kentarou Hirashima1 1 Wakayama Prefectural Museum of Natural History, Funo 370-1, Kainan, Wakayama 642-0001, Japan E-mail: [email protected] 2 Susami Crustacean Aquarium, Esumi 808-1, Susami, Nishimuro, Wakayama 641-3142, Japan 3 Corresponding author (Received 11 January 2015; Accepted 27 October 2015) A new phyllophorid sea cucumber, Thyone susamiensis sp. nov., is described from the intertidal zone of the south coast of Wakayama Prefecture, western Japan. It is distinguished from the most of its congeners by the absence of ossicles in the body wall. Thyone susamiensis has only rosette-like ossicles in the introvert, where table-like ossicles are usually observed in other Thyone species. The usual table-like ossicles were observed only in the peri-oral skin in T. susamiensis. Among conge- ners, this condition has been reported only in Thyone flindersi O’Loughlin, Barmos, and VandenSpiegel, 2012. However, T. flindersi occasionally has table-like ossicles in the anal appendages, where T. susamiensis usually only has rod-like ossicles. The general body colouration and the shape of the calcareous ring are also distinctive, separating species T. susamiensis from T. discolor Sluiter, 1901, T. roscovita Hérouard, 1889, and T. venusta Selenka, 1867, species for which detailed descrip- tion of ossicles are lacking. Key Words: Holothuroidea, Dendrochirotida, Phyllophoridae, Thyone susamiensis sp. nov., new species, Japan. other holothurians even in different families (e.g., species in Introduction the sclerodactyliid genus Havelockia Pearson, 1903), a num- ber of species once placed in Thyone have been transferred The dendrochirotid sea-cucumber genus Thyone Oken, to other taxa, such as Stolus Selenka, 1867. Still, 61 species 1815 (type species: Holothuria fusus Müller, 1776) is one of and two subspecies are currently contained in Thyone (Pau- the oldest in the class Holothuroidea and has a complicated lay 2014), many of which require scrutiny to ascertain their nomenclatural history. Names established in Oken (1815– precise taxonomic placement. 1816), including Thyone, were once ruled to be unavailable Although no full redescription of the type species, T. since the works did not strictly follow the Principle of Bi- fusus (Müller, 1776), has been published, detailed informa- nominal Nomenclature (International Commission on Zoo- tion concerning some of its features was provided by Mad- logical Nomenclature [ICZN] 1956). While Jaeger (1833) sen (1941). According to this and other publications, many subsequently made the dendrochirotid Thyone available, of the currently recognized species of Thyone share certain Lesson (1830) had earlier erected an aspidochirotid holo- common features, which are presently understood as defin- thurian genus with the same name: i.e., the aspidochirotid ing features of the genus: 10 tentacles comprising eight large Thyone Lesson, 1830 had precedence over the dendrochi- and two small (ventral) ones; a long, tubular composite cal- rotid Thyone Jaeger, 1833. Despite the above ICZN ruling, careous ring, in which the anterior first piece of each radial however, Thyone has been in prevailing usage in the sense and inter-radial element is broad with anterior projections, of Dendrochirotidae, rather than Aspidochirotidae. This more posterior to which a mosaic pattern of 1–3 rows of prompted Paulay and O’Loughlin (2012) to ask the ICZN to pieces is usually formed; a bifid posterior part of the radial make Thyone Oken, 1815 available under its plenary power. elements, usually with the beginning of the fork situated Recently, a ruling on Paulay and O’Loughlin’s (2012) Case posterior (sometimes barely anterior) to the rear end of the 3598 was published as OPINION 2367 (ICZN 2015) to adjacent inter-radial element; one Polian vesicle and one make Thyone Oken, 1815 an available and valid name within (sometimes two) stone canals; pedicels covering the entire the Dendrochirotidae, thereby maintaining current usage in body; no pedicels on the introvert; the body elongate and accordance with Article 82.1 of the International Code of cylindrical or tapered towered both ends; multi-perforate Zoological Nomenclature (ICZN 1999). plate-like ossicles or rosette-like ossicles in the tentacles; Being the most speciose genus in the class Holothuroidea, and usually two-pillared table-like ossicles in the body wall Thyone is problematic not only nomenclaturally, but also (these sometimes lacking). taxonomically. Oken’s (1815) original diagnosis was quite The following six nominal species of Thyone sensu lato simple, based on the number of expansible tentacles (10) have been reported from Japanese waters: Thyone benti De- and the shape of the composite calcareous ring (thin and tu- ichmann, 1937 from Sagami Bay (Kuramochi 2006); T. bi- bular). Because this combination of features applies to many cornis Ohshima, 1915 from Suruga Bay (Ohshima 1915); © 2015 The Japanese Society of Systematic