A revision of the xanthid genusPilodiusDana, 1851 (Crustacea: Brachyura: Xanthoidea)

P.F. Clark; B.S. Galil

Currently, 13 subfamilies are recognised in the brachyuran family Xanthidae: Actaeinae, Antrocarcininae, Chlorodiellinae, Cymoinae, Etisinae, Euxanthinae, Kraussiinae, Liomerinae, Polydectinae, Speocarcininae, Xanthinae, Zalasiinae and Zosiminae. This classification has been based on shared adult features like a transversely ovate carapace, well defined dorsal carapace regions, usually with lateral dentition, stout chelipeds and relatively short ambulatory legs. Such characters are now considered to be convergent. Consequently a number of higher xanthid taxa may be artifical and not monophyletic. A broad sample of 147 xanthid species representing 75 out of 124 genera from all 13 xanthid subfamilies were sampled in a multi-gene analysis. Four markers (three mitochondrial and one nuclear) were used and yielded a tree with ca. 30 xanthid clades. Monophyletic support was demonstrated for the Antrocarcininae (although substantially redefined), Cymoinae, and Polydectinae. Almost every other subfamily was para- or polyphyletic. Furthermore, the two other families of the Xanthoidea, Pseudorhombilidae and Panopeidae, were found nested within the Xanthidae. The molecular results were consistent with phylogenetic relationships implied by a suite of novel and/or neglected “ventral” adult characters including sternal characters, position of genital openings and morphology of the first zoea, instead of “dorsal” characters traditionally used to infer xanthid relationships.

A new species of xanthid crab (Brachyura: Xanthidae) is described from the Bohol Sea in the central Philippines. Xanthias joanneae sp. nov. is most similar in morphology to X. maculatus Sakai, 1961 (type locality: Sagami Bay, Japan), particularly in the presence of distinctive ocelli on the carapace and pereopods. It can be separated from this species by the greater number of ocelli on the dorsal surface of the carapace, wider teeth separated by narrow notches on the carapace anterolateral margin, absence of longitudinal ridges on the external surface of the chelar palm, shorter and stouter ambulatory legs, narrower male anterior thoracic sternum, and stouter G1.

The xanthid crab Leptodius exaratus (H. Milne Edwards, 1834) has been widely reported from the Indo-West Pacific, where it is a ubiquitous component of the intertidal and shallow subtidal fauna. The taxonomy of this species, however, is complex and there are many synonyms, most of which have not been adequately treated or discussed in the past. A close examination of a series of specimens throughout its range shows that L. exaratus should be restricted to the western Indian Ocean. Populations from the eastern Indian and western Pacific oceans should be referred to Leptodius affinis (De Haan, 1835), a species originally described from Japan. While their carapaces are superficially similar, the male first gonopod structure is a reliable character to distinguish these two species. Cancer (Xantho) lividus De Haan, 1835, Leptodius nigromaculatus Serène, 1962, as well as some of the varieties of L. exaratus established by Stimpson (1907), are considered to be junior subjective synonyms of L. affinis (De Haan, 1835). Where necessary, lectotypes or neotypes are designated to stabilise the taxonomy of the two species treated herein.

\^n^^jJU^r~ u »-^ CJLy\^^^^\^ J) I < \>-» JOURNAL OF NATURAL HISTORY, 1993, 27, 1119-1206 A revision of the xanthid genus Pilodius Dana, 1851 (Crustacea: Brachyura: Xanthoidea) P. F. CLARK and B. S. GALILt Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK t Israel Oceanographic and Limnological Research Ltd, Tel Shikmona, FOB 8030, Haifa 31 080, Israel {Accepted 30 April 1993) Forty-three species have been assigned to Pilodius. This revision accepts 12 and, in addition, establishes three new species. These 15 species are described, figured, and their distributions mapped. A key to the genus is provided. KEYWORDS: Crustacea, Xanthoidea, Brachyura, Pilodius, identification key, taxonomy, new species. Introduction Alphonse Milne Edwards (1873), Balss (1938a), Serene and Luom (1958, 1959), Forest and Guinot (1961) and Serene (1984) (Table 1) have all attempted revisionary studies of Pilodius Dana, 1851, but none of these authors completely unravelled the complex taxonomic problems associated with this taxon. The independent description of Pilodius under the name Chlorodopsis by A. Milne Edwards created chaos within the Chlorodiinae that was only resolved when Forest and Guinot (1961: 90) applied the rule of priority. In the period between Dana establishing Pilodius and Forest and Guinot finally imposing nominal stability, > 30 species were assigned to the Pilodius/ Chlorodopsis complex. In fact, to date, a total of 43 (Table 2) species have been attributed to this group. This revision retains only 12 (Table 3), and in addition describes three new species. Abbreviations AM, Australian Museum, Sydney; AMNH, American Museum of Natural History, New York; BBM, Bernice Bishop Museum, Honolulu; IRSNB, Institut Royal des Sciences Naturelles de Belgique; MI, Mauritius Institute; MNHN, Museum national d'Histoire naturelle, Paris; NHM, The Natural History Museum, London; NUS, National University of Singapore; OINT, Oceanographic Institute, Nha Trang, Vietnam; QM, Queensland Museum, Brisbane; SAM, South African Museum; SMF, Senckenberg Museum, Frankfurt-am-Main; SWU, Shikoku Women's University, Japan; USNM, United States National Museum, Smithsonian Institution; UMZC, University Museum of Zoology, Cambridge; UY, University of York; ZLKU, Zoological Laboratory, Kyushu University, Japan; ZSI, Zoological Survey of India. The zones of the carapace are divided according to Serene (1984: 18) and all measurements refer to the width of the carapace. 0022-2933/93 $1000 © 1993 Taylor & Francis Ltd. Table 1. Dana (1852) Pilodius P. pilumnoides P. pubescens P. nitidus P. pugil P. scabriculus A. Milne Edwards (1873) Chlorodopsis Ch. pilumnoides Ch. spinipes Ch. areolatus Ch. melanochirus Ch. melanodactylus Serene and Luom (1958) Chlorodopsis Ch. areolata Ch. pilumnoides Ch. scabricula Ch. pugil Ch. granulata Ch. nigrocrinita Ch. spinipes Ch. melanochira Ch. woodmasoni Ch. aberrens Ch. venusta Ch. melanospinis Ch. inoequalis Ch. paulsoni Ch. palaoensis Ch. miersi Ch. philippinensis Ch. melanodactyla Serene and Luom (1959) Chlorodopsis Ch. areolatus Ch. pilumnoides Ch. palaoensis Ch. melanospinis Ch. flava Ch. Ch. Ch. Ch. Ch. Ch. Ch. Ch. Ch. Ch. Ch. to A list of major revisions to Pilodius since Dana (1852). paumotensis nigrocrinita venusta harmsi scabricula granulata pubescens philippinensis pugil spinipes aberrans o Balss (1938a) Pilodius P. pubescens P. flavus P. paumotensis P. harmsi P. granulatus P. nigrocrinitus Chlorodopsis Ch. melanochira Ch. pilumnoides Ch. pugil Ch. melanospinis Ch. areolata Forest and Guinot (1961) Pilodius P. areolatus P. pilumnoides P. pubescens P. pugil P. scabriculus P. granulatus P. nigrocrinitus P. spinipes P. aberrans P. flavus P. paumotensis P. melanospinis P. harmsi P. miersi P. philippinensis Serene (1984) Pilodius P. areolatus P. pilumnoides P. palaoensis P. melanospinis P. flavus P. paumotensis P. nigrocrinitus P. maotieni P. scabriculus P. granulatus P. ajf. spinipes P. pubescens P. philippinensis P. pugil P. spinipes P. serenei P. aberrans P. luomi n 13 w O Revision of Pi/o^ms 1121 Distribution maps: solid dots refer to specimens examined during the study and listed in material examined; circles refer to literature records which could not all be verified. Systematic account Pilodius Dana, 1851 Pilodius Dana, 1851: 126, 1852: 80, 1853: 216; Heller, 1861a: 11, 1861b: 340, A. Milne Edwards, 1863: 284; Heller, 1865: 19; De Man, 1902: 619; Nobili, 1906: 267, 1907: 393; Stimpson, 1907: 57; Stebbing, 1910: 300; Balss, 1934: 227, 1938a: 56; Serene and Luom, 1959: 333; Chen andLan, 1978: 267; Forest and Guinot, 1961: 89; Sakai, 1965: 148; Guinot, 1967: 267; Derijard, 1968: 1244; McNeill, 1968: 72; Serene, 1984: 233; Dai et al, 1986: 304; Dai and Yang, 1991: 327. Chlorodopsis A. Milne Edwards, 1873: 227; Haswell, 1882: 54; De Man, 1887a: 35, 1887b: 281; Cano, 1889: 204; De Man, 1892: 278; Henderson, 1893: 361; Ortmann, 1893: 470; Zehntner, 1894: 151; De Man, 1895: 520; Alcock, 1898; 165; Lanchester, 1900: 737; Borradaile, 1900: 588; Lanchester, 1901: 539; Borradaile, 1902: 261; De Man, 1902: 624; Nobili, 1906: 269; Stebbing, 1910: 300; Klunzinger, 1913: 248; Bouvier, 1915: 279(99); Laurie, 1915: 450; Balss, 1922: 131; McNeill, 1926: 309; Urita, 1926: 11; Ward, 1932: 250; Gordon, 1934: 4; Ward, 1934: 21; Ramadan, 1936: 33; Ward, 1936: 4; Balss, 1938a: 58; Sakai, 1939: 502; Ward, 1939: 10, 1941: 11, 1942: 97; Barnard, 1950: 214; Serene and Luom, 1958: 88, 1959: 336. Description Carapace depressed, transversely oval, regions defined. Front, hardly projecting, more than one-third carapace width; margin divided into two large submedian lobes and two small lateral lobes. Anterolateral margins tri- or quadridentate, equal to or shorter than markedly convergent posterolateral margins. Posterior margin costate, short. Antennule fosset transverse. Basal antennal segment in adult butting into orbital hiatus, excluding short, slender antennal flagellum from orbit. Anterior margin of buccal cavity well defined, covered by third maxillipeds; exhalent channels faintly indicated on posterior region of palate only. Exopod of third maxilliped, shorter than endopod, columnar, distally notched, bearing triangular tooth distally on inner margin. Lateral margins of endopod ischium nearly parallel, proximal margin obliquely truncate, oblique shallow sulcus near setose internal margin; inner distal angle of merus excavate; carpus setose distally on dorsal surface and propodus with setal tuft, dactylus terminally setose. Chelipeds unequal, fingers massive, spoon-shaped. Superior margin of pereiopods spinulate or tuberculate, external surface of pereiopodal carpi furrowed. Modifications of dactylo-propodal articulation well defined, dactylus tip cornute. Genital openings coxal. Male with 5 abdominal segments. Remarks Dana (1851: 125) initially established Pilodius as a genus within the Chlorodinae and then (1852: 80) treated it as a subgenus of Chlorodius Leach. In the same publication he also assigned Chlorodius pilumnoides White to Pilodius and described four new species (see Table 1). Later, Dana (1853: 216) returned the taxon to generic status. Alphonse Milne Edwards (1873: 227) described Chlorodopsis, to which he attributed three species and described two new ones (Table 1). Bouvier (1915: 99) stated that Chlorodopsis was different from other 'chlorodiens' such as Pilodius, Chlorodius, Phymodius A. Milne Edwards and Xanthodes Dana, a view supported by Balss (1934: 227). Ward (1932: 250) also considered Chlorodopsis valid and subsequently named Table 2. A list of the 43 species that have been assigned to the Pilodius complex. Original name Revised name Revisor Chlorodopsis aberrans Rathbun, 1906 Chlorodopsis arabica Laurie, 1915 Chlorodopsis areolatus H. Milne Edwards, 1834 Pilodius armiger Nobili, 1905 Chlorodopsis (areolata) var. brandonensis Ward, 1942 Chlorodopsis espinosus Borradaile, 1902 Pilodius espinosus McNeill, 1968 Pilodius estisoides Takeda and Miyake, 1968 Pilodius flavus Rathbun, 1894 Pilodius fragifer Paulson, 1875 Chlorodopsis frontalis BonsLdsLiie, 1902: 261 Pilodius granulatus Stimpson, 1859 Pilodius harmsi Balss, 1934 Chlorodopsis hawaiiensis Edmondson, 1962 Chlorodopsis inoequalis Klunzinger, 1913 Chlorodopsis kauaiensis Edmondson, 1962 Pilodius luomi Serene, 1971 Pilodius martensis Nobili, 1906 Pilodius maotieni Serene, 1971 Chlorodopsis melanodactylus A. Milne Edwards, 1873 Garthiella aberrans (Rathbun, 1906) Phymodius granulatus (Targioni Tozzetti, 1877) Pilodius areolatus (H. Milne Edwards, 1834) Liocarpilodes armiger (Nobili, 1905) Pilodius areolatus (H. Milne Edwards, 1834) Etisus laevimanus Randall, 1840 Etisus laevimanus Randall, 1840 Validity uncertain sic Actaea fragifer (While,, 1847) Etisus demani Odhner, 1925 sic Liocarpilodes harmsi (Balss, 1934) Pilodius flavus Rathbun, 1906 Phymodius granulatus (Targioni Tozzetti, 1877) Type erroneously assigned to Pilodius Pilodius miersi (Ward, 1936) Chlorodiella cytherea (Dana, 1852) sic Pilodius pubescens Dana, 1852 Titgen (1986: 57) Gordon (1934: 41) Forest and Guinot (1961: 89) Balss (1938a: 45) Tweedie (1950b: 121) Odhner (1925: 83) This study Odhner (1925: 54) Balss (1938: 45) Serene (1971: 914) This study Balss (1938a: 55) This study This study Forest and Guinot (1961 95) Serene and Luom (1959: 316) Q 3 o Chlorodopsis melanochirus A. Milne Edwards, 1873 Chlorodopsis melanospinis Rathbun, 1911 Chlorodopsis miersi Ward, 1936 Chlorodopsis natalensis Ward, 1934 Chlorodopsis natalis Serene, 1984 Pilodius nigrocrinitus Stimpson, 1859 Pilodius nitidus Dana, 1852 Chlorodopsis oahuensis Edmondson, 1962 Chlorodopsis {Cyclodius) ornatus Alcock, 1898 Chlorodopsis {Cyclodius) palaoensis Sakai, 1936 Chlorodopsis paulsoni Klunzinger, 1913 Pilodius paumotensis Rathbun, 1907 Chlorodopsis philippinensis Ward, 1936 Chlorodius pilumnoides White, 1848 Pilodius pubescens Dana, 1852 Pilodius pugil Dana, 1852 Pilodius sakitensis Serene (manuscript name) Pilodius scabriculus Dana, 1852 Pilodius serenei Miyake and Takeda, 1968 Pilodius spinipes Heller, 1861 Pilodius aff. spinipes Serene, 1984 Chlorodopsis venusta Rathbun, 1907 Chlorodopsis woodmasoni Alcock, 1898 Pilodius nigrocrinitus Stimpson, 1859 Pilodius flavus (Rathbun, 1906) Pilodius miersi (Ward, 1936) Liocarpilodes harmsi (Balss, 1934) Pilodius scabriculus Dana, 1852 sic Phymodius nitidus (Dana, 1952) Pilodius paumotensis Rathbun, 1907 Phymodius monticulosus (Dana, 1852) Pilodius pilumnoides (White, 1848) Etisus paulsoni (Klunzinger, 1913) sic Pilodius granulatus Stimpson, 1859 Pilodius pilumnoides (White, 1848) Balss (1938a: 57) This study This study Serene (1971: 914) Serene (1984: 233) Rathbun (1906: 858) Serene (1984: 241) Forest and Guinot (1961: 106) Balss (1938a: 60) Serene (1984: 231) This study Forest and Guinot (1961: 89) CD < Pilodius sic Pilodius sic Pilodius Pilodius Pilodius granulatus Stimpson, 1859 Thus study granulatus Stimpson, 1859 This study spinipes Heller, 1861 scabriculus Dana, 1852 spinipes Heller, 1861 Thus study Forest and Guinot (1961: 89) NobiU (1906: 270) 1124 P. F. Clark and B. S. Galil Table 3. List of valid species currently assigned to Pilodius Dana 1851. (Note that in addition to this list, three new species are described in this study.) Pilodius areolatus (H. Milne Edwards, 1834) PilodiusflavusRathbun, 1894 Pilodius granulatus Stimpson, 1859 Pilodius maotieni Serene, 1971 Pilodius miersi (Ward, 1936) Pilodius nigrocrinitus Stimpson, 1859 Pilodius paumotensis Rathbun, 1907 Pilodius pilumnoides (White, 1848) Pilodius pubescens Dana, 1852 Pilodius pugil Dana, 1852 Pilodius scabriculus Dana, 1852 Pilodius spinipes Heller, 1861 C. melanochirus A. Milne Edwards as the type species. In an attempt to clarify the situation Balss (1938a: 56, 58) assigned six species to Pilodius and five to Chlorodopsis (Table 1). Serene and Luom (Table 1) extensively revised Chlorodopsis. In their generic discussion they considered (1959: 333) Chlorodopsis and Pilodius to be synonymous and recommended that the nomenclatorial confusion should be rectified by applying the rules of priority in favour of Pilodius. But in their concluding comments (p. 336) they ignored their own suggestion and adopted the junior synonym Chlorodopsis. They reasoned that the name Pilodius should be abandoned because its definition was insufficient and, therefore, the less ambiguous description of Chlorodopsis by A. Milne Edwards should be used. Forest and Guinot (1961: 90) opposed the decision made by Serene and Luom because, in their opinion, the definition of Pilodius by Dana was adequate, and it was incorrect to propose derogation of the law of priority by utilization of the name Chlorodopsis. Dana did not select a type species for Pilodius, so Forest and