31 December 2003 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1I6(4):901-919. 2003. On two new genera of pea crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura: Pinnotheridae) from the Indo-West Pacific, with notes on aUied genera

Peter K. L. Ng and Raymond B. Manning*

(PKLN) Department of Biological Sciences, National University of Singapore, Kent Ridge, Singapore 1 19260, Republic of Singapore, e-mail: [email protected]; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163

Abstract.—Two new genera of pea crabs of the family Pinnotheridae, Hol- otheres and Buergeres, members of which are parasites of sea cucumbers, are described from the Indo-West Pacific. Species of Holotheres are characterized by having all the dactyli of the walking legs of females short and subequal in length, with the spatuliform dactylus of the third maxilliped inserted very near or at the base of the propodus. Species of Buergeres are characterized by having the last dactyli of the walking legs of females elongate and longer than those of the other legs, with the subspatuliform dactylus of the third maxilliped inserted distinctly before the base of the propodus. Holotheres and Buergeres are compared to other genera whose members are associated with holothurians, Holothuriophilus Nauck, 1880, and Alain Manning, 1998 (obligate parasites); as well as Pinnaxodes Heller, 1865 (facultative parasites).

In a synopsis of the Pinnotheridae, Man- In addition, the third maxilliped of most of ning (1993a: 127, 128) defined the genus these species has the inner margin usually Pinnotheres Bosc, 1802, sensu stricto as angular, and the carpus of the third maxil- having a subhexagonal or subcircular cara- liped is distinctly shorter than the propodus pace without any surface ornamentation or that is relatively long and slender. In this sculptures, a third maxilliped which has the combination of characters, these species are ischium and merus completely fused (with- very different from almost all other taxa out any trace of a suture) and a three-seg- now classified in Pinnotheres sensu stricto mented palp in which the dactylus is slen- or any of the other non-holothurian sym- der, inserted basally on the propodus; the biotic genera recognized or described by dactyli of the first to fourth walking legs are Manning (1993a, 1993b) and other work- subequal and similar, short and strongly ers. Manning (1993a: 128) did note briefly curved, with the tips simple; and all the in his synopsis, however, that one genus seven male abdominal somites are free, parasitic in holothurians, Holothuriophilus Manning (1993a), however, did not treat Nauck, 1880, was valid, and subsequently most of the species of Pinnotheres which elaborated more on this (Manning 1993b). live in holothurians, all of which have a Currently, 16 species of pinnotherid third maxilliped in which the dactylus is not crabs are known to be parasites of holothu- slender but spatuliform to subspatuliform. rians, and of these, 13 occur only in the Indo-West Pacific (see Schmitt et al. 1973, * Deceased 18 January 2000. Hamel et al. 1999). The eastern Pacific spe- 902 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

cies which are obligate parasites in holo- maxilliped; WLl-4 refer to the first to thurians are both members of Holothurio- fourth walking legs respectively; and Gl to philus Nauck, 1880, although one species the male first pleopod. The terminology fol­ of Opisthopus Rathbun, 1893, is occasion­ lows Manning (1993a). Measurements are ally found in sea cucumbers (Rathbun of the carapace width and length, respec­ 1918, Schmitt et al. 1973). The 13 Indo- tively. Specimens examined are deposited West Pacific species are Alain crosnieri in the Zoological Reference Collection of Manning, 1998 (Indonesia); Ostracotheres the Raffles Museum, National University of holothuriensis (Baker, 1907) (South Austra­ Singapore (ZRC); Senckenberg Museum, lia); Pinnaxodes major (Ortmann, 1894) Frankfurt-am-Main (SMF); National Sci­ (Japan); Pinnotheres deccanensis Chopra, ence Museum, Tokyo (NSMT); Zoological 1931 (southern India); P. flavus Nauck, Museum of the University of Copenhagen 1880 (Philippines); P. halingi Hamel, Ng & (ZMUC); and National Museum of Natural Mercier, 1999 (Solomon Islands); P. holo- History, Smithsonian Institution, Washing­ thuriae Semper, 1880 (Philippines); P. ort- ton, D.C. (USNM). manni Burger, 1895 (Philippines, Peninsu­ lar Malaysia, Singapore); P. pilumnoides Notes on the Material of Otto Burger Nobili, 1905 (Red Sea, Gulf of Aden); P. semperi Burger, 1895 (Java, Sumatra, Su­ A note on the pinnotherid material of lawesi, Singapore); P. setnai Chopra, 1931 Semper, Nauck and Burger in the USNM is (Andaman Islands); P. tenuipes Burger, necessary. In 1906, the USNM received six 1895 (Philippines); and P. villosissimus Do- lots of pinnotherid crabs on exchange from flein, 1904 (Sumatra, Andaman Islands, the University of Gottingen in Germany. New Guinea) (Chopra 1931, Schmitt et al. This material was from the collections 1973, Pregenzer 1988, Van den Spiegel & made by C. Semper and was used for sev­ Jangoux 1989, Hamel et al. 1999). eral studies by Semper (1880), Nauck Most of the species parasitic in holothu- (1880) and Burger (1895), all of whom rians now placed in Pinnotheres sensu lato were at the University of Gottingen. All of can, however, be divided into two distinct this material was reported by Burger "groups" on the basis of the dactyli of their (1895). This shipment included: Pinnoth­ walking legs and form of the third maxil­ eres ajfinis Burger, 1895 (1 ovigerous fe­ liped (see Hamel et al. 1999). Members of male, 9.5 by 8.2 mm, Bohol, Philippines, one "group" (P. semperi, P. flavus, P. vil­ USNM 32435); Pinnotheres exiguus Biir- losissimus and P. setnai) have all the dac­ ger, 1895 (1 ovigerous female, 5.6 by 4.6 tyli of the female walking legs short and mm, Samar, Philippines, USNM 32432); similar in length, and the dactylus of the Pinnotheres flavus Nauck, 1880 (2 oviger­ palp of the third maxilliped inserted at the ous females, 8.4 by 7.8 mm, 10.0 by 9.5 base of the propodus. Members of the sec­ mm, Zamboanga, Philippines, USNM ond "group" {P. ortmanni, P. holothuriae, 32434); Pinnotheres glaber Burger, 1895 (1 P. tenuipes and P. deccanensis) have the ovigerous female, 7.8 by 6.5 mm, Palawan dactyli of the last female walking leg sty- Island, Palaos Islands, USNM 32431); Pin­ liform and much longer than the short and notheres holothuriae Semper, 1880 (2 ovig­ hooked dactyli of the other three pairs of erous females, 10.1 by 9.3 mm, 8.0 by 6.5 legs, and the dactylus of the palp of the mm, Zamboanga, Philippines, USNM third maxilliped inserted well before the 32433); and Xenophthalmus latifrons Biir- base of the propodus. ger, 1895 (1 ovigerous female, 9.5 by 8.1 The present paper serves to diagnose two mm, Bohol, Philippines, USNM 32436). new genera for these two "groups". The Mary Jane Rathbun, then Second Assis­ abbreviations MXP3 refers to the third tant Curator of Invertebrates at the USNM, VOLUME 1 16. NUMBER 4 903

