A new species of Lepidozona (Mollusca, Polyplacophora, Ischnochitonidae) from Okinawa Trough, East China Sea

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Bull. Natl. Mus. Nat. Sci., Ser. A, 39(1), pp. 5 10, February 22, 2013 A new species of Lepidozona (Mollusca, Polyplacophora, Ischnochitonidae) from Okinawa Trough, East China Sea Hiroshi Saito Department of Zoology, National Museum of Nature and Science, 4 1 1 Amakubo, Tsukuba, Ibaraki, 305 0005 Japan E-mail: h-saito@kahaku.go.jp (Received 7 December 2012; accepted 18 January 2013) Abstract A new species of chiton, Lepidozona (Tripoplax) alba is described from Okinawa Trough, East China Sea, at the depth from 576 611 m. The new species is distinguishable from other congeneric species by the flat radiating ribs lacking of granular sculpture on the valves, smooth dorsal girdle scales with long apical spines, and the deep-sea habitat. The new species appears to inhabit a chemosynthetic environment. Key words : chiton, new species, chemosynthetic environment, Minami-Ensei Knoll, Japan. Introduction The Minami-Ensei Knoll, northern Okinawa Trough has been known to have an active hydrothermal vent and a warm seepage at the depth from 670 to 710 m (Okutani et al., 1992). From this vent site, two molluscan taxa, a turbinid gastropod, Thermocolonia jamsteci Okutani and Fujikura, 1990 [ Cantrainea jamsteci (Okutani and Fujikura, 1990) (fide Okutani, 2000)] and a vesicomyid bivalve, Calyptogena solidissima Okutani et al., 1992 [ Calyptogena kawamurai (Kuroda, 1947) (fide Kojima et al., 2006)] were described. During the survey cruise to the Ryukyu Islands area by R/V Hakuho-Maru of the Japanese Agency of Marine Science and Technology Center in 2005, a large number of molluscan shells, including living Calyptogena kawamurai were collected in the neighboring area of the Minami-Ensei Knoll, at the depth of 576 594 m (St. OT-6). Those molluscan samples included a small ischnochitonid chiton and three large disarticulated intermediate valves which were tentatively identified as a species of the genus Lepidozona. Subsequently, this area was surveyed several times by R/V Nagasaki-Maru of Nagasaki University, and a chiton specimen was collected in the latest cruise in November 2012 close to St. OT-6. Examination of those chiton specimens revealed that all belong to a single new species of the genus Lepidozona (Tripoplax), which is described herein. Material and Methods For SEM observation, pieces of girdle and radula were dissected, and macerated with household bleach, rinsed with water, cleaned with an ultrasonic cleaner, dehydrated with ethanol series, then transferred to t-butyl alcohol, and freeze-dried with JEOL-JFD300 freeze drying device. Dried specimens were coated with goldpalladium. SEM images were taken with high vacuum mode with low accelerating voltage, 1.0 kv. Terms for the gill arrangement follow Sirenko (2006). All specimens were deposited in the molluscan collection of the Department of Zoology, National Museum of Nature and Science, Tsukuba (formerly National Science Museum, Tokyo: NSMT).

