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Redescriptions and Attachment Modes of Hypermastus Peronellicola and H

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Redescriptions and Attachment Modes of Hypermastus Peronellicola and H VENUS 69 (1–2): 25–39, 2010 ©Malacological Society of Japan Redescriptions and Attachment Modes of Hypermastus peronellicola and H. tokunagai (Prosobranchia: Eulimidae), Ectoparasites on Sand Dollars (Echinodermata: Clypeasteroida) in Japanese Waters Haruna Matsuda1*, Tatsuo Hamano2, Shigeo Hori3 and Kazuya Nagasawa1 1Graduate School of Biosphere Science, Hiroshima University, Higashi-Hiroshima, Hiroshima 739-8528, Japan 2Institute of Socio-Arts and Sciences, the University of Tokushima, Tokushima 770-8502, Japan 3Hagi Museum, 355, Horiuchi, Hagi, Yamaguchi 758-0057, Japan Abstract: Two species of the eulimid genus Hypermastus are redescribed based on specimens recently collected from sand dollars caught in the Seto Inland Sea and the type specimens: Hypermastus peronellicola (Kuroda & Habe, 1950) from Peronella japonica, and H. tokunagai (Yokoyama, 1922) from Scaphechinus mirabilis. These two eulimid species are very similar in their shell morphology but are distinguished from each other based on characters such as the proportions of shell length to several dimensions of the shell, width/length ratios of each teleoconch whorl, the protruding part of the outer lip margin, and the coloration of the visceral mass that can be seen through the translucent shell in living specimens. H. peronellicola was attached to the host by inserting the proboscis into the host’s body, whereas no proboscis penetration was observed in H. tokunagai. Keywords: eulimids, Hypermastus peronellicola, Hypermastus tokunagai, parasitic gastropod, redescription, sand dollar Introduction Gastropods belonging to the family Eulimidae are known to infest echinoderms (Warén, 1980, 1984; Bouchet & Warén, 1986; Jangoux, 1990). The eulimid genus Hypermastus is a small group that is exclusively parasitic on irregular sea urchins (sand dollars and heart urchins). The genus was established by Pilsbry (1899) and currently contains 30 valid species (e.g. Warén, 1980; Warén & Crossland, 1991; Warén et al., 1994). Although several eulimid species have been erroneously included in the genus Hypermastus in Japan (Higo et al., 1999; Hori, 2000), only two species have been reported from sand dollars: H. peronellicola (Kuroda & Habe, 1950) and H. tokunagai (Yokoyama, 1922). H. peronellicola was originally described as Balcis peronellicola from the clypeasteroid Peronella japonica in Yura Bay, Wakayama Prefecture (Kuroda & Habe, 1950), and transferred to the genus Hypermastus by Warén & Crossland (1991: 100). Matsuda et al. (2008) reported in detail the ecology of H. tokunagai, which is parasitic on Scaphechinus mirabilis in the Seto Inland Sea. This eulimid was originally described as Eulima (Leiostraca) tokunagai Yokoyama, 1922, an Upper Pliocene fossil from Chiba Prefecture (Yokoyama, 1922). Ikebe (2008: 35, fig. 1127) also recorded and illustrated the species from Suiken, Wakayama Prefecture as “Melanella tokunagai (Yokoyama, * Corresponding author: [email protected] 26 H. Matsuda et al. 1922)”. Hypermastus tokunagai and H. peronellicola are, however, very similar to each other in the shell morphology, and it is necessary to compare these eulimids in more detail, and quantify the differences of their morphology for further study of the genus. In 2008, we collected eulimids from two clypeasteroids, P. japonica and S. mirabilis, in coastal waters of the western Seto Inland Sea. Based on examination of the shell morphology of the specimens and comparison with relevant type specimens, we concluded that the species are specifically distinct and represent the two above-mentioned nominal taxa. We present herein their redescription, and discuss ecological relationships between these eulimids and their hosts. Materials and Methods Thirty specimens of eulimids were collected each from Peronella japonica and Scaphechinus mirabilis in the subtidal zone in the western Seto Inland Sea, Yamaguchi Prefecture, from July 2 to September 27, 2008. These specimens were fixed in 70% ethanol or 10% neutralized formalin and some were also brought alive to the laboratory of Hiroshima University to document and compare the coloration of the soft parts that are visible through the translucent shell. The “syntype” (Warén & Crossland, 1991) of Balcis peronellicola was examined at the National Museum of Nature and Science, Tokyo (NSMT). For Eulima (Leiostraca) tokunagai, digital images of the lectotype taken from several directions by Dr. T. Sasaki, the University Museum, the University of Tokyo (UMUT) were examined. Some specimens that were labeled as “Balcis peronellicola” or attached to sand dollars were loaned from the NSMT. In addition, eulimid specimens taken from the beach and from P. japonica and S. mirabilis from other localities (e.g. Ishikawa, Kanagawa, Wakayama, Hiroshima, Tokushima and Okinawa Prefectures) were examined for geographical distribution. In the laboratory, the external morphology was examined under a stereomicroscope. For