considered these specimens to be types. lothurians. Carapace circular to subcircular, Checking with various museums, including often highly setose; posterior carapace mar­ the collections in the University of Gottin- gin distinctly convex. WL subequal right gen, Schmitt et al. (1973) tracked down and left; WL3 similar in length, longer than many of the specimens of Semper, Nauck WL4; female WLl-4 with dactyli short, and Burger but commented that the types similar in length and form. Ischium and of P. exiguus, P. flavus, P. holothuriae and merus of MXP3 completely fused, without X. latifrons were lost or unknown. Curious­ trace of suture; dactylus of palp spatuli- ly, Schmitt did not check his own collec­ form, subequal in length to propodus, in­ tions at the USNM. Apparently, sometime serted near base of propodus. Gl simple, after 1906, at least some of Semper's ma­ curved tube, tapering from base to tip, terial was transferred to the Zoological Mu­ slightly recurved distally. seum, Berlin, where the true types of P. af- Etymology.—The name is derived from finis and P. glaber are still extant (O. Cole­ an arbitrary combination of the name for man, in litt.). There are no types of P. ex­ sea cucumbers Holothuria, and Pinnother­ iguus, P. flavus, P. holothuriae and X. es. Gender masculine. latifrons in the Berlin Museum (O. Cole­ Remarks.—Among taxa wholly symbi­ man, in litt.). The surviving specimens, rep­ otic with holothurians, Holotheres is per­ resenting the bulk of Semper's material in haps closest to Holothuriophilus Nauck, the University of Gottingen, were recently 1880, sensu Manning, 1993b (type species transferred to the Senckenberg Museum. Holothuriophilus trapeziformis Nauck, Michael Tiirkay and Andreas Allspach 1880). The dactylus and propodus of the kindly provided a list of the extant Gottin­ palp of MXP3 of Holothuriophilus species gen pinnotherid specimens now in the SMF. (Fig. 7C), however, is distinctly shorter than Of the species reported from holothurians those of Holotheres species (Figs. IB, 3B). by Semper, Nauck and BUrger, only speci­ In any case, Holothuriophilus is known mens of two species are still extant, P. fla­ only for certain from the eastern Pacific vus and Holothuriophilus trapeziformis. (Manning 1993b). Takeda & Prince Masa- There were also several lots in which the hito (2000) transferred Pinnaxodes mutuen- labels have been lost, but examination of sis (Sakai, 1939), which is known only this material by one of us (PKLN) did not from bivalves, to Holothuriophilus but we uncover specimens of any pinnotherid spe­ do not agree (see "Notes on Pinnaxodes cies parasitic in holothurians. On the basis and Holoturiophilus"). of this material, we also provide herein fig­ Two other genera, Pinnaxodes Heller, ures of P. flavus and P. holothuriae, each 1865, and Opisthopus Rathbun, 1893, have of which are transferred to one of the two some species which are facultatively found new genera here recognized. in holothurians (Rathbun 1918, Schmitt et al. 1973), and have carapaces and MXP3 Taxonomic Account similar to those of Holotheres. Species of Holotheres can be easily separated from Holotheres, new genus Pinnaxodes in having the posterior carapace Type species.—Pinnotheres semperi Biir- margin prominently convex (Figs. lA, J, ger, 1895, by present designation. 2A, 3A) (versus slightly convex to almost Included species.—Five: H. flavus straight. Fig. 6A, E), a proportionally lon­ (Nauck, 1880), H. halingi (Hamel, Ng & ger dactylus and propodus of the MXP3 Mercier, 1999), H. semperi (Burger, 1895), (Figs. IB, 3B versus Fig. 6B, F), the dac­ H. setnai (Chopra, 1931), and H. villosis- tylus of the MXP3 is inserted near the base simus (Doflein, 1904). of the propodus (Figs. IB, 3B) (versus in­ Diagnosis.—Obligate symbionts in ho­ serted % from base of propodus) (Fig. 6B, 904 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Holotheres semperi (Burger, 1895). A-I, neotype male (7.9 by 7.2 mm) (ZRC 1965.11.24.40); J, female (10.7 by 10.2 mm) (ZRC 1965.11.24.41). A, overall view of specimen (setae on right half of body denuded); B, left MXP3, outer view; C, right chela, outer view; D, WLl; E, WL2; F WL3; G, WL4; H, male abdomen; I, left Gl; J, carapace, dorsal view. (Except for A, all structures denuded.) Scales: A, J = 5.0 mm, B-I = 1.0 mm.

