Page 1 VENUS 69 (3–4): 123–133, 2011 ©Malacological Society Of
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VENUS 69 (3–4): 123–133, 2011 ©Malacological Society of Japan The Bivalve Divariscintilla toyohiwakensis n. sp. (Heterodonta: Galeommatidae) from Japan, a Commensal with a Mantis Shrimp Hiroyoshi Yamashita1, Takuma Haga2** and Jørgen Lützen3* 1Association of Conservation Malacology, 3-1-26-103 Matsugaoka, Kugenuma, Fujisawa, Kanagawa 251-0038, Japan 2Department of Geology and Paleontology, National Museum of Nature and Science, 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo 169-0073, Japan 3Biological Institute, University of Copenhagen, Universitetsparken 15, DK-2100, Copenhagen Ø, Denmark Abstract: We describe the morphology of a new species of the bivalve family Galeommatidae, Divariscintilla toyohiwakensis n. sp., which lives in the burrows of the mantis shrimp Acanthosquilla acanthocarpus at Oshinden, Nakatsu, Oita Prefecture, Kyushu, Japan. An unpaired median tentacle and three paired tentacles issue from the mid-mantle fold, which partially covers the shell. A single flower-like organ originates on the anterior surface of the visceral mass. A byssal adhesive gland is observed on the most posterior part of the foot. This is the first record of the genus in Japan and the North Pacific Ocean. Keywords: Acanthosquilla acanthocarpus, flower-like organ, Lysiosquilloidea, morphology, new species, North Pacific Introduction Species of the predominantly tropical/subtropical family Galeommatidae occur chiefly in marine coastal waters. Most species are cryptic and live attached to the undersurfaces of stones, shale and dead coral pieces. Other species are associated with burrowing crustaceans in that they hide in their host’s burrows. Three species of Ephippodonta Tate, 1889 live commensally with thalassinidean shrimps (Tate, 1889; Matthews, 1893; Lützen & Nielsen, 2005) and one, Ambuscintilla praemium Iredale, 1936, lives in burrows made by a prawn. A few species of Scintillona Finlay, 1927 occur in galleries made by crabs (Morton, 2008). Eleven species belonging to Divariscintilla Powell, 1932 and a small number of other genera are known or suspected to live in the burrows of mantis shrimps (Mikkelsen & Bieler, 1992). While excavating the burrows of the mantis shrimp Acanthosquilla acanthocarpus (Claus, 1871) on a Japanese tidal beach, Mr. Taichi Wada (T.W.) discovered a species of Divariscintilla that is new to science and represents the first record of the genus in Japan and the Pacific Ocean north of Australia. The present paper describes the shell, the morphology of the soft parts, and some aspects of the host association of the new species. * Corresponding author: [email protected] ** Present address: Marine Biodiversity Research Program, Institute of Biogeosciences, Japan Agency for Marine-Earth Science and Technology, 2-15 Natsushima-cho, Yokosuka, Kanagawa 237-0061, Japan 124 H. Yamashita et al. Materials and Methods The bivalves were sampled at extreme low water from the burrows of Acanthosquilla acanthocarpus at Oshinden (part of the Suo-nada Sea, Seto Inland Sea; northwestern coast of Kyushu), in Nakatsu, Oita Prefecture, Japan. The burrow of A. acanthocarpus in sandy mud sediment sufficiently solid that it can be excavated with a shovel while maintaining its form and revealing the bivalves’ habitat (Fig. 1C). Live bivalves were maintained in a Petri dish filled with seawater at room temperature for behavioral studies. For histological studies, six specimens collected on 1 July 2007 were preserved in 70% ethanol and five specimens collected on 27 August 2007 were preserved in 4% formaldehyde. Two formalin-preserved bivalves (3.5 mm and 3.6 mm in shell length) were postfixed and decalcified in Bouin’s fluid, embedded in araldite, cut into 2-µm-thick sagittal and transverse section series, and stained with toluidine blue. Fifteen specimens collected on 22–23 July 2009 were selected for conchological observations and processed according to Fukuda et al. (2008) to remove the valves from the soft parts, of which three sets were preserved in pure ethanol for future molecular work. Twenty-six living specimens collected on 22–23 July 2009 were anaesthesized by immersion in 7.5% MgCl·6H20 in freshwater solution for one hour, fixed in 10% buffered seawater-diluted formalin for 48 hours, rinsed in tap water, and preserved in 70% ethanol. For scanning electron microscopy, specimens were passed through an ethanol to t-buthyl alcohol series, freeze-dried on a JFD-310 (JEOL) freeze drier device, and coated with gold. Scanning electron micrographs were produced on a JSM-T330A (JEOL) at the Department of Geology and Paleontology, National Museum of Nature and Science. Abbreviations: FMNH – Field Museum of Natural History, Chicago; NSMT – Department of Zoology, National