some specimens from the western Seto Inland Sea, the shape of the protoconch and imperceptible microsculpture on the surface were observed with a JSM-6390LV scanning electron microscope (SEM). Measurements of the shell were made from digital images that were taken by an Olympus DP20 digital camera with an Olympus SZX10 compound microscope. Measurement planes generally followed Hori et al. (2002: figs. 1–3). In addition, dimensions of the outer lip margin, which are regard as important taxonomical characters in eulimids (Warén, 1984; Bouchet & Warén, 1986), were measured. Accordingly the following measurements were made (Fig. 1): Entire view of shell (Fig. 1A): shell length (SL), body whorl length (BL), aperture length (AL), aperture width (AW), shell width (SW); aperture (Fig. 1B): columella width (CW), aperture anterior extension (distance from anteriormost end of columellar lip and anteriormost end of aperture) (AE), aperture lateral protrusion (distance from adapical end to lateral end of outer lip margin) (AP); posterior portion of shell (Fig. 1C): protoconch length (PL), protoconch width (PW), length of the first to eighth teleoconch whorl (TL1–TL8), width of the first to eighth teleoconch whorl (TW1–TW8); outer lip margin (Fig. 1D): chord of the outer lip margin (distance from the posterior extremity to the anterior extremity of the outer lip margin) (OL), length of protrusion of outer lip margin (distance from OL to the outer lip margin) (LP), distance from posterior extremity of outer lip margin to widest point of LP (L), distance from OL to outer lip margin at 1/4 of OL (L1), distance from OL to outer lip margin at 1/2 of OL (L2); distance from OL to outer lip margin at 3/4 of OL (L3). The specimens whose outer lip margin was damaged were not used for measurements of the margin. The relationship among certain shell dimensions (e.g. SL and BL, SL and AL, SL and AW, SL and SW) was treated with general equation Y = aXb, and parameters of equivalent allometric regression log Y = log a + b log X were analyzed by ANCOVA. Redescriptions of Hypermastus peronellicola and H. tokunagai 27 Fig. 1. Shell dimensions measured in this study. A. Entire view of shell. B. Aperture. C. Posterior portion of shell. D. Lateral view of outer lip margin. Taxonomy Family Eulimidae Philippi, 1853 Genus Hypermastus Pilsbry, 1899 Hypermastus peronellicola (Kuroda & Habe, 1950) (Figs. 2A, 3A, C, D) Balcis peronellicola Kuroda & Habe, 1950: 60; Habe, 1952: 77, pl. 6, fig. 17; Yamamoto, 1963: 81–83, pl. 1, figs. 1–3; Habe, 1976a: 158; Habe, 1976b: 2; Inaba & Oyama, 1977: 95, pl. 5, fig. 16. Hypermastus peronellicola (Kuroda & Habe, 1950) — Warén & Crossland, 1991: 100–101, fig. 7. Melanella peronellicola (Kuroda & Habe, 1950) — Higo et al., 1999: 58, 189; Hori, 2000: 348–349, pl. 173. Redescription: Shell tall, slender [SW/SL = 0.25–0.32 (0.29 ± 0.02; n = 30)], conical, and straight (Fig. 3A-a). External surface glossy, translucent, almost perfectly smooth and polished without any coloration, sometimes with thin, non-aligned incremental scars, variably situated on each whorl. Protoconch convex, colorless, consisting of slightly more than 2.5 whorls, with visible height of 0.28–0.38 mm (0.35 ± 0.02; n = 30), depending on concealment by teleoconch whorl (Fig. 3A-b). Protoconch whorls separated from teleoconch by moderately impressed incremental scar. Teleoconch whorls slightly convex, 5.9–9.0 in number, depending on shell size. Teleoconch whorls gradually increasing in diameter towards body whorl, demarcated by shallow suture. Width/length ratio of each whorl gradually decreasing toward body whorl [TW1/TL1 = 0.99 ± 0.07 (n = 30), TW2/TL2 = 0.89 ± 0.05 (n = 30), TW3/TL3 = 0.87 ± 0.04 (n = 30), TW4/TL4 = 0.83 ± 0.04 (n = 30), TW5/TL5 = 0.80 ± 0.04 (n = 28), TW6/TL6 = 0.81 ± 0.05 (n = 8), TW7/ TL7 = 0.79 ± 0.04 (n = 5), TW8/TL8 = 0.67 (n = 1)]. Body whorl relatively large, constituting almost half of shell length [BL/SL = 0.46–0.55 (0.52 ± 0.02; n = 30)], with round periphery and base. Aperture pear-shaped, offset laterally [AP/AW = 0.07–0.24 (0.17 ± 0.04; n = 30)] and extending slight anteriorly beyond base [AE/AL = 0.07–0.16 (0.12 ± 0.02; n = 30)]. Columella 28 H. Matsuda et al. Fig. 2. Hypermastus peronellicola and H. tokunagai. A. Lectotype of H. peronellicola, NSMT-Mo 39847, SL = 4.32 mm. B. Lectotype of H. tokunagai, UMUT-CM 21021, SL = 7.17 mm. a, ventral view of shell; b, posterior portion of shell; c, lateral view of outer lip margin. narrow [0.07–0.17 mm (0.11 ± 0.03; n = 30)], and columellar parietal wall slightly curved or slightly angulated at transition. Outer lip margin sigmoid in lateral view, prosocline, protruding just above mid-point [L/OL = 0.32–0.48 (0.40 ± 0.04; n = 29)] (Fig. 3A-c). Ventral end of suture line almost straight. In live specimens, apical portion of visceral mass reddish orange or brown seen through translucent shell, more clearly in larger specimens (Fig. 3G). Foot relatively large and functional. Small black eyes located at outer base of tentacles visible through shell. Proboscis fully retractile. Operculum oval, thin, transparent and colorless.
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