Guinot subsequently designated Chlorodopsis pilumnoides White, 1848 as the type species. Forest and Guinot (1961: 89) listed the species assigned to Pilodius Dana, 1852 under their original names (Table 1). Of the 43 species listed (Table 2), eight were misidentified, and a further six were assigned to other genera within the subfamily Chlorodiinae because the original description of Pilodius by Dana 1852 was inadequate. On the basis of this description. Serene (1984: 233) suggested that the exclusion of the antennal flagellum from the orbital hiatus by the prolongation of the basal antennal segment is a character for separating Pilodius from other genera of the Chlorodiinae. However, Crosnier pointed out (footnote. Serene 1984: 233) that this prolongation develops with age and may not be useful for identifying juveniles. Furthermore, Crosnier noted that some species of Pilodius, i.e. P. paumotensis Rathbun, have only a very feeble prolongation, even in mature specimens, whereas species of other genera, such as Phymodius ungulatus H. Milne Edwards, can have a slight prolongation of the basal antennal segment. Despite this limitation the prolongation of the basal antennal segment remains the sole character for distinguishing Pilodius from other Chlorodiinae. Type status The type of the genus is Pilodius pilumnoides, and the specimens are extant in The Natural History Museum. Gender masculine. Revision of Pi/o^iMS 1125 Pilodius areolatus (H. Milne Edwards, 1834) (Figs l A - G , 31 A, 40A, 44B) Chlorodius areolatus H. Milne Edwards, 1834: 400; White, 1847: 18; Adams and White, 1849: 41; Hess, 1865: 135. Chlorodopsis areolatus: A. Milne Edwards, 1873: 231, pi. VIII, fig. 8; Hilgendorf, 1879: 790; Richters, 1880: 140 (Ust), 148; Haswell, 1882: 54; Miers, 1884: 517, 532; Muller, 1887: 474 (list); Whitelegge, 1889: 227 (list); De Man, 1890: 54; Ortmann, 1893: 470; Alcock and Anderson, 1894: 200 (list); Bouvier, 1915: 278, figs 30, 31; Balss, 1922: 131; Ward, 1932: 251; Michel, 1964: 24. Chlorodopsis areolata: Alcock, 1898: 166; Lenz, 1905: 354, pi. 47, fig. 8; Laurie, 1906: 405; NobiU, 1906: 269; Rathbun, 1906: 858; Nobili, 1907: 396, pi. 2, fig. 3; Caiman, 1909: 705 (Ust); Lenz, 1910: 551; Stebbing, 1910: 300; Klunzinger, 1913: 250; Sendler, 1923: 38; Odhner, 1925: 36; Hale, 1929: 70; Montgomery, 1931: 443; Edmondson, 1933: 250, fig. 152b, 1946: 296, fig. 178f; Balss, 1935: 139, 1938a: 62; Miyake, 1939: 215, 237; Sakai, 1939: 502, pi. 97, fig. 3; Tweedie, 1947: 27 (Hst), 1950b: 121; Barnard, 1950: 214, figs 39d,e; Holthuis, 1953: 15; Sakai, 1956: 40 (Appendix); Guinot, 1958: 176, figs 21a, b; Serene and Luom, 1958: 96, fig. 2, pis lA, Via, 1959, fig. 5a; Edmondson, 1962: 269, fig. 19a; Sankarankutty, 1962: 138, figs 24, 25; Garth, 1964: 140 (Ust); Ooishi, 1964: 199; Kensley, 1970: 104 (Ust). Pilodius areolatus: Forest and Guinot, 1961: 90; Guinot, 1962: 237, 1964b: 66; Sankarankutty, 1966: 48, 50; Guinot, 1967: 267; Derijard, 1968: 1244; Serene, 1968: 80 (Ust); Ooishi, 1970: 93; Sakai, 1976: 460, pi. 164, fig. 1; Takeda and Miyake, 1976: 110 (list); Takeda and Nunomura, 1976: 62 (list), 72; Peyrot-Clausade, 1977a: 27 (Ust); Serene, 1977: 51 (list); Chen and Lan, 1978: 267, fig. 8:2, pi. 8, fig. 30; Takeda, 1978: 40; Kensley, 1981: 45; Serene, 1984: 241, figs 143c, 144, pi. 33C; Dai et al., 1986: 305, pi. 43(2), fig. 165B(1); Takeda, 1989: 165, 178 (table); Dai and Yang, 1991: 328, pi. 43(2), fig. 165B(1). Chlorodius perlatus: MacLeay, 1838: 59; Krauss, 1843: 31. Actea perlata: Ward, 1942: 88. Xantho dehaanii Krauss, 1843: 29, pi. I, fig. 2. Etisodes caelatus Dana, 1852: 77; 1853: 188, 1855, pi. 9, fig. 4; Whitelegge, 1897: 131. Chlorodopsis areolata var. brandonensis Ward, 1942: 97, pi. 6, fig. 3. Actaeodes qffinis Dana, 1852: 78; 1853: 197, 1855: pi. XI, fig. 3; Stimpson, 1907: 43. Actaeodes tomentosus: Miers, 1886: 135 (part); Lanchester, 1900: 734 (part). Actaea affinis: A. Milne Edwards, 1865: 263; HasweU, 1882: 45; Whitelegge, 1889: 226; Borradaile, 1900: 583, 1902: 254; Grant and McCuUoch, 1906: 11; Rathbun, 1906: 852; Stimpson, 1907: 43; Rathbun, 1907: 42; 1911: 219, 1914: 658; Balss, 1922: 121; Edmonson, 1923: 15. Phymodius ungulatus: Boone, 1934: 143 (part); Serene and Luom, 1958: 96; Serene, 1984: 241. non H. Milne Edwards, 1834. Description Regions of carapace well defined, separated by deep grooves, 2M divided into two longitudinally, 3M tripartite, anterior lobe not reaching anterior margin of 2M, densely covered with minute setae, leaving free only pearliform granules. Submedian frontal lobes arched, minutely denticulate, separated by 'U'-shaped indentation, lateral lobes triangular, granulate. Anterolateral margin quadrilobate, lobes rounded, granulate. Anterior margin of cheliped merus granulate. External surface of cheliped carpus with clusters of pearliform granules, one furrow parallel with palmar joint. External surface of chela minutely setose, proximally set with large rounded granules, lower margin smooth. Pereiopods thickly fringed with plumose setae; merus, carpus with conical tubercles on superior margin; carpus, propodus with pearliform granules externally. Material examined Indian Ocean Sta. B4, Kilifi; coll. W. Baumeister, 1984; 1 S 15 m m (SMF no reg.). Near Watamu, South of Malindi—Sta. Ke-1; coll. H. G. Muller, 24/vii-7/viii/1989; 26 1 2 - 8 m m 1126 P. F. Clark and B. S. Galil (SMF no reg.)—Sta. Ke-2; Kenya; coll. H. G. Muller, 24/vii-7/viii/1989; 26 16-5-9mm, 19 ovig. 11-5mm, 2 $ 11-5-lOmm (SMF no reg.)—Sta. Ke-5; coll. H. G. Muller, 24/vii-7/viii/1989; 5S 23-17 mm, 1 9 ovig. 15 mm, 1 $ 11 mm (SMF no reg.). Saline-les-Basins, Reunion; Sta. Re-1; coll. H. G. Muller, 20/1/1989, 0-5-1 m; \S 12 mm, 1$ 11mm (SMF no reg.)—Sta. Re-7; Rifflagune; coll. H. G. Muller, 21-22.1.1989, 0-5-l-5mm; 26 20-5-14mm, 3 ? 13-9mm (SMF no reg.)—Sta. Re-31; coll. H. G. Muller, 30/i-4/ii/1989; 4c? 14-11mm, 1$ 15 mm, 5 2 ovig. 13-5-9 mm, 1 9 15-9 mm (SMF no reg.)—Sta. Re-35; coll. H. G. Muller, 3-5/ii/1989, 0-5-1 m; 4c? 14-9mm, 19 ovig. 11mm, 19 14mm (SMF no reg.)—Sta. Re-43; Riffdach; coll. H. G. Muller, 5/ii/1989, 0-5-1 m; 1 ? 11 mm (SMF no reg.). Coetivy, Seychelles; coll. Sealark, 1905; 6 c? 20-5-10 mm, 2 2 ovig. 13-12-5 mm, 5 9 16-5-lOmm; formerly Actaea affinis det. M. J. Rathbun (UMZC Aug. 7, 1910)— Peros, Coin, Chagos Archipelago; 1 c? 23-5 mm (UMZC Aug. 7, 1910)—Salomon Is.; 5 c? 18-11 mm, 2 9 ovig. 12 mm; redet. T. Odhner 1922 as Chlorodopsis areolatus (UMZC Aug. 7, 1910). Maldive Is.; Fadiffolu Atoll; coll. J. S. Gardiner; Ic? 14mm, 19 12mm; formerly Actaea affinis det. L. A. Borradaile, redet. T. Odhner 1922 as Chlorodopsis areolatus (UMZC June 20, 1900)—Minijol Atoll; 1 9 9-5 mm (UMZC June 20, 1900)—Goifurfehendu Atoll; 19 12 mm (UMZC June 20, 1900)—Hulule, Male; 8 c? 18-9-5 mm, 3 9 ovig. 13-5-12 mm, 2 9 13-5-10mm (UMZC June 20, 1900). Weligama, Sri Lanka; coll. Low-Beer, 31/2/1912; 2c? 18-13 mm, 19 ovig. 13 mm (SMF no reg.). Cocos Keeling Is.; coll. C. A. Gibson-Hill, 1941; 2c? 20-5-16mm, 2 9 15-13-5 mm (NUS 1965.11.11.28-32). Cocos Keeling Is.; coll. J. Covacevich, February 1986; 3c? 14-5-lOmm, 109 10mm (QM W. 12385). Christmas Is.; coll. C. A. Gibson-Hill, 1940; Ic? 7mm (NUS 1965.11.11.38). Japan Bise Village, Motobu-Cho, Okinawa; coll. P. Ng; 1 9 ovig. 18-5 mm, 3 9 17-15 mm (NUS 1992:4986-4989)—Horikawa, Tamagusuka Village; 3 c? 17-13 mm, 19 10mm (NUS 1992:4990-4993)—Kunri-Hama Beach, Sesoko Is.; 19 ovig. 13 mm, 19 11 mm (NUS 1992:4984^985)—Ic? 8mm (NUS 1992:4994). Southeast Asia Philippine Is.; coll. From H. Cuming's collection; 1 c? damaged, 1 9 19 mm; formerly det. Chlorodius areolatus by Adams and White, redet. Phymodius ungulatus by A. Milne Edwards; reg. was 857a, b now (NHM 1843:6). Australia Masthead, Queensland; 1 c? 22-5 mm; formerly Actaea affinis det. F. E. Grant and A. R. McCulloch (AM G. 5845). Port Jackson, New South Wales; 1 c? 22 mm; formerly Actoea affinis det. W. A. Haswell (AM G. 5534). Heron Is., Queensland; coll. T. C. Marshall, 29 August 1939; 1 c? 21-5 mm (QM W. 983)—coll. A. A. Cameron, 21 July 1941; 1 c? 27 mm (QM W. 1406). Lady Elliot Is., Queensland; coll. P. Davie and D. Potter, 8 August 1985; 1 c? 17 mm (QM W. 15444)—13 August 1985; det. J. Short, Revision of Pi/orfms 1127 16 18 mm, 1? 16-5 mm (QM W. 15439)—coll. B. Sanker, 13 August 1985; 15 17mm (QM W. 15448)—coll. P. Davie and D. Potter, 16 August 1985, 5m; 16 12 mm, 1 9 ovig. 17 mm, 1 $ 15 mm (QM W. 15449)—coll. P. Davie and J. Lowe, 17 August 1985; 1 c? 11 mm (QM W. 15442)—coll. P. Davie and D. Potter, 17 August 1985; 16 18 mm, 1 9 15 mm (QM W. 15447). South Is., Queensland; coll. P. Davie and J. Short, 25 May 1987; det. P. Davie, 30 May 1987, 16 18 mm (QM W. 13133). Wreck Reef, nr Porpoise Cay, Queensland; coll. J. Short, 7 May 1988, 2 m; det. J. Short, 19 13 mm (QM W. 15435)—coll. J. Short, 7 May 1988, 2 m; det. J. Short, 16 12 mm (QM W. 15438)—coll. J. Short and S. Mullens, 10 May 1988, 0-5 m; 4 9 19-10 mm (QM W. 15452)—coll. J. Short and S. Mullens, 14 May 1988; 56 23-13 mm (QM W. 15453). Lady Elliot Is., Queensland Australia; coll. P. Davie and D. Potter, 16 August 1988, 5 m; 1 9 14 mm (QM W. 15436). Middleton Reef, Tasman Sea; coll. J. Short and R. McKay, 9 May 1987; 16 15 mm (QM W. 13022). Pacific Ocean Palau Is.; coll. C. Semper, 1862; 2 9 14-10-5mm (SMF 323a) (Ex. Museum Gottingen)—det. A. Sendler; \6 22 mm (SMF 1712). Sta. 20, Platier du Phare Amedee, Nouvelle-Caledonie; coll. 17 September 1978; 2 c? 22-20mm, 3 9 17-15mm (MNHN no reg.). Oahu, Hawaiian Is.; 10c? 17-10mm, 5 9 ovig. 19-13-5 mm, 2 9 13-5-11mm (BBM 510403). Sta. FPM-3, Maharepa, 2-6 km west of Flughafen, Moorea; coll. H. G. Miiller, iii/ 1988, 0-5 m; 4c? 17-10mm, 3 9 ovig. 16-5-14mm, 19 15 mm (SMF no reg.). Society Is., Polynesia; FPM-13, Afareaitu; coll. H. G. Muller, 29.iii.1988,0-0-5 m; 3 c? 21-19mm, 1 9 ovig. 13mm, 4 9 18-13mm (SMF no reg.)—Bora Bora; coll. H. G. Muller, 27/ii-6/iii/1988, 0-5-1 m; 30 specimens (SMF no reg.). Temae, 17°29'S, 149°46'W, Polynesia; coll. H. G. Muller, 31/iii/1988, 0-0-5m; 5c? 18-11mm, 3 9 16-12 mm, 2juvs (SMF no reg.). Viti Levu Is., Fiji; 1 c? 24 mm, 19 20 mm (SMF 1711) (Ex. Museum Godeffroy). Rotuma Is., Fiji; coll. J. S. Gardiner; 1 c? 14 mm, 1 9 ovig. 11 mm, 4 9 15-5-11 mm; formerly Actaea affinis det. L. A. Borradaile, redet. T. Odhner 1922 as Chlorodopsis areolatus (UMZC Sept. 15, 1897). Remarks Crabs identified by Borradaile (1900) and Rathbun (1907, 1911, 1914) as Actaea affinis and deposited in the University Museum of Zoology, Cambridge were redetermined as Pilodius areolatus by T. Odhner. Miers (1886: 135) erroneously synonymized the Actaeodes affinis Dana (1853: 197) with Actaeodes tomentosus (H. Milne Edwards), see Montgomery (1931). A number of species are considered junior synonyms of Pilodius areolatus, Xantho dehaanii Krauss by Miers (1884), Chlorodius perlatus MacLeay by A. Milne Edwards (1873), Etisodes caelatus Dana by Haswell (1882), Actaeodes affinis Dana (assigned to Actea by A. Milne Edwards) by Montgomery (1931) and Actea perlatus and Chlorodopsis areolatus brandonensis both of Ward by Barnard (1950). The Chlorodius areolatus material of Adams and White (1849) was assigned by A. Milne Edwards (1873: 232) to Phymodius ungulatus, however, this material was examined during this revision and the identification of Adams and White was confirmed. 1128 P. F. Clark and B. S. Galil Distribution Reported from the Red Sea and east coast of Africa to Mangareva, Polynesia, Pacific Ocean (Fig. 16). Type status The Type from Nouvelle-Hollande (Australia) is no longer extant. Pilodius cephalalgicus sp. n. (Figs2A-G, 31B, 40B) Description HOLOTYPE male. Regions of carapace well defined, 2M divided into two longitudinally, areoles granular, granules acuminate, apically comute on 1L-5L, set with short, coarse, dark setae. Transverse row of adjoining pearliform granules medially on IP; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes arched, prominently denticulate, separated by deep 'U'-shaped indentation, lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth multispinose, apically comute, terminal spine accompanied by ancillary spines. Anterior margin of cheliped merus prominently spinose. External surface of cheliped carpus with cornute, conical tubercles, coarsely setose, ill-defined furrow parallel with palmar joint. External surface of chela set with short, coarse, dark setae; comute, conical tubercles on upper margin, rounder inferiorly; lower margin granulate and dark coloration restricted to fixed finger. Pereiopods with coarse dark setae, merus, carpus prominently spinose on superior margin. Material examined Malayan Peninsula Sta. 20, 6°55'N, 102°45'E; coll. Serene, 24 July 1965; 1 c? 20 mm, HOLOTYPE (MNHN MP B.20935)^c? 14-10mm, 19 11 mm, PARATYPES (MNHN MP B.24030)— Sta. 15, Tilok Kekhe, Perhentian Besar, 6°55'N, 102°45'E; 21 July 1965, \S 8mm, PARATYPE (MNHN MP B.24031)—IcJ 10 mm, 1 juv., PARATYPES (MNHN MP B.24029)—1 juv., PARATYPE (MNHN MP B.24034)—Sta. 18, 6°55'N, 102°45'E; 23 July 1965; IS 8mm, PARATYPE (MNHN MP B.24032)—Sta. 31, Poulo Babi Nyong, 2°30'N, 103°57'E; 21 August 1965; \S 16mm, PARATYPE (MNHN MP B.20936). Remarks Pilodius cephalalgicus sp. n. is morphologically similar to P. pilumnoides, because the anterior margin of the cheliped mems is spinose, but it differs from the latter by the dark coloration on the cheliped being restricted to the fixed finger. The pieopods of these two species are distinct. Distribution Restricted to the east coast of the Malayan Peninsula (Fig. 17). Type status Type series extant and deposited in the Museum national d'Histoire naturelle, Paris. Holotype from 6°55'N, 102°55'E, Malayan Peninsula. Revision of Pi/o^ms 1129 Etymology Cephalalgicus from the Greek Kephalalgikos, meaning causing a headache, referring to the severe taxonomic problems the authors encountered while revising Pilodius. Gender masculine. Pilodius concors sp. n. (Figs 3A-G, 32A, 40C) Chlorodopsis pilumnoides: Balss, 1938b: 56; Tweedie, 1950a: 92; Guinot, 1958: 179, fig. 25a, b. non White, 1848. Pilodius nigrocrinitus: McNeill, 1968: 73 (part), non Stimpson, 1859. Description HoLOTYPE male. Regions of carapace well defined, 2M divided into two longitudinally. Areoles prominently granular, set with short coarse setae; on 1L-5L granules larger, apically cornute. Transverse row of adjoining pearliform granules on IP; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes arched, prominently denticulate, separated by deep 'U'-shaped indentation, lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth multispinose, apically cornute, terminal spine accompanied by ancillary spines. Anterior margin of cheliped merus prominently spinose. External surface of cheliped carpus with large cornute tubercles, coarsely setose, furrow parallel with palmar joint. Upper margin of chela with cornute tubercles, external surface smooth. Pereiopods with coarse dark setae, merus, carpus, propodus prominently spinose on superior margin. Material examined Australia Hock Reef, East of Hayman Is., Queensland; coll. F. A. McNeill and J. K. Howard, May 1953; IS 37mm, PARATYPE (AM P. 12306). Great Barrier Reef, Queensland; coll. Great Barrier Reef Expedition 1928-29; \S 18 mm, PARATYPE; formerly det. F. A. McNeill Pilodius nigrocrinitus (NHM 1937.9.21.104-123). Cape York, Queensland; 1S 25-5 mm, PARATYPE; was reg. 319.68 (MNHN MP B.2394). Coconut Beach, west side of Lindeman Is.; Queensland; coll. P. Davie and J. Short, 26 March 1987, 16m; \S 22-5mm, PARATYPE ( Q M W.12914) (part). Southeast Asia Poulo Condore; coll. Mr Germain (410-68); 16 19 mm, PARATYPE; formerly det. D. Guinot Pilodius pilumnoides (MNHN MP B. 13755). Horsburgh Lighthouse, Singapore, South China Sea; coll. A. Monteiro, April 1938; \S 62 mm, HOLOTYPE; formerly det. Tweedie Chlorodopsis pilumnoides (NUS 1965.11.11.147)—Sultan Shoal; coll. M. W. F. Tweedie, December 1933; 1$ 61mm, PARATYPE (NUS 1965.11.11.149)—Sentosa Reef; coll. P. K. L. Ng, 13 December 1989; 26 44-37-5 mm, 19 32-5 mm, PARATYPES (NUS 1989.3428-3430). Tekek Bay, Pulau Tioman, Malaysia; coll. P. K. L. Ng, June 1983; \6 58 mm, PARATYPE (NUS 1985.1506). Edam Is. (Damar Besar), Djakarta, Java, Indonesia; coll. J. Brock; 26 23-21 mm, PARATYPES; formerly det. De Man as Ch. melanochira; Ex. Museum Gottingen (SMF 592a) (part). 