Zoology 142 Yusuke Yamana et al. T. multipes Augustin, 1908 from Uraga Strait and Sagami oral skin on the abdominal-lateral side; 3) the skin of the Bay (Augustin 1908) and Suruga Bay (Ohshima 1915); T. oesophagus on the abdominal-lateral side; 4) the anterior, 5) parva Ohshima, 1915 from off the Ojika Peninsula (Ohshi- middle, and 6) posterior parts of the skin of the introvert in ma 1915); T. punctata Ohshima, 1915 from off Cape Sata the inter-radius, on the abdominal-lateral side; 7) the ante- (Ohshima 1915); and T. sacellus (Selenka, 1867) from off rior, 8) middle, and 9) posterior parts of the body-wall in- Kyushu (Mitsukuri 1912). Today, T. multipes, T. parva, and tegument in the inter-radius, on the abdominal side; 10–12) T. punctata are accepted as Allothyone multipes, Paracu- the integument of the same regions of the body on the dor- cumaria parva, and Stolus punctatus, respectively (Panning sal side; 13) pedicels of the middle parts of the abdominal 1949), and T. sacellus is considered a synonym of Stolus buc- side; and 14) the body-wall integument and 15) appendages calis (Stimpson, 1855) (Thandar 1990). However, accord- around the anus, both on the abdominal-lateral side. Exam- ing to Augustin’s (1908) description, Allothyone multipes ination of ossicles from parts 2) and 3) was done only for actually does not display the diagnostic features of Allothy- three specimens because the necessary dissection resulted in one Panning, 1949 (q.v.), so it may really belong to Thyone. major damage to the specimens. Furthermore, Thyone benti actually should be included in The tissue samples were dissolved in sodium hypochlorite Eupentacta Deichmann, 1938 or Havelockia Pearson, 1903 (NaClO, 5%) under a compound microscope (Nikon Op- (see Discussion). Therefore, currently, only Thyone bicornis tiphot). Because no obvious differences were found among and Allothyone multipes are the only confirmed or suspected the ossicles of the 10 specimens examined, those of three representatives of Thyone in Japanese waters. Thyone bicor- specimens were counted and measured on the basis of pho- nis was found at a depth of 86 m in Suruga Bay, Shizuoka, tographs. We classified the ossicles into five types: two types during the North-western Pacific survey by the U.S. Fish of ‘rosette’ (i.e., rosette-like ossicle, extremely branched Commission Steamer Albatross (Ohshima 1915). Allothyone and perforated), one type of ‘table’ (ossicles consisting of multipes was found at a depth of 350 m in the Uraga Strait a multi-perforate ‘disc’ and a skeleton-constructing ‘spire’ between Chiba and Kanagawa, and at a depth of 150 m in that grows perpendicularly from the disc), one type of ‘rod’ Sagami Bay, Kanagawa (Augustin 1908). Recently, we found (rod-like ossicles armed distally with lumps or branches or a species of Thyone that is distinct from both T. bicornis and few perforations), and one type of ‘plate’ (multi-perforate A. multipes on an intertidal shore of the Pacific Ocean far- flat ossicles). For every tissue sample, we measured a total ther southwest in Japan. We compared this species to all its of 10–100 ossicles to the nearest micrometer. The type speci- congeners through a careful survey of the literature, which mens of the new species and glass slides of their ossicles resulted in the conclusion that this shore-dwelling species is have been deposited in the Invertebrate Collection (INV) new to science. of the Wakayama Prefectural Museum of Natural History (WMNH), in Kainan, Wakayama, Japan. Materials and Methods Thyone susamiensis sp. nov. (Figs 1–7) Ten specimens of the new species were collected by the authors between December, 2013, and January, 2014, from Material examined. Holotype, WMNH-2014-INV-2 the type locality, viz., the intertidal zone of a boulder shore (anaesthetized; length 127 mm, width 20 mm). Paratypes: (33°30′21″N, 135°35′24″E) in the town of Susami on the WMNH-2014-INV-1 (anaesthetized; length 111 mm, south coast of Wakayama Prefecture, Japan. width 17 mm); WMNH-2014-INV-3 (anaesthetized; length Five specimens were fixed in 80% ethanol after anaesthe- 121 mm, width 19 mm); WMNH-2014-INV-4 (anaesthe- tization; the remaining five were fixed without anaesthesia. tized; length 94 mm, width 18 mm); WMNH-2014-INV-5 Anaesthetization was performed in a menthol solution: (anaesthetized; length 74 mm, width 8 mm); WMNH- 0.37 g of menthol was dissolved into 100 mL of 99% etha- 2014-INV-6 (unanaesthetized; length 54 mm, width nol, which was diluted to 3.0% with seawater. The specimens 22 mm); WMNH-2014-INV-7 (unanaesthetized; length were injected with 0.5–2.5 mL of this diluted solution and 51 mm, width 9 mm); WMNH-2014-INV-8 (unanaesthe- immersed in the diluted solution for 2.0–6.0 h; when the an- tized; length 71 mm, width 23 mm); WMNH-2014-INV-9 aesthesia had taken effect, the tentacles emerged in response (unanaesthetized; length 58 mm, width 25 mm); WMNH- to a gentle squeeze of the body.