Museum of Nature and Science, Tokyo (formerly National Science Museum, Tokyo); SEM – scanning electron microscopy; SH – shell height; SL – shell length. Systematics Superfamily Galeommatoidea J. E. Gray, 1840 Family Galeommatidae J. E. Gray, 1840 Genus Divariscintilla Powell, 1932 Type species: Divariscintilla maoria Powell, 1932, by original designation. Divariscintilla toyohiwakensis n. sp. (Figs. 1A–E; 2–4) Description Shell: Shell small, ranging 2.1–4.1 mm (M ± SD: 3.71 ± 0.46, n = 16) in SL and 1.8–3.3 mm (M ± SD: 2.86 ± 0.21, n = 16) in SH, roundly triangular, equivalve, thin, laterally compressed, and slightly inequilateral with beaks in front of midline (Fig. 2A–B). Very shallow indentation (vi in Fig. 2A) occurs at midventral margin in both valves. Numerous close-set very fine commarginal striae at external valve margins. Shell colorless, translucent. Brownish digestive gland visible through shell in living specimen (Fig. 1). Very thin, almost transparent periostracum covers valves. Shell surface smooth and glossy (Fig. 1A, C–E); interior bears weak radial ribs along edges, resulting in formation of finely denticulate margin, especially posteriorly (Fig. 2A). Shell capable of complete closure, not gaping at ends. Adductor muscle scars indistinct and subequal (Fig. 2A). Pallial line entire and indistinct (Fig. 2A). Hinge line short. One cardinal tooth in each valve; lateral teeth absent (Fig. 2D). Cardinal teeth abutting, not interlocking. Shallow socket (cs in Fig. 2D) on anterior part of left hinge plate accommodating right cardinal tooth. A New Species of Divariscintilla from Japan 125 Fig. 1. Divariscintilla toyohiwakensis n. sp. and its in situ occurrence. A. Living individual with soft parts extended. B. Camera lucida drawing from left. C. Longitudinal section of burrow of Acanthosquilla acanthocarpus; arrow and dotted line indicate a colony of D. toyohiwakensis n. sp. and sea bottom surface, respectively. D. Close-up of a colony. E. Live individual hanging down from wall of aquarium. F. Host, Acanthosquilla acanthocarpus. Abbreviations: amf, anterior mid-mantle fold; bag, byssus adhesive gland; bms, burrow of mantis shrimp; by, byssus; ec, excurrent siphon; ft, foot; ic, incurrent aperture; imf, inner mantle fold; mf, point of mantle fusion; pd, prodissoconch II; pfe, posterior extension of foot; pmf, posterior mid-mantle fold; pt1–pt3, pallial tentacle pairs 1–3; sh, shell; ut, unpaired tentacle. A, D–E are not scaled. 126 H. Yamashita et al. Fig. 2. Valves of Divariscintilla toyohiwakensis n. sp., scanning electron micrograph. A. Internal view of left valve. B. External view of left valve. C. Dorsal view of right valve. D. Internal view of umbonal area of conjoined valves. E. Close-up of prodissoconch II. Arrowheads indicate demarcation between prodissoconch II and dissoconch. A–C and E, holotype (NSMT-Mo 77209); D, paratype (NSMT-Mo 77210). Abbreviations: aam, anterior adductor muscle scar; apr, anterior pedal retractor muscle scar; cs, cardinal tooth socket; ct, cardinal tooth; el, external ligament; il, internal ligament; ny, nymph; pam, posterior adductor muscle scar; pe, periostracum; pl, pallial line; ppm, pedal protractor muscle scar; ppr, posterior pedal retractor muscle scar; re, resilifer; rh, ridge of hinge plate; vi, ventral indentation. Internal ligament (il in Fig. 2D) opisthodetic, short and stout. Resilifers distinct in both valves (re in Fig. 2D). External ligament (el in Fig. 2D) amphidetic, supported by nymphae. Nymphae present at both anterior and posterior margins of umbo. Nymph of right valve prominent, overlying that of left valve (ny in Fig. 2D, E). Rather long and transparent ridges (rh in Fig. 2D) present in hinge plate laterally to opaque cardinal tooth region in both valves. Prodissoconch II brownish-yellow, 320–400 µm long (n = 5), and abruptly demarcated from dissoconch in few specimens observed that were not corroded (Fig. 2E). Prodissoconch I not confirmed. Soft parts: Shell more or less internalized by profuse development of mid-mantle fold that reaches maximum extension anteriorly and posteriorly (Figs. 1B; 3A, C). Anterior edges of extended inner mantle folds form wide and hood-like incurrent region; posterior edges fused except for short excurrent siphon (Fig. 1B). Numerous small, rounded papillae (rp in Fig. 3A) A New Species of Divariscintilla from Japan 127 Fig. 3. Scanning electron micrograph of Divariscintilla toyohiwakensis n. sp., freeze-dried, paratype (NSMT- Mo 77215). A–C. Left, ventral and dorsal views, respectively. D. Close-up of posterior extension of foot showing byssal threads and byssal adhesive gland (outlined in B). E. Close-up of unpaired tentacle (outlined in C). F. Close-up of pallial tentacle #2 (outlined in A). Abbreviatioins: amf, anterior mid-mantle fold; bag, byssal adhesive gland; bg, byssal