Nat. Hist. Res., Vol. 8 No. 1: 33-63, March 2004

New Records of Four Grapsoid (Crustacea: : Brachyura) from , with Notes on Four Rare Species

Tomoyuki Komai1', Takashi Nagai2!, Akifumi Yogi3!, Tohru Naruse2', Yoshihisa Fujita2' and Shigemitsu Shokita2' 1: Natural History Museum and Institute, Chiba 955-2 Aoba-cho, Chuo-ku, Chiba 260-8682, Japan 2lDepartment of Marine and Environmental Sciences, Graduate School of Engineering and Science, University of the Ryukyus 1 Senbaru, Nishihara-cho, Okinawa 903-0213, Japan 3;Okiden Sekkei Co., Ltd. 5-2-1 Makiminato, Urasoe-shi, Okinawa 901-2131, Japan

Abstract Four species of semiterrestrial grapsoid crabs are recorded from Japanese waters for the first time based on material from the Ryukyu Islands. These are: Pachygrapsus planifrons De Man, 1888 (); Nanosesarma andersoni (De Man, 1888), Sesarmoides kraussi (De Man, 1887), and Stelgistra stormi (De Man, 1895) (). Supplementary information is provid­ ed for four other little known species, three sesarmids Clistocoeloma villosum (A. Milne-Edwards, 1869), Metasesarma obesum (Dana, 1851) and Perisesarma semperi (Burger, 1893), and one varunid Ptychognathus capillidigitatus Takeda, 1984. The generic assignment of C. villosum is briefly discussed. The range of P. capillidigitatus is extended south to Iriomote Island.

Key words: Crustacea, Decapoda, Brachyura, , , new record, Ryukyu Islands, Japan

treated in this paper: one grapsid, Pachygrap­ Introduction sus planifrons De Man, 1888; five sesarmids, The fauna of the coastal re­ Clistocoeloma villosum (A. Milne-Edwards, gions of the Ryukyu Islands, particularly 1869), Metasesarma obesum (Dana, 1851), that of mangroves and estuaries, has been Nanosesarma andersoni (De Man, 1888), Peri­ relatively poorly investigated in spite of an semperi (Burger, 1893), Sesarmoides expected high diversity. Nakasone and Irei kraussi (De Man, 1887), and Stelgistra stormi (2003) compiled information on the known (De Man, 1895); and one varunid Ptychogna­ blackish water and mangrove fauna of thus capillidigitatus Takeda, 1984. Of them, the Ryukyu Islands, but indicated that their N. andersoni P. planifrons, Sesarmoides krau­ faunal inventory was still incomplete. Docu­ ssi and Stelgistra stormi are new to the Japa­ mentation of coastal fauna is particularly im­ nese fauna. A minor southerly range exten­ portant for conservation, as coastal regions sion is also recorded for the varunid Ptycho­ are readily affected by human activities. gnathus capillidigitatus. Since 1994, the authors have made extensive collections of decapod in various Material and methods locations in the Ryukyu Islands, i.e. Amami, The specimens examined in this study are Okinawa, Kerama, Kume-jima, Miyako, Ishi- deposited in the Natural History Museum gaki and Iriomote islands. The present paper and Institute, Chiba, Japan (CBM), and the is intended as the first of a series of papers Ryukyu University Museum, Fujukan, Oki­ that provide new taxonomic information on nawa, Japan (RUMF). The carapace width, the grapsoid crab families from the Ryukyu abbreviated as cw, was taken from the widest Islands. The following eight species are position, which is varies according to species.

— 33 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

The partially fused anterior three thoracic tidal, under stone, 11 July 1998, coll. T. sternites are termed here the "anterior ster­ Komai, 1 male cw 6.7 mm, 1 female cw 7.2 nal plate." Synonymies are restricted to the mm (CBM-ZC 7166). Kuroshima Island: Iko original citation and references providing Pier, upper intertidal, under stone, 11 July significant information on morphology and/ 1998, coll. T. Komai, 1 male cw 8.7mm or distribution. (CBM-ZC 5439). Identification of species belonging to the Comparative material. Thailand, Phuket: Grapsidae, Sesarmidae and Varunidae is Patong Beach, upper intertidal. 10 November sometimes very difficult. Therefore, detailed 1995, coll. T. Komai, 4 males cw 6.6-7.9 mm, descriptions are given for the five little 4 ovigerous females cw 5.7, 8.0 mm 'CBM-ZC known species, Pachygrapsus planifrons, 6078); Cape Panwa, beach in Phuket Marine Clistocoeloma villosum, Nanosesarma ander- Biological Centre, upper intertidal. under soni, Perisesarma semperi and Sesarmoides stone, 16 October 1990, coll. T. Komai. 5 ovi­ kraussi, for clear documentation of the spe­ gerous females cw 4.3-6.6 mm 'CBM-ZC cific identity and for providing information 6319). on supposedly important characters not fully Description. Carapace (Figs. 1. 2A< trape­ mentioned in previous literature (i.e., anten­ zoidal in dorsal view, greatest width between nae, epistome, buccal cavity, suborbital exorbital teeth about 1.3 times carapace region and thoracic sternites). Illustrations length. Front about 0.6 exorbital width, pro­ of selected parts are given for Stelgistra duced anteriorly as thin eave overhanging stormi, Metasesarma obesum and Ptychogna- antennae, anterior margin very faintly sinu­ thus capillidigitatus in order to facilitate iden­ ous; preorbital angle broadly rounded. tification of these species and to supplement Dorsal surface of carapace naked, shining. the previous descriptions. with 2 widely separated postfrontal lobe? on either side of midline, mesial lobe with Taxonomic accounts convex anterior margin; regions poorly in­ dicated; 1 lateral branchial striae originating Superfamily Grapsoidea from exorbital tooth. Upper orbital margin Family Grapsidae concave, slightly oblique; lower orbital margin (Fig. 2B) sinuous in anterior view, Pachygrapsus planifrons De Man, 1888 smooth; inner orbital tooth triangular, some­ (Figs. 1A, 2) what directed laterally, not reaching to pre­ Pachygrapsus planifrons De Man, 1888: 368, orbital angle of front. Exorbital angle pro­ pi. 10, fig. 2; Tesch, 1918: 77 ; Ward, 1934: duced as a sharp tooth directed ante- 25 ; Edmondson, 1959: 173, figs. 10b, lla-e; rolaterally. Lateral margin slightly concave, Garth, 1965: 30, figs. 15, 16; Dai and Yang, and slightly converging posteriorly, without 1991: 510, fig. 261-1, pi. 65, fig. 4. trace of epibranchial tooth. ? Pachygrapsus planifrons - Hartnoll, 1975: Antennal peduncle (Fig. 2B) obliquely set; 307, table 1. second segment with elongate dorsolateral projection; flagellum relatively long. Material examined. Ishigaki Island: Shira- Epistome (Fig. 2B) with sharp lateral ridge ho, upper intertidal, under stone, 1 July 2001, just inferior to antennule on either side. An­ coll. T. Komai, 2 males cw 7.7, 9.0 mm terior part of buccal cavity concave, without (CBM-ZC 7051); same locality, 14 July 1998, median ridge. coll. T. Komai, 1 male cw 7.6 mm (CBM-ZC Ischium of third maxilliped with pro- 7131); Tonoshiro, upper intertidal, under ximomesial angle somewhat produced; stone, 10 July 2000, coll. T. Komai, 3 males merus shorter than ischium, with strongly cw 7.9-92 mm (CBM-ZC 7407). Iriomote produced mesial margin, anterolateral Island: Hoshizuna Beach, under stone, upper margin broadly rounded; exopod slender, intertidal, 5 July 2001, coll. T. Komai, 4 males with well-developed flagellum. cw 6.0-7.6 mm, 2 females cw 6.9, 7.8 mm Male chelipeds (Fig. 1A) subequal, moder­ (CBM-ZC 7067); Haemida Beach, upper inter­ ately large, robust. Merus (Fig. 2C) with

— 34 - - New Records of Grapsoid Crabs from Japan

Fig. 1. A, Pachygrapsus planifrons De Man, 1888, male from Tonoshiro, Ishigaki Island (cw 9.2 mm; CBM-ZC 7407); B, Clistocoeloma villosum (A. Milne Edwards, 1869), female from Hoshidate, Iriomote Island (cw 15.3 mm; CBM-ZC 7095); C, Metasesarma obesum (Dana, 1851), male from Hoshizuna Beach, Iriomote Island (cw 16.4 mm; CBM-ZC 7403); D, Nanosesarma andersoni (De Man, 1887), male from Hoshidate, Iriomote Island (cw 8.0 mm; CBM-ZC 7091); E, Perisesarma semperi (Burger, 1893), male from Hoshidate, Iriomote Island (cw 27.8 mm; CBM-ZC 7192); F, Stelgistra stormi (De Man, 1895), ovigerous female from Hoshizuna Beach, Iriomote Island (cw 16.2 mm; CBM-ZC 7378); G, Sesarmoides kraussi (De Man, 1887), male from Shirahama, Iriomote Island (cw 23.7 mm; CBM-ZC 7401).

— 35 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

0.5 mm 0.5 mm

36 New Records of Grapsoid Crabs from Japan inner ventral margin bearing row of sharp telson subtriangular, slightly longer than teeth increasing in size distally; dorsal sur­ basal width, lateral margins straight and ter­ face rounded, rugose; outer surface slightly minal margin rounded. Female abdomen rugose. Carpus (Fig. 2D) with inner angle (Fig. 21) broad, subcircular in general outline; produced as large sharp tooth; dorsal surface telson very wide, about 4.5 times as wide as with scattered, longitudinal or oblique short long. ridges. Palm (Fig. 2E) with microscopically First gonopod (Fig. 2J) relatively slender, granular surfaces; no trace of ventral ridge nearly straight in mesial view; terminal pro­ on outer surface. Tips of fingers somewhat cess (Fig. 2K) elongate, bearing several irreg­ spoon-shaped, each bordered by corneous ular lobules. claw, bearing subterminal tuft of short setae. Coloration. Carapace mottled with brown. Fixed finger straight, cutting edge with row Cheliped generally brown, paler ventrally, of small, rounded teeth. Dactylus weakly fingers also paler. Ambulatory legs ob­ curved, shorter than palm, surfaces micro­ scurely banded by brown. scopically granular, cutting edge with very Distribution. Widely distributed in the low teeth; narrow hiatus between fingers. Indo-Pacific Ocean: Tanzania, Mergui Archi­ Female chelipeds (Fig. 2F) smaller than pelago (type locality), Phuket, Christmas male chelipeds. Fixed finger with distinct Island, Indonesia, Hainan Island in the South longitudinal ridge on outer surface. Sub- China Sea, southern Ryukyu Islands (Ishi­ terminal setae on fingers longer than those of gaki, Iriomote, and Kuroshima islands), males. Dactylus slightly longer than palm. Davao Gulf of Mindanao in the Philippines, Ambulatory legs (Figs. 1A, 2G) moderately Palau Islands, Hawaii, and Clipperton Islands. long, fourth pereopod longest. Coxa of (De Man, 1888; Tesch, 1918; Tweedie, 1936; fourth pereopod with dense tufts of soft Ward, 1941; Edmondson, 1959; Garth, 1965; setae. Each merus armed with sharp sub- Hartnoll, 1975; Takeda, 1989; Dai and Yang, distal tooth on both anterior and posterior 1991; this study). margins, merus of fourth pereopod about Ecological notes. This species was com­ 2.2-2.3 times as long as wide; dorsal surfaces monly found in the upper intertidal zone of of meri naked, but with traces of short trans­ rocky areas. The crabs were active during verse ridges; anterior margins sharply edged, the day. When disturbed, they very quickly each with row of corneous spinules (only few run away. spinules present on fifth pereopod); posterior Remarks. The present specimens from the margin not sharply edged. Distal three seg­ Ryukyu Islands agree well with the previous ments slender. Carpi smooth on dorsal sur­ descriptions of Pachygrapsus planifrons by De faces. Propodi each with few long corneous Man (1888), Garth (1965) and Dai and Yang spinules on inner margin. Dactyli 0.6-0.8 (1991) and with the comparative material times as long as propodi, each with double from Phuket, Thailand. This species is char­ row of long corneous spinules on inner acterized by a suite of characters, including margin and some subterminal corneous spi­ the rather smooth dorsal surface of the cara­ nules on outer surface. pace having a few obliquely transverse Thoracic sternum smooth, naked. striae, the merus of the fifth pereopod armed Male abdomen (Fig. 2H) moderately broad­ only with one tooth on the posterior margin, ly triangular in general outline; outer surface the posteriorly convergent lateral margins of smooth, naked; sixth somite slightly shorter the carapace lacking an epibranchial tooth, than fifth, with straight lateral margins; and the possession of a brush-like tuft of

Fig. 2. Pachygrapsus planifrons De Man, 1888. A-E, G, H, J, K, male from Tonoshiro, Ishigaki Island (cw 9.2 mm; CBM-ZC 7407); F, I, female from Hoshizuna Beach, Iriomote Island (cw 6.9 mm; CBM-ZC 7067). A, carapace and eyes, dorsal view; B, anterior part of carapace and cephalic appendages, left side, anterior view; C, ischium and merus of left cheliped, ventral view; D, carpus of left cheliped, dorsal view; E, F, left chela, outer view; G, left fifth pereopod, dorsal view; H, I, abdomen, ventral (outer) view; J, left first gonopod, lateral view; K, same, distal part, mesial view.