1130 P. F. Clark and B. S. Galil Remarks Specimens referred to by Balss (1938b), Tweedie (1950a) and Guinot (1958) as P. pilumnoides, and by McNeill (1968) as P. nigrocrinitus (part) pertain to P. concors sp. n. Pilodius concors sp. n. is morphologically similar to P. maotieni, P. cephalalgicus sp. n. and P. pilumnoides in having 1L-5L with conical tubercules, but it differs from these species by having the lower external surface of the cheliped smooth. The pleopods can also be used to separate these species. Etymology From the Latin concors, to be of one mind, for the authors' decade of scientific cooperation. Gender masculine. Distribution Off the southern coast of Vietnam (South China Sea) to the northwest coast of Queensland, Australia (Fig. 18). Type status Type series extant and the holotype is deposited in the National University of Singapore. Type locality is Horsburgh Lighthouse, Singapore, South China Sea. Pilodius flavus Rathbun, 1894 (Figs 4A-G, 32B, 40D, 41A) PilodiusflavusRathbun, 1894: 239, 1906: 860,fig.21; Edmondson, 1925: 43; Balss, 1938a: 57; Miyake, 1939: 215. Chlorodopsisflava:Serene and Luom, 1959: 330, fig. 2C, 5F, pi. IB, pi. IIIB. Chlorodopsis melanospinis Rathbun, 1911: 226, pi. 18 fig. 11; Balss, 1938a: 62; Serene and Luom, 1958:108, pi. I, fig. D, pi. Ill, fig. b, pi. IV, fig. c, 1959, fig. 2M. Pilodius melanospinis: Guinot, 1964b: 67, 1967: 268; Serene, 1968: 80 (list), 1984: 242, figs 143e, 146, pi. XXXIIIE; Dai and Yang, 1991: 329, pi. 43(4), fig. 166(1). Pilodius pubescens: De Man, 1902: 619. non Dana, 1852. IPilodius pubescens: Nobili, 1907: 395. non Dana, 1852. Chlorodopsis melanodactylus: Miers, 1884: 531 (part). Chlorodopsis pilumnoides: Laurie, 1906: 406. non White, 1848. Chlorodopsis hawaiiensis Edmondson, 1962: 273, fig. 21a-e. Description Regions of carapace well defined, 2M partly divided into two longitudinally, granulose, granules conical, prominent laterally, dorsal surface covered with long bristly setae. Two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes broad, denticulate, separated by 'U'-shaped emargination, lateral lobes small, bluntly triangular. Extraorbital angle prominently spinose. Anterolateral margin quadridentate, teeth distinct, spinose, second and third teeth often with ancillary spines. Anterior margin of cheliped merus spinose. External surface of cheliped carpus with long conical tubercles, setose. External surface of chela with large tubercles, conical near upper margin, rounder medially, lower margin smooth. Pereiopods with long bristly setae, merus, carpus, propodus with produced slender spines on superior margin. Re\ision of Pilodius 1131 Material examined Indian Ocean Aldabra; coll. Calypso 1954; \S 8 mm; formeriy Pilodius melanospinis det. D. Guinot, 1964 (MNHN MP B.13733)—1 ? 9 mm; formeriy Pilodius melanospinis det. D. Guinot, 1964 (MNHN MP B. 13734). coll. A. J. Bruce; 1 9 8-5 mm; formeriy Pilodius melanospinis det. R. Serene (MNHN MP B.8015). Sta. 191, Etoile Is., Amirante Group; coll. P. R. W. Coppinger, HMS Alert, April 1882, 13 fms; 2S 14-8-5mm, 19 ovig. 9mm, \9 10mm; formerly Chlorodopsis melanodactylus det. Miers (NHM 1882: 24). Saya de Malha Bank, (c. 10°30'S, 61°30'E), Western Indian Ocean; coll. J. S. Gardiner, Sealark, 1905, 26-29 fms; IS 13mm, 1$ 13mm, 3 juv.; formerly Chlorodopsis melanospinis det. Rathbun, types; pres. Gardiner (NHM 1912.2.10.5357). Sta. C.19, Saya de Malha Bank (c. 10°30'S, 61°30'E), Western Indian Ocean; coll. J. S. Gardiner, Sealark, 7 September 1905, 29 fms; \S 17 mm, 1$ 12 mm, 3 juv.; formerly Chlorodopsis melanospinis det. M. J. Rathbun, types (USNM 41268). Sta. C.16, Saya de Malha Bank, (c. 10°30'S, 61°30'E), Western Indian Ocean; coll. J. S. Gardiner, Sealark, 6 September 1905, 26 fms; 1$ 15 mm, 6 juv.; formerly Chlorodopsis melanospinis det. M. J. Rathbun; types; (UMZC Aug. 7, 1910). Mauritius; coll. Peyrot-Clausade 1974; 3>6 9-7-5 mm, 3 $ ovig. 10-8-5 mm, 19 9-5-9 mm; formerly Pilodius melanospinis det. R. Serene; (MNHN MP B.6696). Ceylon (Sri Lanka); coll. W. A. Herdman; 7c? 15-7mm, 4 9 ovig. 14-11mm, 69 11-7 mm; formerly Chlorodopsis pilumnoides det. R. D. Laurie; pres. W. A. Herdman (NHM 1907.5.22.241^5). Colombo, Sri Lanka; Ex. Colombo Museum; \S 12mm (NUS 1970.1.20.2.). Southeast Asia Macclesfield Bank (c. 15°50'N, 114°20'E), South China Sea; coll. P. Bassett-Smith, HMS Egeria, 44 fms; 1S 12 mm; pres. Lords of the Admiralty (NHM 1893.11.3.45)— 32-44 fms; \6 8-5mm; det. H. Balss June 1932 (NHM 1893.11.3.102)—HMS Penguin; 3c? 10-8 mm, 19 11mm, 2 juv. (NHM 1932.7.7.29). Peari Bank, 2 mis and 349° from Zal Is., Sulu Archipelago; coll. B. R. Wilson, Pele, 10 fms, 22/2/1964; \S 13mm, 19 13mm ovig., 1 9 8mm (MNHN MP B. 10485). Ternate, Moluccas, Indonesia; coll. W. Kukenthal, 1894; \S ovig. (SMF 19984). Arafura Sea; coll. P. Bassett-Smith; HMS Penguin; \S Admiralty (NHM 1932.7.7.29). 11-5 mm, 19 11mm 14 mm; pres. Lords of the Australia Wreck Reef, nr Porpoise Cay, Queensland; coll. J. Short and S. Mullens, 11 May 1988, 12m; \S 12mm (QMW. 15157). 1132 P. F. Clark and B. S. Galil Pacific Ocean Nouvelle Caledonie; Lagon, Sta. 67, Bale de Prony, He Ouen, 22°26'S, 166°29'E; coll. B. Richer de Forges, ORSTOM, 21 m; 1 c? 11 mm (MNHN no reg.)—Sta. 343, Grand Recif Sud, 22°49'S, 166°48'E; 32m; \S 24mm (MNHN no reg.)—Sta. 480, Lagune Nord, 18°56'S, 163°29'E; 2 March 1985, 31m; \6 13mm (MNHN no reg.)—Sta. 554, Grand Recif Sud, 22°50'S, 166°55'E; 16 July 1985,27 m; \S 13 mm (MNHN no reg.). Hawaiian Is.; Sta. 189, 1-5 miles off Pokai Bay, Oahu; coll. Pele Exp., 30 August 1959; IS 10mm; formerly Chlorodopsis hawaiiensis det. Edmondson (BBM S 7043)— Anahole, Kauai; coll. Pele Exp., September 1959; 1 9 10-5 mm; formerly Chlorodopsis hawaiiensis det. Edmondson (BBM S 6878). NW Hawaiian Is.; Pearl and Hermes Reef; coll. Pietschmann, February 1928; IS 14mm (BBM S 3047)—French Frigate Shoals; coll. Thaanum, June 1923; \S 11 mm, 1 ? 12 mm (BBM S 1287). Hawaiian Is.; Sta. 3469, Kaiwi Channel, 21°14'51"N, 157°43'30"W; coll. Albatross, 1891, 14 fms; 1? 9 mm; det. M. J. Rathbun, HOLOTYPE ( U S N M 17317)—Sta. 3970, French Frigate Shoal; coll. Albatross Hawaiian Exploration, 1902, 29 May 1902; 1$ 10-5 mm; det. M. J. Rathbun (USNM 29536)—Sta. 3968, French Frigate Shoal; 1 9 10-5mm (USNM 29535)—Sta. 4150; Modu, Manu Is. 5 August 1902; 1? 9mm (USNM 29539)—Sta. 4148; 19 13 mm, 1 juv. (USNM 29538)—Sta. 4158; 12 13mm, 1 juv. (USNM 29540)—Sta. 4159; 7 August 1902; 1? 10mm (USNM 29541)—Sta. 4162; 8 August 1902; 19 13 mm (USNM 29542). Remarks Rathbun (1911) noted that Chlorodopsis melanospinis resembled Pilodius flavus, but its carapace was not as deeply areolated and devoid of spines on the dorsum and upper margin of the orbit. In contrast. Serene (1984) regarded the differences in areolation as difficult to appreciate, but the presence or absence of spines on the orbital margin as providing a good character for differentiation. He further commented that for the males of these two species the pleopods were of the same type (Serene and Luom, 1959: fig. 2c, m). A study of the male chelipeds by Serene led him to conclude that the black coloration extends further back on to the propodus in the case of P. flavus, but not all in the case of Ch. melanospinis. After examination of Pilodius flavus and Chlorodopsis melanospinis specimens, we regard them as one species. We agree with some of the observations made by Serene, i.e. that differences in areolation are not significant and that pleopod morphology is identical, but after examination of a large number of specimens the difference in the spinulation of the orbital margin as originally described by Rathbun appears to be just variation. Furthermore, the coloration of the male chelipeds is the same in both species, with the larger cheliped of the male appearing to develop in specimens > 12 mm carapace width. The P. pubescens of De Man (1902) were deposited in Senckenberg Museum and were examined by Balss (1938a), who redetermined them as P. flavus. Balss (1938a) also considered the material that Nobili (1907) tentatively identified as IP. pubescens as P. flavus. During this study, material identified by Edmondson (1962) as Chlorodopsis hawaiiensis was examined and proved to be P. flavus. Distribution From Aldabra Is., Indian Ocean to Hawaiian Is., Pacific Ocean (Fig. 19). Not recorded from the coast of east Africa or the Red Sea. Revision of Pi/orfms 1133 Type status Holotype is extant, see material examined. United States National Museum, Smithsonian Institution. Type locality, off Sandwich Is. (Hawaii), 21°14'51"N, 157°43'30"W. Pilodius granulatus Stimpson, 1859 (Figs 5A-G, 33A, 4IB) Pilodius granulatus Stimpson, 1859: 34, 1907: 58, pi. VII, fig. 2. Chlorodopsis granulatus: Miers, 1884: 216 (part), plate XXI, fig. a'. Chlorodopsis philippinensis Ward, 1941: 11. Pilodius serenei Miyake and Takeda, 1968: 393, figs 3, 4; Takeda and Nunomura, 1976: 62 (fist), 73. non Chlorodopsis granulatus: Miers, 1884: 216, pi. XXI, fig. A (part); Sakai, 1936a: 164, pi. 49, fig. 1, 1939: 503, fig. 41, pi. 62, fig. 1, 98, fig. 6; Serene and Luom, 1959: 307, fig. lA, fig. 2E, F, pi. I, fig. D, pi. Ill, fig. c, F = Pilodius miersi Ward, 1936. non Chlorodopsis granulatus: Nobili, 1907: 46 = Pilodius paumotensis Rathbun, 1907. non Pilodius granulatus: Sakai, 1965: 148, pi. 73,fig.6, 1976:460, pi. 164,fig.3; Serene, 1984: 240; Dai et al, 1986: 306, pi. 43(3), fig. 165B(2); Dai and Yang, 1991: 329, pi. 43(3), fig. 165B = Pilodius miersi Ward, 1936. Description Regions of carapace well defined, separated by wide, smooth grooves, areoles granular, set with tufts of setae. 2M divided into two longitudinally. IP uniformly granulate; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes rounded, denticulate, separated by 'V-shaped emargination, lateral lobes triangular, denticulate. Anterolateral margin quadrispinose, teeth multispinose. Anterior margin of cheliped merus tuberculate. External surface of cheliped carpus with large tubercles, minutely setose, wide furrow parallel with palmar joint. External surface of chela with large tubercles, conical near upper margin, rounder medially, lower margin smooth. Pereiopods setose, merus, carpus distinctly spinose on superior margin. Material examined Southeast Asia Tolo Harbour, Hong Kong, South China Sea; coll. Albatross, Philippine Expedition 1907-8, 2-6 fms; 1 9 10 mm; det. F. A. Chace (USNM 152640). No. 201, Singapore; coll. R. W. Coppinger, HMS Alert; pres. Lords of the Admiralty; \S 11mm; formerly Chlorodopsis granulatus det. E. J. Miers, Chlorodopsis melanochirus det. H. Balss, 1934 (NHM 1882.24). No locaUty; coll. D. S. Johnson; 96 12-7mm, 19 8mm (NUS 1985:1142-1151)—Ic? 11 mm (NUS 1985:1152). Reef flat at Baran Darat, Singapore; coll. D. S. Johnson, 2 December 1952; 26 8-7-5 mm, 2 9 ovig. 9-7 mm, 2 9 7-6mm (NUS 1985:1125-11300)—Lower Beach, Labrador; 7 December 1953; Ic? 5 mm (NUS 1985:13110)—Lower Beach, Labrador; 7/12/53; \6 5mm (NUS 1985:1311)—Pulau Hartu; 21 November 1953; 19 ovig. 7mm, 39 10-5-5mm (NUS 1985:1138-1141)—Pulau Pawai; 26 7-5-7mm, 19 6mm ovig. (NUS 1985:1135-1137)—Raffles Lighthouse; 12 November 1958; 26 13-9-5 mm. 19 10-5 mm (NUS 1970.3.13.1153-1155)—6 March 1985; \6 10mm, 19 ovig. 6mm, 19 6-5mm (NUS 1985.1122-1124). Sultan Shoal, Singapore; coll. 1933; 36 11-7-5 mm, 3 9 10-7 mm (NUS 1977.7.5.26-310). Pulau Senang, Singapore; coU. M. W. F. Tweedie, November 1934; 56 11-9-5 mm, 5 9 10-8 mm (NUS 1134 P. F. Clark and B. S. Galil 1965.11.11.80-89)—Raffles Lighthouse; July 1937; 6S 14-9mm, 1? 9mm (NUS 1965.11.11.119-125)—Horsburg Lighthouse; 1936; \S 10mm; formerly Pilodius sakitensis (manuscript name) det. R. Serene (NUS 1969.11.24.12)—1 2 9 mm (NUS 1969.11.24.13). Pulau Hartu, Singapore; coll. P. Ng, March 1984; \S 10mm, 1$ ovig. 9mm, 19 9mm (NUS 1987:2525-2527)—Pulau Kukor; 30/12/86; IS 109-5mm, 3 9 9-8mm (NUS 1987:2533-2537)—Pulau Kukor; IS 9mm, 19 ovig. 8 mm, 29 9-6 mm (NUS 1987:2538-2541)—Ic? 8 mm, 1 juv. (NUS 1987:25422543)—Sentosa Reef; June 1983; 19 ovig. 7-5mm (NUS 1985:1875)—Ic? 10mm; (NUS 1985:1786)—December 1985; 19 9mm (NUS 1987:2531)—Labrador; 1980; \S 9-5mm (NUS 1985:1502)—Labrador Beach; February 1987; 2S 10-6mm, 2 9 8-7 mm, 4 juv. (NUS 1987:2544-2551). Pulau Jong, Singapore; coll. Beverly Goh, 26 August 1986; 19 6mm (NUS 1987:2529)—24 October 1986; Ic? 8mm (NUS 1987:2530). Gulf of Davao, Mindanao, Philippines; coll. Mr Godfred R. Oesch, 8 March 1936; 1S 9-5 mm, 1 9 ovig. 7 mm; formerly Chlorodopsis philippinensis det. Ward; Types (AMNH 8376)—28 June 1936; \S 9-5 mm, 1 9 ovig. 7 mm (AMNH 8319)—6 July 1936; 1 9 ovig. 9mm; 1 9 9mm (AMNH 8298)—coll. Dr WilHam G. van Name, 14 November 1937; 1 c? 8 mm (AMNH 8581)—1S 10 mm (AMNH 8340). Palawan; coll. R. Serene, June 1963; \S 10mm (MNHN MP B.6781). Ngadarak reef, Palau Is. (7°17'30"N, 134°28'30"E); coll. 22-31 May 1939; 1^ 11 mm; formerly Pilodius serenei det. Miyake and Takeda; HOLOTYPE ( Z L K U 2979)—\S 9 mm; formerly Pilodius serenei paratype (ZLKU 2980). Pulau Soegi, nr Sumatra, Indonesia; coll. M. W. F. Tweedie, 1936; 5S 11-9 mm, 69 8-5-7 mm; formerly Pilodius sakitensis (manuscript name) det. Serene (NUS 1969.11.24.1-11). Indonesia; 5S 10-8-5mm (NUS 1969.11.22.7-11). Pulau Sakit, Indonesia; coll. R. Serene, 1963; 1 c? 11 mm; formerly Pilodius sakitensis (manuscript name) det. Serene (NUS 1969.11.24.17)—\S 11-5 mm; formerly Pilodius sakitensis (manuscript name, type) (NUS 1969.11.24.16). Arafura Sea; coll. P. Bassett Smith, HMS Penguin; pres. Lords of the Admiralty; 1 9 9mm(NHM 1932.7.7.30). Australia Lizard Is., Queensland; coll. P. Davie and J. Short, 24 May 1987, l-5m; \S 13 mm (QM W. 15469). No. 176, Port Darwin, Northern Territory; coll. R. W. Coppinger, HMS Alert, May 1881; pres. Lords of the Admiralty; \S 10 mm (NHM 1882.7). No locality; \S 11 mm (SMF 7131). Remarks Stimpson (1859: 34) described this species only briefly, without reference to a figure. Miers (1884: 216) identified Stimpson's species as Chlorodopsis granulatus from material collected in Singapore and Australia, and gave a full description of the male specimen from Port MoUe (no. 95), Australia, which was figured (plate XXI fig. A, a). After the death of Stimpson in 1872, his report on the Crustacea (Brachyura and Anomura) collected by the North Pacific Exploring Expedition 1853-56, was discovered and Rathbun, realizing the importance of the historical document. Revision of Pi/o^ms 1135 published the manuscript (Stimpson, 1907). In this paper Stimpson gave an extended description of P. granulatus and figured (plate VII, fig. 2) the type from Hong Kong Is. The male cheliped coloration figured and described by Stimpson differs from that of Miers, and this has caused much confusion. Stimpson (1907: 59) described the male cheliped of P. granulatus as possessing a broad encircling band of brown near the bases of the fingers. His figure does not show this brown band present on the dorsal margin of the propodus, i.e. the coloration does not circumvent the palm of the cheliped. Miers described cheliped colour variation in the HMS Alert material collected by R. W. Coppinger from Australia. The cheliped of the male from Port MoUe (no. 95) had fingers coloured deep brown, and this coloration extended over the greater part of the inner and outer surfaces of the palm (see Miers, plate XXI, fig. a). The coloration encompasses the palm of the male cheliped. This type of cheliped was exhibited by another male from Port Molle (no. 103). Miers further described a male from Port Darwin as a variety (see plate XXI, fig. a'). Re-examination of this specimen revealed that the faded coloration pattern of the cheliped propodus was identical to that of P. granulatus Stimpson, 1859. The HMS Alert material from Australia consists of two species, P. granulatus Stimpson and a new species, subsequently described by Ward (1936: 4) as P. miersi. In the reference collection of the National University of Singapore are type specimens attributed to Pilodius sakitensis by Serene. However, this name appears only in a manuscript found by Crosnier, Paris Museum, and the type material is here ascribed to P. granulatus. Specimens of Chlorodopsis philippinensis Ward (1941), Pilodius philippinensis Serene (1971) and P. serenei Miyake and Takeda (1968) were examined during this present study and all are identified as P. granulatus. Distribution From Hong Kong and Singapore to the east coast of Queensland, Australia (Fig. 20). Type status The Type of Pilodius granulatus Stimpson is no longer extant (Manning, personal communication), it was destroyed in the Chicago fire of 1871. Type locality Hong Kong. Pilodius maotieni Serene, 1971 (Figs 6A-G, 33B, 41C) Chlorodopsis pilumnoides: Serene and Luom, 1958: 102 (part), pi. IC, Ilia, IVB; 1959: 302 (part), fig. 2A, 5G, pi. 3A. non White, 1848. Pilodius maotieni Serene, 1971: 913; 1984: 238, fig. 152. Description Regions of carapace well defined, separated by wide, smooth grooves. 