F), the ischium and merus are completely Campos et al. 1998: fig. 3A, C; Takeda & fused without any trace of a suture (Figs. Masahito 2000: fig. 4B) (see also Campos IB, 3B versus Fig. 6B, F), and the male et al. 1998, Takeda & Prince Masahito telson is semicircular (Figs. IH, 2G) (versus 2000). Opisthopus has the same male ab­ usually medially expanded laterally to form domen form as Holotheres, but can be sep­ rounded to subpentagonal structure, cf. arated by the same carapace and MXP3 VOLUME 1 16. NUMBER 4 905

characters discussed above for Pinnaxodes eral regions, completely obscuring outline (see Campos et al. 1998, Campos & Man­ and surface; intestinal region slightly de­ ning 2000). pressed. Anterolateral margin of male al­ Of the five species here included in Hol­ most straight, lateral margin gently convex, otheres, the types of two species, H. sem- posterolateral margin gently concave; an­ peri (Burger, 1895) and H. flavus (Nauck, terolateral and lateral margin of female 1880), are considered to be lost. The tax­ more confluent, gently convex; sub-bran­ onomy of only H. semperi and H. flavus are chial surface densely setose. MXP3 with is­ discussed herein. Holotheres setnai has chium and merus completely fused, without been well described and figured by Chopra trace of suture, subovate, inner margin sub- (1931). Holotheres halingi has been de­ angular submedially; exopod relatively scribed and figured in detail by Hamel et stout, outer margin convex, with well de­ al. (1999). Holotheres villosissimus (see veloped setae on flagellum; palp 3-seg- Doflein 1904, Van den Spiegel & Jangoux mented, propodus elongate, conical, sub- 1989) is not well known and must be re- equal in length to dactylus; dactylus spa- described when fresh specimens become tuliform, articulates at base of propodus. available. Chelipeds subequal; surfaces of merus, car­ pus, palm and fingers densely covered with Holotheres semperi (Burger, 1895) setae, those on inner surfaces relatively lon­ Fig. 1 ger; palm relatively stout; fingers shorter than palm; cutting edges of dactylus and Pinnotheres semperi Burger, 1895:382, pi. propodus each with sub-basal large tooth 9 fig. 28, pi. 10 fig. 27.—Lanchester, and numerous denticles before and after it; 1900:761.—Chuang, 1961:189, pi. 94.— distalmost part of fingers excavated on in­ Schmitt et al., 1973:85. (See Schmitt et ner surface. WL relatively stout, WL2 > al. (1973:85) for additional references). WL3 > WLl > WL4 in length (basis-is- Neotype.—Male (7.9 by 7.2 mm) (ZRC chium to tip of dactylus); dactyli of WLl- 1965.11.24.40) in respiratory tree of Holo- 4 relatively long, sharply tapering, all sub- thuria scabra, coll. S. H. Chuang, 1950s. equal in length; dorsal and ventral margins Other material examined. — 1 female of meri, carpi and propodi of all legs dense­ (10.7 by 10.2 mm) (ZRC 1965.11.24.41), ly lined with setae of various lengths, com­ same data as neotype. 1 male (10.5 by 10.0 pletely obscuring margins. Male abdomen mm), 1 female (4.8 by 4.3 mm) (ZRC with all 6 segments distinct, freely articu­ 1965.11.24.38-39), in cloaca of Holothuria lating, telson semicircular; lateral margins scabra, coll. S. H. Chuang, 1950s. 2 males of segments 3-6 gently concave. Female (7.7 by 6.8 mm, 7.0 by 6.6 mm), 2 females abdomen with all segments freely articulat­ (11.2 by 9. mm, 10.7 by 9.0 mm) (all spec­ ing. Gl relatively slender, gently sinuous, imens badly damaged and carapace crushed tip distinctly curving upwards. in parts) (ZRC 1965.11.24.34-37), in clo­ Biology.—Holotheres semperi has been aca of Holothuria scabra, coll. S. H. reported from inside Holothuria fursociner- Chuang, 1950s. 1 male (8.4 by 7.9 mm) ea and H. scabra (see Schmitt et al. 1973). (ZRC 1965.11.24.16), 6-12 fathoms, from Remarks.—The identity of Pinnotheres sea cucumber, Pulau Tekong, Singapore, semperi Burger, 1895, is a problem. In de­ coll. M. W. F. Tweedie, 1960s. scribing the species on the basis of one Diagnosis.—Male carapace subquadrate, male and one female. Burger (1895:382, pi. width to length ratio ca. 1.1; female cara­ 9 fig. 28, pi. 10 fig. 27) described and fig­ pace subcircular, width length ratio 1.1-1.2; ured a female specimen with a subhexagon- dorsal surface gently convex, covered with al carapace highly setose over its carapace dense layer of short setae longer along lat­ and legs, well produced triangular frontal 906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