— 37 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita setae at the tip of the cheliped fingers. Re­ Chiromantes villosum - Nomoto et al, 1999: 9, gardless of sex, these characters immediately pi. 1-6 ; Kishino et al, 2001a: 17, pi. 2, 2; separate P. planifrons from other known East Kishino et al, 2001b: 127; Shokita et al, Asian species of Pachygrapsus Randall, 1840, 2002: 78, photo 4A-1. P. crassipes Randall, 1840, P. fakaravensis Rathbun, 1907, P. minutus A. Milne-Edwards, Material examined. Miyako Island: Ohura 1873, and P. plicatus (H. Milne Edwards, Bay, mangrove swamps, upper intertidal, 1837). The first gonopod of the male is also mud bottom under stone, 22 December 1996, distinctive in having a multi-lobed terminal coll. T. Komai, 1 male cw 10.5 mm (CBM-ZC process. 3347); Irie, mangrove swamps, upper inter­ Our specimens represent the first record of tidal, mud bottom under stone, coll. T. Komai, P. planifrons from East Asian waters, al­ 1 male cw 12.8 mm (CBM-ZC 3352). Iriomote though this species is widely distributed in Island: Yonada-gawa estuary, Hoshidate, the tropical Indo-West Pacific. In Iriomote burrow under boulder, upper intertidal, 10 and Ishigaki islands, this species is common July 2001, coll. T. Komai, 2 females cw 8.0, in the upper intertidal zone of rocky areas. 15.3 mm, 1 ovigerous female cw 16.4 mm Two other grapsid species, Metopograpsus (CBM-ZC 7095); Uehara Beach, under coral thukuhar (Owen, 1839) and M. messor (Fors- rocks, upper intertidal, 16 July 2000, coll. T. kal, 1775), are found sympatrically. Pachy­ Komai, 1 male cw 10.3 mm, 1 female cw 8.7 grapsus planifrons can be easily mistaken for mm (CBM-ZC 7405); , upper juveniles or young individuals of Metopo­ margin of mangrove forest, 26 February grapsus species, and perhaps such confusion 2003, coll. T. Nagai and A. Yogi, 1 male cw is why this species has not been previously 12.6 mm, 1 female, cw 14.7 mm (RUMF-ZC- recorded from Japanese waters. 129). Comparative material Clistocoeloma mer- Family Sesarmidae guiense De Man, 1888: Sungei Buloh man­ groves, Singapore, May 1998, coll. T. Komai, Clistocoeloma villosum (A. Milne Edwards, 3 males cw 10.0-11.4 mm, CBM-ZC 6106. 1869) Clistocoeloma sinense Shen, 1933. Obitsu- (Figs. IB, 3) gawa Estuary, Kaneda, Kisarazu, Chiba Pre­ Sesarma villosum A. Milne-Edwards, 1869: 31; fecture, Japan, 7 May 2000, coll. T. Furota, 2 Rathbun, 1907: 35. males cw 12.1, 13.0 mm, 5 females cw 11.0— Sesarma villosa-De Man, 1887: 644; Lenz, 16.0 mm, CBM-ZC 6453. 1910:560. Description. Carapace (Figs. IB, 3A) rectan­ Sesarma (Sesarma) villosa-De Man, 1895: gular; greatest width across middle of cara­ 153; 1898: pi. 29, fig. 30, 30a~e. pace, width about 1.2 times length. Front Sesarma (Holometopus) villosa - Tesch, 1917: 0.60-0.65 times as wide as exorbital width, 208, pi. 17, fig. 2 ; Sendler, 1923 : 35. moderately deflexed, with faint median Sesarma (Holometomus) villosum - Crosnier, notch; preorbital angle rounded. Dorsal sur­ 1965: 55, figs. 75, 76, 77a, 78 ; McNeil, 1968: face of carapace with numerous very short 79. stiff setae often grouped in small patches, Clistocoeloma villosum - Davie, 2002: 221. always covered with mud and sediment;

Fig. 3. Clistocoeloma villosum (A. Milne-Edwards, 1869). A, B, F-H, K, female from Hoshidate, Iriomote Island (cw 16.4 mm; CBM-ZC 7095); C-E, L, M, male from Irie, Miyako Island (cw 12.8 mm; CBM-ZC 3347). A, carapace and eyes, dorsal view (setae omitted on left side); B. anterior part of carapace and cephalic appendages, left side, anterior view (setae partially omitted); C, F, left chela, outer view; D, same, dorsal view; E, inner dorsal margin of left palm, showing pectinated crest, outer view; G, carpus and chela of left cheliped, dorsal view; H, left fifth pereopod, dorsal view; I, anterior sternal plate, ventral view (setae omitted); J, abdomen; K, sixth abdominal somite and tclson, ventral view (setae omitted); L, left first gonopod, lateral view; M, same, distal part, mesial view (setae removed).

— 38 — New Records of Grapsoid Crabs from Japan

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— 39 T. Komai, T, Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita post-frontal lobes inconspicuous; regions on bercle transversely ovate, proximal tubercles dorsal surface poorly indicated; lateral bran­ each with slightly compressed summits; with chial striae absent. Exorbital angle weakly microscopic striae running parallel to long­ produced, blunt. Lateral margin of carapace itudinal axis of dactylus; outer surface of slightly sinuous with anterior part slightly dactylus smooth; cutting edge with row of convex, without trace of epibranchial tooth. blunt teeth; narrow hiatus between fingers. Upper orbital margin concave, slightly Chelipeds of females (Fig. IB) somewhat oblique; lower orbital margin (Fig. 3B) smaller than those of males. Palm (Fig. 3F, G) minutely granular, covered with very short with dorsal carina finely tuberculate, but setae; inner orbital tooth rather large, broad­ lacking pectinated corneous teeth. Dactylus ly rounded, but not reaching to preorbital (Fig. 3F, G) with 12-16 very small, rounded angle of front. tubercles on proximal 0.6, tubercles becom­ Antennule and antenna (Fig. 3B) conti­ ing obsolete toward tip. guous. Basal antennular segment sub- Ambulatory legs (Figs. IB, 3H) moderately rectangular. Antenna set obliquely; second short, fourth pereopod longest, each covered segment with elongate distolateral process with short stiff setae. Coxae with short stiff overreaching distal margin of third segment; setae on ventral surfaces, but lacking tufts of flagellum relatively long, extending into long setae. Each merus armed with low, orbit. blunt subdistal projection on anterior Epistome (Fig. 3B) concave on outer sur­ margin, but unarmed on posterior margin, face; ventral margin smooth, protuberant merus of fourth pereopod about 2.2-2.3 times medially, extending onto anterior part of as long as wide; dorsal surfaces of meri with buccal cavity as short ridge. sparse very short setae and scattered small Third maxilliped with well-developed patches of short setae; anterior and posterior flagellum on exopod. margins not sharp, each with numerous short Male chelipeds (Fig. 3C, D) subequal or stiff setae. Carpi each with 1 or 2 lines of slightly unequal, moderately robust. Merus short stiff setae on dorsal surface. Propodi with inner ventral margin weakly tubercu- stout, each with numerous short stiff setae late, but without conspicuous tooth; dorsal and rows of longish setae on surfaces, but margin sharply carinate, with row of very without corneous spinules. Dactyli 0.7-0.8 short stiff setae and/or row of granules; outer times as long as propodi, lacking corneous surface apparently smooth. Carpus with spines, but with covering of short stiff setae inner angle weakly produced in rounded and rows of longish setae. lobe, inner margin not clearly delimited; Anterior sternal plate (Fig. 31) broadly tri­ dorsal surface with scattered small patches angular, with concave lateral margins; sur­ of short setae; inner face narrow, unarmed. face slightly concave, with few short setae. Palm with 1 sharp, partially pectinated ridge Fourth sternite with short to long setae par­ over entire length of dorsal surface (Fig. 3E), ticularly numerous along margins; suture be­ composed of 20-30 small corneous teeth tween third and fourth sternites lined by medially and becoming small tubercles dis- long setae. Abdominal cavity in males nearly tally and proximally; outer surface of palm reaching to suture between third and fourth convex, smooth, without trace of ventral sternites. ridge; inner surface with weak elevation Male abdomen (Fig. 3J) wide, abruptly nar­ lined with row of small tubercles adjacent to rowed at base of telson; surface covered with dorsal margin, otherwise smooth; ventral sparse very short setae. Sixth abdominal surface (including fixed finger) also smooth; somite almost as long as fifth somite, with fixed finger straight, terminating in weakly strongly convex lateral margins. Telson dis­ bifid corneous claw, cutting edge with row of tinctly longer than basal width, lateral mar­ small, rounded teeth. Dactylus slightly gins evenly convex and terminal margin longer than palm, terminating in simple cor­ rounded. neous claw; dorsal surface bearing 20-25 tu­ Female abdomen with telson (Fig. 3K) bercles becoming lower toward tip; each tu­ evenly rounded, as long as basal width and

40 — New Records of Grapsoid Crabs from Japan

longer than mid-line length of sixth abdomi­ cies to Chiromantes Gistel, 1848, perhaps fol­ nal somite. lowing the subgeneric assignment to Holo­ First gonopod (Fig. 3L) stout, nearly st­ metopus by Tesch (1917) and Crosnier (1965) raight, subdistal portion slightly inflated; ter­ [for nomenclatural amendments from Holo­ minal process (Fig. 3M) short, truncate dis- metopus to Chiromantes, see Holthuis (1977)]. tally, with shallow notch distally. Ng and Liu (1999) commented that the real Coloration. Generally muddy color; palm of Chiromantes might be restricted for C. cheliped purplish; facial setae dark brown. haematocheir (De Haan, 1833) and also sug­ Distribution. Widely distributed in the gested that Chiromantes villosum was more Indo-Pacific: Madagascar, Aceh, Sumatra, likely to be a species of Clistocoeloma on ac­ central and southern Ryukyu Islands count of the features of the carapace and (Miyako, Iriomote, Okinawa, and Amami- setose surfaces. In his catalogue of Austra­ Ohshima islands), in lian brachyuran crabs, Davie (2002) referred mainland of Japan, New Guinea, Queensland, this species to Clistocoeloma, although no de­ Australia, Caroline Islands, and Samoa Is­ tailed comments were given. We compared lands (type locality) (A. Milne-Edwards, 1869; our specimens with specimens of Clisto­ De Man, 1895; Rathbun, 1907; Tesch, 1917; coeloma merguiense De Man, 1888 and C. Sendler, 1923; Crosnier, 1965; McNeil, 1968; sinense Shen, 1933. Our examination has Nomoto et al, 1999; Kishino et al., 2001a; shown that this species is rather similar to Shokita et al, 2002; this study). the species of Clistocoeloma than those of Ecological note. Clistocoeloma villosum Chiromantes (sensu Serene and Soh, 1970; Ng occurs under stones of landward edges of and Liu, 1999) in the following points: the mangrove forests. Soil of the habitats is carapace and ambulatory legs are covered slightly wet, occasionally covered with fallen with short stiff setae, occasionally forming leaves. This species sometimes occurred prominent patches; and the palm of the cheli­ sympatrically with hippolytid shrimp Mer- ped is provided with a single pectinated crest guia oligodon (De Man, 1888) and sesarmid running the over entire length of the dorsal Sesarmoides kraussi in Iriomote Islands, and surface. One of the diagnostic characters of while in Okinawa Island, it was found togeth­ Clistocoeloma is the orbit excluding the er with Parasesarma acis Davie, 1993 (see Ng antennular fiagellum with an inner orbital et al, 2001 for possible synonymy with Para­ tooth that extends to the preorbital angle of sesarma tripectinis (Shen, 1940)). the front (e.g. Tesch, 1917; Serene and Soh, Remarks. The specimens at our disposal 1970). However, it has been found that the agree with the previous accounts of this spe­ development of the inner orbital tooth is var­ cies [De Man (1895), as Sesarma (Sesarma) iable in Clistocoeloma merguiense and C. villosa; Tesch (1917), as Sesarma (Holo- sinense, and thus the antennular fiagellum is metopus) villosa; Crosnier (1965), as Sesarma occasionally in touch with the orbit in the (Holometopus) villosum; Nomoto et al. (1999) latter two species. The generic significance and Kishino et al. (2001a), as Chiromantes of this antennule-orbit character is question­ villosum]. able. Therefore, we follow Davie (2002) to assign the present species to Clistocoeloma. The generic assignment of this species has been subject to disagreement. Originally, The genus Clistocoeloma contains the fol­ this species was assigned to Sesarma Say, lowing seven species other than C. villosum: 1817 (A. Milne Edwards, 1869), and later C. amamaparense Rahayu and Takeda, 2000; Tesch (1917) and Crosnier (1965) referred it C balansae A. Milne-Edwards, 1873 (type to the subgenus Holometopus H. Milne Ed­ species of the genus); C. lanatum (Alcock, wards, 1853. Tesch (1917) mentioned that S. 1900); C. merguiense; C sinense; C. suvaense villosum was clearly different from species of Edmondson, 1951; and C. tectum (Rathbun, Clistocoeloma A. Milne-Edwards, 1873 in the 1914). Some species, such as C. balansae, C. structure of the orbit. Japanese authors lanatum and C. suvaense, needs to be redescr- (Nomoto et al, 1999; Kishino et al, 2001a, ibed, as the previous descriptions are insuffi­ 2001b; Shokita et al, 2002) referred the spe­ cient. Nevertheless, C. villosum may be read-