2M divided into two longitudinally. Areoles granular, set with short, dark, coarse setae interspaced with longer, finer ones. On 1L-5L granules larger, conical, apically cornute. Transverse row of adjoining pearliform granules on IP; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes rounded, denticulate, separated by 'V-shaped emargination, lateral lobes triangular, denticulate. Anterolateral margin quadridenticulate, teeth multispinose. Anterior margin of cheliped merus tuberculate. External surface of cheliped carpus with dark conical tubercles, setose. External surface of chela with dark tubercles, large, 1136 P. F. Clark and B. S. Gain conical, well spaced near upper margin, smaller, rounder and closer set inferiorly. Lower margin smooth. Pereiopods setose, merus, carpus distinctly spinose on superior margin. Material examined Southeast Asia Vietnam; \S 20 mm; formerly Chlorodopsis pilumnoides det. Serene and Luom (OINT E.41.911, Rte 1850)—coll. Nguyen Van Luom, 5 May 1958; 19 16mm (OINT E.41.912,Rtel850)—1? 16-5mm;reg.E.41.913,Rte 1850—1 $ ovig. 16mm(OINT E.41.914, Rte 1850). IS 14mm (OINT E.42.210, Rte 1863). coll. Serene; \S 17 mm; formerly Chlorodopsis pilumnoides det. Serene and Luom (MNHN E.34.581, MP B.9312)—1(? 5 mm (MNHN E.42.161, MP B.9388)—Ic? 7 mm (MNHN E.42.763)—Bale de Nhatrang; \S 17 mm. HOLOTYPE; coll. Serene and Luom, 1959 (MNHN E.41.915, MP B.9313). Indochine; 1 9 10mm (MNHN E.1573, MP B.9411). Macclesfield Bank; coll. P. Bassett-Smith, HMS Penguin, 25-28 fms; 1 S 18 mm, 1 9 ovig. 16mm (NHM 1892.8.28.238-40); HMS Egeria, 44 fms; IS 20mm (NHM 1893.11.3.45-6). Pearl Bank, 2 mis and 349° from Zal Is., Sulu Archipelago; coll. B. R. Wilson, Pele, 10 fms, 22/2/1964; \S 16mm (MNHN MP B.10485). Pacific Ocean Sta. 483, Lagon Nord, 19°01'S, 163°32'E, Nouvelle-Caledonie; coll. B. Richer de Forges, ORSTOM, 2 March 1985, 33 m; \S 21 mm (MNHN no reg.). Remarks Serene (1971: 913) described P. maotieni from a specimen originally determined by Serene and Luom (1958, 1959) as C. pilumnoides. The pleopod of P. maotieni was illustrated by these authors (1958, pi. IVb, 1959, fig. 2A) and Serene (1948: 238, fig. 152). Distribution Eastwards from Nhatrang, Vietnam to New Caledonia (Fig. 21). Not recorded from the Indian Ocean. Type status Holotype is extant, see material examined Museum national d'Histoire naturelle, Paris. Type locality, Nhatrang, Vietnam. Pilodius miersi (Ward, 1936) (Figs 7A-G, 34A, 4ID) Chlorodopsis miersi Ward, 1936: 4, pi. II, figs 1-3. Chlorodopsis granulatus: Miers, 1884: 216 (part), pi. XXI, fig. A, a; Sakai, 1956: 40 (Appendix), non Stimpson, 1859. Revision of FiVo^JMS 1137 Chlorodopsis granulata: Sakai, 1936a: 164, pi. 49, fig. 1, 1939: 503, Text-fig. 41, pi. LXII, fig. 1, pi. XCVII, fig. 6; Serene and Luom, 1959: 307, fig. lA, fig. 2E, F, pi. I fig. D, pi. Ill fig. C, F. non Stimpson, 1859. Pilodius granulatus: Sakai, 1965: 148, pi. 73,fig.6, 1976: 460, pi. 164, fig. 3; Dai et al, 1986: 306, pi. 43(3), fig. 165(B); Dai and Yang, 1991: 329, pi. 43(3), fig. 165B(2). non Stimpson, 1859. Pilodius luomi Serene, 1971: 913, 1984: 240. Description Regions of carapace well defined, 2M divided into two longitudinally, 3M tripartite, anterior lobe surpassing anterior margin of 2M. Areoles with rounded granules, minutely setose. IP uniformly granulate; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes arched, denticulate, denticles diminishing in size laterally, separated by 'U'-shaped emargination, lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth basally tuberculate, apical spine curved. Anterior margin of cheliped merus tuberculate. External surface of cheliped carpus with clustered granules, furrow parallel with palmar joint. External surface of chela with conical tubercles on upper margin, diminishing in size inferiorly, lower margin smooth. Pereiopods setose, merus, carpus with conical tubercles on superior margin. Material examined Japan Kushimoto, Japan; coll. Kono, 23 February 1970; \S 13 mm, 19 9-5 mm; det. Pilodius pilumnoides by T. Sakai (SWU TS. 00006). Magari-Zaki, Japan; coll. 13 August 1956; 4c? 10-7mm, 1 9 8mm; det. Pilodius granulatus (SWU TS. 00025). Southeast Asia Vietnam; coll. R. Serene; \S 10 mm; formerly Chlorodopsis granulatus det. R. Serene; reg. (MNHN MP B.9391; formerly E.42.206). Horsburgh Lighthouse, off Singapore, South China Sea; coll. April 1934; 3 c? 8 mm, 3 9 9-8 mm; formerly Pilodius luomi, det. R. Serene; paratypes (NUS 1965.11.11.39-44). Pulau Kukor, Singapore; coll. P. Ng, 30 December 1986; 2S 10-9-5 mm, 3 9 9-5-9 mm (NUS 1987:2533-2537)—Pulau Kukor; \S 9 mm, 3 9 9-6 mm (NUS 1987:2538-2541)—1 c? 8 mm, 1 juv. (NUS 1987:2542-2543)—Pulau Hantu; March 1984; 1S 10mm, 1 9 ovig. 9mm, 1 9 9mm (NUS 1987:2525-2527)— Sentosa; \S 10mm (NUS 1985:1786)—Sentosa; December 1985; 19 9mm (NUS 1987:2531)—Labrador Beach; February 1987; IS 10-6mm, 2 9 8-7mm, 4 juv. (NUS 1987:2544-2551). Pulau Jong, Singapore; coll. Beverly Goh, 24 August 1986; \S 8mm (NUS 1987:2530)—26 August 1986; 19 6mm (NUS 1987:2529). Grande Is., off Southern Luzon, Philippines; coll. Albatross, Philippine Expedition 1907-8, 8 January 1908; 1 9 10-8-5 mm; formerly Chlorodopsis granulatus, det. M. Ward (USNM 65288)—Jolo Lt., Jolo; 29 fms, 15 January 1908; 2 9 11-9-5 mm (USNM 65291)—Sta. 5159, Tinakta Is., Sulu Archipelago; 10 fms, 21 February 1908; 16 9 mm, 2 9 8 mm (USNM 65290)—Sta. 5559, Cabalian Point, Jolo Is.; 13 fms, 18 September 1909; 1 9 13 mm; det. M. Ward (USNM 65287). 1138 P. F. Clark and B. S. Gain Australia Geoffrey Bay, Magnetic Is., Queensland; coll. D. R. Fielder, 20 June 1963; 1S 10 mm (AM P 16505). Hayman Is., Whitsunday Group, Queensland; coll. F. A. McNeill, May 1933; IS 13-3mm (AM P 10411). Lindeman Is., Queensland; M. Ward exchange; IS 12-5mm; formerly Chlorodopsis miersi det. M. Ward; types (NHM 1940.2.23.4)—2 J 17-15 mm, 2 9 15 mm types (NHM 1937.7.15.21-23). No. 95, Porto Molle, Queensland; coll. R. W. Coppinger, HMS Alert, May 1881; \S 14mm, 1? 12 mm; formerly Chlorodopsis granulatus det. E. J. Miers, Chlorodopsis melanochirus det. H. Balss 1934, Chlorodopsis miersi det. M. Ward (NHM 1881:31)—no. 103; 3 c? 12-7 mm (NHM 1881:31). Southern side of Seaforth Is., near Lindeman Is., Queensland; coll. P. Davie and J. Short, 24 March 1987; \S 13mm (QM W. 14404); 2 9 12-9 mm (QM W. 14405)—Home Beach; 25 March 1987; ?>S 10-5-lOmm; det. J. Short (QM W. 14417)—Coconut Beach; 26 March 1987; \S 12mm(QMW. 14512)—2d 12-10mm,49 10-5-8mm, 5 juv. (QM W. 12934)—1 9 12 mm (QMW. 14466). Remarks Miers (1884: 216) identified material from Australia, collected by HMS Alert, as Chlorodopsis granulatus (Stimpson). Examination of the specimens revealed the presence of two species, Pilodius granulatus Stimpson (plate XXI, fig. a') and a new species (plate XXI, fig. A, a), later described by Ward (1936: 4) as P. miersi. The cheliped figured by Miers (plate XXI, fig. A, a) features pigmentation that encompasses the propodus, and corresponds exactly with the type material of Ward (NHM reg. 1937.7.15.21-23, 1940.2.23.4) from Lindeman Is., Queensland, Australia. Figures by Sakai and by Serene and Luom under the name of P. granulatus are of P. miersi. P. luomi paratypes of Serene (1971: 913, 1984: 240) deposited in the National University of Singapore (reg. 1965.11.11.30^4) were examined and the pigmentation pattern of the male chelipeds matches the P. miersi of Ward. Further, Dai and Yang's illustration (1991: 328, pi. 43(3), fig. 165B.2) of the first pleopod and their description of the cheliped manus with a black circular band, pertain to P. miersi not to P. granulatus. Distribution Singapore eastwards to Queensland, Australia and Japan (Fig. 22). Not recorded from the Indian Ocean. Type status Type series is extant, see material examined in the Natural History Museum. Type specimens in the Australian Museum, Sydney were not examined. Type localities are cited as Singapore and Lindeman Is., Queensland, Australia. Pilodius moranti sp. n. (Figs 8A-D; 34B, 42A) Description HoLOTYPE male. Regions of carapace well defined, 2M divided into two longitudinally, smooth furrows dividing setose granulose areoles, granules more prominent laterally. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules on 2P. Submedian frontal lobes arched, minutely denticulate, medially produced, separated by narrow sinus, lateral lobes triangular, granulate. Anterolateral Revision of PJ/o^fms 1139 margin quadridentate, teeth multispinose, two median teeth wider, more prominent. Anterior margin of chehped merus proximally tuberculate. External surface of cheliped carpus with two setae-fringed furrows parallel with palmar joint. External surface of chela closely set with conical tubercles, smaller, obsolescent inferiorly, lower margin smooth, longitudinal furrow proximally near upper margin. Pereiopods thickly fringed with long setae; merus, carpus prominently spinose on superior margin. Posterioir surface of pereiopodal carpus medially with deep setae-fringed furrow, its borders granulate. Etymology This species is named for Harry Morant, Australian bushman, poet and soldier, who, in spite of being innocent, was executed during the Boer War in Pietersburg on 27 February 1902. Gender masculine. Material examined Australia Wreck Reef, nr Porpoise Cay, Queensland; coll. J. Short and S. Mullens, 10 May 1988, 0-5m; IS 21mm; PARATYPE ( Q M W . 15454)—14 May 1988; 46 16-5-9mm; PARATYPES (QM W. 15456)—2 9 19-14 mm; PARATYPES (QM W. 15142)—Ic? 24 mm; PARATYPE (QM W. 15143)—0-5 m; 1 9 22 mm; PARATYPE (QM W. 15455). NE coast of Austraha?; coll. Mr Rayner, Surgeon, HMS Herald; 16 12-5 mm, 1 9 13 mm; PARATYPES; purch. Mr Warwick (NHM 1862.53). Lord Howe Is., New South Wales; coll. P. R. Redley, April 1910; IS 24mm; HOLOTYPE (AM P. 2371)—coll. Icely, 1889; IS 27-5mm; PARATYPE (AM P. 366). Middleton Reef, Tasman Sea; coll. J. Short and R. McKay, 9 May 1987; 1 9 20 mm; (QM W. 13025)—1 9 9 mm. PARATYPE (QM W. 13026). PARATYPE Remarks Pilodius moranti keys out as close to P. granulatus and P. spinipes in having 1L-5L lacking conical tubercles, but it differs from these two species because the carpus of the cheliped has two anterior furrows. These three species have distinct pleopods. Distribution Only recorded from the east coast of Queensland, Australia and the Tasman Sea. (Fig. 23). Type status Type series is extant and deposited in the Queensland Museum, Australia. Pilodius nigrocrinitus Stimpson, 1859 (Figs 9A-G, 35A, 42B-C) Pilodius nigrocrinitus Stimpson, 1859: 34, 1907: 58, pi. 7, fig. la,b; Balss, 1938a: 57; Forest and Guinot, 1961: 89; Sakai, 1965: 148, pi. 73, fig. 5; McNeill, 1968: 73 (part); Sakai, 1976; 461, text-fig. 248a, b, pi. 164, fig. 2; Takeda and Nunomura, 1976: 73; Serene, 1984: 239; Dai etal., 1986: 307, pi. 43(5), fig. 166(2); Dai and Yang, 1991: 330, pi. 43(5), fig. 166(2). Chlorodopsis nigrocrinitus: Urita, 1926: 11. 1140 P. F. Clark and B. S. Galil Chlorodopsis nigrocrinita: Sakai, 1936a: 163, pi. 49, fig. 2, 1939: 504, text-fig. 42, pi. LXII, fig. 2, XCVII, fig. 2; Tweedie, 1950a: 92; Sakai, 1956: 40 (Appendix); Guinot, 1958: 179, fig. 24a, b; Serine and Luom, 1958: 112, pis IIA, Ille, IVe, 1959: 304, figs 2B, 5B, pi. lA; Sakai, 1960: 70, pi. 35, 2. Chlorodopsis melanochirus A. Milne Edwards, 1873: 228, pi. VIII, fig. 5; Haswell, 1882: 55; Ortmann, 1893: 471; Nobifi, 1899: 30; Stephenson et al, 1931: 59; Ward, 1932: 251. Chlorodopsis melanochira: De Man, 1887b: 281, 1892: 278, 1895: 520; Alcock, 1898: 168; De Man, 1902: 624, 1929: 1 (list); Serene and Luom, 1958: 117, pis IIB, Illd, lYd; Forest and Guinot, 1961: 89. Chlorodopsis pilumnoides: Sankarankutty, 1962: 138, figs 38, 39. non White, 1848. Pilodius spinipes: McNeill, 1968: 74. non Heller, 1861. non Pilodius nigrocrinitus: McNeill, 1968: 73 (part)=P. concors sp. n. Description Regions of carapace well defined, covered with coarse dark setae. 2M partly divided into two longitudinally, minutely granulate, granules more prominent laterally. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules on 2P. Submedian frontal lobes arched, minutely denticulate, separated by 'U'-shaped indentation; lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth multispinose, spines subequal, cornute. Anterior margin of cheliped merus tuberculate, tubercles diminishing in size distally. External surface of cheliped carpus coarsely setose, with cornute conical tubercles, lacking longitudinal furrow parallel with palmar joint. External surface of chela coarsely setose proximally, with conical tubercles proximally, smaller, obsolescent inferiorly, lower margin smooth. Pereiopods thickly fringed both with plumose setae and dark coarse setae, merus, carpus prominently tuberculate on superior margin. Material examined Indian Ocean Goh Phuket, Thailand, Andaman Sea; coll. Gallardo, 4 February 1966; pres. R. Serene; \S 10mm (USNM 184261). Great Cocos Is.; coll. Investigator, \S Oct. 17, 1900, 28). 13-5 mm; pres. Indian Museum (UMZC Southeast Asia Sta. 15, Tilok Kehe, Perhentian Besar, 6°55'N, 102°45'E, Malaysian Peninsula; coll. R. Serene, 21/7/65; 1 ? 