6 Hiroshi Saito Systematics Class Polyplacophora Gray, 1821 Order Chitonida Thiele, 1910 Family Ischnochitonidae Dall, 1889 Genus Lepidozona Pilsbry, 1892 Subgenus Tripoplax Berry, 1919 Type species: Trachydermon trifidus Carpenter, 1864, by original designation Lepidozona (Tripoplax) alba sp. nov. (Figs. 1 17) Material examined. Holotype: NSMT-Mo 78478, body length 32 mm, near Minami-Ensei Knoll area, northern Okinawa Trough, R/V Nagasaki-Maru St. J6 1, 28 32.080 N, 127 01.894 E 28 34.126 N, 127 02.546 E, 605 611 m, on sunken wood, 16 Nov. 2012. Paratypes: 1 small individual, NSMT-Mo 78479, body length 9.8 mm, near Minami-Ensei Knoll, Okinawa Trough, R/V Hakuho-Maru St. OT-6, 28 32.22 N, 127 01.84 E 28 31.05 N, 127 01.49 E, 576 594 m, on dead shell, 13 May 2005; 3 intermediate valves, NSMT-Mo 78480, valve width 10.6 mm, 23.2 mm and 24.0 mm, same locality with another paratype, NSMT-Mo 78479. Type locality. Minami-Ensei Knoll area, northern Okinawa Trough, 28 32.080 N, 127 01.894 E 28 34.126 N, 127 02.546 E, 605 611 m. Description of holotype. Animal large, 32 mm in body length, 19 mm in body width, oval (Figs. 1, 2). Valves moderately elevated, carinated in dorsum, hardly beaked, slightly convex in side slopes. Dorsal elevation 0.35 in valve IV, which lowering towards tail valve (Figs. 5 8). Color of valves and girdle white, partially covered with blackish foreign deposit. Head valve semicircular, with shallow apical notch, almost straight in anterior slope (Fig. 4). Intermediate valves broadly rectangular; anterior margin slightly convex at both sides of somewhat forwardly produced jugal area; side margin weakly rounded, hind margin nearly straight or slightly concave; apices weakly protruding; lateral areas raised (Figs. 5 7). Tail valve less than semicircular, narrower than head valve, which length is nearly half the width; mucro well defined, located slightly behind centre (Fig. 8); postmucronal slope slightly concave. Head valve, lateral areas of intermediate valves and posterior area of tail valve cut by shallow sulci into low, flat, smooth radiating ribs: 24 in head valve, 3 4 in lateral areas, 26 in postmucronal area; sulci in posterior area very weak. Pleural areas sculptured with rather irregularly arranged longitudinal riblets, 18 20 in each side, and fine commarginal riblets; crossing points of both riblets often granularly raised. Sculpture of jugal area weak, finely reticulate with low oblique riblets (Figs. 4 6, 8). Articulamentum white; sutural laminae wide, short, more or less rounded, connected across shallow sinus by short, slightly concave jugal plate, weakly notched at both sides (Figs. 5, 6, 8, 9). Insertion plates squarish, short, outer surface rough with minute grooves. Slit formula 16/2 3/10. Girdle of moderate width (Fig. 1), covered with loosely imbricating, obliquely implanted, bent, almost smooth scales, up to 370 μm in width, 300 μm in length, with long, longitudinally grooved apical spine (Fig. 12). Surface of scales near girdle margin with numerous weak longitudinal riblets. Marginal fringe composed of long, slender, straight, blunt-topped, smooth needles, ca. 170 22 μm, sheathed on a long chitinous stalk (Fig. 13), very small, rather coarsely grooved, also stalked spicules, 40 58 10 μm, and wedge shaped, obliquely grooved, blunttopped spicules, ca. 150 40 48 μm (Fig. 14). Ventral side of girdle paved with partly overlapping rows of needles which are similar to the long stalked needle at the girdle margin, 160 190 ca. 20 μm (Fig. 15). Gills holobranchial, adanal, with space, 38 ctenidia on each side. Posterior 7 10th ctenida largest (Fig. 2). Radula 11 mm in length with 38 transverse

New Chiton from Okinawa Trough Figs. 1 11. Lepidozona alba n. sp. 1, 2, 4 9, holotype, NSMT-Mo 78478; 3, paratype, NSMT-Mo 78479; 10 11, paratypes, NSMT-Mo 78480. 1, 2, Intact animal, dorsal and ventral view. 3. Small animal, dorsal view. 4, Head valve, dorsal view. 5, Valve II, dorsal view. 6, 7, Valve IV, dorsal and frontal views. 8, 9, Tail valve, dorsal and ventral views. 10, 11, Sculpture of intermediate valves. Scales: 10 mm for 1 and 2; 5 mm for 3 11. 7

8 Hiroshi Saito rows of mineralized teeth. Central tooth narrow, weakly keeled at base, widening at top, with wide cutting edge. Centro-lateral (first lateral) teeth lower than central tooth, with small accessory plate at antero-dorsal corner, grooved at inner surface. Major lateral (2nd lateral) teeth bicuspid; inner cusp much longer than outer one, pointed at tip; outer cusp blunt at tip. Shaft of major lateral with long petaloid process that is almost attaining near the tip of the inner cusp. Major uncinus teeth with fairly wide cusp (Figs. 16, 17). Figs. 12 17. Lepidozona alba n. sp., holotype, NSMT-Mo 78478. 12, Scales on perinotum. 13, Dorsal girdle margin. 14, Close up of dorsal girdle margin. 15, Spicules on hyponotum, inner surface. 16, Radula, anterior 16 20th rows. 17, Close up of central part of radula, anterior 10 12th rows. Scales: 100 μm for 12, 13, 15 17; 50 μm for 14.