41 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita ily distinguished from the other seven spe­ postfrontal lobes present on either side of cies by the absence of epibranchial teeth or distinct median groove, mesial lobe distinct, lobes on the carapace. The latter species all but lateral lobe only faintly denned. Front have one or two epibranchial teeth on the deep, vertically deflexed, somewhat spatu- carapace. late, covering antennae in anterior view, frontal width more than half of greatest Metasesarma obesum (Dana, 1851) carapace width; free edge slightly convex in (Figs. 1C, 4) anterior view, preorbital angle slightly pro­ Sesarma obesum Dana, 1851: 252; 1852; 356; duced, rounded; frontal width about 0.6 of 1855: pi. 22, fig. 10. exorbital width. Lateral margin of carapace Metasesarma rousseauxi H. Milne Edwards, unarmed, convex in anterior part, slightly 1853: 188; 1854: 158, pi. 10, fig. 1, la-c; converging backwards in posterior part; ex- Crosnier, 1965: 73, figs. 166-120; Ng and orbital tooth directed forward. Upper orbital Davie, 1995: 39, fig. 5a~c; Watanabe and margin strongly oblique; lower orbital Wada, 2002: 112, figs. 3, 4. margin (Fig. 4B, C) granulate, with numerous Metasesarma granulans Heller, 1862: 522. very short setae; inner orbital tooth triangu­ Metasesarma rugulosa Heller, 1865: 65. lar, extending to inner side of preorbital lobe Sesarma (Metasesarma) rousseauxii - Pilai, in adults, thus antenna completely excluded 1951:37. from orbit. Sharp carina extending from ex- Chiromantes obesum - Ng and Liu, 1999: 229. orbital tooth into orbit. Metasesarma obesum - Ng and Schubert, Antennule and antenna (Fig. 4C) contigu­ 2003: 426, figs. 1-3. ous, visible only in ventral view. Basal seg­ ment of antennule transversely elongate sub- Material examined. Iriomote Island: rectangular; flagellum greatly reduced. An­ Yonada-bashi, Hoshidate, shoreline grass­ tenna small, set obliquely; second segment land, coll. T. Komai, 1 male cw 8.7 mm with distolateral process reaching distal (CBM-ZC 7139); Sonai, terrestrial, under boul­ margin of fourth segment; flagellum absent. der, 10 July 2002, coll. T. Komai, 1 male cw Epistome (Fig. 4B, C) nearly flat on outer 11.3 mm, 1 ovigerous female cw 12.8 mm surface, with dense covering of short setae; (CBM-ZC 7402); Hoshizuna Beach, terrestrial, ventral margin smooth, protuberant medial­ under coral rocks, 13 July 2002, coll. T. ly, extending onto anterior part of buccal Komai, 2 males 15.3, 16.4 mm, 2 females 15.5, cavity as short ridge. 16.7 mm (CBM-ZC 7403). Miyako Island: Third maxilliped with exopod bearing ru­ Nikawadori, supralittoral coral pebble beach, dimentary flagellum in adults; outer margin 25 July 2002, coll. T. Naruse et al, 1 male cw of merus regularly convex 8.2 mm (RUMF-ZC-128). Male chelipeds (Fig. 1C) subequal. Ischium Description. Carapace (Fig. 1C, 4A) sub- with small granules on inner margin. Merus quadrate, relatively deep; dorsal surface with dorsal surface strongly convex, distinct­ naked, but covered with minute, low, squami- ly ridged; inner ventral margin smooth, with­ form tubercles on gastric region and minute out distinct tooth. Carpus with dorsal sur­ granules on cardiac and mesial part of bran­ face nearly smooth, inner angle somewhat chial region; posterolateral part of branchial produced, but not forming tooth or spine. region with numerous short oblique striae; 2 Chela (Fig. 4D, E) massive; dorsal surface of

Fig. 4. Metasesarma obesum (Dana, 1851). A-C, F-H, L, female from Hoshizuna Beach, Iriomote Island (cw 16.7 mm; CBM-ZC 7403); male from same lot, D, E, I, J, K (cw 16.4 mm). A, carapace and eyes, dorsal view (ornamentation on dorsal surface omitted); B, anterior part of carapace, left side, anterior view; C, anterior part of carapace and cephalic appendages, ventral view (setae omitted); D, F, left chela, outer view; E, same, dorsal view; G, left fifth pereopod, dorsal view; H, same, dactylus, dorsal view; I, coxae of left second to fourth pereopods (= first to third ambulatory legs), ventral view; J, anterior sternal plate, ventral view (setae partially omitted); K, abdomen, ventral (outer) view (setae omitted); L, sixth abdominal somite and telson, ventral view (setae partially omitted).

— 42 - New Records of Grapsoid Crabs from Japan

— 43 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita palm nearly smooth, inner margin distinctly First gonopod illustrated by Ng and Davie ridged, faintly granular; outer surface of (1995) and Watanabe and Wada (2002). palm convex, smooth; inner surface concave Coloration. See Ng and Davie (1995). dorsally, bearing some large but low tuber­ Distribution. Widely distributed in the cles; fingers moderately long, tips pointed; Indo-Pacific: Zanzibar, Madagascar, Lacca- cutting edges each with rudimentary teeth; dive Islands, Mergui Archipelago, Kerala no subterminal tuft of brush-like setae. Dac- (Travancore), India, Sri Lanka, Christmas tylus curved, without row of tubercles on Island, Indonesia, Sabah (type locality; neo- dorsal surface. type designation by Ng and Schubart (2003)), Chelipeds of females somewhat smaller Philippines, Taiwan, southern Ryukyu Is­ than those of males. Carpus rugose with lands (Iriomote and Miyako islands), Guam, short rows of granules on dorsal surface. New Guinea, and Tahiti (Dana, 1851; H. Milne Palm (Fig. 4F) with short longitudinal rows Edwards, 1853; De Man, 1888; Tesch, 1917; of granules on dorsal surface adjacent to Pilai, 1951; Crosnier, 1965; Ng and Davie, inner margin; fingers with row of small but 1995; Ng et al, 2001; Watanabe and Wada, distinct teeth on cutting edges. Dactylus 2002; Ng and Schubart, 2003; this study). straight, with small granules on dorsal sur­ Ecological notes. This species occurs in face proximally terrestrial boulder area somewhat remote Ambulatory legs (Fig. ID, 4G) relatively from the beach line. The crabs hide under short, rather slender. Meri each with 1 blunt stone or coral rock during the day. subdistal projection on anterior margin, but Remarks. Ng and Schubart (2003) dis­ unarmed on posterior margin. Carpi each cussed that Sesarma obesum Dana, 1851, was with distinctly carinate anterior margin; a senior synonym of Metasesarma rousseauxi dorsal surfaces each with 1 or 2 longitudinal H. Milne Edwards, 1853, and transferred Ses­ carinae. Propodi each with bristles arranged arma obesum to the genus Metasesarma H. in 3-4 irregular rows on outer surface and Milne Edwards, 1853. However, considering bristles and small corneous spines arranged the extensive geographical range of the spe­ in 2 rows on inner surface. Dactyli (Fig. 4H) cies, Ng and Schubart (2003) still suggested subequal in length to propodi, each with 3 that Metasesarma obesum may involve more rows of corneous spines or bristles on outer than one species and that Metasesarma rous­ surface and 2 rows of corneous spines or seauxi (type locality: Zanzibar) may yet prove bristles on inner surface; dorsal surface shal- to be different from Metasesarma obesum lowly concave, with row of short bristles ad­ (type locality: northern Sabah). Our adult jacent to inner margin. Tufts of long setae on specimens agree very well with the neotype anteroventral surfaces of coxae of third and of M. obesum figured by Ng and Schubart fourth pereopods (Fig. 41). (2003). In this study, we follow Ng and Anterior sternal plate transversely elon­ Schubart's (2003) account in applying the gate rectangular (Fig. 4J); surface nearly flat, name Metasesarma obesum for the Japanese with few short stiff setae. Fourth sternite material. with line of short stiff setae along margins. This species was recently reported from Abdominal cavity not reaching suture be­ Japanese waters for the first time by Wata­ tween third and fourth sternites in males. nabe and Wada (2002, as M. rousseauxi) based Male abdomen (Fig. 4K) triangular, rela­ on material from Iriomote Island. Our speci­ tively broad for sesarmids, distinctly narrow­ mens agree very well with the previous ac­ ed at base of telson; sixth somite slightly counts of M. obesum (cf. Crosnier, 1965; Ng longer than fifth somite, with strongly and Davie, 1995; Watanabe and Wada, 2002; convex lateral margins; telson longer than as M. rousseauxi), except for a small specimen broad and distinctly longer than sixth ab­ collected from Miyako Island (cw 8.2 mm; dominal somite. RUMF-ZC-128). The specimen from Miyako Female telson (Fig. 4L) evenly rounded, as Island has the antenna incompletely ex­ long as wide and shorter than mid-line length cluded from the orbit and a well-developed of sixth abdominal somite. flagellum on the exopod of the third maxilli-