8mm (MNHN MP B.24033). Aor Is. (Pulau Aur), Malaysia, South China Sea; coll. M. W. F. Tweedie, 1938; 26 12-11 mm, 19 11 mm, 1 9 ovig. 10-5 mm, 19 10-5 mm; det. M. W. F. Tweedie as Chlorodopsis nigrocrinita, 1949 (NUS 1950: 92). Pangkor Laut, Perak, Malaysia; coll. D. S. Johnson, 29 October 1953; \S 9mm, 19 ovig. 9mm, 19 8mm (NUS 1985.1357-1359)—Tg. Dalan, Tioman; coll. P. Ng, 22 June 1983; 1 S 7-5 mm (NUS 1985.1877)—Buru Dalam, Telok, Pulau Tioman; 23 June 1983; \S 11 mm, 19 10mm (NUS 1985.1507-1508). Kelantan, Malaysian Peninsula; coll. Skeat Expedition; 4cJ 11-10 mm, 4 9 ovig. 11-8-5 mm, 1 9 8 mm (UMZC Nov. 30, 1899). Damar Besar (=Edam Is.), Djakarta, Java, Indonesia; coll. J. Brock; 3<J 16-15 mm; formerly det. J. G. De Man as Chlorodopsis melanochira (SMF 592a) (Ex. Museum Gottingen). Ternate, Molluccas, Indonesia; coll. W. Kukenthal; formerly det. J. D. De Man as Chlorodopsis melanochirus (SMF 1716). Indopacific; coll. J. Brock; 36 Revision of Pj/o^ms 1141 16-15-5 mm; formerly det. J. D. De Man as Chlorodopsis melanochira (SMF 592a) (Ex. Museum Gottingen). Aor Is.; coll. R. Serene, 1934; AS 14-lOmm, 1$ ovig. 10mm, 2$ 10mm (SMF 7137). Australia Murray Is., Queensland; coll. C. Headley and A. R. McCulloch, August-October 1907; 3 c? 13-7 mm; 2 9 10-8 mm (AM P.7538)—Cape York; coll. A. R. McCulloch, November 1920; 19 ovig. 17 mm (AM P.4983)—Masthead Is.; coll. D. B. Fry, January 1911; AS 16-12mm, 39 13-5-12mm (AM P.2585)—Holbom Is., Port Denisen; coll. E. H. Rainsford, June 1921; 4c? 15-10-5 mm (AM P.5321)—July 1918; 56 19-11 mm, 2 9 14-11 mm (AM P.4176)—Holbom Is., Port Denisen; November 1921; 4c? 17-11 mm, 19 ovig. 13-5mm, 2 9 14-12-5mm (AM P.5556)—1 9 14mm (AM P.5557)—St Crispin Reef, Port Douglas; coll. A. R. McCulloch, 1918; 1 c? 9 mm (AM P.4308)—Lady Elliot Is.; coll. Mrs C. Wright, 1964; 7c? 16-5-12mm, 2 9 14-12mm (AM P. 17259)—Swain Reef; coll. Swain's Reef Expedition 1962; 3c? 15-5-11-5 mm, 4 9 11-8 mm (AM P. 17208)—One Tree Is., Capricom Group; coll. D. J. G. Griffin and A. Wellington, 14 October 1968; Ic? 11-5 mm, 19 11mm (AM P. 17260)—Beach Rock; coll. D. J. G. Griffin, 11 October 1972; 3c? 15-lOmm (AM P. 18913)—One Tree Is.; 2 October 1972; 1 c? 14-5 mm (AM P. 18914)—coll. D. J. G. Griffin, 6 November 1972; 2c? 14-13 mm (AM P. 18915)—coll. H. F. and P. Recher, 15 October 1972; 2c? 17-5-13mm (AM P.17265)—Townsville; Ic? 18-5mm, 19 14 mm; pres. D. R. Fielder (AM P. 17277). The Boulder Tract, Low Isles, Queensland; coll. Great Barrier Reef Expedition 1928-29; 1 c? damaged, 1 9 ovig. 8 mm; formerly Pilodius spinipes det. F. A. McNeill (NHM no reg.); Queensland—1 c? 10 mm, 19 8mm (NHM 1937.9.21.104-123)—1 c? 13-5mm, 19 11mm; McNeill (NHM 1937.9.21.104-123)—Low Isles; 20 March 1929; 2c? 14mm (NHM 1937.9.21.104123)—12c? 14-5-8mm, 129 12-8mm (NHM 1937.9.21.104-123)—4 April 1929; 19 ovig. 9mm; det. F. A. McNeill (NHM 1937.9.21.104-123)—Batt Reef; Ic? 14mm, 2c? 13-5-11 mm (NHM 1937.9.21.104-123)—Three Isles; 5 May 1929; 19 13mm (NHM 1937.9.21.104-123)—6 May 1929; \6 11mm (NHM 1937.9.21.104123)—8 May 1929; Ic? 8mm, 19 ovig. 9mm (NHM 1937.9.21.104-123)—2c? 11-511mm (NHM 1937.9.21.104-123)—Ic? 14mm (NHM 1937.9.21.104-123)— Asterina Spit; 19 November 1928; Ic? 11-5mm (NHM 1937.9.21.104-123)—Shingle Rampart; 1 c? 11 mm (NHM 1937.9.21.104-123)—22 March 1929; 1 9 ovig. 14mm, 1 juv.; (NHM 1937.9.21.104-123)—8 March 1929; 1 c? 13 mm (NHM 1937.9.21.104123). Murray Is., Torres Straits, Queensland; coll. F. W. Moorhouse, 10/11/1930; 1 9 14mm (QM W.310)—Lindeman Is.; coll. 14 July 1934; 2 9 13-12mm (QM W.513)—Orpheus Is.; coll. A. A. Cameron, 27 June 1940; 19 13 mm (QM W.1097). Mt Adolphus Is., Queensland; coll. 16 July 1974; Ic? 14mm (QM W.12482); Dugon Is.; coll. 17 July 1974; 2c? 19-15 mm (QM W. 12487)—Lizard Is.; coll. P. Davie and J. Short, 7 June 1987; 2c? 12-10mm, 2 juv. (QMW. 15467)—Lizard Is., 30 June 1987; Ic? 10mm, 4 9 10-9mm (QM W. 13137)—Lady Elliot Is.; coll. P. Davie and D. Potter, 11 August 1985; Ic? 12mm, 1 9 8mm (QM W. 15441)—coll. P. Davie and J. Lowe, 17 August 1985; 3 c? 10-8 mm, 1 juv. (QM W. 15440)—Palfrey Is., P. Davie and J. Short, 24 May 1987; 2c? 12-11 mm, 1 9 ovig. 10mm, 3 9 9-6mm; det. P. Davie, 30 May 1987 (QM W. 13132). Cape York, Queensland; coll. R. Schutte 1876; 2 9 1142 P. F. Clark and B. S. Galil 16 mm, 1 ? 13 mm (SMF 403a) (Ex. Museum Gottingen). Austraha; coll. R. Schutte; 1 S 13 mm (SMF no reg.) (Ex. Museum Gottingen). Pacific Ocean Solomon Is.; coll. Capt. Farrel; 2S 12-11 mm (AM P.1755)—Peu, Vanikoro, Santa Cruz Group; coll. Traughton and Livingstone; 1 <? 11 mm (AM P.9096). Noumea Harbour, New Caledonia; coll. A. R. McCulloch, September 1924; AS 16-9 mm, 3 9 11-5 mm, 2 $ 10-9 mm (AM P.7497)—Anse Vata, Noumea; coll. A. F. Bassett-Hull, August 1925; ?>S 12-10mm (AM P.8268)—coll. 1926; \6 14-5 mm, 3 9 12-5-9 mm (AM P.8880). Sta. 2, Platier Ricaudy, Nouvelle-Caledonie; coll. 31 August 1978, ORSTOM; 26 16mm (MNHN no reg.)—Sta. 6, Platier de Namie; coll. 3 September 1978; \S 12mm, 2 9 8mm (MNHN no reg.)—Sta. 20, Platier du Phare Amedee; coll. 17 September 1978; 1 9 15 mm (MNHN no reg.)— Sta. 28, Platier Rocher a'la voile; coll. 29 September 1978; 2c? 14mm (MNHN no reg.)—Sta. 31, Platier de I'ilot Maitre; coll. 3 October 1978, \S 16 mm (MNHN no reg.)—Sta. 33, coll. 5 October 1978, IS 15mm (MNHN no reg.)—Mangrove d'Arama; coll. September 1984; \S 15 mm (MNHN no reg.). Samoa; 2S 13-12 mm (SMF 1714) (Ex. Museum Gottingen). No locality; \S 12-5 mm (SMF no reg.) (Ex. Museum Gottingen). \S 9 mm (ZSI 8100/6)—IcJ 12mm, 19 ll-5mm(ZSI 1958/7). Remarks Stimpson (1859: 34) based his description off*, nigrocrinitus on a female, but this species was subsequently described and figured by A. Milne Edwards (1873: 117, pi. VIII, fig. 5) under the name Chlorodopsis melanochirus. A report written by Stimpson (1907) and published posthumously by Rathbun, included a figure (pi. Villa, b) of the female holotype. This figure was used by Balss (1938: 57) to establish that C. melanochirus is a junior synonym off. nigrocrinitus. Sakai (1939: 504, Text-fig. 42) illustrated the left pleopod of P. nigrocrinitus from its ventral aspect. The accuracy of this drawing was confirmed by Serene and Luom (1958, pi. IVe, 1959: 314, fig. 2B) who figured the ventral view of the right pleopod. The left pleopod of C. melanochira is figured in ventral aspect by Serene and Luom (1958, pi. IVd) and corresponds to that presented by Sakai (1939: 504, Text-fig. 42). Distribution Ranging from the eastern Indian Ocean (Andaman and Nicobar Is.) to Japan, the Society and Hawaiian Is. in the Pacific Ocean (Fig. 24). Not recorded west of longitude 90°E. Type status Stimpson based his description of P. nigrocrinitus on a female which is no longer extant because it was destroyed in the great fire of Chicago 1871. Type locality was Simoda, Japan. Re\ision of Pilodius 1143 Pilodius paumotensis Rathbun, 1907 (Figs lOA-G, 35B, 43A) Pilodius paumotensis Rathbun, 1907: 52, pi. 8, fig. 2, 2a, 2b; 1911: 227; Baiss, 1938a: 58; Holthuis, 1953: 25; Guinot, 1962: 237, fig. 14a, b; Takeda and Miyake, 1968: 7, pi. 1, fig. f; Serene, 1968: 80 (list); Peyrot-Clausade and Serene, 1976: 1357, pi. 4D; Takeda and Miyake, 1976: 110 (list); Peyrot-Clausade, 1977a: 27; Ribes, 1978: 127; Serene, 1984: 241, figs 143d, 145, pi. 33d. Chlorodopsis granulatus: Nobili, 1907: 46. non Stimpson, 1859. Chlorodopsis oahuensis: Edmondson, 1962: 270. Chlorodopsispilumnoides: Barnard, 1955: 3 (part); Michel, 1964: 24; Guinot, 1967: 268 (part). non White, 1848. Description Regions of carapace nearly indistinct, dorsal surface shagreened, with short pubescence interspaced with long setae. 2M entire. IP uniformly granulate; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes arched, minutely denticulate, separated by 'U'-shaped emargination; lateral lobes triangular, denticulate. Anterolateral quadridentate, two median teeth largest. Anterior margin of cheliped merus proximally tuberculate. External surface of cheliped carpus with furrow parallel with palmar joint. External surface of chela with short pubescence interspaced with long setae, tuberculate. Pereiopods setose, merus, carpus with conical tubercles on superior margin. Material examined Indian Ocean Delagoa Bay (Baia de Lourenco Marques), Mogambique; \S Chlorodopsis pilumnoides det. K. H. Barnard (SAM A. 10855). 9 mm formerly Reunion; coll. M. Peyrot-Clausade; 5 S 9-5 mm, 3 2 ovig. 7-5-5 mm, 2 9 5-5^-5 mm; det. R. Serene (MNHN MP B.6697). Tulear, Madagascar; coll. M. Peyrot-Clausade; 10c? 9-5-4 mm, 3 $ ovig. 6-5-6 mm; det. R. Serene (MNHN MP B.19260)—Itampolo, 3 c? 7-5-5 mm, 3 ? 8-5 mm, 3 juv. (MNHN MP B.6780). Mauritius; 1 S 1 mm; formerly Chlorodopsis pilumnoides; (MI no reg.). Re-35, Saline-les-Bains, Reunion; coll. H. G. Muller, 3-5/ii/1989; \$ 9-5 mm (SMF no reg.). Addu Atoll, Maldives; coll. S. Gerlach, 2 January 1958, Xarife Expedition 1957-58; \S 8 mm; det. D. Guinot (MNHN MP B. 13751). Peros, Coin, Chagos Archipelago; coll. J. S. Gardiner, Sealark 1905; 1$ 8-5 mm; formerly Chlorodopsis spinipes det. M. J. Rathbun (UMZC Aug. 7 1910). South Pacific Guam Is., Mariana Is.; coll. 1945; IS 9-5-9mm; det. J. S. Garth; (USNH 134593)—1 juv. (USNM 134594). 1144 P. F. Clark and B. S. Galil Likiep Atoll, Nado Is., Marshall Is.; coll. S. F. MacNeil, 1951-52; 3cJ 6-5 mm, 1 9 ovig. 7-5 mm, 3 juv.; det. L. B. Holthuis (USNM 93999). Tamana, Gilbert Is.; coll. L. Bock, 16 September 1917; 1? 9mm (MNHN MP B.13753). Waianae, Oahu, Hawaiian Is.; coll. Edmondson, 1920; 3 9 7-5-5 mm, 1 juv.; formerly Chlorodopsis oahuensis det. Edmondson (BBM S.923)—1^ 8 mm (BBM S.5287). Marutea,Tuamotu Archipelago; coll. G. Seurat, 1905; \S 5mm; det. D. Guinot, 1962; formerly Chlorodopsis granulatus det. G. Nobili, 1906 (MNHN MP B. 13754). Fakarava Is., Tuamotu Archipelago; coll. Albatross, 12 October 1899; 16 5 mm; det. M. J. Rathbun; Smithsonian Exchange, reg. 33323 (MNHN MP B. 13752). Bora Bora, Society Is.; coll. Bredin Expedition, 25 April 1957; 2c? 7-5-5mm, 19 6 mm; det. J. S. Garth (USNM 156070). Eua Is., Tonga; coll. Albatross, 28 November 1899; \6 8 mm; J. S. Garth (USNM 134591). Makemo, Tuamotu Archipelago; coll. Albatross, 21 October 1899; 16 8 mm, 19 7mm; det. M. J. Rathbun, Types (USNM 32852)—Outer Reef, Fakarava Is.; 12 October 1899; 1 juv.; det. J. S. Garth (USNM 134592). Remarks Examination of Chlorodopsis oahuensis Edmondson (1962) and Ch. pilumnoides Barnard (1955, part) and Michel (1964) proved the specimens to be P. paumotensis. Guinot (1962) examined a juvenile male identified by Nobili (1907) as Chlorodopsis granulata, but she considered it to be Pilodius paumotensis. Distribution Ranging from the coast of Mo9ambique, Indian Ocean to Makemo Is. in the Pacific Ocean (Fig. 25). Type status Type series is extant, see material examined United States National Museum, Smithsonian Institution. Type locality Fakarava Id. and Makemo, Pautmotus. Pilodius pilumnoides (White, 1848) (Figs l l A - G , 36A, 42D) Chlorodius pilumnoides White, 1847: 18 {nom. nud.); White, 1848a: 226, 1848b: 286; Adams and White, 1849: 41, tab. IX, fig. 3. Pilodius pilumnoides^: Dana, 1853: 221, 1855, pi. XII, fig. lOa-c. Pilodius pilumnoides: Sakai, 1976: 461, text-fig. 249 a-b-b'; Garth and Kim, 1983: 686; Serene, 1984, figs 143j, 150, pi. 34E; Takeda, 1989: 166, 178 (table). Chlorodopsis pilumnoides: De Man, 1887a: 35, 1887b: 281; Cano, 1889: 204; Alcock, 1898: 167; Lanchester, 1900: 737; Rathbun, 1923: 108; McNeill, 1926: 309; Gordon, 1934: 47, fig. 26a; Miyake, 1936: 509 (Ust); Balss, 1938a: 59, pi. 1, fig. 4, 1938b: 56; Sakai, 1939: 505, Text-fig. 43; Holthuis, 1953: 16; Sakai, 1956: 40 (Appendix); Serene and Luom, 1958: 102 (part), 1959: 302 (part), 5G, pis lA, IIIA; Sakai, 1976: 461, Text-fig. 249. Chlorodopsis (Cyclodius) palaoensis Sakai, 1936b: 167, pis XIII, fig. 2, XIV fig. 1; Miyake, 1939: 215. non Chlorodopsis pilumnoides: Laurie, 1906: 406=PilodiusflavusRathbun, 1894. Revision of Pilodius 1145 non Chlorodopsis pilumnoides: Serene and Luom, 1958: 102 (part), pis IC, Ilia, IVb, 1959: 314 (part), figs 2A=Pilodius maotieni Serene, 1971. non Chlorodopsis pilumnoides: Sankarankutty, 1962: 138, figs 38, 39=Pilodius nigrocrinitus Stimpson, 1859. non Chlorodopsis pilumnoides: Barnard, 1955: 3; Guinot, 1967: 26S=Pilodius spinipes Heller, 1861 and P. paumotensis Rathbun, 1907. non Chlorodopsis pilumnoides: Balss, 1938b: 56; Tweedie, 1950a: 92, Guinot, 1958: 179, fig. 25a, h=Pilodius concors sp. n. non Chlorodopsis pilumnoides: Garth, 1964: l40=Pilodius scabriculus Dana, 1852. non Chlorodopsis pilumnoides: Michel, 1964: 24=Pilodius paumotensis Rathbun 1907. Description Regions of carapace well defined, 2M divided into two longitudinally, areoles prominently granular, granules acuminate, apically comute on 1L-5L, set with short, coarse, dark setae. Transverse row of adjoining pearliform granules medially on IP; two parallel transverse rows of adjoining pearliform granules on 2P, anterior row medially disjunct. Submedian frontal lobes arched, prominently denticulate, separated by deep 'U'-shaped indentation, lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth multispinose, apically cornute, terminal spine accompanied by ancillary spines. Anterior margin of cheliped merus prominently spinose. External surface of cheliped carpus with cornute, conical tubercles, coarsely setose, ill-defined furrow parallel with palmar joint. External surface of chela set with short, coarse, dark setae; cornute, conical tubercles on upper margin, rounder inferiorly; lower margin smooth and dark coloration expands on to palm. Pereiopods with coarse dark setae, merus, carpus prominently spinose on superior margin. Material examined Southeast Asia Singapore; coll. From H. Cuming's collection; 1 c? 33 mm; Type, det. A. White (NHM 1843.6). Bohol (sphalm. Rohol), Philippine Is.; coll. From H. Cuming's collection; 19 15 mm; Type; det. A. White (NHM 1843.6). West-end Little Sta., Cruz Is., Zamboanga, Mindanao, Philippines; coll. Albatross, 26 May 1908; 1 9 damaged; det. J. S. Garth and H. S. Kim. Pulau Aor, South China Sea; coll. M. W. F. Tweedie, June 1938; 16 34 mm, Chlorodopsis pilumnoides, det. Tweedie (NUS 1965.11.11.148). Banda-Neira, Dutch East Indies (Indonesia); coll. 5 m, 24 February 1929; 16 16 mm; formerly Chlorodopsis pilumnoides det. I. Gordon, 1934 (IRSNB 9223). Australia Cape York, Queensland; 26 23-20 mm (MNHN MP B.2393) (319.68). Bowen, Queensland; 16 23 mm; det. M. Ward, June 1931 (QM W.151)—Lindeman Is.; 26 23-17 mm; formerly Chlorodopsis pilumnoides, det. M. Ward, 14 July 1934 (QM W.518) (part). Lizard Is., Queensland; coll. P. Davie and J. Short, 16m; 26 1413 mm, 1 9 15 mm (QM W.13142)—Coconut Beach, west side of Lindeman Is.; 26 March 1987; 4c? 24-15 mm, 19 14mm (QM W. 12914) (part)—Blue Lagoon, Lizard Is.; 5 June 1987, 5-7m; 26 14-7mm (QM W. 15462)—Research Pt.; 24 May 1987; 1146 P. F. Clark and B. S. Galil IS 15-5 mm; det. P. Davie (QM W. 13143)—First Beach, Lizard Is.; South of Osprey; 30 May 1987; IS 17mm (QM W. 15468). Dugong Is., Queensland; 17 July 1974; 1S 23 mm (QM W. 12487)—Orpheus Is.; coll. A. A. Cameron, 27 July 1940; 1S 22mm, 19 23 mm (QM W.1096). Lindeman Is., Queensland; coll. M. Ward; 1S 35 mm, det. Ward(NUS 1965.11.11.150). Ohne Fundort; 1 S 20 mm (Ex. Museum Gottingen) (SMF no reg.)—3S 14-17-5 mm; 592a (part). Remarks Balss (1938) wrongly synonymized P. pubescens Dana, 1852 and Chlorodopsis melanodactylus A. Milne Edwards, 1873 with P. pilumnoides. In the same paragraph he also synonymized Chlorodopsis palaoensis Sakai, 1936. Since the description of Sakai is based on two females, this synonymy of Balss is difficult to confirm because the black coloration on the cheliped of females is not diagnostic (see discussion). Further, the types of Ch. palaoensis could not be located. The specimen of P. pilumnoides identified by Gordon lacked (originally) chelipeds, making certain determination difficult. Material assigned to P. pilumnoides by Balss (1938b), Tweedie (1950a) and Guinot (1958) all belongs to P. concors sp. n. A specimen of Ch. pilumnoides identified by Serene and Luom was later redescribed by Serene (1971) as P. maotieni. All specimens identified by Garth (1964) were reidentified as Pilodius scabriculus and the P. pilumnoides of Takeda (1989) cannot be confirmed. Distribution From Singapore to the Ryukyu Islands, Japan and Fiji (Fig. 26). Records from the Indian Ocean are dubious; specimens of R pilumnoides identified by Barnard (1955) were redetermined as P. spinipes and P. paumotensis, and those of Laurie (1906) pertain to P. flavus. The Andaman Islands and Mergui Archipelago material of Alcock (1898) were not made available for study. Type status Type series extant, see material examined The Natural History Museum, London. Type locality Singapore and Philippines. Pilodius pubescens Dana, 1852 (Figs 12A-G, 36B, 43B) Pilodius pubescens Dana, 1852: 80; 1853: 217, 1855, pi. 12,fig.6a-d; Takeda and Nunomura, 1976: 62 (list), 73; Dai et al., 1986: 307, pi. 43(6), fig. 166(3); Dai and Yang, 1991: 331, pi. 43(6), Fig. 166(3). Chlorodopsis pubescens: Serene and Luom, 1959: 316, fig. IB, 2D, 5C. Chlorodopsis melanodactylus: A. Milne Edwards, 1873: 229, pl. 7, fig. 7, 7a; Miers, 1884: 531 (part); Nobili, 1899: 258, 1900: 498; Caiman, 1900: 12; Ward, 1932: 251; Tweedie, 1950b: 121; Holthuis, 1953: 16; Guinot, 1958, fig. 23a, b. Chlorodopsis melanodactyla: De Man, 1902: 624; Gordon, 1934: 47; Serene and Luom, 1958: 125, pl. ID, Illf, IVg. Chlorodopsis pilumnoides: Balss, 1938a: 60 (part), non White, 1848. non Pilodius pubescens: De Man, 1902: 619=P.flavusRathbun, 1894. non IPilodius pubescens: Nobili, 1907: 395=P.flavusRathbun, 1894. non Pilodius pubescens: Serene, 1977: 51 (list)=/'. spinipes Heller, 1861. non Chlorodopsis melanodactylus: Miers, 1884: 531 (part)=P. spinipes Heller, 1861. non Chlorodopsis melanodactyla: Lenz, 1905: 255=P. spinipes Heller, 1861. Revision of Pj/o^ms 1147 Description Regions of carapace nearly indistinct, surface granulate, evenly covered with short fine setae. 2M entire. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules posteriorly on 2P. Submedian frontal lobes arched, denticulate, separated by 'U'-shaped emargination; lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth well defined, curved, terminating in chitinized spine. Anterior margin of cheliped merus spinose. External surface of cheliped carpus tuberculate, with furrow parallel with palmar joint. External surface of chela with wellspaced, large rounded tubercles on upper margin, diminishing in size inferiorly. Pereiopods setose, merus, carpus distinctly spinose on superior margin. Material examined Southeast Asia Vietnam; \S 11-5 mm; formerly Chlorodopsis melanodactylus and Chlorodopsis pubescens det. Serene and Luom (MNHN E 41.925 MP B.9387). Moluccas, Indonesia; coll. Rumphius Exp., 27 January 1975; 1? 10mm; det. Th. Monod and R. Serene (MNHN MP B.9921). Ternate, Moluccas; coll. W. Kukenthal; Ic? 8mm (SMF 1715). Banda Neira; 24 November 1929; \S 6-5mm formerly Chlorodopsis melanodactyla, det. I. Gordon, 1934 (IRSNB 9223). Indonesia; coll. R. Serene, 4 April 1963; \S 10-5 mm; det. R. Serene, 10 April 1963 (NUS 1969.11.24.31). Australia Murray Is., Queensland; coll. A. C. Haddon, 1888; 1S 16-5 mm; formerly Chlorodopsis melanodactylus, det. W. T. Caiman; pres. University College of Dundee (NHM 1954.9.14.74)—Torres Straits; \6 12mm (NHM 1954.9.14.73). Site 30, Elizabeth Reef; coll. 10 December 1987; 19 10 mm (AM P.38260)—Site 37; 12 December 1987; 1 9 13 mm (AM P.38256). North West Island, Capricorn Group, Queensland; coll. M. Ward, December 1929; \S 11-5-13 mm; formerly Chlorodopsis melanodactylus det. M. Ward (NHM 1931.4.14.19-20). Lizard Is., Queensland; coll. P. Davie and J. Short, 3 June 1987, l-2m; \6 13mm, 19 10mm (QM W.15464)— Wreck Reef, Coral Sea, nr Porpoise Cay; coll. J. Short and S. Mullens, 11 May 1988, 12m; IS 13mm, 1 9 10-9mm, 1 juv. (QM W. 15451)—coll. J. Short and B. Batley, 2-3m; \S 9mm, 19 10mm (QM W. 15450). Pacific Ocean Eastern Seas; coll. F. M. Rayner, HMS Herald; \S 13 mm; formerly Chlorodopsis melanodactylus det. E. J. Miers; (NHM 1862.53). Nouvelle-Caledonie; coll. M. Balansa; \S 15 mm; formerly Chlorodopsis melanodactylus det. A. Milne Edwards, Type and Chlorodopsis pubescens det. Serene and Luom (MNHN MP B. 13758)—7c? 15-10 mm, 3 9 ovig. 12-9-5 mm, 6 9 14-9-5 mm (MNHN MP B. 13757). Langon, Sta. 3, Touaouroo, Nouvelle-Caledonie; coll. 2 September 1978; 19 13mm (MNHN no reg.)—Sta. 20, He Amedee; coll. 17 September 1978, \S 10-5 mm, 4 9 9-8 mm (MNHN no reg.)—Sta. 79, Bale du Prony, He Ouen, 22°29'S, 166°29'E; coll. B. Richer de Forges, ORSTOM, 16m, 14 January 1987; \S 10mm, 19 13 mm (MNHN no reg.)—Sta. 82; 10 m; 1 9 11 mm (MNHN no reg.)—Sta. 159; 17m; \S 12mm, 2 9 ovig. 12-lOmm, 19 7mm (MNHN no reg.)—Sta. 160; 10m; 2S 14-11 mm, 5 9 ovig. 12-lOmm (MNHN no reg.)—Sta. 283; Secteur de Noumea, 22°27'S, 166°24'E; 13m; \S 12-lOmm (MNHN no reg.)—Sta. 480, Lagune Nord, 1148 P. F. Clark and B. S. Galil 18°56'S, 163°29'E; 31m, 2 March 1985, IS 11 mm, 19 10mm ovig., 1? 8mm (MNHN no reg.)—Sta. 483; 33 m, 2 March 1985; 1 c? 21 mm (MNHN no reg.)—Sta. 554, Grand Recif Sud, 22°50'S, 166°55'E; 16 July 1985, 27m; 16 13-8mm, 19 11mm (MNHN no reg.)—Sta. 899, Lagune Est, 20°14-2'S, 164°2515'E; 16m, 14 January 1987; 16 9-5 mm, 19 11 mm (MNHN no reg). Ovalua Is., Viti Levu, Fiji; coll. F. M. Rayner, HMS Herald; 26 6mm (NHM 1858.172). Namuba, Viti Levu, Fiji; coll. S. Bock; 1 6 9-5 mm; Formerly Chlorodopsis pilumnoides det. H. Balss; Munich Museum Exchange (NHM 1934.6.27.5). Viti, Fiji; 1 9 12-10 mm (SMF 1713) (Ex. Gottingen). Samoa; coll. Mus. Godeffroy; 26 11-9-5 mm, 19 10 mm; pres. University College Dundee (NHM 1955.1.5.9-10). Ohura, Polynesia; coll. G Seurat, 1905; 1 <? 5 mm; det. G Nobili, 1906 (MNHN MP B.13756). Sta. FPH-8a, Temae, Society Is., Polynesia; coll. H. G Muller, 30.iii.l988; 1 c? 11 mm (SMF no reg.). Remarks Chlorodopsis melanodactylus was described and figured by A.. Milne Edwards (1873) from New Caledonia, and it was this name that became widely used in the literature. Miers (1884: 531) identified Chlorodopsis melanodactylus from Africa or Eagle Island (now P. spinipes), Seychelles (now P. spinipes), Etoile Island (now P. flavus) and Oriental Seas (now P. pubescens). The Chlorodopsis melanodactyla of Lenz (1905) from Zanzibar pertains to P. spinipes. Balss (1938a: 59) synonymized both P. pubescens Dana and "^Chlorodopsis melanodactylus A. Milne Edwards with C. pilumnoides (White, 1848), a decision not supported by Tweedie (1950: 12) who claimed (as did Miers, 1884: 531) that the pigmentation found on the male cheliped of P. pubescens extended only very slightly on to the palm (propodus), a character that separated the species from P. pilumnoides. Although this character is adequate for separating these two species, part of Miers' material and Tweedie's (Serene, 1984: 244) was not Pilodius pubescens. A male syntype of C. melanodactylus was examined by Serene and Luom (1959: 316), and they concluded that the species of A. Milne Edwards agreed perfectly with the description and plates of P. pubescens Dana, 1852. S. Bock collected a single male from Namuba, Viti Levu, Fiji, which was identified by Balss (1938a: 60) as C pilumnoides and subsequently formed part of an exchange by the Munich Museum with the NHM. This male was redetermined as P. pubescens. Balss (1938a: 57) also synonymized the IPilodius pubescens of Nobili 1907: 395 with Pilodius flavus. Distribution From Nhatrang Bay, Vietnam to Samoa (Fig. 27). Not recorded from the Indian Ocean. Specimens recorded as Pilodius pubescens from Seychelles (Serene, 1977) and P. spinipes and the record of Tweedie (1950b) from the Cocos Keeling Is. must be considered dubious. Type status The type material was destroyed at the time of the Chicago fire, because Stimpson had on loan from the Smithsonian Institution a large collection of Crustacea including many type specimens of James D. Dana. Sooloo (Sulu) Sea, or Balabac Passage, were quoted by Dana as the type locality. Revision of Pilodius 1149 Pilodius pugil Dana, 1852 (Figs 13A-G, 37A, 43C) Pilodius pugil Dana, 1852: 80; 1853: 219, 1855, pi. 12,fig.8a-i; Heller, 1865: 19; Laurie, 1906: 406; Forest and Guinot, 1961: 91; Guinot, 1964b: 67, 1967: 268; Serene, 1968: 80 (list); Peyrot-Clausade, 1977a: 27, 1977b: 213; Chen and Lan, 1978: 267 pi. 4, fig. 14; Serene, 1984: 242, figs 143f, 147, pi. XXXIIIF; George and George, 1987: 238 (list). Table 5; Dai et al, 1986: 308, pi. 44(1), fig. 166(4); Dai and Yang, 1991: 332, pi. 44(1), Fig. 166(4). Chlorodopsis pugil: Nobili, 1907: 395; Gordon, 1934: 48; Ramadan, 1936: 33 (part); Balss, 1938a: 61; Sakai, 1939: 506; Ward, 1939: 10; Miyake, 1939: 216; Holthuis, 1953: 17; Sakai, 1956: 40 (Appendix); Guinot, 1958: 180; Serene and Luom, 1959: 319,figs2K, 3A, 5D, pi. II, fig. c, pi. Ill, figs D, D'. Chlorodopsis spinipes: A. Milne Edwards, 1873: 230 pi. 8, fig. b; De Man, 1887b: 282, 1892 278; Ortmann, 1893: 471; Henderson, 1893: 361; Zehntner, 1894: 151; Alcock, 1898: 169 Caiman, 1900: 12; Borradaile, 1900: 588, 1902: 261; De Man, 1902: 626; Rathbun, 1907 50, pi. 2,fig.5; Lenz, 1910: 551; Rathbun, 1911: 226; Ward, 1932: 251; Miyake, 1936:509 Serene and Luom, 1958: 135; pi. 1, fig. 13, pi. 4,fig.h; Sankarankutty, 1961: 121 (Ust), 129 Michel, 1964: 24. non Heller, 1861. Pilumnus globosus Boone, 1934: 152, pi. 78. non Chlorodopsis pugil: Klunzinger, 1913, pi. 6, fig. 18; Ramadan, 1936: 33 (part)=Pilodius spinipes Heller, 1861. Description Regions of carapace well defined by broad smooth deep-cut grooves, 1L-4L conical shaped, 2M entire, dorsal surface shagreened, with few scattered setae. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules posteriorly on 2P. Submedian frontal lobes trapezoid, crenulate, separated by distinct 'U'shaped groove, lateral lobes acuminate tubercles. Anterolateral margin tridentate, three equally prominent, large hook-shaped teeth lacking ancillary spines or granules. Anterior margin of cheliped merus prominently spinose. External surface of cheliped carpus and propodus studed with large, conic tubercles, smaller and obsolescent ventrally on propodus. Pereiopods fringed with long setae, superior margin of carpus, merus, propodus prominently spinose. Material examined Indian Ocean Abu Latt Is., Saudia Arabi; Red Sea; coll. Calypso, 1952; 19 7 mm, 1 juv.; det. D. Guinot, 1962 (MNHN MP B. 16483)—Red Sea; 19 6-5mm(MNHNMPB.16487). Aldabra; coll. Norman Sloan, 3 March 1978; 3 9 13-10-5 mm; det. R. Serene 1980 (NHM 1986: 211)—Passe Femme; coll. J. D. Taylor, 29 November 1967, \S 10-58 mm; det. R. Serene 1980; pres. Royal Society Expedition, Aldabra 1967-68 (NHM 1986: 206)—Main Channel; coll. J. D. Taylor, 10 December 1967; 1 juv. (NHM 1986: 208)—16 12-5-9-5 mm (NHM 1986: 210)—North side. Main Channel; 11 December 1967; 19 12 mm (NHM 1986: 212)—West side. East Channel; 30 September 1967; \S 13-12mm,49 14-9 mm (NHM 1986: 213). Aldabra; coll. A. J. Bruce; Ic? 14mm, 1 9 17 mm; det. R. Serene, 9 March 1980 (MNHN MP B.8014). He Mayotte, Archipel des Comores; coll. M. Marie, 1903; 3c?, 5 9 ; det. R. Serene (MNHN MP B.6782). Tamatave, Madagascar; coll. J. Millot; 2 9 ovig. 9-7 mm, 3 9 6-10 mm; formerly Chlorodopsis spinipes, det. H. Balss, redet. A Crosnier; (MNHN MP B.8016). 1150 P. F. Clark and B. S. Galil Mauritius; I d 13mm; formerly Chlorodopsis spinipes (NHM 1986: 420). Mauritius; \S 10 mm (MI no reg.), formerly Chlorodopsis spinipes, det. C. Michel. Mauritius; coll. P. Carie, 1910; \S 11mm; formerly Chlorodopsis spinipes, det. E. Bouvier; redet. J. Forest and D. Guinot 1960 (MNHN MP B.16480)—Port Louis; 2c? 11 mm, 2 $ 10-8-5 mm; redet. D. Guinot, 1960 (MNHN MP B. 16482). Mauritius, coll. M. Peyrot-Clausade; 5 9 11-6 mm, 3 juv. (MNHN MP B.6698). Sta. Re-1, Riffdach, Saline-les-Bains, Reunion; coll. H. G. MUller, 0-5-1 m, 20 January 1989; \6 14mm (SMF no reg.)—Sta. Re-U, Saline-les-Bains; 0-5-1 m, 25 January 1989; \e 14mm, 1 9 10mm, 1 juv. (SMF no reg.)—Sta. Re-35; 3-5 February 1989 \S 10 mm, 5 9 13-8 mm (SMF no reg.)—Sta. Re-31; 30 January-4 February 1989 3S 13-lOmm, 1 9 10-5 mm (SMF no reg.)—Sta. Re-35; 0-5-1 m, 3-5 February 1989 IS 10mm, 5 9 13-8mm (SMF no reg.). Egmont Reef, Chagos Archipelago; coll. J. S. Gardiner, Sealark 1905; 1 9 10-9 mm; formerly Chlorodopsis spinipes det. M. J. Rathbun (UMZC Aug. 7 1910)—Salomon; 2 9 11-5-9 mm (UMZC Aug. 7 1910)—Hulule, Male Atoll, Maldive Is.; coll. J. S. Gardiner; ?>S 11-5-9 mm, 2 9 ovig. 8-7 mm, 19 9-5 mm; formerly Chlorodopsis spinipes, det. L. A. Borradaile (UMZC no reg.)—Minikoi, Laccadive Is.; 19 11 mm (UMZC June 20 1900)—Funafuti, Ellice Is.; 15-25 fms; 2S 11-6mm (UMZC Jan. 11 1897). Gulf of Mannar; coll. W. A. Herdman, 1902; 1 S 14 mm; det. R. D. Laurie; pres. Miss Herdman (NHM 1934.1.16.85). Japan Ishigaki Is., Japan; 1 9 10-7 mm (SMF reg. 7733). Southeast Asia Phuket, Thailand; coll. R. Serene, 4 February 1966; 5 9 1970.3.13.8-12). 13-5-8 mm (NUS Pulau Mantabuan West, Sabah, Borneo; coll. J. D. and J. George, 30 August 1980; 1 juv.; pres. Semporna Marine Park Survey, World Wildlife Fund, Malaysia (NHM 1985: 325). Philippines; coll. 1859-1864; 19 11 mm (SMF 324a) (Ex. Museum Gottingen). Temate, Molluccas, Indonesia; coll. W. Kiikenthal; 1 S 12 mm; formerly Chlorodopsis spinipes det. J. D. De Man (SMF 1717). Amboina; coll. Strubell, 1891; \S 10 mm (SMF 1651). Ambon, Moluccas, Indonesia; coll. R. Serene, 1975; \S 8-5 mm, 1 9 7 mm, 1 juv. 4-5 mm; det. R. Serene (MNHN MP B.8026). Australia Murray Is., Torres Straits, Queensland; coll. A. C. Haddon, 1888; 19 11-5 mm; redet. R. Serene; formerly Chlorodopsis spinipes, det. W. T. Caiman; pres. University College Dundee (NHM 1954.9.14.75). North West Islet, Capricorn Group, Queensland; coll. F. A. McNeill, 1933; 1 9 9mm (AM P. 10375)—One Tree Is.; coll. J. C. Yaldwyn, November-December 1966; 1 9 ovig. 9-5 mm (AM P. 15967). Capri Cay, Swain Reef, Queensland; coll. October 1962, Swain Reefs Expedition; 1S 9-5 mm (AM P. 16961). North West Is., Capricorn Group, Queensland; coll. M. Ward, December 1929; 1 9 9mm; formerly Chlorodopsis spinipes; pres. M.Ward (NHM 1931.4.14.21). Revision of Pilodius 1151 Pacific Ocean New Caledonia; coll. 1903; 2 9 ovig. 7 mm, 1 9 10 mm (MNHN) formerly Chlorodopsis spinipes, det. A. Milne Edwards. Sta. 551, Grand Recif Sud, 23°00'S, 166°59'E, 9 m; coll. B. Richer de Forges, ORSTOM, 15 July 1985; 3 c? 12-6-5 mm, 1 ? 9 mm, 1 9 7 mm (MNHN no reg.). Gilbert Is.; \S 13 mm; formerly Chlorodopsis pugil, det. H. Balss; Munich Museum Exchange (NHM 1934.6.27.1). Samoa; Pres. S. J. Whitmee; \S 10mm (AM P.3171), exchange with NHM 1912 original reg. 1877:35. Samoa Is.; 3c? ll-10-5mm, 19 12-5-lOmm; Pres. S. J. Whitmee (NHM 1877:35). French Polynesia; coll. G. Seurat, 1905; 1S 6-5 mm, 19 6 mm; formerly Chlorodopsis pugil, det. G. NobiU, 1906 (MNHN MP B. 16486); Lagoon de Has; Ic? 8 mm (MNHN MP B.16477)—Ic? 8 mm; det. J. Forest and D. Guinot, 1960 (MNHN MP B. 16479)—2c? 9-7 mm, 8 9 12-7 mm; det. G. Nobih, 1906 (MNHN MP B. 16487)— Moorea, Tahiti; coll. M. Peyrot-Clausade; 2c? 11-8 mm, 19 10mm; det. R. Serene (MNHN MP B.6699)—coll. O. Odinetz; Ic? 8 mm, 4 9 11-7-5 mm (MNHN MP B.13598)—Taka Poto; 2c? 9-7 mm, 29 9mm, 5 juv.; det. D. Guinot, 1982 (MNHN MP B. 17067). Hikueru, Archipel des Tuamotu; coll. G. Rouson; 1 9 ovig. 9 mm; det. J. Forest and D. Guinot (MNHN MP B. 16478). Upolu; 1 c? 11 mm, 19 11 mm (SMF 1694) (Ex. Museum Gottingen). Temae, 17°29'S, 149°46'W, Society Is., Polynesia; coll. H. G. Miiller, 0-0-5 m; 31 March 1988; 21 specimens (SMF no reg.)—FPM-8a, 2m, 30 March 1988; 1 c? 11 mm, 2c? 9mm, 8 juv. (SMF no reg.)—FPM 1 lb, Tiahura Reef, Moorea; 1 m, 30 March 1988; 2c? 9-7 mm (SMF no reg.)—FPM-13, Afareaita; 0-0-5m, 29 March 1988; Ic? 10mm (SMF no reg.). Remarks The confusion between Pilodius pugil and P. spinipes was first noted by Nobili (1906, 1907). He stated that the description of Chlorodopsis spinipes given by Alcock (1898) was similar to the figure of P. pugil by Dana (1855: pi. 12, fig. 8a). Ramadan (1936) erroneously synonymized the Chlorodopsis spinipes of Nobili (1906), Laurie (1915) and Balss (1924) and the Chlorodopsis woodmasoni of Alcock (1898) with Ch. pugil. All this material belongs to P. spinipes. Serene and Luom (1959: 320) assigned the Pilumnus globosus of Boone (1934) to P. pugil. Distribution Balss (1938a: 62) remarked that only P. spinipes occurred in the Red Sea, and he considered P. pugil to be an oriental species. However, Guinot (1964: 68) identified three small Pilodius specimens collected in the Red Sea as P. pugil and showed that while P. spinipes was indeed restricted to the Indian Ocean, P. pugil had a wider distribution which extended from the Red Sea to the Tuamotu Archipelago (Fig. 28). Type status Dana's type specimen is no longer extant; it was destroyed in the great fire of Chicago while on loan to Stimpson. Type locality Upolu, Samoa, Pacific and Balabac Passage, north of Borneo. 1152 P. F. Clark and B. S. Galil Pilodius scabriculus Dana, 1852 (Figs 14A-G, 37B, 43D) Pilodius scabriculus Dana, 1852: 80, 1853: 220, 1855, pi. 12,fig.9; Nobili, 1907: 394; Forest and Guinot, 1961: 91, fig. 83a,b, fig. 84, 86; Guinot, 1967: 268; Serene, 1968: 80 (list); Peyrot-Clausade, 1977a: 27, 1977b: 213 (list); Thomassin, 1978: 64 (Appendix 3); Serene, 1984: 244; Dai et al, 1986: 309, pi. 44(2), fig. 166(5); Dai and Yang, 1991: 332, pi. 44(2), fig. 166(5). Chlorodopsis scabriculus: Edmondson, 1923: 17, 1925: 43. Chlorodopsis melanodactylus: Tweedie, 1950b: 121, pi. XVII d, e. Chlorodopsis spinipes: Sankarankutty, 1962: 139, fig. 40, 41. non Heller, 1861. Chlorodopsis venusta Rathbun, 1907: 49, pi. 1, fig. 5, 1911: 226; Caiman, 1909: 705 (list); Ward, 1941: 11; Tweedie, 1947: 27 (Ust); Serene and Luom, 1958: 131, pi. II, fig. C, pi. Ill fig. e, pi. IV, fig. f. Chlorodopsis venusta: Balss, 1938a: 53 (part). Pilodius etisoides Takeda and Miyake, 1968b: 186, fig. 2, pi. 8 C, D. Chlorodopsis pilumnoides: Garth, 1964: 140 (list), non White, 1848. non Chlorodiella venusta: Balss, 1938a: 53 (part) = Liocarpilodes integerrimus (Dana, 1852). non Chlorodopsis scabricula: Rathbun, 1906: 859, 1907: 50, pi. 1, fig. 3, pi. 9, fig. 5 = Phymodius nitidus (Dana, 1852). Description Regions of carapace nearly indistinct, better defined laterally, dorsal surface shagreened, sparsely setose. 