margin, WL4 substantially shorter than the male figured by Burger (1895: pi. 9 fig. rest of the legs, and MXP3 in which the 28a) is an important one. The females on dactylus of the palp is spatuliform and as hand all have very narrow, non-protruding long as the propodus. A pair of specimens front, with the margin distinctly sinuous, was subsequently reported by Lanchester and the eyes are not visible in dorsal view (1900:761) who noted: "The specimens (Fig. IJ). In contrast, males have a more which Dr. Burger has described came from produced front, appearing subtruncate when the cloaca of Hoi. fusco-cinerea; whereas viewed dorsally (Fig. lA). Even when a the specimens we obtained from that Ho- specimen is angled such that the anterior lothurian are, I find, referable to P. ortman- part of the carapace arches upwards, the ni. The host, from which came Dr. Burger's frontal margin never appears triangular. The example of the latter species, was, I sup­ frontal margins of uncleaned male speci­ pose, not known, as he makes no reference mens, however, are densely lined with setae to it. In both these species, the animal was and when covered with fine sediment, the found high in the respiratory tree, forming "front" does appear to be more triangular an enormous gall; whether it would be able in shape, especially when observed dry and to extricate itself from this, and move, at the setae clump medially. None of the ex­ will, up or down the respiratory tree, I do amined female specimens, however, resem­ not know, but it has the appearance of a ble the specimen figured by Burger, even fixture, nolens volens'\ The species was when they are not cleaned. It would thus next reported by Chuang (1961:189) who appear that Biirger might have figured an noted that the species occurs in the respi­ uncleaned, undenuded male rather than a ratory tree of the sea cucumber, Holothuria female. It is also pertinent to note that the scabra: "A pair of them consisting of a carapace shapes of the present males agree male and a female may occur in one single well with Burger's figure, although the pre­ sea cucumber. The relatively thick carapace, sent specimens do not appear to be as se­ about 1 cm in diameter, is white in colour tose. The distinct pattern of setae figured by and has a circular outline. The cylindrical Burger is certainly not at all apparent in the legs and both chelipeds are hairy". present specimens. Biirger (1895:382) not­ In their synopsis of the Pinnotheridae, ed that the male he had, measured 7.0 mm Schmitt et al. (1973) stated that the male by 6.8 mm, and the female 6.5 by 6.5 mm. and female syntypes of H. semperi (muse­ All the present specimens examined are um number not stated), which were origi­ broader than long. Lanchester (1900:761), nally in the University of Gottingen, are no however, reported that his specimens were longer extant. The Gottingen material, as longer than broad (male 10.5 by 11.0 mm, discussed earlier, was examined by one of female 7.75 by 8.0 mm). Whether these us (PKLN), and it does not contain any measurements were accurate or if they were specimens which can be identified with this taken from cleaned specimens could not be species. A check with the Berlin Museum ascertained. The very short WL4 is note­ confirms that the types of H. semperi are worthy, as none of the present specimens not there (O. Coleman, in litt.). They are on hand (male or female) has such a short also not in the USNM, and are thus almost WL4 as figured by Burger (1895: pi. 9 fig. certainly lost. 28a). It seems likely though, that the WL4 of Burger's specimen appears shorter than Comparisons of the present series of it actually is because the leg was bent specimens from Singapore and the descrip­ slightly upwards when it was drawn. The tion and figures of the types by Burger MXP3 figured by Burger (1895: pi. 10 fig. (1895), however, reveal a number of dis­ 27) agrees well with those of the present crepancies. The relatively prominently pro­ specimens, although the dactylus of the duced triangular frontal margin of the fe­ VOLUME 116. NUMBER 4 907