— 44 New Records of Grapsoid Crabs from Japan

;. -c. These discrepancies are most probably male cw 7.3 mm, 1 female cw 7.8 mm -:ze-related, as that specimen is much smaller (CBM-ZC 7486). :r.an other specimens examined. Description. Carapace (Fig. ID, 5A) trape­ The genus Metasesarma is represented by zoidal with lateral margins somewhat con­ ".wo species, M. aubryi A. Milne Edwards, verging backward; dorsal surface nearly 1869 and M. obesum. Although M. aubryi has smooth, but with scattered short stiff setae been recorded from Taiwan (Ng et al, 2001), (setae more numerous in females than in there has been no record of M. aubryi as yet males), and having several long oblique from Japanese waters. Future collections striae on lateral part; 2 postfrontal lobes may eventually reveal the presence of M. present on either side of midline, both low, aubryi in the Ryukyu Islands. The two spe­ but distinctly demarcated. Front vertically cies are distinguished by several morpholog­ deflexed, but not covering antennae in ical features (Ng and Davie, 1995): the frontal anterior view; preorbital angle rounded. width is less than half the greatest carapace Lateral margin with trace of epibranchial width in M. aubryi, rather than more than it tooth (Fig. 5B). Upper orbital margin strong­ in M. obesum; the lateral margin of the ex- ly oblique; lower orbital margin (Fig. 5C) orbital angle is slightly diverging backward smooth; inner orbital tooth triangular, some­ in M. aubryi, while subparallel to the long­ what directed laterally, reaching only half­ itudinal axis of the carapace in M. obesum; way between lower orbital margin and pre­ the lateral margin of the merus of third max- orbital angle of front. illiped bears a broad and low lobe in M. Antennule and antenna (Fig. 5C) contigu­ aubryi, rather than regularly convex in M ous. Basal segment of antennule subrectan- obesum; and the terminal process of the first gular. Antenna set obliquely; second seg­ gonopod is more strongly bent in M. aubryi ment with elongate distolateral process than in M. obesum. reaching distal margin of third segment; Metasesarma obesum is easily distinguished flagellum relatively long, extending into from other local sesarmid crab by the verti­ orbit. cally deflexed front covering the very Epistome (Fig. 5C) nearly flat on outer sur­ narrow antennal region in anterior view, the face, without covering of dense setae; ventral antenna excluded from the orbit, the un­ margin granular, protuberant medially, ex­ armed lateral margin of the carapace, the tending onto anterior part of buccal cavity as rudimentary exopodal flagellum of the third short ridge. maxilliped, and the absence of pectinated Third maxilliped with exopod bearing crests on the cheliped palm. well-developed flagellum. Male chelipeds (Fig. ID) subequal. Ischium Nanosesarma andersoni (De Man, 1887) with 1 conspicuous tooth on inner margin. (Figs. ID, 5) Merus with dorsal surface strongly convex, Sesarma andersoni De Man, 1887: 657; 1888: bluntly ridged; inner ventral margin minute­ 172: pi. 12, figs. 1-4; Alcock, 1900: 418. ly denticulate, with prominent subdistal Sesarma (Parasesarma) andersoni - Tesch, tooth. Carpus with dorsal surface rugose 1917: 129. with short ridges, inner angle produced in Nanosesarma andersoni - Tweedie, 1950: 310. weak tooth, with row of long stiff setae prox­ Nanosesarma (Beanium) andersoni - Serene imal to tooth. Chela (Fig. 5D, E) with low but and Soh, 1970: 394. distinct ventral ridge on smooth outer sur­ Beanium andersoni - Tan and Ng, 1994: 82. face; dorsal surface of palm rugose with lon­ gitudinal or obliquely longitudinal granular Material examined. Iriomote Island: ridges of various length and 1 obliquely Yonada-gawa estuary, Hoshidate, mangrove transverse pectinated crest (Fig. 5F); inner swamps, burrow on dead wood piece, coll. T. surface of palm nearly smooth, lacking dis­ Komai, 1 male cw 8.0 mm (CBM-ZC 7091); tinct ridge; fingers moderately long, tips not Funaura, mangrove swamps, burrow on dead particularly expanded, but spoon-like, cut­ wood piece, 15 July 2003, coll. T. Komai, 1 ting edges each with distinct triangular teeth

— 45 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

46 New Records of Grapsoid Crabs from Japan

ver entire length; no subterminal tuft of ites lined by row of long setae. Abdominal brush-like setae; patches of dense, short setae cavity in males nearly reaching to suture r. bases of both fingers. Dactylus (Fig. 5G) between third and fourth sternites. r.early straight, with 14 tubercles on dorsal Male abdomen (Fig. 5M) triangular, rela­ -urface; tubercles increasing in size but be­ tively narrow for sesarmids, not abruptly coming lower toward tip of dactylus, proxi­ narrowed at base of telson; sixth somite mal 6 tubercles somewhat longer in trans­ nearly as long as fifth somite, with weakly verse axis, other tubercles basally circular. convex lateral margins; telson nearly sub- Female chelipeds subequal, generally simi­ ovate, slightly longer than broad and dis­ lar to male chelipeds but somewhat smaller. tinctly longer than sixth abdominal somite. Palm (Fig. 5H, I) with irregular rows of very Female telson (Fig. 5N) evenly rounded, small tubercles, but lacking pectinated crests; distinctly wider than long and longer than ventral ridge on outer surface more coarsely mid-line length of sixth abdominal somite. tuberculate than in males. Dactylus with First gonopod (Fig. 50) moderately slen­ longitudinal row of 8 very small tubercles on der, nearly straight in mesial view; corneous proximal half of dorsal surface. terminal process (Fig. 5P) somewhat elon­ Ambulatory legs (Fig. ID, 51) moderately gate, bent at right angle, with shallow notch long, stout; dorsal surfaces of meri, carpi and distally. propodi weakly to somewhat rugose. Meri Coloration. Generally brown in dorsal very broad, foliaceous, each with 1 subacute view. Distal parts of propodi of ambulatory subdistal tooth on anterior margin and 2 or 3 legs paler. strong subdistal teeth on strongly expanded Distribution. Mergui Archipelago (type lo­ posterodorsal margin, posterodorsal distal cality); Malay Peninsula, and southern margin with row of several tiny teeth. Carpi Ryukyu Islands (Iriomote Island) (De Man, each with patch of short plumose setae on 1888; Tan and Ng, 1994; this study). anterior surface and 1 sharp median ridge on Ecological notes. The present specimens dorsal surface. Propodi each with patch of were collected from cavities made by wood thick short plumose setae on dorsal surface boring sphaeromatid isopod on decayed and with several corneous spinules on outer wood in mangrove swamps. distal margin. Scattered longish, darkly pig­ Remarks. The genus Nanosesarma Twe- mented setae on outer and inner margins of edie, 1950, was partially revised by Serene carpi and propodi. Dactyli about half of pro­ and Soh (1970), who divided the genus into podi in length, each with scattered short two subgenera, Nanosesarma sensu stricto setae on outer margin and 1 pair of slender and Beanium Serene and Soh, 1970. Later, corneous spinules arising mid-length of dac­ Tan and Ng (1994) considered full generic tylus on inner margin (Fig. 51). No promi­ status for the two taxa, although they men­ nent tufts of long setae on coxae of ambulato­ tioned that a revision of Nanosesarma sensu ry legs. lato by P. Davie was in progress at that time. Anterior sternal plate triangular (Fig. 5L); As pointed out by Holthuis (1978), however, surface slightly to somewhat concave, with the designation of the type species of the two few short setae. Fourth sternite with sparse subgenera by Serene and Soh (1970) was in­ setae; suture between third and fourth stern- correct. Tweedie (1950) clearly stated that

Fig. 5. Nanosesarma andersoni (De Man, 1887). A-G, J-M, O, P, male from Hoshidate, Iriomote Island (cw 8.0 mm; CBM-ZC 7091); H, I, N, female from Funaura, Iriomote Island (cw 7.8 mm: CBM-ZC 7486). A, carapace and eyes, dorsal view; B, exorbital tooth and anterolateral margin of carapace, dorsal view; C, anterior part of carapace and cephalic appendages, left side, anterior view; D, left chela, outer view; E, same, dorsal view; F, pectinated crest on palm, outer view; G, dactylus of left chela, dorsal view; H, right chela, outer view; I, same, dorsal view; J, left fifth pereopod, dorsal view; K, same, dactylus and distal part of propodus, dorsal view; L, anterior sternal plate, ventral view; M, abdomen, ventral (outer) view (setae omitted); N, sixth abdominal somite and telson, ventral view (setae omitted); O, left first gonopod, mesial view; P, same, distal part, mesial view.

— 47 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita the type species of Nanosesarma was Sesarma (Tweedie, 1950). The possession of a single andersoni, while Serene and Soh (1970) in­ pectinated crest on the dorsal surface of the dicated that Sesarma minutum De Man, 1887 cheliped palm and the presence of a short was the type species of Nanosesarma and as­ transverse ridge posterior to the upper orbit­ signed Sesarma andersoni to the subgenus al margin seem to distinguish N. andersoni Beanium. Therefore, the assignment of Ses­ from N. nunongi. In N. nunongi, there are two arma andersoni to Nanosesarma sensu stricto pectinated crests on the dorsal surface of the is correct. The species assigned to Beanium palm; the carapace lacks a short transverse by Serene and Soh (1970) should be referred ridge posterior to the upper orbital margin. to Nanosesarma, and a new generic name From the other local species of the Sesarmi- should be proposed for the species assigned dae, N. andersoni can be easily separated by to Nanosesarma by Serene and Soh (1970). the ambulatory meri armed with strong However, as suggested by Tan and Ng (1994), spines on the posterior margins and the non- a thorough revision of Nanosesarma sensu setose carapace lacking an epibranchial lato is needed to assess the taxonomic posi­ tooth. tion of the species assigned to Nanosesarma sensu stricto or Beanium. This is beyond the Perisesarma semperi (Burger, 1893) scope of this study and is currently being (Figs. IE, 6, 7) completed by P. Davie. Sesarma semperi Burger, 1893: 630, pi. 21, fig. Our specimens from Iriomote Island agree 1; Tweedie, 1950: 342, fig. le. generally with the previous descriptions of Sesarma (Perisesarma) semperi - De Man, Nanosesarma andersoni by De Man (1888, as 1902: 542. Sesarma andersoni) and Alcock (1900, as Ses­ Sesarma (Chiromantes) semperi - Tesch, 1917: arma andersoni). De Man (1888) mentioned 198. that the lateral margins of the carapace were Sesarma {Chiromantes) semperi semperi - nearly parallel, scarcely converging back­ Campbell, 1967: 4 (key). ward, while his illustration (De Man, 1888, Perisesarma semperi - Rahayu and Davie, plate 12, fig. 1) clearly shows that a trapezo­ 2002: 605. idal carapace with the lateral margins some­ Perisesarma semperi semperi - Nakasone and what converging backward. In our speci­ Irei, 2003: 272 (key), 275. men, the lateral margins of the carapace are slightly converging backwards. Alcock Material examined. Iriomote Island: Funa- (1900) stated that the lateral margins of the ura, mangrove swamps, upper intertidal, 23 carapace of this species were slightly con­ March 1997, coll. T. Komai, 2 males cw 17.7, verging backwards. We assume that De Man 26.0 mm, 1 female 21.7 mm (CBM-ZC 3761); (1888) was inaccurate in stating the shape of Yonada-gawa estuary, Hoshidate, mangrove the carapace. The geographical range of this swamps, 6 July 1998, coll. T. Komai, 1 male species is substantially extended north from cw 27.8 mm, 1 female cw 15.5 mm (CBM-ZC the Malay Peninsula. 7192); Funaura, mangrove swamps, upper in­ Nanosesarma andersoni is most similar to tertidal, 8 July 1998, coll. T. Komai, 1 male cw N. nunongi (Tweedie, 1950) in having a non- 16.8 mm (CBM-ZC 7231). setose dorsal surface of the carapace and the Comparative material. absence of a conspicuous epibranchial tooth (De Haan, 1835). Kume-jima Island: Tomari,

Fig. 6. Perisesarma semperi (Burger, 1893). A-E, I, J, L, M, male from Hoshidate, Iriomote Island (cw 27.8 mm; CBM-ZC 7192); F, G, K, lemale from same lot (cw 15.5 mm). A, carapace and eyes, dorsal view (setae omitted on left side); B, anterior part of carapace and cephalic appendages, left side, anterior view (setae partially omitted); C, F, chela, outer view (C, right; F, left); D, G, distomesial part of dorsal surface of palm (D, right; G, left), showing pectinated crests, dorsal views; E, H, dactylus of chela, dorsal view (E, right; H, left; setae omitted); I, left fifth pcrcopod; dorsal view; J, abdomen, ventral (outer) view (setae omitted); K, sixth abdominal somite and tclson, ventral view (setae omitted); L, left first gonopod, mesial view; M, same, distal part, lateral view (setae removed).