2M entire. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules posteriorly on 2P. Submedian frontal lobes rounded, minutely denticulate, separated by shallow notch; lateral lobes triangular, denticulate. Anterolateral margin quadridentate, teeth granulate, anteriormost smallest, lacking curved apical spine. Anterior margin of cheliped merus distinctly spinose. External surface of cheliped carpus tuberculate, with furrow parallel with palmar joint. External surface of chela with conical tubercles on upper margin, smaller and obsolescent inferiorly, lower margin shagreened. Pereiopods sparsely setose, merus, carpus with conical tubercles on superior margin. Material examined Indian Ocean lies Glorieuses; coll. J. Millot, September 1958; \S 10mm, 1 2 9 mm; det. R. Serene (MNHM MP B.6733). He Europa; coll. P. Fourmanoir; 1 $ 11-5 mm; det. R. Serene (MNHN MP B.6732). Nosy Famy, Madagascar; coll. A. Crosnier, February 1962; 2S 11-5-11mm; det. R. Serene (MNHN MP B.6734). Coin Peros, Chagos Archipelago; coll. J. S. Gardiner, Sealark, 1905; \S 12mm; formerly Chlorodopsis venusta det. M. J. Rathbun; (UMZC Aug. 7 1910)—Salomon Is.; 1 9 8-5 mm, 1 juv. (UMZC Aug. 7 1910)—Coetivy; 2c? 9-5-5-5mm, 2 ? 8 mm, 1 juv. (UMZC Aug. 7 1910). Cocos Keeling Island, Indian Ocean; coll. C. A. Gibson-Hill; 3c? 10-5-9-5mm, 29 11-9 mm; formerly Pilodius melanodactyla det. M. W. F. Tweedie, redet. R. Serene 20/11/69 (NUS 1969.11.11.126-130). Christmas Island, Indian Ocean; coll. C. A. Gibson-Hill 1940; 19 8-5 mm; det. R. Serene 20/11/69 (NUS 1969.11.24.30). Christmas Island; coll. C. W. Andrews, Rewision of Pilodius 1153 1908; 4S 10-7-5 mm, 5 ? 9-5-6mm; formerly Chlorodopsis venusta det. W. T. Caiman, redet. J. Forest and D. Guinot (NHM 1909.5.19.54-58). Australia Thirteen miles SE of Cape Capricorn, Queensland; coll. Endeavour, \S 5mm (AM E.3146). Wreck Reef, nr Porpoise Cay, Queensland; coll. J. Short and B. Matheson, 9 May 1988, 0-5 m; 3 (? 12-8 mm, 5 9 10-7 mm, 2 juv. (QM W. 15443)—coll. J. Short and S. Mullens, 14 May 1988; \S 8 mm; det. P. Davie, 3 June 1988 (QMW. 15149)— 15 May 1988; \S 13 mm (QM W.15446). Green Is.; coll. Wm. A. Bartes; 1^ 8 mm; formerly Pilodius pilumnoides det. J. S. Garth (USNM 156078). Pacific Ocean Yap Is., Caroline Is., coll. R. W. Hyatt; \S 6 mm; formerly Pilodius pilumnoides det. J. S. Garth; (USNM 156077)—IcJ 15 mm; (USNM 156080)—Ic? 7 mm (USNM 156081). 8-13-3, Ifaluk Atoll, Caroline Is.; coll. D. Abbott and F. M. Bayer 1953; \S 4mm; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.)—39-C, Ifaluk Atoll; fragments (USNM no reg.)—74-B; 1S 5-5 mm (USNM no reg.)—73-G-3; 1S 3-5 mm (USNM no reg.). Teffaao, Huahine Is., 16°43'S, 151°00'W; coll. 2 May 1957; IS 9-5-6mm, 6 ? ovig. 8-7 mm ovig., 2 ? 7-6 mm, 13 juv., formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.). Saipan Id., Mariana Is.; coll. A. H. Banner 1945; 26 8 mm, 1 ? 8mm, 2 9 damaged, 20 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.)—1 9 6-5 mm (USNM 156079); 3c? 8-6mm, 69 ovig. 7-5-5mm; 1 9 5mm (USNM no reg.)—2c? 8-7 mm, 1 9 8 mm, 1 9 7 mm, 3 juv. (USNM no reg.)—1 S 7 mm, 2 juv. (USNM no reg.); \S 8 mm; (USNM no reg.)—1 c? 7 mm, 2 juv. (USNM no reg.). Arno Atoll, Marshall Is., Micronesia; coll. R. W. Hyatt, June 1950; 1 9 8 mm, 2 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM 134597)—2c? 10-9 mm, 3 9 9-7 mm (USNM no reg.)—Rigili Is., Eniwetok Atoll; coll. H. S. Ladd, May 1952; 1S damaged, 19 8 mm, 19 damaged (USNM no reg.)—East Rigili Is.; coll. J. P. E. Morrison 30 May 1952; \S 10mm (USNM no reg.). Canton Id., Phoenix Is., Polynesia; coll. C. A. Ely, January 1942; 1 9 7 mm; formerly Pilodius pilumnoides det. J. S. Garth (USNM 134595). Enyu Id., Bikini Atoll; coll. F. M. Bayer, 1 August 1947; 1 S 9 mm; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.)—Namu Id.; coll. M. W. Johnson, 3 April 1946; 3 c? 9-6 mm, 6 9 8 mm, 3 juv. (USNM no reg.)—coll. M. W. Johnson, 3 April 1946; 1 S 8 mm, 1 9 9 mm (USNM no reg.)—Enurikku Is.; 1 c? 8 mm (USNM no reg.). Namu Id.; Ic? 8mm (USNM no reg.)—Ic? 7-5mm, 1 9 7mm (USNM no reg.). West of Point Hauru, Moorea, Society Is., Polynesia; coll. 8 May 1957; 2 9 ovig. 7-6 mm, 12 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.)— Moorea; coll. Bredin Exp., 8 May 1957; 16 juv. (USNM no reg.). West of Motu Tapee, Bora Bora; coll. 25 April 1957; 2c? 7 mm, Ic? damaged, 1 9 8 mm; formerly Pilodius pilumnoides det. J. S. Garth; (USNM no reg.)—1 juv. 1154 P. F. Clark and B. S. Galil (USNM no reg.)—Bora Bora; 1 6 10mm, 1 $ ovig. 8mm, 6 juv. (USNM no reg.)— 1 9 9 mm, 1 $ 8 mm; 6 juv. (USNM no reg.). Raiatea, Tetaro Id.; coll. 29 April 1957; 1 $ 9-5 mm, 1 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.). Vaiorea Is.; coll. 1 May 1957; 1 9 7 mm, 2 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.). Palmyra Is.; 1 9 8mm (MNHN MP B. 16490). Temoe, Tuamotu; coll. L. G. Seurat, 22 April 1903; \S 7-5 mm, 19 9mm; det. G. Nobili, 1906 (MNHN MP B. 16494). Hao; coll. L. G. Seurat, 1905; 1 9 10mm; det. G. Nobili, 1906; (MNHN MP B.16489). Tahiti; coll. G. Ranson; 1 9 6 mm; det. Forest and Guinot 1960 (MNHN MP B. 16495). Hikueru; coll. G. Ranson; IS 7mm; 2 9 9-8mm; det. Forest and Guinot 1960 (MNHN MP B. 16496). Fakahina, Tuamotu; coll. L. G. Seurat, 1905; 1S 10 mm; det. G. Nobili, 1906 (MNHN MP B.16497)—1 juv. (MNHN MP B.16492)—Marutea, \S 10-5mm, 2 9 11-8mm (MNHN MP B. 16491)—Marutea; 1 9 6-5 mm ovig., 1 juv. (MNHN MP B.16493). Tickahau Atoll, Tuamotu Is., Polynesia; coll. Bredin Exp., 11 April 1957; \S 7mm, 2 9 7 mm, 15 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.)— 1 9 ovig. 6mm (USNM no reg.)—12 April 1957; 1 9 ovig. 9mm, 1 9 8mm (USNM no reg.)—11 April 1957; 1 juv. (USNM 156082)—13 April 1957; 1 9 5mm (USNM 156083)—Maiai Is.; 14 April 1957; \S 10-5mm 1 9 8mm ovig., 1 juv. (USNM no reg.). No data; 1 c? 8 mm, 2 juv.; formerly Pilodius pilumnoides det. J. S. Garth (USNM no reg.). Remarks Material identified by Rathbun from Honolulu (1906) and Tahiti (1907) as Chlorodopsis scabricula was assigned to Phymodius nitidus by Forest and Guinot (1961). Balss (1938) synonymized Chlorodiella asper Edmondson, 1925 with Chlorodiella venusta; however. Serene 1984: 263 assigned this material of Edmondson to Liocarpilodes integerrimus (Dana, 1852). Forest and Guinot (1971) considered Chlorodopsis venusta and Chlorodiella venusta as junior synonyms of Pilodius scabriculus. Specimens identified by Garth (1964) as P. pilumnoides belong to P. scabriculus. The Chlorodopsis spinipes of Sankarankutty (1962) and the Pilodius etisoides of Takeda and Miyake (1968) were assigned by Serene (1984: 245) to Pilodius scabriculus. Distribution Recorded from the Island of Europa, west of Madagascar, to Polynesia (Fig. 29). Not recorded from mainland east Africa, Red Sea or India. Type status Type specimen destroyed in the great fire of Chicago while on loan to Stimpson. Type locality Balabac Passage, South China Sea. Revision of Pilodius 1155 Pilodius spinipes Heller, 1861 (Figs 15A-G, 38A,B, 39A, 44A, C) Pilodius spinipes Heller, 1861a: 11-12, 1861b: 340-341, pi. Ill, fig. 22; Guinot, 1964a: 12, 1964b: 68, 1967: 268; Derijard, 1968: 1244; Peyrot-Clausade, 1977a: 27; Serene, 1984: 243 (part), figs 143g, 148, pi. XXXIV A, B. Pilodius aff. spinipes Serene, 1984: 244, fig. 143h, pi. XXXIV C. Chlorodopsis spinipes: De Man, 1881: 98; Alcock, 1898: 169; Nobili, 1901b: 14, 1906: 270; Laurie, 1915; 455 pi. XLIII, fig. 3a-d; Balss, 1924: 11; Monod, 1938: 132; Serene and Luom, 1959: 321-323, 338,figs2E, 3B, 4B, 5E, pi. IIB, IIIH; Guinot, 1958: 178fig.22a,b. Chlorodopsispilumnoides: Barnard, 1955: 3 (part); Guinot, 1967: 268 (part), non White, 1848. Chlorodopsis pugil: Klunzinger, 1913: 248 (152) pi. VI(X), figs 18a-c; Ramadan, 1936: 33 (part), non Dana, 1852. Chlorodopsis melanodactylus: Miers, 1884: 531 (part). Chlorodopsis melanodactyla: Lenz, 1905: 355. Chlorodopsis woodmasoni Alcock, 1898: 170, 1899, pi. 37,fig.7; Borradaile, 1902: 261; Lenz, 1910: 551; Rathbun, 1911: 226; Bouvier, 1915: 280(103), fig. 29; Michel 1964: 24. Chlorodopsis pubescens: Serene, 1977: 51 (list), non Dana, 1852. Etisus spinipes: Paul'son, 1875: 39, pi. VI, fig. I. Chlorodius niger: Ruppell, 1830: 20 (part). Chlorodiella nigra: Serene, 1984: 285 (part). non Chlorodopsis spinipes: A. Milne Edwards, 1873: 230 pi. VIII, fig. 6; De Man, 1887a: 282, 1892: 278; Ortmann, 1893: 471; Henderson, 1893: 361; Zehntner, 1894: 151; Alcock, 1898: 169; Caiman, 1900: 12; Borradaile, 1900: 588, 1902: 261; De Man, 1902: 626; Rathbun, 1907: 50, pi. n, fig. 5; Lenz, 1910: 551; Rathbun, 1911: 226; Ward, 1932: 251; Miyake, 1936: 509; Serene and Luom, 1958: 135 pi. IB, IVh; Sankarankutty, 1961: 121 (list), 129; Michel, 1964: 24 = Pilodius pugil Dana, 1852. non Chlorodopsis spinipes: Sankarankutty, 1962,fig.40,41 = Pilodius scabriculus Dana, 1852. non Pilodius spinipes: Serene, 1984: 243 (part) = Phymodius granulatus (Targioni Tozzetti, 1877). non Pilodius spinipes: McNeill, 1968: 74 = Pilodius nigrocrinitus Stimpson, 1859. Description Regions of carapace well defined, deep cervical groove in mesogastric region, surface unevenly granular, sparsely setose. 2M partly divided into two longitudinally. IP uniformly granulate; sinuous transverse row of adjoining pearliform granules posteriorly on 2P. Submedian frontal lobes rounded, denticulate, separated by 'U'shaped notch, external lobes small, spine-like. Anterolateral margin quadridentate, teeth well defined, clearly separated, curved, furnished with"ancillary spines distally, this being most pronounced in 2nd tooth. Anterior margin of cheliped merus spinose. External surface of cheliped carpus with large, conic tubercles, furrow parallel with palmar joint. External surface of chela closely set with long conical tubercles, smaller inferiorly, lower margin granulate. Pereiopods sparsely setose, superior margin of carpus, merus, propodus spinose. Material examined Indian Ocean Gulf of Suez, Red Sea; 16 13 mm, 1 9 9 mm, pres. R. MacAndrews (NHM 1869: 49). Dahab, Egypt; Gulf of Aqaba, Red Sea; coll. Manihine 14 February 1949; 16 14 mm (NHM 1951.1.17.34)—Sharm el Sheik; 2 February 1949; IS 13mm, 19 ovig. 13-5mm (NHM 1951.1.17.35). Biological Station, Ghardaqa, Egypt; Red Sea; coll. R. Gumey; 1$ 16 mm; pres. R. Gurney (NHM 1986: 419). 1156 P. F. Clark and B. S. Galil Red Sea; IS 12mm; formerly Chlorodius niger det. Ruppell (SMF 1724). Sharm Yenbo (Jambo), Saudi Arabia; Red Sea; coll. S.M.S. Pola, December 1895; 1 2 ovig. 10 mm; formerly Chlorodopsis spinipes det. Balss; Munich Museum Exchange (NHM 1934.6.27.2). Abu Latt Is., Saudi Arabia; Red Sea; coll. Calypso, 1952; IS 7-5mm, 2 ? ovig. 12-lOmm; det. D. Guinot (MNHN MP B.16507)—0-5-1 m; \S 14mm, 19 ovig. 11mm, 1$ 10-5 mm (MNHN MP B. 16508). Ras Zeili, Red Sea; coll. R. Ph. Dollfus; 19 9mm, 1 juv.; det. Th. Monod 1932 (MNHN PM B.13718). Sta. Vc, off Beacon Is., Khor Dongonab, Sudan; Red Sea; coll. Cyril Crossland, 26 April 1905; \S 7-5mm, 2$ 7-5-7mm; formerly Chlorodopsis spinipes det. R. D. Laurie; pres. Miss Herdman (NHM 1934.1.17.78)—Sta. VIII D, Suakin Harbour; 1905; 1 2 10mm (NHM 1934.1.17.79). Sta. 33, Sudan; Red Sea; coll. R. G. Hartnoll; \S 11mm; pres. R. G. Hartnoll; det. A. C. Edwards (NHM 1962.9.12.38). Perim and Obock, Djibouti, Red Sea; coll. F. P. Jousseaume, 1897; IS 14mm, 1 9 ovig. 15 mm, 19 13 mm; det. E. L. Bouvier (AM P.5402), exchange with Paris Museum September 1921, original reg. MP B.8057. Perim and Obock, Djibouti; Red Sea; coll. F. P. Jousseaume, 1897; 2AS 18-7-5 mm, 1 9 ovig. 12-5 mm, 21 9 17-8 mm, 2 damaged specimens; formerly Chlorodopsis spinipes det. Bouvier, verif. G Nobili 1905; 7 specimens Phymodius granulatus det. P. F. Clark (MNHN MP B.8057). Djibouti; coll. M. Maindron; 1S 15-5 mm; det. G NobiH 1905 (MNHM MP B.13715). Djibouti; 1S 16-5-11 mm (MNHN MP B. 13713)—coll. H. Coutiere; 1 9 10mm; det. G NobiH 1905 (MNHN MP B.13717)—lies Musha, Djibouti; coll. Mission Ch. Gravier, 1904; IS 10mm; det. G NobiH 1905 (MNHN MP B.13716). Aden; 4S 12-7-5mm, 3 9 12-7-5 mm; pres. Capt. J. W. Yerbury; (NHM 1884: 25)—2S 12-5-12mm, 5 9 12-8mm (NHM 1885.14). Sapper Bay, Aden; coH. Capt. K. England, October 1966; IS II mm, 29 10-9-5mm; pres. Capt. K. England (NHM 1974: 101)—Ras Tarshyne; IS 15mm, 19 10-9-5mm (NHM 1974: 112). Khasab Bay, Oman; coU. P. F. S. Cornelius 4 December 1971; 2 9 7-6-5 mm (NHM 1974: 142)—Khor Ghubb AH, North Oman; 18 December 1971, 10 fms; IS 12 mm, 19 ovig. 8 mm, 3 9 8-5-7 mm; 1 juv. 5 mm (NHM 1974: 89)—off Bukha; 26 November 1971, 10 fms; IS 14mm, 29 10-5-9mm (NHM 1974: 90)—Khasab Bay, off Ras Ghyyh Masud; 24 November 1971; 1 d 9 mm (NHM 1974: 143). Mirbat, Oman; IS 15-14mm (UY)—det. P. Hogarth; 1 juv. (UY)—Muscat al Bustan; 1 9 10mm (UY)—Sadh; 2S 11-10mm, 2 9 9-5-6-5 mm (UY)—Ic? 10-5mm, 19 ll-5mm (UY)—19 ovig. 9-5mm; 19 10mm, 1 juv. (UY)—^c? ll-5-5mm, 3 juv. (UY)—Raysat; IS 8-5mm (UY)—3c? 10-5-9mm; 19 10-5mm, 1 juv. 6mm (UY)—1 d 7 mm (UY). Island off Entre-Dent, Aldabra; coll. J. D. Taylor 19 April 1967; \S 16 mm; pres. Royal Society Expedition, Aldabra 1967-68 (NHM 1986: 201). Aldabra; coll. A. J. Bruce; IS 16-14mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.6779). lies Glorieuses; coll. A. Crosnier, 29 January 1973; 1S 16 mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.8024). Revision of Pilodius 1157 African or Eagle Is., Amirante Group; coll. R. W. Coppinger March 1882, HMS Alert 10 fms; IS 16mm; formerly Chlorodopsis melanodactylus det. E. J. Miers; pres. Lords of the Admiralty; (NHM 1882: 24)—no. 194; 4-12 fms; 4 c? 12-9 mm, 2 9 13-9 mm (NHM 1882: 24). Seychelles; 1? 9 mm; pres. N. Polunin (?) (NHM 1974: 583). Gan, Maldives; coll. P. S. Davies, 20 August 1964; \S 14 mm (NHM 1966:2:1:30). Mahe, Seychelles; 1S 12 mm; det. Balss (MNHN MP B. 16506). Praslin Reef, Seychelles, coll. J. S. Gardiner, Sealark 1905; 19 ovig. 14mm; formerly Chlorodopsis woodmasoni det. M. J. Rathbun; (UMZC Aug. 7, 1910)—Coetivy; 2S 19-12mm, 19 14mm; (UMZC Aug. 7, 1910)—Lagoon, Salomon, Chagos Archipelago; 12 fms; 16 10-5-9 mm (UMZC Aug. 7, 1910)—Diamant, Peros; 16 fms; 19 10mm (UMZC Aug. 7, 1910)— Hulule, Male Atoll, Maldive Is.; 36 16-10 mm, 1 9 14-5 mm; formerly Chlorodopsis woodmasoni det. L. A. Borradaile (UMZC June 20, 1900)—16 7 mm, 1 9 9-5 mm, 1 juv. (UMZC June 20, 1900)—Goidu, Gorfurfehendu Atoll; 46 15-7-5 mm, 1 9 ovig. 12 mm (UMZC June 20, 1900). Mayotte, Archipel des Comores; coll. M. Marie; 16 14mm, 19 13-5 mm; det. D. Guinot (MNHN MP B. 16505)—coll. A. Crosnier, September 1959; 26 15-514-5 mm, 4 9 14-10 mm; formerly Pilodius aff. spinipes det. Serene (MNHN MP B.8025). Delagoa Bay (Baia de Lourenco Marques), Mozambique; 1 9 ovig. 15 mm; formerly Chlorodopsis pilumnoides det. K. H. Barnard (SAM A 10855). Madagascar; 1 9 ovig. 14 mm, 1 9 14 mm; det. H. Balss (MNHN MP B. 16499) and (MNHN MP B. 19261)—coll. 1903; ?>6 16-5-15 mm, 5 9 ovig. 14-5-10mm; det. H. Balss (MNHN MP B.16504)—3c? 15-16-5mm, 5 9 ovig. 14-5-lOmm; det. H. Balss (MNHN MP B. 16504)—Nosy Be; coll. A. Crosnier, January 1958; 5 c? 18-11mm, 19 ovig. 16 mm, 2 9 13-12 mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.8017)—Tamatave; Ic? 14-10-5 mm; det. H. Balss (MNHN MP B. 16498)—Tulear; coll. G. Petit; 1 6 18 mm; det. H. Balss; formerly Pilodius aff. spinipes det. Serene (MNHN MP B.8018)—Tulear; coll. M. Peyrot-Clausade; 3 c? 18-7-5 mm, 1 9 13 mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.8019)—Tulear; Madagascar; coll. 17 May 1898; 1 9 14 mm; det. Balss (MNHN MP B.16500)—Tulear; coll. A. Grandidier 1903; Ic? 10mm, 2 9 12 mm (MNHN MP B. 19263)—Tulear; coll. P. Fourmanoir; Ic? 14-5 mm; formerly Actea depressa det. K. H. Barnard, Pilodius aff. spinipes det. R. Serene (MNHN MP B.8020)—Fort Dauphin; coll. R. Decary 1931; Ic? 11-5mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.8021)—Fort Dauphin; coll. A. Crosnier, 5 March 1973; 2c? 13-5-9-5 mm, 2 9 14-11mm; formerly Pilodius aff. spinipes det. R. Serene (MNHN MP B.8022). Mauritius; coll. P. Carie, 1913; 19 10 mm; formerly Chlorodopsis woodmasoni det. E. L. Bouvier (MNHN MP B. 13721)—Grand Port; 2c? 11-9mm (MNHN MP B. 16503)—5 c? 16-6 mm, 4 9 14-11mm; formerly Chlorodopsis woodmasoni det. E. L. Bouvier, Pilodius aff. spinipes det. R. Serene (MNHN MP B.8023)—2(? 