palp he figured appears slightly less spatu- ing more ovate in the female (Fig. lA ver­ liform, the dactylus is subequal in length to sus Fig. IJ). The form and proportions of the propodus, and the outer margin of the the chela and WLl-4 are not sexually di­ fused ischium-merus is more rounded. The morphic. figure, however, is too schematic. The male abdomen figured by Burger (1895: pi. 9 fig. Holotheres flavus (Nauck, 1880) 28b) is also too schematic to be of much Figs. 2, 3 use, although its general shape and propor­ Pinnotheres flavus Nauck, 1880:23, 66.— tions agree with those of the present spec­ Burger, 1895:383, pi. 9 fig. 29, pi. 10 fig. imens. 29, 35.—Schmitt et al., 1973:45. (See In describing H. setnai from the Anda­ Schmitt et al. (1973:45) for additional man Islands (as Pinnotheres setnai), Cho­ references). pra (1931:313) commented that his species differed from H. semperi in the carapace Lectotype.—Male (8.6 by 7.9 mm) (SMF being more arched; the front is truncate 951a), Bohol, Ubay, Philippines, coll. C. with the anterior margin of the front de­ Semper, 1863-1864. pressed medially (not triangular); the large Paralectotypes.—2 males (7.9 by 7.6 chela is more slender, with the palm pro­ mm, 8.0 by 7.6 mm), 2 females (10.3 by portionately longer and the fingers having 10.0 mm, 8.9 by 8.0 mm) (SMF 951b), well developed teeth; WL4 is very much same data as lectotype. 2 ovigerous females smaller than the others; and the propodus (10.0 by 9.5 mm, 8.4 by 7.8 mm) (USNM of the palp of MXP3 is longer than the dac­ 32434), Zamboanga, Philippines, coll. C. tylus (not subequal in length). Of the above Semper, 1876. differences, as has been discussed earlier, Diagnosis.—Female carapace subcircu- only the proportions of the chela are really lar, width to length ratio ca. 1.1; female car­ valid. On the basis of Chopra's (1931) ex­ apace subcircular, width length ratio ca. 1.1; cellent description and figures, it is clear dorsal surface gently convex, covered with that H. setnai and H. semperi are very short setae which obscures outline. Antero­ close. Chopra, however, did not figure the lateral margin arcuate, posterolateral mar­ carapace of the male. On the basis of Cho­ gin gently convex; sub-branchial surface pra's (1931) figures, H. setnai can also be setose. MXP3 with ischium and merus separated from H. semperi by the relatively completely fused, without trace of suture, broader and more sinuous female frontal subovate, inner margin subangular subme- margin, the more slender ischio-merus of dially; exopod relatively stout, outer margin MXP3, and the carpus of the chelipeds not convex, with well developed flagellal setae; possessing a deep longitudinal cleft. Both palp 3-segmented, propodus elongate, sub- species should thus be regarded as separate conical, subequal in length to dactylus; dac­ for the time being. tylus spatuliform, articulates at base of pro­ The pair of specimens figured by Chuang podus. Chelipeds subequal; inner surfaces (1961: pi. 94: Figs. 2 and 3) is now in the of merus, carpus, palm and fingers covered ZRC (ZRC 1965.11.24.40-41), and the with setae; palm relatively stout; fingers male, 7.9 by 7.2 mm (ZRC 1965.11.24.40), shorter than palm; cutting edges of dactylus is here designated as the neotype of Pin­ and propodus each with sub-basal large notheres semperi Burger, 1895. The speci­ tooth and several denticles before it. WL mens are very consistent in all the charac­ relatively stout, WL2 > WL3 > WLl > ters discussed here. Other than the sexually WL4 in length (basis-ischium to tip of dac­ dimorphic features of the abdomen and re­ tylus); dactyli of WLl-4 relatively long, productive structures, the only major dif­ sharply tapering, all subequal in length; ference is in the shape of the carapace, be­ dorsal and ventral margins of meri, carpi 908 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ^r*r-%.

Fig. 2. Holotheres flavus (Nauck, 1880). Lectotype male (8.6 by 7.9 mm) (SMF 951a). A, overall view of specimen (setae on right half of body denuded); B, right chela, outer view; C-F, WLl-4 respectively; G, male abdomen; H, left Gl. (Except for A, H, all other structures denuded.) Scales: A = 5.0 mm, B-H = 1.0 mm. VOLUME 1 16. NUMBER 4 909