— 48 — New Records of Grapsoid Crabs from Japan

49 T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

D °0 O

a •nQ Q|

^

Fig. 7. A, C, Perisesarma semperi (Burger, 1893), male from Hoshidate, Iriomote Island (cw 27.8 mm; CBM-ZC 7192); B, D, Perisesarma bidens (De Haan, 1835), male from same locality (cw 22.0 mm; CBM-ZC 3753). A, B, exorbital tooth and anterolateral margin of carapace, dorsal view; C, D, carpus of right cheliped, dorsal view.

upper intertidal, 12 June 1995, coll. T. Komai, Antennule and antenna (Fig. 6B) separated 4 males cw 13.9-14.5 mm, 2 females cw 17.7, by incomplete septum. Basal segment of an­ 21.5 mm (CBM-ZC 3131). Iriomote Island: tennule subrectangular. Antenna set ob­ Hoshidate, Yonada-gawa river mouth, upper liquely; second segment with elongate dis- intertidal, 23 March 1997, coll. T. Komai, 2 tolateral process reaching distal margin of males cw 17.0, 22.0 mm, 1 female 16.3 mm fourth segment; flagellum relatively short, (CBM-ZC 3753); Hoshidate, mangrove but extending into orbit. swamps, upper intertidal, 6 July 1998, coll. T. Epistome (Fig. 6B) concave on outer sur­ Komai, 2 females cw 16.7, 17.2 mm (CBM-ZC face, covered with very short setae; ventral 7404). margin granular, protuberant medially, ex­ Description. Carapace (Figs. IE, 6A) rect­ tending onto anterior part of buccal cavity as angular; greatest width between exorbital short ridge. angles about 1.3 times carapace length. Third maxilliped with exopod bearing Front 0.55-0.60 times as wide as exorbital well-developed flagellum. width, moderately deflexed, with broad Male chelipeds (Fig. IE) subequal or slight­ median notch and shallow concavity accom­ ly unequal, large, robust. Merus with ante- modating antennal peduncle just mesial to roventral margin tuberculate, bearing large, preorbital angle; preorbital angle weakly pro­ subdistal triangular tooth; dorsal margin duced and bluntly pointed. Dorsal surface of sharply carinate, tuberculate, with small sub- carapace with tufts or short transverse rows distal tooth; posterior surface with numerous of short stiff setae (tufts of setae sparse on transverse rows of granules. Carpus (Fig. 7C) posterior part); two distinct post-frontal lobes with inner angle not produced, inner margin on either side of midline, mesial lobe broader granular; dorsal surface with scattered short than lateral lobe, separated by deep furrow; to moderately long, transverse rows of small regions moderately well indicated; lateral granules or tubercles; inner surface with lon­ branchial ridges rather obsolete, each ac­ gitudinal row of tiny granules on midline. companied by row of short stiff setae. Ex­ Palm (Fig. 6C, D) with 2 obliquely transverse orbital angle sharply pointed. Lateral pectinated crests each followed by row of margin of carapace slightly concave, with small, rounded tubercles and few tufts of small subacute epibranchial tooth defined by short stiff setae distally on dorsal surface; relatively shallow V-shaped notch (Fig. 7A). first crest composed of 13-15 long corneous Upper orbital margin faintly sinuous; lower teeth; second crest well developed, shorter orbital margin (Fig. 6B) minutely granular, than first, with 7-9 wider corneous teeth; 1 with small, triangular inner orbital tooth not additional row of small tubercles pro- reaching to preorbital angle of front. ximomesial to second crest; dorsal inner

50 New Records of Grapsoid Crabs from Japan rr.argin of palm bordered by small, rounded slightly concave, heavily setose. Fourth •.ubercles, distal portion faintly dentate; outer sternite of male with dense soft setae on an­ surface of palm convex, naked, covered with terior part. Abdominal cavity in males not very small, low, rounded tubercles, but with­ reaching to suture between third and fourth out trace of ventral ridge; inner surface also sternites. covered with small, low tubercles; ventral Male abdomen (Fig. 6J) moderately wide. surface (including fixed finger) with sharper Telson as long as broad at base, evenly tubercles; fixed finger straight, cutting edge rounded, slightly shorter than sixth abdomi­ terminating in trifid corneous claws, with nal somite. Sixth abdominal somite longer row of small, rounded or pointed teeth. Dae- than fifth somite, strongly convex at antero­ tylus (Fig. 6C, E) slightly longer than palm, lateral angle; width across anterolateral terminating in bifid corneous claw; dorsal angles distinctly greater than basal width of surface bearing 6-8 tubercles widely spaced telson. and becoming lower toward tip; each tuber­ Female abdomen with telson (Fig. 6K) cle dome-like, with microscopic striae run­ evenly rounded, shorter than basal width ning parallel to longitudinal axis of dactylus; and shorter than mid-line length of sixth ab­ inner dorsal margin of dactylus with row of dominal somite. small, rounded tubercles in proximal half; First gonopod (Fig. 6L) rather slender, sli­ outer surface with scattered small to very ghtly curved; corneous terminal process (Fig. small tubercles proximally; cutting edge 6M) somewhat elongate, bent at about 30°, with row of triangular or blunt teeth; narrow with deep, narrow notch distally. hiatus between fingers. Coloration. Carapace and ambulatory legs Female chelipeds proportionally smaller generally dark brown. Merus and carpus of than those of males, but generally similar in chelipeds also dark brown; outer surface of structure. Palm (Fig. 6F, G) with 1 pectinated chelae dark reddish brown. crest composed of corneous teeth (corneous Distribution. Singapore, Labuan, Malaysia, teeth more slender than those of males) and 2 Philippines, southern Ryukyu Islands (Ishi- granular ridges subparallel to inner dorsal gaki and Iriomote islands), and southwestern margin; inner surface less granular. Dactylus Irian Jaya, Indonesia. (Burger, 1893; De Man, (Fig. 6F, H) with 7 or 8 dome-like tubercles, 1902; Tesch, 1917; Ward, 1941; Tweedie, distal-most tubercle very low, obsolete; inner 1950; Tan and Ng, 1994; Rahayu and Davie, dorsal margin with short row of small tuber­ 2002; Nakasone and Irei, 2003; this study). cles proximally. Ecological notes. This species occurs in Ambulatory legs (Figs. IE, 61) moderately coastal or sheltered mangrove swamps, but it short, fourth pereopod longest. Meri broad, is not common in the Ryukyu Islands. The foliaceous, each armed with sharp subdistal congeneric P. bidens was occasionally found tooth on anterior margin, but unarmed on sympatrically. posterior margin, merus of fourth pereopod Remarks. According to Rahayu and Davie about twice as long as wide; dorsal surfaces (2002), the following 16 species are assigned of meri of third and fourth pereopods each to the genus Perisesarma De Man, 1895 at with short, occasionally tuberculate or gran­ present: P. bengalense Davie, 2003; P. bidens ular, transverse ridges. Carpi each with 2 (De Haan, 1835); P. brevicristatum (Campbell, distinct carinae on dorsal surface. Propodi 1967); P. cricotus Rahayu and Davie, 2002; P. stout, with dense stiff setae on outer surfaces darwinensis (Campbell, 1967); P. dussumieri and row of darkly pigmented corneous (A. Milne Edwards, 1853); P. eumolpe (De spines on inner distal margin. Dactyli 0.7-0.8 Man, 1895); P. foresti Rahayu and Davie, times as long as propodi, devoid of corneous 2002; P. guttata (A. Milne Edwards, 1869); P. spines or spinules, but with some long­ haswelli (De Man, 1887); P. indiarum (Twe­ itudinal rows of short stiff setae. No promi­ edie, 1940); P. lividum (A. Milne Edwards, nent tufts of long setae on coxae of ambulato­ 1869); P. longicristatum (Campbell, 1967), P. ry legs. messa (Campbell, 1967); P. onychophorum (De Anterior sternal plate triangular; surface Man, 1895); and P. semperi (Burger, 1893). In

— 51 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

Japanese waters, this genus has long been bidens. represented only by P. bidens until Nakasone Sesarmoides kraussi (De Man, 1887) and Irei (2003) recorded this species (as Peri- (Figs. 1G, 8) sesarma semperi semperi) from Ishigaki and Iriomote Islands. However, Nakasone and Sesarma kraussi De Man, 1887: 652 ; 1888: Irei (2003) did not retain voucher specimens. 193, pi. 14, figs. 1-3 ; Alcock, 1900: 425. The presence of this species in Japanese Sesarma (Sesarma s.s.) kraussi - Tesch, 1917: waters is here confirmed based on specimens 164 ;Tweedie, 1936:51 ;1954: 119;Guinot from Iriomote Island. and Crosnier, 1964: 212, figs. 1-3, 6. Perisesarma semperi is characterized by Sesarmoides kraussi - Serene and Soh, 1970: having six to eight, similarly shaped dactylar 403; Ng, 2002: 434 (key). tubercles, which are circular and widely spaced. In most other congeneric species, the Material examined. Iriomote Island: Shira- dactylar tubercles count more than eight. hama, outer edge of mangrove swamp, upper Perisesarma onychophorum also has relatively intertidal, burrow under stone, 13 July 2002, few dactylar tubercles, counting seven to coll. T. Komai, 1 male cw 23.7 mm (CBM-ZC nine (Tesch, 1917), but P. onychophorum can 7401); Kuira-gawa river mouth, mangrove be immediately distinguished because the swamps, 14 July 2003, coll. T. Saki, 1 male cw distalmost dactylar tubercle is greatly elon­ 23.4 mm (CBM-ZC 7417); Yukishida-gawa gate and occupies nearly one-quarter of the river mouth, mangrove forest, 19 October length of the dactylus. 2001, coll. T. Naruse, 1 young male cw 5.8 It is useful to compare P. semperi and P. mm (RUMF-ZC-130); Urauchi River, land­ bidens, as the two species sometimes occur ward edge of mangrove forest, 26 February sympatrically in the Ryukyu Islands. Other 2003, coll. T. Nagai and A. Yogi, 1 male cw than the number of dactylar tubercles, the 9.3 mm, 1 female cw 13.6 mm (RUMF-ZC- following characters can be used to discrimi­ 131). Ishigaki Island: Miyara-gawa river nate the two species. The notch defining the mouth, 29 June 2003, coll. A. Nomoto, 1 epibranchial tooth is shallower in P. semperi young male cw 9.3 mm (CBM-ZC 7533). (Fig. 7A) than in P. bidens (Fig. 7B), thus the Comparative material. Phuket, Thailand: epibranchial tooth of P. semperi is less clearly Ao Nam Bor, landward edge of mangrove defined. The striae on the lateral part of the swamp, under stone, 25 October 1995, coll. T. dorsal surface of the carapace are less clearly Komai, 1 male cw 14.3 mm, 1 juvenile cw 6.9 demarcated in P. semperi (Fig. 7A) than in P. mm (CBM-ZC 2333). bidens (Fig. 7B). The outer surface of the Description. Carapace (Fig. IF, 8A) trapezo­ palm of the male chela is covered with very idal with lateral margins notably diverging small tubercles in P. semperi, rather than backwards; greatest width between post­ coarse tubercles in P. bidens. The tubercles erolateral corners about 1.3 times carapace on the dorsal surface of the carpus are also length. Front about 0.4 times as wide as ex- much smaller in P. semperi (Fig. 7C) than in P. orbital width, weakly deflexed, with broad bidens (Fig. 7D), and this character is useful median notch; preorbital angle rounded. for both male and female. Finally, the color Dorsal surface apparently shining, but mi­ of the chela is different between the two nutely punctate, with sparse very short setae; species. In P. semperi, the chela is dark red­ post-frontal lobe on either side of midline dish brown, while it is yellowish brown in P. low, not divided; regions moderately well in-

Fig. 8. Sesarmoides kraussi (De Man, 1887). Male from Shirahama, Iriomote Island (cw 23.4 mm; CBM-ZC 7401). A, carapace and eyes, dorsal view (setae omitted on left side); B, anterior part of carapace and cephalic appendages, left side, anterior view; C, left antennule, antennal and inner orbital tooth, anterior view; D, left chela, outer view; E. carpus and chela of left cheliped, dorsal view; F, left fifth pereopod. dorsal view; G, anterior sternal plate, ventral view (setae partially omitted); 11, abdomen, ventral (outer) view (setae omitted); I, left first gonopod, mesial view; J, same, distal part, lateral view (setae removed).