New Chiton from Okinawa Trough 9 Variation of valve sculpture. The three isolated intermediate valves exhibit considerable variation in the sculpture. Figure 10 shows a valve with a very weak sculpture, some parts are nearly smooth, whereas the valve in Figure 11 has a coarser reticulate sculpture. The sculpture of the third, smallest valve is intermediate between other two valves. Rather irregularly arranged longitudinal ribs with granular raises in the sides of pleural areas and the flat radial ribs without granules are common features shared in all specimens. In the smallest specimen (Fig. 3), the valve sculpture is much smoother than those of the larger specimen. Only a faint reticulation is found in the pleural areas. Etymology. alba, Latin, meaning white, for the color of the animal. Remarks. Among species of the subgenus Tripoplax occurring in the Northwestern Pacific area, the present species most resembles Lepidozona andrijaschevi (Yakovleva, 1952) by having flat radiating ribs in the head valve, lateral areas of the intermediate valves and postmucronal area of the tail valve. However, the present species differs from L. andrijaschevi by absence of granules on flatter radiating ribs, and the deepsea habitat. For the flat ribs without granules, the present species also resembles Lepidozona trifida (Carpenter, 1864) and Lepidozona attuensis Clark, 2000, but it can be distinguishable from either species by having long apical spine on the dorsal girdle scales, far distant distribution area, and the deep-sea habitat. L. trifida is distributed along the west coast of North America from Puget Sound, Washington, north to Shumagin Island, Alaska, in subtidal zone, and L. attuensis has been known only from Attu Island, Aleutian Islands, Alaska, in intertidal zone. Schwabe and Sellanes (2004) described Lepidozona balaenophila [ Lepidozona (Tripoplax) balaenophila Schwabe and Sellanes, 2004] from whale bones at the depth of 240 m off Conception, Chile. They suggested that L. balaenophila feeds on whale bones or is a potential member of a chemosynthetic community. They also reported Tripoplax cowani Clark, 2008 [ Lepidozona (Tripoplax) cowani (Clark, 2008)] from a methane seep area off Conception, Chile, at the depth of 922 m (Schwabe and Sellanes, 2010). The present species also appears to live in a chemosynthetic environment, because all the specimens were collected together with a bivalve, Calyptogena kawamurai which is characteristic to chemosynthetic community. Okutani (2011) described a new large lucinid bivalve, Elliptiolucina ingens from the same site with the present paratypes, and suggested that it is associated with a chemosynthetic environment. Further investigations are needed for both cases of chitons to clarify their exact habitats and the relationships with the chemosynthetic communities. However, it should be noted that three species of the same subgenus have been found in such environments. The deep-sea habitat of Tripoplax has been noted by Clark (2008). He raised Tripoplax as a full generic rank based on the deep-sea habitat together with the distribution range in northern, colder area and some morphology such like smaller dorsal scales and finer sculpture of the tegmentum other than the multiple slitting of the intermediate valves, which suggested the genus to be a natural (monophyletic) group. The features of the present new species may support his assertion, but as he remarked, further studies, especially molecular analysis, are needed to test his hypothesis and its relationship to Lepidozona (sensu Clark, 2008). Associated organisms on the sunken wood on which the holotype was attached included large number of acmaeid limpet, Pectinodonta sp. (T. Sasaki, pers. comm.) and teledinid bivalves, a few numbers of a small snail, probably a columbelliid and a small white galatheid (Crustacea), which can be seen in an image of the sunken wood taken onboard. Acknowledgements The author is grateful to Dr. Suguru Ohta, Professor Emeritus of the University of Tokyo and Dr. Jun Hashimoto, Professor of Nagasaki University, for providing such interesting specimens

10 Hiroshi Saito at his disposable. The author is indebted to the captains and crew of R/Vs Hakuho-Maru and Nagasaki-Maru for their kindest help in sampling. The author is also grateful to Dr. Hiroshi Namikawa of National Museum of Nature and Science for sampling and careful fixation of the holotype, Mr. Enrico Schwabe of Bavarian State Collection of Zoology, Munich, and Dr. Takashi Okutani, Professor Emeritus of Tokyo University of Fisheries, for their invaluable comments in reviewing the manuscript. References Clark, R. L. 2008. Two new chitons of the genus Tripoplax Berry, 1919 from the Monterey Sea Canyon. American Malacological Bulletin, 25: 77 86. Kojima, S., E. Tsuchida, H. Numanami, K. Fujikura and T. Okutani 2006. Synonymy of Calyptogena solidissima with Calyptogena kawamurai (Bivalvia: Vesicomyidae) and its population structure revealed by mitochondorial DNA sequences. Zoological Science, 23: 835 842. Okutani, T. 2000. Marine Mollusks in Japan. 1175 pp. Tokai University Press, Tokyo. Okutani, T. 2011. Bizarre lucinid bivalves from southwestern Japan, including a new species, and relatives in adjacent waters. Venus, 69: 115 122. Okutani, T., J. Hashimoto and K. Fujikura 1992. A new species of vesicomyid bivalve associated with hydrothermal vents near Amami-Oshima Island, Japan. Venus, 51: 225 233. Schwabe, E. and J. Sellanes 2004. A new species of Lepidozona (Mollusca: Polyplacophora: Ischnochitonidae), found on whale bones off the coast of Chile. Iberus, 22: 147 153. Schwabe, E. and J. Sellanes 2010. Revision of Chilean bathyal chitons (Mollusca: Polyplacophora) associated with cold-seeps, including description of a new species of Leptochiton (Leptochitonidae). Organisms Diversity & Evolution, 10: 31 55. Sirenko, B. 2006. New outlook on the system of chitons (Mollusca: Polyplacophora). Venus, 65: 27 49.