Fig. 3. Holotheres flavus (Nauck, 1880). Paralectotype female (10.0 by 9.5 mm) (USNM 32434). A, overall view of specimen; B, left MXP3 (denuded), outer view; C, right cheliped, outer view; D-G, WLl-4 respectively, denuded. Scales = 1.0 mm. and propodi of all legs lined with setae of Remarks.—Holotheres flavus was orig­ various lengths, obscures margins. Female inally described by Nauck (1880) (as Pin­ abdomen with all segments freely articulat­ notheres flavus) on the basis of an un­ ing. known number of specimens from an un­ Biology.—Burger (1895) noted that the specified site or sites in the Philippines, species was collected from an unidentified obtained from collections there by Semper holothurian. Nothing else is known. (Nauck, 1880:5), although he did not 910 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON explicity state so when describing the spe­ Diagnosis.—Obligate symbionts in ho- cies. Burger (1895:383) subsequently re­ lothurians. Carapace circular to subcircular, ported six males and six females collected broader than long. WL subequal right and by Semper from Zamboanga in the Phil­ left; WL4 dactylus of female much longer ippines. De Man (1887:720, 721), who re­ than short, hooked dactyli of WLl-3. Is­ appraised Nauck's taxa, provided mea­ chium and merus of MXP3 completely surements for only one male and one fe­ fused, without trace of suture; dactylus of male (without specific location), suggest­ palp subspatuliform, subequal in length to ing he only had two specimens. Schmitt propodus, inserted distal to base of propo- et al. (1973:45) commented that there dus (Vs to midway from base). Gl a simple were "2 syntypes: ZIMB 67/297 [Gottin- tube, tapering from base to tip, slightly re­ gen Museum], no longer extant; type lo­ curved distally. cality: 'Philippinen'". How Schmitt et al. Etymology.—The genus is named in hon­ (1973) knew there were only two syntypes or of Otto Burger, whose key paper in 1895, was not stated, perhaps these were the laid the foundation for much of what has specimens used by De Man (1887). As been done recently in pinnotherid taxono­ discussed earlier, two of the females from my. The genus name is derived from an ar­ Gottingen were actually given to the bitrary combination of Burger's name and USNM, and the SMF has taken over from Pinnotheres. The gender is masculine. Gottingen three male and two female Remarks.—The key diagnostic features specimens of P. flavus. There are no spec­ of Buergeres are the form of the MXP3 imens on this species in the Berlin Mu­ (dactylus subspatuliform, inserted well be­ seum (O. Coleman, in litt.). Since Sem- fore base of propodus), and relative length per's material was used by Nauck, Burger of the female WL4 (dactylus much longer and De Man, it is reasonable to believe than WLl-3). No other group of pinnoth- that all the material on hand are syntypes. erids associated with holothurians has these We hereby select a male (8.6 by 7.9 mm, characters. With regards to the form of the SMF 951a) as the lectotype of Pinnoth­ MXP3 and WL4, Buergeres is perhaps eres flavus Nauck, 1880. closest to the American Tumidotheres Cam­ Holotheres flavus is similar to H. sem- pos, 1989, but in members of the latter ge­ peri, but can be distinguished by its more nus, the carapace is less rounded with the quadrate male carapace with the lateral front more distinctly produced, and the pro­ margins straighter, appearing subparallel podus and dactylus of MXP3 are propor­ (Fig. 2A) (versus more rounded. Fig. lA), tionately shorter (see Campos 1989). Tum­ the dorsal surface of the carapace covered idotheres is also wholly symbiotic with var­ with pubescence only along the margins ious bivalves while Buergeres is an obligate (Fig. 2A) (versus uniformly all over, Fig. parasite of holothurians. The relatively sub­ lA), and most significantly, the Gl is spatuliform dactylus of MXP3 also allies it more strongly curved, appearing distinct­ with Atlantic and eastern Pacific taxa like ly C-shaped (Fig. 2H versus Fig. II). Afropinnotheres Manning, 1993, Nepin- notheres Manning, 1993, Alainotheres Manning, 1993, and Raymondia Campos, Buergeres, new genus 2002 (see Manning 1993a, Campos 2002) Type species.—Pinnotheres ortmanni but they are quite different in the form of Burger, 1895, by present designation. the carapace and legs. Some genera of pin- Included species.—Four: B. deccanensis notherids have adult females with the WL3 (Chopra, 1931), B. holothuriae (Semper, longest (e.g., Arcotheres Manning, 1993, 1880), B. ortmanni (Burger, 1895), and B. see Campos & Manning 2001); or WL2 tenuipes (Burger, 1895). longest (e.g., the Atlantic Zaops Rathbun, VOLUME 1 16. NUMBER 4 911