— 52 New Records of Grapsoid Crabs from Japan

— 53 T. Koraai, T. Nagai, A. Yogi, T. Naruse. Y. Fujita and S. Shokita dicated; lateral branchial striae weak, granu­ tion) in adults. Dactylus longer than palm, lar, occasionally lined with very short setae. straight in young individuals, weakly curved Exorbital angle sharply pointed, but not in adults; dorsal surface almost smooth strongly produced. Lateral margin with 2 except for proximal part bearing few very blunt epibranchial teeth, both defined by U- small tubercles; cutting edge similarly armed shaped notch; second tooth much smaller as fixed finger, but adults bearing several than first tooth. Upper orbital margin faintly rudimentary teeth between 2 prominent sinuous, slightly oblique; lower orbital teeth; broad hiatus between fingers in adults. margin (Fig. 8B) composed of 2 ridges, upper Ambulatory legs (Figs. 1G, 8F) long; fourth ridge sharply crested, minutely granular, not pereopod noticeably elongated, 2.3-2.7 times reaching level of base of antenna, lower ridge as long as carapace width. Each merus less sharp, not granular but setose; inner armed with sharp subdistal tooth on anterior orbital tooth very small, subtriangular. margin, but unarmed on posterior margin, Antennule and antenna (Fig. 8B) contigu­ merus of fourth pereopod about 3.5 times as ous. Basal segment of antennule relatively long as wide; dorsal surfaces of meri of third large, subtriangular. Antenna set vertically; and fourth pereopods each with sparse gran­ second segment with short distolateral pro­ ules or very short ridges. Carpi each with 2 cess not reaching distal margin of third seg­ faint longitudinal ridges on dorsal surface. ment; flagellum relatively short, but extend­ Propodi slender, with 1 or 2 black corneous ing into orbit. spinules at inner distal margin; setation Epistome (Fig. 8B) concave on outer sur­ different among pereopods; propodus of face, with sparse setae; ventral margin second and third pereopods with mat of smooth, protuberant medially, extending dense short setae on both outer and inner onto anterior part of buccal cavity as short surfaces, that of fourth pereopod with ridge, narrow longitudinal band of very short setae Male chelipeds (Fig. 1G) subequal or slight­ on distal halves of both outer and inner sur­ ly unequal, large, robust. Merus with inner faces, that of fifth pereopod with mat of ventral margin nearly smooth or faintly gra­ dense short setae on whole outer surface and nular, lacking prominent tooth; dorsal narrow band of short setae on distal half of margin sharply carinate, faintly tuberculate, inner surface. Dactyli 0.7-0.8 times as long but without conspicuous tooth; outer surface as propodi, devoid of corneous spines; outer with numerous short transverse ridges. and inner surfaces each covered with short Carpus (Fig. 8E) with inner angle not mark­ setae mixed with several longish setae; dorsal edly produced; dorsal surface with few short and ventral surfaces each with 2 longitudinal to moderately long ridges; inner surface bands of short setae on either side of naked narrow, smooth. Palm (Fig. 8D, E) lacking midline. Prominent tufts of long setae pre­ pectinated crests on dorsal surface; dorsal sent on anteroventral surfaces of coxae of inner margin of palm bordered with low, third and fourth pereopods. branched ridge; outer surface of palm Anterior sternal plate (Fig. 8G) trapezoidal; convex, almost smooth, but with sparse very surface somewhat concave, with sparse short short setae, without trace of ventral ridge; setae. Fourth sternite of male with dense soft ventrodistal part of outer surface with 14-16 setae on anterior part. Abdominal cavity in low, rounded tubercles, extending onto fixed males not reaching to suture between third finger as short row; inner surface with and fourth sternites. scattered low, rounded tubercles; ventral sur­ Male abdomen (Fig. 8H) relatively wide, face (including fixed finger) nearly smooth. somewhat narrowed at base of telson. Sixth Tips of fingers pointed, terminating in simple somite, with strongly convex lateral margins, corneous claw. Fixed finger straight in distal margin slightly concave, length 0.44- young individuals, weakly curved in adults, 0.45 times as wide as basal width, longer cutting edge with row of small teeth in than fifth somite. Telson distinctly longer young individuals, with 2 small, but promi­ than basal width and slightly longer than nent teeth (distal tooth at subterminal posi­ sixth abdominal somite, lateral margins

— 54 — New Records of Grapsoid Crabs from Japan

v. eakly convex, terminal margin rounded. the fixed finger. Variations in the develop­ First gonopod (Fig. 81) stout, almost ment of the row of tubercles on the palm is straight; corneous terminal process (Fig. 8J) probably size-related. short, slightly bent, broadly rounded or trun­ Sesarmoides kraussi is most similar to S. cate distally, without notch. borneensis (Tweedie, 1950) in the features of Coloration. Large individuals entirely dark the carapace, chelae, and ambulatory legs. purplish brown in dorsal view; outer surface Sesarmoides kraussi can be distinguished of chelae generally light yellowish brown, from S. borneensis by minor differences in the but becoming darker toward dorsal surface. chela and male abdomen (Ng, 2002). The Small immature specimen bearing reddish tubercles on the fixed finger of the chela are pereopods, with propodus and proximal half more numerous (eight to 13) and less elon­ of dactylus of fourth pereopod being whitish. gate in S. kraussi, rather than less numerous Distribution. Mergui Archipelago (type lo­ (three to five) and more elongated in S. cality), west coast of Malay Peninsula, and borneensis. The sixth somite of the male southern Ryukyu Islands (Ishigaki and Iri- abdomen is slightly shorter than half the omote islands) (De Man, 1888; Alcock, 1900; proximal width in S. kraussi, slightly longer Tweedie, 1936; Tan and Ng, 1994; this study). in S. borneensis. In our specimens, the Ecological notes. The specimens from Iri- number of tubercles on the outer surface of omote Island were found burrowing in well the chela is 14-16, and the sixth abdominal drained areas of sandy mud near the seaward somite of male is 0.44-0.45 times as long as edge of mangrove swamps. As mentioned the proximal width. Therefore, we refer the before, Clistocoeloma villosum was occasion­ two adult specimens from Iriomote Island to ally found in the same habitat. Sesarmoides kraussi with little hesitation. An Remarks. Our specimens agree generally additional character that may be useful in with the previous descriptions of Sesarmo- discriminating S. kraussi and S. borneensis is ides kraussi by De Man (1888, as Sesarma the shape of the dactyli of the ambulatory kraussi) and Guinot and Crosnier (1964, as legs, which appears less slender in S. kraussi Sesarma (Sesarma) kraussi) except for the than in S. borneensis (see Ng, 2002, fig. 1). much stouter chela. In his key to species of Among the local sesarmid species, this spe­ the genus Sesarmoides, Ng (2002) mentioned cies is readily recognized by the trapezoidal that the outer surface of the fixed finger of carapace with markedly posteriorly diverg­ the chela bears a distinct longitudinal row of ing lateral margins, the presence of two dis­ small conical tubercles in S. kraussi. Guinot tinct epibranchial teeth, the lack of pectinat­ and Crosnier (1964) also mentioned the pres­ ed crests on the dorsal surface of the palm of ence of a row of conspicuous tubercles on the the chela and the noticeably elongate third outer surface of the fixed finger. However, in and fourth pereopods. Some other charac­ the present specimens, the number and shape ters, such as the relatively large basal seg­ of the tubercles are rather variable. In adult ment of the antennular peduncle, vertically male specimens examined here, no distinct standing antennal peduncle broadly in touch row of small conical tubercles is seen on the with the inner part of the orbit, and the sharp outer surface of the fixed finger, but there are suborbital crest, and the trapezoidal anterior patch of low, rounded tubercles at the base of thoracic plate are also useful in distinguish­ the fixed finger, two or three of which extend ing S. kraussi from other sesarmid species in onto the fixed finger in a very short row. The local waters. young male specimen from Ishigaki and Iri- Sesarmoides currently contains 14 species omote islands (cw 5.3-9.3 mm) do not have from the Indo-Pacific region (Ng, 2002), but conspicuous tubercles on the outer surface of none has been reported from Japanese the palm. The young male specimen from waters. Phuket (cw 14.3 mm) has a row of five low tubercles on the proximal part of the fixed Stelgistra stormi (De Man, 1895) finger; the much smaller male specimen from (Figs. IF, 9) the same lot (cw 6.9 mm) lacks tubercles on Sesarma (Sesarma) stormi De Man, 1895: 148;

— 55 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

Fig. 9. Stelgistra stormi (De Man, 1895). A-E, G, female from Hoshizuna Beach, Iriomote Island (cw 16.2 mm; CBM-ZC 7378); F, male from same lot (cw 11.7 mm). A, carapace and eyes, dorsal view (dorsal ornamentation omitted on left side); B, anterior part of carapace and cephalic appendages, left side, anterior view; C, left chela, outer view; D, left fifth pereopod, dorsal view; E, same, dactylus and distal part of propodus, dorsal view; F, coxae of left second to fourth pereopods, ventral view; G, sixth abdominal somite and telson, ventral view.

— 56 — New Records of Grapsoid Crabs from Japan

1898: 702, pi. 29, fig. 29. move according to the changing of tidal Sesarma (Holometopus) stormi - Tesch, 1917: level. Ng and Liu (1999) stated that the crab 200, 237. hides in deep crevices or holes in rock during Stelgistra stormi - Ng and Liu, 1999: 234, figs. the day. However, we encountered active 5-10. crabs climbing on the rock surface during the day. Material examined. Iriomote Island: Hoshi- Remarks. In the key to the Indo-Pacific zuna Beach, crevice on rock, supra-littoral, 5 genera of Sesarmidae, Serene and Soh (1970) July 2001, coll. T. Komai, 2 males cw 11.1, defined the genus Holometopus for species 13.3 mm (CBM-ZC 7062); same locality, 12 characterized by the antenna excluded from July 2002, 1 male cw 11.7 mm, 1 ovigerous the orbit, the lateral margin of the carapace female cw 16.2 mm (CBM-ZC 7378); same lo­ devoid of the trace of an epibranchial tooth, cality, 21 July 2001, coll. M. Osawa, 1 male cw and the dorsal surface of the palm of the male 14.3 mm, 1 ovigerous female cw 16.0 mm chela bearing only one pectinated ridge, or iCBM-ZC 7416). having none at all. Although Serene and Soh Description. See Ng and Liu (1999). (1970) did not list species assigned to Holo­ Supplemental description. Front with metopus, Serene (1968) had earlier listed 13 broadly rounded preorbital angle (Fig. 9B). species Holometopus (as a subgenus of Ses­ Inner orbital tooth triangular, falling far arma), including Sesarma stormi known at short of preorbital angle of front. Suborbital that time by the type series from northern ridge bordered by small tubercles. Sumatra. Holthuis (1977) clarified nomencl- Antennal region (Fig. 9B) narrow, partially atural confusion in sesarmid genera. He overhung by deflexed front. Basal segment showed that Chiromantes is a senior syno­ of antennular peduncle transversely oblong, nym of Holometopus, as the type species of only ventral portion visible in frontal view. the two genera was the same, Grapsus Antennule and antenna separated by narrow haematocheir De Haan, 1833. Recently, Ng hiatus, but without septum between them. and Liu (1999) redescribed Sesarma stormi Antenna set obliquely; second segment and reassigned it to the newly established deeply concave on outer surface, bearing genus Stelgistra based on a single syntype elongate ventrolateral process; antennal and supplemental specimens newly dis­ flagellum short. covered from southern Taiwan. Epistome (Fig. 9B) concave on outer sur­ The present specimens from Iriomote face, ventral margin bordered by rounded Island closely agree with the detailed redes- tubercles, protuberant medially, extending cription of Stelgistra stormi by Ng and Liu onto anterior part of buccal cavity as short (1999). Therefore, we have no hesitation in ridge. assigning our specimens to this species. As Female abdomen moderately broad; telson mentioned before, this species has been re­ (Fig. 9G) roundly subtriangular, slightly ported only twice, from the type locality in longer than mid-line length of sixth abdomi­ the northern part of Sumatra, and from the nal somite. southern part of Taiwan. Our specimens Coloration. See Ng and Liu (1999). extend the geographical range of this rare Distribution Aceh, northern Sumatra, species slightly to the north. Indonesia (type locality), Renting National As Ng and Liu (1999) noted, Stelgistra Park, Pingtong County, southern Taiwan, stormi is similar to species of Chiromantes in and southern Ryukyu Islands (Iriomote the characters mentioned by Serene and Soh Island,); supra-littoral (De Man, 1895; Ng and (1970). Stelgistra stormi can be easily distin­ Liu, 1999; this study). guished from species currently assigned to Ecological notes. As Ng and Liu (1999) Chiromantes by the spoon-shaped tips of the mentioned, this species occurred only on fingers of the chela (Figs. IF, 9C). Further­ highly eroded dead coral heads in the supra- more, the markedly trapezoidal carapace littoral or intertidal zone 1 to 1.5 m above the (Fig. 9A), the broadly foliaceous meri of the surface of the water. The crab seemed to ambulatory legs (Fig. 9D), the dactyli of the

— 57 - T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

ambulatory legs armed with a row of corne­ 117, figs. 1-4; Marumura, 1994: 65, fig. 1; ous spinules on the inner margins (Fig. 9E), Nomoto et al, 1999: 6, pi. 1-3; Kishino et al. and the presence of dense tufts of soft setae 2001b : 127 ; Nakasone and Irei, 2003: 274, on the coxa of the third and fourth somites fig. 48. (Fig. 9F) are also useful in recognizing this species. The carapace of most species of Material examined Iriomote Island: Ura- Chiromantes is squarish, except for C. ob- uchi, river mouth of unnamed small river, tusifrons Dana, 1851, which has a markedly under stone, 12 July 1998, coll. T. Komai, 1 trapezoidal carapace. The presence of the male cw 5.5 mm (CBM-ZC 7135); Nishi- dense tufts of soft setae on the third and Gehda-gawa river mouth, under stone, 11 fourth pereopods is also known in C. eulim- July 2002, coll. T Komai, 1 male cw 4.7 mm ene (De Man, 1898), C. obtusifrons, and C. (CBM-ZC 7406); Gehda-gawa river mouth, 16 ortmanni (Crosnier, 1965) (cf. Ng and Liu, July 2000, coll. T. Komai, 3 males cw 5.7-9.1 1999). mm, 2 ovigerous females cw 5.3, 5.8 mm (CBM-ZC 7415). Family Varunidae Comparative material. Fukuro, Kushimoto, Kii Peninsula, under stone, 24 October 1984, Ptychognathus capillidigitatus Takeda, coll. K. Wada, 2 males cw 5.9, 7.3 mm, 1 1984 female cw 7.6 mm (CBM-ZC 5208). (Fig. 10) Description. See Takeda (1984). Ptychognathus capillidigitatus Takeda, 1984: Supplemental description. Carapace (Fig. 10

0.5 mm

Fig. 10. Ptychognathus capillidigitatus Takeda, 1984. A, B, D, F, ovigerous female from Gehda-gawa river mouth, Iriomote Island (cw 5.3 mm; CBM-ZC 7415); C, E, G, male from same lot (cw 9.1 mm). A, carapace, eyes and antennae, dorsal view; B, C, left lower orbital margin, ventral view; D, left chela, outer view; E, F, sixth abdominal somite and telson, ventral view; G, left first gonopod, lateral view.