12-11-5 mm; formerly Chlorodopsis woodmasoni det. E. L. Bouvier (MNHN no reg.)—Ic? 10mm; formerly Chlorodopsis woodmasoni (MI no reg.). Andamans; coll. Investigator, \6 A. Alcock, Type (ZSI 2032). 13 mm; formerly Chlorodopsis woodmasoni det. 1158 P. F. Clark and B. S. Galil Remarks Nobili (1906: 270, 1907: 395) commented on the confusion that had existed over the descriptions of three species Pilodius pugil Dana, 1852, P. spinipes Heller, 1861 and Chlorodopsis woodmasoni Alcock, 1898. The Ch. spinipes description by Alcock agreed with Dana's figure (1855: pi. 12, fig. 8a) of Pilodius pugil and fitted perfectly with Nobili's examples. After examining numerous specimens of P. spinipes from the Red Sea, the type locality of Heller's species, Nobili concluded that they accorded well with Ch. woodmasoni of Alcock, but Nobili was unable to confirm this statement as Heller's types were no longer extant in the Vienna Museum. However, Serene (1984: 243), after examining Alcock's syntypes of Ch. woodmasoni, supported Nobili by saying that this species was identical to P. spinipes. The extreme variation found in P. spinipes (pi. 2A, B) was shown by Serene (pis XXXIVa, b), from specimens collected in Djibouti (MP B.8057). Further, the material assigned by Serene (1984: 243) to Pilodius aff. spinipes (pi. 9A) was examined and it was found to be within the morphological variation exhibited by P. spinipes Heller. Serene suggested that Chlorodopsis melanodactyla (now as Pilodius spinipes) of Miers (1884: 531) from Seychelles, Barnard's (1955: 3) Pilodius pilumnoides (ovigerous 9 now P. spinipes) from Delagoa Bay (Baia de Lourenco Marques) Mo9ambique and McNeill's P. spinipes from Queensland (1968: 74), Australia (now P. nigrocrinitus) should be re-examined. Michel's Chlorodopsis spinipes and C. woodmasoni from Mauritius (1964: 24) were redetermined as Pilodius pugil Dana, 1852 and Phymodius drachi Guinot, 1964 respectively. Distribution Pilodius spinipes is restricted to the Red Sea and Indian Ocean (Fig. 30). The redetermination of McNeill's material leaves Alcock's Mergui Archipelago recording {Chlorodopsis woodmasoni) (1898: 171) as eastern-most Indian Ocean record. Type status A recent inquiry (Pitkin, personal communication) confirms that the type specimen is not extant in the Vienna Museum. Type Locality: Red Sea. Discussion Previous revisors of Pilodius Dana, 1851 were unsuccessful in resolving the complex taxonomic problems associated with this taxon. Many factors have contributed to the situation where 43 species (Table 2) have been described and referred to Pilodius. A major part of the problem is that only male specimens can be positively identified to species. Male pleopod morphology had great value as an effective differentiating character between the many old and new species established and male cheliped coloration pattern provide further stable identification features. Other characters are more variable; for example the difference in cheliped morphology between mature and immature specimens of the same species is marked, (Figs 40D, 41 A, 42B,C), also variation of exoskeleton spinature in males can be extreme (Figs 38A, B, 39A). Females of Pilodius are almost impossible to identify to species because their cheliped coloration patterns are different from those of the male counterpart (Figs 4 0 A ^ , 44AC?, 44B §, 44C 9), but are almost uniform between species (Figs 44B 9, 44C 9). Takeda and Miyake (1968b) based their description off*, etisoides on a single ovigerous female and, although this specimen was examined during the present study, the validity of the species cannot be confirmed in the absence of a male. Rathbun based 1159 Revision of Pilodius the description of Pilodius flavus on an immature female holotype measuring 9 m m in carapace width. She later described the same species under a different name, Chlorodopsis melanospinis, from a mature male specimen (Fig. 40D) and a number of paratypes. Examination of available P. flavus and Ch. melanospinis material enabled the present authors to synonymize the two species. This decision, although correct under the rules of nomenclature, makes the type worthless as a morphological representative of the species P. flavus. The ovigerous cotype of Ch. kauaiensis described by Edmondson from Kauai, Waimea, Hawaii (Bernice P. Bishop Museum reg. S 6876), was examined and the prolongation of the basal antennal segment was absent. Consequently, the antennal flagellum was not excluded from the orbital hiatus and therefore this species has been assigned to Pilodius erroneously. In Table 2, 14 species are junior synonyms. The creation of some of these could have been avoided if the available type material of other species had been examined first. For example, Chlorodopsis (Cyclodius) palaoensis Sakai 1938 is Pilodius pilumnoides (White, 1847), the types of which are extant and available for study. Unavailability of material may have caused many problems; the type specimens P. pubescens, P. pugil and P. scabriculus, P. granulatus and P. nigrocrinitus, were destroyed in the great fire in Chicago, 1871. This caused confusion between the identifications of P. spinipes and P. pugil and precluded the precise identity of P. granulatus. Two brachyuran collections are not available for study, the material of Sakai and Alcock, both are extant. The examination of Ch. melanochira, Ch. pilumnoides, Ch. spinipes and ICh. nigrocrinata as determined by Alcock (1898) would have solved many distribution and taxonomic problems, however the Zoological Survey of India did not make this valuable material available to the authors of this current study. la lb Ic Key to species of Pilodius 2M entire 2M partly divided into 2 longitudinally . 2M divided into 2 longitudinally 2a(la) 2b (la) Anterolateral margin of carapace 3-dentate Anterolateral margin of carapace 4-dentate 3a(2b) . 2 . 5 . 6 pugil . 3 . . 3b(2b) 2P with two parallel transverse rows of adjoining pearliform granules, anterior row medially disjunct P. paumotensis 2P with single sinuous transverse row of adjoining pearliform granules . . 4 4a(3b) 4b (3b) Carapace densely covered with short setae Carapace sparsely covered with setae 5a(lb) Carapace and pereiopods profusely furnished with dark bristly setae; upper margin of pereiopodal merus, carpus and propodus set with conical tubercules P. nigrocrinitus Carapace and pereiopods covered with long, light-coloured setae; upper margin of pereiopodal merus, carpus and propodus set with long slender spines P. flavus 5b(lb) P. pubescens P. scabriculsus 6a(lc) 6b(lc) 3M tripartite 3M entire 7 7a(6a) 7b (6a) Anterior lobe of 3M not reaching anterior margin of 2M Anterior lobe of 3M surpassing anterior margin of 2M 8a(6b) 8b(6b) 1L-5L with cornute conical tubercles 1L-5L lacking cornute conical tubercles. 9a (8a) Lower external surface of cheliped smooth . . . . . P. areolatus P. miersi . 9 . 12 P. concors sp. n. (Fig. 40C) 1160 9b (8b) P. F. Clark and B. S. Galil Lower external surface of cheliped granulate 10a (9b) Anterior margin of cheliped merus spinose 10b (9b) Anterior margin of cheliped merus granulate 1 la(9b) Dark coloration restricted to fixed finger only of chela. lib(9b) Dark coloration extends on to manus of chela 12a (8b) 12b (8b) Cheliped carpus with two furrows anteriorly Cheliped carpus with a single furrow anteriorly . . . 10 (Figs 40B, 41C, 42D) 11 P. maotieni P. cephalalgicus sp. n. (Fig. 40B) . P. pilumnoides (Fig. 42D) P. moranti sp. n. 13 13a (12b) 2P with two parallel transverse rows of adjoining pearliform granules, anterior row medially disjunct P. granulatus 13b (12b) 2P with single sinuous transverse row of adjoining pearliform granules P. spinipes Revision of Pilodius 1161 FIG. 1. Pilodius areolatus (H. Milne Edwards, 1834); NHM reg. 1931.7.24.95-100; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1162 FIG. 2. P. F. Clark and B. S. Galil Pilodius cephalalgicus sp. n.; MNHN reg. MP B 20935; Holotype; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius ^ o ^ 1163 OA 5 mm FIG. 3. Pilodius concors sp. n.; NUS reg. 1965.11.11.147; Holotype; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1164 FIG. 4. P. F. Clark and B. S. Galil Pilodius flavus Rathbun, 1894; USNM reg. 41268; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius 1165 FIG. 5. Pilodius granulatus Stimpson, 1859; figs (A-C) reg. NUS reg. 1985.1142-1151, figs (D, E) NUS reg. 1969.11.22.7-11; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. P. F. Clark and B. S. Galil 1166 A FIG. 6. Pilodius maotieni Serene, 1971; MNHN no reg.; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius 1167 FIG. 7. Pilodius miersi (Ward, 1936); figs (A-C) MNHN reg. MP B.9391, figs (D-G) NHM reg. no. 1937.7.15.21-23, Type; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1168 P. F. Clark and B. S. Galil FIG. 8. Pilodius moranti sp. n.; figs (A-C) AM reg. P.2372; figs (D-G) AM reg. P.2371 Holotype; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius FIG. 9. 1169 Pilodius nigrocrinitus Stimpson, 1859; figs (A-C) AM reg. P. 17208, figs (D-G) NHM reg. 1937.9.21.104-123; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1170 P. F. Clark and B. S. Galil FIG. 10. Pilodius paumotensis Rathbun, 1907; USNM reg. 32852; Type; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius -TvyxS^. 1171 ./X^^xr9^./r<i FIG. 11. Pilodius pilumnoides (White, 1848); figs (A-C) NHM reg. 1843.6, figs (D-G) NUS reg. 1965.11.11.148; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1172 P. F. Clark and B. S. Galil FIG. 12. Pilodius pubescens Dana, 1852; NHM reg. 1954.9.14.74; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius 1173 FIG. 13. Pilodius pugil Dana, 1852; NHM reg. figs (A-C) 1877.35, figs (D-E) NHM reg. 1986:213; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1174 P. F. Clark and B. S. Galil I mm FIG. 14. Pilodius scabriculus Dana, 1852; fig. (A) MNHN reg. MP B.6732, figs (B-G) MNHN reg. MP B.6734; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. Revision of Pilodius /\_ 1175 I mm FIG. 15. Pilodius spinipes Heller, 1861; NHM reg. 1986:201; (A) frontal margin, (B) left anterolateral margin, (C) propodus and dactylus of left 5th pereiopod, (D, E) ventral view of left pleopod, (F, G) dorsal view of left pleopod. 1176 P. F. Clark and B. S. Galil 5,^* 60 FIG. 16. 90 120 150 1180 Distribution of Pilodius areolatus (H. Milne Edwards, 1834). 90 FIG. 17. 120 jIBO !180 Distribution of Pilodius cephalalgicus sp. n. Revision of Pilodius 1177 FIG. 18. Distribution of Pilodius concors sp. n. 90 FIG. 19. |120 150 !180 Distribution of Pilodius flavus Rathbun, 1894. 1178 P. F. Clark and B. S. Galil 60 90 FIG. 20. 120 1150 ;180 Distribution of Pilodius granulatus Stimpson, 1859. FIG. 21. Distribution of Pilodius maotieni Serene, 1971. !150 Revision of Pilodius 90 FIG. 22. 120 150 1179 180 Distribution of Pilodius miersi (Ward, 1936). FIG. 23. Distribution of Pilodius moranti sp. n. 1180 P. F. Clark and B. S. Galil 90 120 150 180 FIG. 24. Distribution of Pilodius nigrocrinitus Stimpson, 1859. FIG. 25. Distribution of Pilodius paumotensis Rathbun, 1907. 60 90 120 150 !180 150 Revision of Pilodius 60 90 FIG. 26. FIG. 27. 120 150 1181 180 Distribution of Pilodius pilumnoides (White, 1848). Distribution of Pilodius pubescens Dana, 1852. 150 P. F. Clark and B. S. Galil 1182 S(^f FIG. 28. 90 FIG. 29. Distribution of Pilodius pugil Dana, 1852. 120 1150 180 Distribution of Pilodius scabriculus Dana, 1852. Revision of Pilodius FIG. 30. Distrihution of Pilodius spinipes HeWev, 1861. 1183 1184 P. F. Clark and B. S. Galil FIG. 31. Dorsal view; (A) Pilodius areolatus (H. Milne Edwards, 1834), NHM reg. 1931.7.24.95-100; (B) Pilodius cephalalgicus sp. n., MNHN reg. MP B.20935, Holotype. Revision of Pilodius FIG. 32. 1185 Dorsal view; (A) Pilodius concors sp. n., NUS reg. 1965.11.11.147, Holotype; (B) Pilodius flavus Rathbun, 1894, USNM reg. 41268. 1186 FIG. 33. P. F. Clark and B. S. Galil Dorsal view; (A) Pilodius granulatus Stimpson, 1859, NUS 1965.11.11.119-125; (B) Pilodius maotieni Serene, 1971, MNHN no reg. Sta. 483. 1187 Revision of Pilodius ^ B^^^Br^ '^^^^^99 ^i^^H ^>^^^^^^H FIG. 34. Dorsal view; (A) Pilodius miersi (Ward, 1936), NHM reg. 1937.7.15.21-23; (B) Pilodius moranti sp. n., AM reg. P.2371, Holotype. 1188 FIG. 35. P. F. Clark and B. S. Galil Dorsal view; (A) Pilodius nigrocrinitus Stimpson, 1859, AM reg. P.17259; (B) Pilodius paumotensis Rathbun, 1907, MNHN reg. MP B.6697. Revision of Pilodius FIG. 36. 1189 Dorsal view; (A) Pilodius pilumnoides (White, 1848), NHM reg. 1843.6, Type; (B) Pilodius pubescens Dana, 1852, NHM reg. 1931.4.14.19-20. 1190 FIG. 37. P. F. Clark and B. S. Galil Dorsal view; (A) Pilodius pugil Dana, 1852, NHM reg. 1934.6.27.1; (B) Pilodius scabriculus Dana, 1852, MNHN reg. MP B.9383. Revision of Pilodius 1191 FIG. 38. Dorsal view; (A, B) Pilodius spinipes Heller, 1861, MNHN reg. MP B.8057. 1192 P. F. Clark and B. S. Galil FIG. 39. Dorsal view; (A) Pilodius aff. spinipes Serene, 1984, MNHN reg. MP B.8017. Revision of Pilodius ^ ^ ^ ^ ^ H ^ \"'^ ^M>. ^ W^y- t^''^--5^|i 1193 if-/' "''^'^^^B /'t^i^M| '- 'li^lH \ i '• -.„ -..-,. rj^^ ""'^i ^ ^ ^ FIG. 40. External surface of 6 chelipeds; (A) Pilodius areolatus (H. Milne Edwards, 1834), NHM reg. 1931.7.24.95-100; (B) Pilodius cephalalgicus sp. n., MNHN reg. MP B.20935, Holotype; (C) Pilodius concors sp. n., NUS reg. 1965.11.11.147, Holotype; (D) Pilodius flavus Rathbun, 1894, USNM reg. 41268 (mature 6). P. F. Clark and B. S. Galil 1194 E^l - 4.:*A ^B^' '^ ^s^l^*i ". ^^m m^'^^K^' te: ^^^^^tm El ^^S^p' ^^^^^^^:r.^^'^' FIG. 41. External surface of 6 chelipeds; (A) Pilodius flavus Rathbun, 1894, NHM reg. 1907.5.22.241^5 (immature c^); (B) Pilodius granulatus Stimpson, 1859, NUS reg. 1965.11.11.119-125; (C) Pilodius maotieni Serene, 1971, MNHN no reg. Sta. 483; (D) Pilodius miersi (Ward, 1936), NHM reg. 1937.7.15.21-23, Type. Revision of Pilodius 1195 FIG. 42. External surface of S chelipeds; (A) Pilodius moranti sp. n., AM reg. P.2371, Holotype; (B) Pilodius nigrocrinitus Stimpson, 1859, AM reg. P. 17259, (immature S); (C) Pilodius nigrocrinitus Stimpson, 1859, AM reg. P. 17259, (mature 6); (D) Pilodius pilumnoides (White, 1848), NHM reg. 1843.6, Type. 1196 P. F. Clark and B. S. Galil FIG. 43. External surface of 6 chelipeds; (A) Pilodius paumotensis Rathbun, 1907, MNHN reg. MP B.6697; (B) Pilodius pubescens Dana, 1852; (C) Pilodius pugil Dana, 1852, NHM reg. 1934.6.27.1; (D) Pilodius scabriculus Dana, 1852, MNHN reg. MP B.9383. Revision of Pilodius 1197 FIG. 44. External surface of S chelipeds; (A) Pilodius spinipes Heller, 1861, MNHN reg. MP B.8057; external surface of ? chelipeds; (B) Pilodius areolatus (H. Milne Edwards, 1834), NHM reg. 1931.7.24.95-100; (C) Pilodius spinipes Heller, 1861, MNHN reg. MP B.8017. Acknowledgements We thank our colleagues who entrusted us with material, without their help and support this paper would never have been completed: Harold Feinberg (AMNH); Roger Springthorpe (AM); K. Wouters (IRSNB); Beatrice Burch (BBM); Teru Aki Lchida (ZLKU); Claude Michel (MI); Alain Crosnier (ORSTOM, c/o MNHN); Peter Ng (NUS); Vu Do Quynh (OINT); Peter Davie (QM); Michael Turkay (SMF); Michelle van der Merwe (SAM); Ray Manning (USNM); Ray Symonds (UMZC); Peter Hogarth (UY); Katsushi Sakai (SWU); Linda Pitkin (NHM), while on study leave to Vienna Museum combed the reference collection for the type of Pilodius spinipes Heller, we thank her for this effort. We are grateful to Phil Hurst, NHM Photo Unit, for the photographs reproduced in this paper. 1198 P. F. Clark and B. S. Galil References ADAMS, A. and WHITE, A., 1849, Crustacea, in A. Adams (ed). The Zoology of the voyage of HMS Samarang under the command of Captain Sir Edward Belcher during the years 1843-1846 (London: Reeve, Benham, and Reeve) (part II): viii + 33-66, pis 7-13. [For dates of publication see Sherborn, C. D, 1922: cxi.] ALCOCK, A . , 1898, Materials for a Carcinological Fauna of India. No. 3. 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