1900, and Waldotheres Manning, 1993; see MXP3 with ischium and merus completely Manning 1993a, 1993b; Campos and Man­ fused, without trace of suture, subovate, in­ ning 2000); but in the case of Buergeres, ner margin subangular at distal Vr, exopod WLl-3 are more or less subequal in length, relatively stout, outer margin gently con­ with the dactylus of WL4 longest and more vex, with well developed flagellal setae; prominently elongate than the rest. In any palp 3-segmented, propodus elongate, lon­ case, none of the species in these genera are ger than dactylus; dactylus spatuliform, ar­ associated with holothurians. ticulation point anterior to base of propo­ Of the four species presently referred to dus. Chelipeds subequal; inner surfaces of Buergeres, two, B. holothuriae (Semper, merus, carpus, palm and fingers densely 1880) and B. ortmcmni (Burger, 1895) are covered with relatively long setae; palm rel­ treated here in detail. One of these, B. dec- atively stout; fingers shorter than palm; cut­ canensis (Chopra, 1931) was well described ting edges of dactylus and propodus each and figured by Chopra (1931) and its affin­ with sub-basal large tooth and 1 or 2 den­ ities are not in doubt. Buergeres tenuipes ticles before it, rest of cutting edges (Burger, 1895) (type locality Ubay, Philip­ smooth, almost blade-like; tips hooked. WL pines) was described apparently from only relatively slender, WL2 > WLl > WL3 > one female specimen, but on the basis of WL4 in length (basis-ischium to tip of dac­ Burger's (1895:371, pi. 9 fig. 11, pi. 10 fig. tylus); dactyli of WLl-3 relatively short, 11) figures of the third maxillipeds and sharply tapering; dactylus of WL4 very walking legs, we have little doubt that it long, ca. 3 times as long as those of other belongs in this genus as well. The type of legs; dorsal margins of meri of all legs rel­ B. tenuipes is apparently lost (see Schmitt atively densely lined with setae of various et al. 1973:89) and we have not been able lengths, becoming sparse or absent along to locate it in any of the depositories we distal one-quarter; ventral margins of pro­ have checked. podus of all legs relatively densely lined with setae on distal half. Female abdomen Buergeres ortmanni (Burger, 1895) with all 6 segments distinct, freely articu­ Fig. 4 lating, telson semicircular. Male unknown. Biology.—This species had been collect­ Pinnotheres ortmanni Burger, 1895:384, pi. ed from Holothuria fursocinerea (see 9 fig. 30, pi. 10 fig. 28.—Schmitt et al., Lanchester 1900). The present specimens 1973:61. (See Schmitt et al. (1973:61) for were obtained from an unknown host. additional references). Remarks.—Burger (1895) described Neotype.—Female (11.4 by 9.5 mm) Buergeres ortmanni (as Pinnotheres ort­ (ZRC 2003.59), in holothurian in coral reef, manni) on the basis of one ovigerous fe­ exposed at edge of reef, Pulau Salu, north­ male originally obtained by Semper from eastern Singapore, coll. R. U. Gooding, 28 "Aibukit" in the Philippines. Burger's April 1967. specimen cannot be located in the SMF, Other material examined.—2 females USNM or Berlin Museum, and is presumed (both 12.5 by 11.8 mm) (ZRC 1965. lost, as Schmitt et al. (1973:61) had indi­ 11.24.14-15), coll. Singapore, no other cated. Lanchester (1900) subsequently re­ data. ported the species from Singapore. No other Diagnosis.—Female carapace almost cir­ specimens are known. The present speci­ cular, slightly wider than long, width to mens in the ZRC from Singapore agree length ratio 1.1-1.2; dorsal surface smooth, with the type description and figures well gently convex; intestinal region slightly de­ and we are confident they are conspecific. pressed. Lateral margins gently convex; To stabilize the taxonomy of this species sub-branchial surfaces densely setose. (especially since it is the proposed type spe- 912 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cies of Buergeres), we hereby select a neo- of all legs lined with setae. Female abdo­ type for Pinnotheres ortmanni Burger, men with all 6 segments distinct, freely ar­ 1895, a female specimen measuring 11.4 by ticulating, telson semicircular. Male un­ 9.5 mm from Singapore (ZRC 2003.59). known. Biology.—This species has been collect­ Buergeres holothuriae (Semper, 1880) ed from Stichopus variegatus (see Biirger Fig. 5 1895), although the record from Holothuria scabra by Semper may be incorrect (see Pinnotheres ortmanni Semper, 1880:105.— Schmitt et al. 1973:44). Burger, 1895:381, pi. 9 fig. 27, pi. 10 fig. Remarks.—Semper (1880) described 26.—Schmitt et al., 1973:49. (See Buergeres holothuriae (as Pinnotheres hol­ Schmitt et al. (1973:49) for additional othuriae) from an unspecified number of references). specimens from Zamboanga on Mindanao Type material.—Lectotype: ovigerous fe­ island in the Philippines. Burger (1895) male (10.1 by 9.3 mm) (USNM 032433a), who used Semper's specimens for his study, Zamboanga, Philippines, coll. Semper, reported that he had three males and eight 1876. Paralectotype: 1 ovigerous female females available, all of which are regarded (8.0 by 6.5 mm) (USNM 032433b), same as syntypes. Of these, the USNM has two data as lectotype. females. The SMF does not have any spec­ Diagnosis.—Female carapace almost cir­ imens. Schmitt et al. (1973:49) had incor­ cular, slightly wider than long, width length rectly stated that the types are no longer ratio 1.1—1.2; dorsal surface smooth, gently extant. The larger female specimen (10.1 by convex. Lateral margins gently convex; 9.3 mm, USNM 032433a) is here designat­ sub-branchial surfaces gently setose. MXP3 ed as the lectotype of Pinnotheres holothu­ with ischium and merus completely fused, riae Semper, 1880. without trace of suture, subcircular, inner margin rounded; exopod relatively stout, Notes on Allied Taxa Symbiotic outer margin gently convex, with well de­ in Holothurians veloped flagellal setae; palp 3-segmented, propodus elongate, longer than dactylus; The generic affinities of Ostracotheres dactylus spatuliform, articulation point holothuriensis (originally described as Pin­ slightly anterior to base of propodus. Che- notheres holothuriensis) should be re-ex­ lipeds subequal; inner surfaces of merus, amined in view of recent systematic devel­ carpus, palm and fingers covered with rel­ opments in the family. The species was de­ atively long setae; outer surface of chela scribed from South Australia and has a two- densely covered with short pubescence; segmented palp (Baker 1907, Pregenzer palm relatively stout; fingers shorter than 1988), but its carapace form is rather un­ palm, basal one-third covered with dense usual. Two other species. Pinnotheres pil- pubescence; cutting edges of dactylus and umnoides and Ostracotheres holothuriensis, propodus each with numerous denticles ex­ have been reported from hosts other than cept for almost unarmed distal Vs. WL rel­ holothurians (see Schmitt et al. 1973). The atively slender, WL2 > WLl > WL3 > other species mentioned, all placed in the WL4 in length (basis-ischium to tip of dac­ genus Pinnotheres thus far, are obligate par­ tylus); dactyli of WLl-3 relatively short, asites of holothurians and are not known distinctly hooked; dactylus of WL4 very from other hosts. long, ca. 5 times as long as those of other Pinnotheres pilumnoides Nobili, 1905, legs; dorsal and ventral margins of meri of known only from the Red Sea and Gulf of all legs lined with setae of various lengths; Aden, is similar to Holotheres, with short ventral margins of propodus and dactylus WL, the last apparently not elongate (al- VOLUME 1 16, NUMBER 4 913