58 — New Records of Grapsoid Crabs from Japan

A i nearly quadrate, greatest width across ex- bun, 1914); P. barbatus (A. Milne Edwards, orbital angles or first epibranchial teeth; lat­ 1873); P. glaber Stimpson, 1858; P. hachijo- eral margins with 2 small, blunt epibranchial ensis Sakai, 1955; P. ishii Sakai, 1939; P. joh- teeth, posterior half slightly converging annae Rathbun, 1914; P. pusillus Heller, 1865; backwards. Tubercles on suborbital border and P. takahashii Sakai, 1939. In addition to more conspicuous in females (Fig. 10B) than the 10 named species, there are still four in males (Fig. IOC). undescribed species from Okinawa Island Female chela (Fig. 10D) with conspicuous (Nakasone and Irei, 2003). tuberculate ventral ridge on outer surface; Ptychognathus capillidigitatus is readily dis­ fixed finger with scattered short setae on tinguished from the 14 East Asian congeners outer surface in basal half; fingers each with by a suite of characters: the carapace with subterminal tuft of long stiff setae: dactylus the greatest width across the exorbital or with few short setae basally on outer surface first epibranchial tooth; the presence of two and low tubercles on dorsal surface. epibranchial teeth behind the exorbital tooth; Male telson (Fig. 10E) about twice as long the chela having a tuft of long setae at the as wide, lateral margins nearly parallel, ter­ proximal part of the fixed finger in males and minal margin broadly rounded. Female ab­ having a patch of very short setae at the domen moderately broad; telson triangular same position in females; the fingers of the (Fig. 1 OF), about 1.8 times as long as wide chela having subterminal tufts of brush-like and distinctly longer than sixth abdominal setae; and the rather inconspicuous frontal somite. median ridge consisting of a single row of First gonopod (Fig. 1OG) almost straight in small, low granules. The shape of the cara­ ventral view, slightly curved dorsally in lat­ pace and the possession of the subterminal eral view, moderately slender, terminal cor­ tuft of setae on each dactylus and fixed finger neous process relatively long, partially ob­ of the chela are particularly useful in recog­ scured by tufts of setae; subterminal process nizing this species from the rest of the local obsolete, with tuft of long setae. crab fauna. Coloration. Generally dark brown in dorsal Acknowledgments view. Distribution. So far known only from We thank M. Marumura, A. Nomoto of Japan: Kii Peninsula (type locality) and cen­ Kokudokankyo Co. Ltd., M. Osawa of the Na­ tral to southern Ryukyu Islands (Amami- tional Science Museum, Tokyo, T. Saki, and Oshima, Okinawa, and Iriomote islands) K. Wada of Nara Women's University for (Kishino etal, 2001; Nakasone and Irei, 2003; donating us specimens used in this study. this study). We also thank P. J. F. Davie of the Queens­ Ecological notes. This species occurred at land Museum, P. K. L. Ng of the National Uni­ the mouth of small rivers strongly influenced versity of Singapore, K. Wada and J. Okuno by seawater. Crabs were found to hide under of the Coastal Branch, Natural History stone. Reproductive characteristics were Museum and Institute, Chiba, for reviewing reported by Fukui and Wada (1986). the manuscript. Remarks. Our specimens from Iriomote Island agree well with the original descrip­ References tion of Ptychognathus capillidigitatus by Alcock, A. 1900. Materials for a carcinological Takeda (1984) and comparative material fauna of India. No. 6. The Brachyura Catamet- from topotypic locality (Kii Peninsula). opa, or Grapsoidea. J. Asia. Soc. Bengal 69: 279- The taxonomy of the genus Ptychognathus 456. is still in need of considerable study. Other Burger, O. 1893. Beitriige zur Kenntniss der Gat- than P. capillidigitatus, the following nine tung Sesarma. Zool. Jahrb., (Syst.) 7: 613-631. named species have been reported from East Campbell, B. M. 1967. The Australian Sesarminae Asian waters (Sakai, 1976; Dai and Yang, (Crustacea: Brachyura). Five species of Sesarma 1991; Miyake, 1998; Nakasone and Irei, 2003): (Chiromantes). Mem. Queensland Mus. 15(1): 1- P. affinis De Man, 1895; P. altimanus (Rath- 19.

— 59 — T. Komai, T. Nagai, A. Yogi, T. Naruse, Y. Fujita and S. Shokita

Crosnier, A. 1965. Crustaces Decapodcs Grapsidae spiciis superiorum, qui summum in India Bataya et Ocypodidae. Faune de Madagascar 18: 1-143. Imperium tenent, suscepto, annis 1823-1830 col- Dai, A.-Y. and S.-L. Yang. 1991. Crabs of the China legit, notis, observationibus et adumbrationibus Seas. 608 pp., 74 pis. China Ocean Press, Beijing. illustravit. 243 pp, pis. 1 55, A-Q, 2. Lugduni- Dana, J. D. 1851. Conspectus Crustaceorum quae in Batavorum, Leiden. Orbis Terrarum circumnavigatione, Carolo Hartnoll, R. G. 1975. The Grapsidae and Ocypodi­ Wilkes e Classe Reipublicae Foederatae Duce, dae (Decapoda: Brachyura) of Tanzania. J. Zool., lexit et descripsit J. D. Dana. Pars VI. Amer. J. Sci. London 177:305-328. Arts (2) 11(32): 268-274. Heller, C. 1862. Neue Crustaceen gesammelt wah- Dana, J. D. 1852. Crustacea, Part 1. In United States rent der Weltumseglung der K. K. Fregatte Exploring Expedition, during the years 1838, Novara. Verhand. K. Zool.-Bot. Gesell. Wicn 12: 1839, 1840, 1841. 1842, under the command of 519-528. (not seen) Charles Wilkes, U.S.N. 13(2), pp. 686-1618. C. Heller, C. 1865. Crustaceen. Reise der osterreich- Sherman, Philadelphia. ischen Fregatte "Novara" um die Erde, in den Dana, J. D. 1855. Crustacea, Atlas. In United States Jahren 1857, 1858, 1859, unter den Befehlen des Exploring Expedition, during the years 1838, Commodore B. von Wiillerstorf-Urbair. Zoologi- 1839, 1840, 1841, 1842, under the command of scher Theil. Vol. 2. 280 pp., 25 pis. Kaiserlich- Charles Wilkes, U.S.N. 14, pp. 1-27, pis. 1-96. C. Koniglichen Hof-und Staatsdruckerei, Wien. Sherman, Philadelphia. Holthuis, L. B. 1977. The Grapsidae, Davie, P. J. F. 1993. A new species of sesarmine and Palicidae (Crustacea: Decapoda: Brachyura) crab (Brachyura: Grapsidae) from Japan and of the Red Sea. Israel J. Zool. 26: 141-192. Taiwan, previously known as Sesarma erythro- Holthuis, L. B. 1978. A collection of decapod Crust­ dactyla Hess, 1865. Crust. Res. 22: 65-74. acea from Sumba, Lesser Sunda Islands, Indone­ Davie, P. J. F. 2002. Crustacea: : Eu- sia. Zool. Verhand. 162: 1-55, pi. 1. carida (Part 2): Decapoda-Anomura, Brachyura. Kishino, T., T. Yonezawa, A. Nomoto, S. Kimura In Wells, A. and W. W. K. Houston (eds.), Zoolog­ and K. Wada. 2001a. Twelve rare species of bra­ ical Catalogue of Australia. Vol. 19.3B. xvi+ 641 chyuran crabs recorded in the brackish waters of pp. CSIRO Publishing, Australia. Amami-Oshima Island, Kagoshima Prefecture, Davie, P. J. F. 2003. A new species of Perisesarma Japan. Nankiseibutu 43(1): 15-22. (in Japanese) (Crustacea: Brachyura: Sesarmidae) from the Bay Kishino, T., A. Nomoto, S. Kimura, T. Yonezawa of Bengal. Raff. Bull. Zool. 51(2): 393-398. and K. Wada. 2001b. Brachyuran crab species Edmondson, C. H. 1951. Some central Pacific crus­ recorded in the brackish waters of Amami- taceans. Occ. Pap. Bernice P. Bishop Mus. 20: Oshima Island, Kagoshima Prefecture, Japan. 183-243. Nankiseibutu 43(2): 125-131. (in Japanese, with Edmondson, C. H. 1959. Hawaiian Grapsidae. Occ. English summary) Pap. Bernice P. Bishop Mus. 22(10): 153-202. Lenz, H. 1910. Crustaceen von Madagaskar, Ost- Forskal, P. 1775. Dcscriptiones animalium, avium, afrika und Ceylon. In Voeltzkow, A. (cd.), Reise in amphibiorum, piscicum, insectorum, vermium. Ostafrika in den Jahren 1903-1905, pp. 539-576, 19 + xxxii— 164 pp. Halfniae. pis. 1-4. Stuttgart, (not seen) Fukui, Y. and K. Wada. 1986. Distribution and Man, J. G. de. 1887. Ubersicht der Indo-pacifischen reproduction of four intertidal crabs (Crustacea, Arten der Gattung Sesarma Say nebst einer Brachyura) in the Tsuda River Estuary, Japan. Kritik der von W. Hess und E. Nauck in den Mar. Ecol. Prog. Ser. 30: 229-241. Jahren 1865 und 1880 beschriebenen Deca- Garth, J. S. 1965. The brachyuran decapod crusta­ poden. Zool. Jahrb. (Syst.) 2: 639-689. ceans of Clipperton Island. Proc. Calif. Acad. Sci. Man, J. G. de. 1888. Report on the podophthalmus 33: 1-46. Crustacea of the Mergui Archipelago, collected Gistel, J. N. F. X. 1848. Naturgeschichte des Thier- for the trustees of the Indian Museum, Calcutta. reichs, fur hohere Schulen. 216 pp, 32 pis. (not J. Linn. Soc. London 22: 1-312. seen) Man, J. G. de. 1895. Bericht tiber die von herrn Guinot, D. and A. Crosnier. 1964. Caracteres et Schiffscapitan Storm zu Atjeh, an den west- afflnitcs de deux Sesarma, S. longipes Krauss et S. lichen Kiisten von Malakka, Borneo und Celebes kraussi De Man (Crust. Decap. Brachyura). Bull. sowie in der Java-See gesammelten Decapodcn Mus. natn. Hist, nat., 2c ser., 36(2): 211-221. und Stomatopoden. Zool. Jahrb. (Syst.) 9: 75-218. Haan, W. de. 1833-1850. Crustacea. In von Siebold, Man, J. G. dc. 1898. Bericht fiber die von herrn P. F. (ed.), Fauna Japonica sive descriptio anima­ Schiffscapitan Storm zu Atjeh, an den west- lium, quae in itinere per Japoniam, jussu et au- lichen Kusten von Malakka, Borneo und Celebes

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sowie in der Java-See gesammelten Dccapoden new genus forS. stormi. Zool. Studies 38(2): 228- und Stomatopoden. Sechster (Schluss-) Theil. 237. Zool. Jahrb. (Syst.) 10: 677-708, pis. 28-38. Ng, P. K. L. and C. D. Schubart. 2003. On the ident­ Man, J. G. de. 1902. Die von Herrn Professor Kiiken- ities of Sesarma obesum Dana, 1851, and Sesarma thal im Indischen Archipel gesammelten Deka- eydouxi H. Milne Edwards, 1853 (Crustacea, Dec­ poden und Stomatopoden. In Kukenthal, W. (ed.), apoda, Brachyura, Sesarmidae). Zoosystema 25 Ergebnisse einer zoologischen Forschugsreise in (3): 425-437. den Molukken und Borneo. Abh. Senckenberg Ng, P. K. L., C.-H. Wang, P.-H. Ho and H.-T. Shih. naturforsch. Gcs. 25: 467-929, pis. 19-27. 2001. An annotated checklist of brachyuran Marumura, M. 1994. New localities of two rare crabs from Taiwan (Crustacea: Decapoda). Natn crabs of the family Grapsidae. Nankiseibutu 36 Taiwan Mus. Spec. Publ. Ser. 11: 1-86. (1): 65-66. (in Japanese) Nomoto, A., S. Yodo, S. Kimura, T. Kishino, M. McNeil, F. 1968. Crustacea, Decapoda & Stomato- Sakano and K. Wada. 1999. Six rare brachyuran poda. Sci. Rep. Great Barrier Reef Exped. 7: 1-98. species of the family Grapsidae, recorded from Milne-Edwards, A. 1869. Note sur quelques nou- the Kinokawa river estuary, Prefec­ velles especes de genre Sesarma Say. Nouv. Arch. ture. Nankiseibutu 41(1): 5-9. (in Japanese) Mus. Hist. nat„ Paris 5: 25-31. Owen, R. 1839. Crustacea. In The zoology of cap­ Milne-Edwards, A. 1873. Recherches sur la faune tain Becchey's voyage; compiled from the collec­ carcinologique de la Nouvellc-Caledonie. Deux- tions and notes made by captain Beechey, the ieme Partic. Groupe des Cyclometopes Neptu- officers and naturalist of the expedition, during nens. Nouv. Arch. Mus. Hist. nat. 9: 155-332. the voyage to the Pacific and Bering Straits per­ Milne Edwards, H. 1837. Histoire naturelle des formed in His Majesty's ship Blossum, under Crustaces, comprenant l'anatomie, la physiologie command of captain F. W. Beechey, R.N.F.R.S. in et la classiflcaution de ces animaux. Vol. 2. 532 the years 1825, 26. 27, and 28, pp. 77-92, pis. 24- pp., atlas, pis. 1-42. Paris. 28. H. G. Bohn, London. Milne Edwards, H. 1853. Memoires sur la famille Pilai, N. K. 1951. Decapoda (Brachyura) from Tra- des Ocypodiens, suite. Ann. Sci. Nat. ser. 3 (Zool.) vancore. Bull. Res. Inst. Univ. Travancore Ser. C, 20: 163-228, pis. 6-11. 2(1): 1-46. Milne Edwards, H. 1854. Notes sur quelques Crus­ Rahayu, D. L. and P. J. F. Davie. 2002. Two new taces nouveaux ou pen connus conserves dans la species and a new record of Perisesarma (Deca­ collection du Museum national d'Histoire natur­ poda, Brachyura, Grapsidae, Sesarminae) from elle. Arch. Mus. natn. Hist. nat. 7: 145-192, pis. Indonesia. Crustaceana 75(3-4): 597-607. 9-16. Rahayu, D. L. and M. Takeda. 2000. A new species Miyake, S. 1998. Japanese Crustacean Decapods of the genus Clistocoeloma (Crustacea: Decapoda: and Stomatopods in Color, Volume 2: Brachyura Grapsidae) from Irian Java, Indonesia. Bull Natn. (Crabs), viii + 277 pp. 64 pis. Third printing. Hoi- Sci. Mus., Tokyo, Ser. A. 26(2): 35-41. kusha, Osaka, (in Japanese) Randall. J. W. 1840. Catalogue of the Crustacea Nakasone, Y. and M. Irei. 2003. Grapsidae. In brought by Thomas Nuttal and J. K. Townsend, Nishida, M., N. Shikatani and S. Shokita (eds.), from the west coast of North America and the The flora and fauna of inland waters in the Sandwich Islands. Acad. Nat. Sci. Philadelphia J. Ryukyu Islands, pp. 272-275. Tokai University 8: 106-147. Press, Tokyo, (in Japanese) Rathbun. M.J. 1907. Reports on the scientific re­ Ng, P. K. L. 2002. New species of cavernicolous sults of the expedition to the tropical Paciiic, in crabs of the Genus Sesarmoides from the western charge of Alexander Agassiz, by the U.S. Fish­ Pacific, with a key to the genus (Crustacea: Deca­ eries Commission steamer "Albatross", 1899- poda: Brachyura: Sesarmidae). Raff. Bull. Zool. 50 1900. IX. Reports on the scientific results of the (2): 419-436. expedition to the tropical Pacific, in charge of Ng, P. K. L. and P. J. F. Davie. 1995. The terrestrial Alexander Agassiz, by the U.S. Fisheries Com­ sesarmine crabs of the genera Metasesarma and mission steamer "Albatross", 1904-1905. X. The (Crustacea: Decapoda: Brachyura: Brachyura. Mem. Mus. Comp. Zool. Harvard 35 Grapsidae) of Ujung Kulon, west Jawa, Indo­ (2): 23 74, pis. 1-9. nesia. Tropical Biodiversity 3(1): 29 43. Rathbun, M.J. 1914. New species of crabs of the Ng, P. K. L. and H.-C. Liu. 1999. The taxonomy of families Grapsidae and Ocypodidae. Proc. U.S. Sesarma tangi Rathbun, 1931 and S. stormi De Natn. Mus. 47: 68-85. Man, 1895 (Crustacea: Decapoda: Brachyura: Sakai, K. 2003. The Japanese names for the Japa­ Grapsidae: Sesarminae), with establishment of a nese crabs. Taxa 15-30. (in Japanese)

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Sakai, T. 1939. Studies on the crabs of Japan. IV. Tan, C. G. S. and P. K. L. Ng. 1994. An annotated Brachygnatha, Brachyrhyncha. Yokendo Ltd., check list of mangrove brachyuran crabs from Tokyo, pp. 365-741, pis. 42-111. Malaysia and Singapore. Hydrobiologia 285: 75- Sakai, T. 1955. Further notes on the brachyuran 84. Crustacea of the Hachijo Island. Rec. Oceanogr. Tesch, J.J. 1917. Synopsis of the genera Sesarma, Works Jap. 2(1): 193-202. Metasesarma, Sarmatium and Clistocoeloma with Sakai, T. 1976. Crabs of Japan and adjacent seas. a key to determination of the Indo-Pacific spe­ 725 pp. (English text), 461 pp. (Japanese text), cies. Zool. Meded. 3(2-3): 127-260. 151 pis. Kodansha Ltd., Tokyo. Tesch, J.J. 1918. The Decapoda Brachyura of the Say, T. 1817-1818. An account of the Crustacea of Siboga Expedition. I. Hymenosomidae, Retro- the United States. J. Acad. Nat. Sci. Philadelphia plumidae, Ocypodidae, Grapsidae and Gecarcini- 1(1-2): 57-63, 65-80 (pi. 4), 97-101, 155-160, dae. Siboga Exped. Monogr. 39c 82: 1-148, pis. 161-169 [1817]; 235-253, 313-319, 374-380, 1-6. 381-401,423-441 [1818], Tweedie, M. W. F. 1936. On the crabs of the family Sendler, A. 1923. Die Decapoden und Stomato- Grapsidae in the collection of the Raffles poden der Hanseatischen Siidsee-Expedition. Museum. Bull. Raff. Mus. 12: 44-71. Abh. Sencken. Naturf. Ges. 38: 21-47, pis. 5, 6. Tweedie, M. W. F. 1940. New and interesting Ma­ Serene, R., 1968. The Brachyura of the lndo Pacific laysian species of Sesarma and Utica (Crustacea Region. Prodromus for a check list of the non- Brachyura). Bull. Raff. Mus. 16: 88-113. planctonic marine fauna of South East Asia. Sin­ Tweedie, M. W. F. 1950. Notes on grapsoid crabs gapore National Academy of Science, Special from the Raffles Museum. Bull. Raff. Mus. 18: Publication No. 1: 33-120. 310-324, pi. 7. Ward, M. 1934. Notes on a collec­ Serene, R. and C. L. Soh. 1970. New Indo-Pacific tion of crabs from Christmas Island, Indian genera allied to Sesarma Say, 1817 (Brachyura, Ocean. Bull. Raff. Mus. 9: 5-28, pis. 1-3. Decapoda, Crustacea). Treubia 27(4): 387-416. Ward, M., 1941. New Brachyura from the Gulf of Shen, C.-J. 1933. Description of a new species of Davao, Mindanao, Philippine Islands. Amer. Mus. crab of the genus Clistocoeloma from China. Ann. Novitates 1104: 1-15. Mag. Nat. Hist. (10) 12: 52-59. Watanabe, T. and K. Wada. 2002. Neosarmatium Shen, C.-J. 1940. On the collection of crabs of South laeve (A. Milne-Edwards, 1869) and Metasesarma China. Bull. Fan. Mem. Inst. Biol. (Zool.) 10(2): rousseauxi (Dana, 1851) (Crustacea, Brachyura, 69-104. Sesarmidae): new record for Japan. Nankiseibutu Shokita, S., T. Nagai, Y. Fujita, T. Naruse, A. lto, T. 44(2): 111-113. (in Japanese, with English sum­ Nagamatsu, T. Yamazaki, M. Shinjo and Y. mary) Nagata. 2002. Distribution and abundance of (Accepted 15 February 2004) crustaceans in the Ohura River mangrove swamp of Okinawa Island, Japan. In A Compre­ hensive Study for Mangrove Ecosystem in Oki­ nawa, pp. 73-86. Research Institute for Sub- tropics, Naha. (in Japanese with English summa­ ry). Stimpson W. 1858. Prodromus descriptionis ani- malium evertebratorum, quae in Expeditione ad m # § #" • ft # PI21 • m m m x3> • Oceanum Pacificum Septentrionalem, a Repub- 21 2) lica Federata missam Cadwaladaro Ringgold et BE m m • m m * >A • mmm& Johanne Rodgers ducibus, observavit et descrip- sit. Pars V. Crustacea Ocypodoidea. Proc. Acad. Nat. Sci. Philadelphia 10: 93-110. T260-8682 ^ffr^KWW 955-2 Takeda, M. 1984. A new crab of the family Graps- T903-0213 -/tMfi+sItraiiHT^PIS 1 idae from Japan. Bull. Natn. Sci. Mus., Tokyo, 3) Series A (Zool.) 10(3): 117-120. myrtmmf Takeda, M. 1989. Biogeographical distribution of 1=901-2131 mmmm^kmKTB 2ti§ crabs from the Palau Islands. In Ohba, H., I. Hayami and K. Mochizuki (eds.), Current aspects of biogeography in West Pacific and East Asian ^-SI^fg^tSW^fflSlfSTfeS. ^fificfcn regions. The University Museum, The Universi­ T, JilT© 4 fH^-fS'ir'^'S : Pachygrapsusplanifrons ty of Tokyo, Nature and Culture, No. 1: 45-56. De Man, 1888 {A 7 ii-W- SfftW y'x v A 7 *'-);

— 62 — New Records of Grapsoid Crabs from Japan

Xanosesarma andersoni (De Man, 1888) (^v^r-f if Edwards, 1869) {^•y'ri if-W- y'yfil-), - f''I-: ST $F ? -f + t / ^ > y -1" *" -); Sesarmoides Metasesarma obesum (Dana, 1851) (^ y zf -i if—f4- kraussi (De Man, 1887) (^ V "T i ii-W- Wfc-7 * ^ 7 h t" ^ y 'r -i if —), Perisesarma semperi - ti'^i •s'r-IJ]' — ); Stelgistra stormi (De Man, 1895) (Burger, 1893) (-< y >T 4 if-Q: 7 9 >r J if-mzpftmt& fcfflT.tr, IQ, pfim^M^ti S^-rr"tcigP|$nTfci9 (If7cfti2, 1999; jgg|5 • ft tztilz, ±Effl«aT*i5 h. $b!^, Clistocoeloma villosum (A. Milne-

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