Fig. 4. Buergeres ortmanni (Burger, 1895). A-G, neotype female (11.4 by 9.5 mm) (ZRC); H, female (12.5 by 11.8 mm) (ZRC 1965.11.24.14). A, overall view of specimen; B, H, left MXP3 (denuded), outer view; C, right chela (denuded), outer view; D-G, WLl-4 respectively (denuded). Scales: A = 5.0 mm, B-H = 1.0 mm. though this is not at all clear from Nobili's tinctly shorter compared to other members (1905, 1906) descriptions, with the dactylus of the group, and the inner margin of the of MXP3 subspatuliform and inserted near fused ischio-merus is rounded and not an­ the base of the propodus. The dactylus of gular (Nobili 1905:308, fig. 12). As such, MXP3 of P. pilumnoides, however, is much in some ways, P. pilumnoides resembles shorter than the propodus which is also dis­ members of the eastern Pacific Holothurio- 914 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Buergeres holothuriae (Semper, 1880). Lectotype female (10.1 by 9.3 mm) (USNM 032433a). A, overall view of specimen; B, left MXP3 (denuded), outer view; C, left MXP3 (denuded), inner view; C, left chela, outer view; D, WL3 (partially denuded); E, WL4 (partially denuded). Scales = 1.0 mm. philus (sensu Manning 1993b) although slope depths of 399-405 and 457-461 m; they are clearly not congeneric. We prefer the male abdomen comprises only six so­ to exclude P. pilumnoides from the present mites, with the second and third fused; and discussion until we have an opportunity to the apex of male first pleopod is complex examine specimens of the species. Certain­ rather than simple. ly, as things are, P. pilumnoides does not fit into any of the Indo-West Pacific pin- Notes on Pinnaxodes notherid genera now recognized. and Holothuriophilus Alain Manning, 1998, can be distin­ guished from both Holotheres and Buer­ The following material has been exam­ geres, as well as P. holothuriensis and P. ined for the following notes. Pinnaxodes pilumnoides, in the following characters: chilensis: 2 females (14.4 by 12.7 mm, 13.4 the spatuliform dactylus of MXP3 is in­ by 12.4 mm, carapace broken) (ZMUC), serted medially rather than proximally on parasite, in rectum of echinoid Loxechinus the propodus; the carapace is subcircular, albus (Molina), Chile. Pinnaxodes major: 1 ornamented with few setae dorsally; WL2 female (13.1 by 11.1 mm) (ZMUC), inland and 3 are the longest; the species lives at sea, from bivalve, coll. K. Hatonaka, 7 May VOLUME 1 16. NUMBER 4 915

Fig. 6. A-D, Pinnaxodes major Ortmann, 1894, female (15.4 by 13.0 mm) (ZRC 1999.39a); E-H, Pinnax­ odes mutuensis Sakai, 1939, female (14.2 by 12.6 mm) (NSMT). A, E, dorsal views of carapaces; B, F left MXP3 (outer view); C, G, WL3; D, H, WL4. (All structures denuded.) Scales = 1.0 mm.

1935; 2 females (14.7 by 12.5 mm, 15.4 by naxodes mutuensis: 1 female (14.2 by 12.6 13.0 mm) (ZRC 1999.39), 4 females mm) (NSMT), from Mytilus gallprovinci- (NSMT), from Atrina pectinata, Hokkaido, alis, northern Japan, coll. 19 May 1982. Japan, coll. Tsukiji Market, Tokyo, P. K. L. Some comments on the genus Pinnaxo­ Ng & M. Takeda, 3 February 1999. Pin­ des Heller, 1865 (type species Pinnaxodes 916 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hirtipes Heller, 1865, a subjective junior cating the two segments, most clearly seen synonym of Pinnaxodes chilensis (H. Milne under transmitted light. This is the same as Edwards, 1837)), are necessary as one in P. chilensis, although the suture between Indo-West Pacific species, P. major Ort- the ischium and merus in the present spec­ mann, 1894, has been reported from a ho- imen cannot be easily seen (Fig. 7A), being lothurian (Sakai 1939, 1976), although its very faint. However, the MXP3 of P. major preferred host seems to be bivalves. Man­ and P. mutuensis differ from that of P. chi­ ning (1993b:524, 525) separated Pinnaxo­ lensis in that the latter has the dactylus and des from Holothuriophilus on the basis of propodus proportionately shorter (Fig. 7A, the carapace shape (greatest width posterior B). Campos et al. (1998) had commented to the midlength, in Holothuriophilus it is that P. gigas and P. floridensis have the anterior to the midlength), relatively slender male telsons "basally expanded", forming and elongate dactyli of the WL (short and a more rounded or pentagonal structure stout in Holothuriophilus), and the presence (Campos et al. 1998: fig. 3A, C). This is of a suture between the ischium and merus also the case for P. major (see Takeda & of MXP3 (absent in Holothuriophilus). Prince Masahito 2000: fig. 4B), but not P. Campos et al. (1998:374) discussed the tax­ mutuensis (see Takeda & Prince Masahito onomy of the American P. gigas Green, 2000: Fig. 5L). In addition, while P. gigas, 1992, and P. floridensis Wells & Wells, P. floridensis and P. major have their fronts 1961, including illustrations, and suggested distinctly produced anteriorly (Fig. 6A; that Opisthopus Rathbun, 1893, be regarded Campos et al. 1998: fig. 2A, C; Takeda & as a separate genus (it had been a junior Prince Masahito 2000: fig. 4A), the front synonym of Pinnaxodes). barely extends beyond the orbits in P. mu­ Takeda & Prince Masahito (2000) dis­ tuensis (Fig. 6E; Takeda & Prince Masahito cussed the taxonomy of Pinnaxodes chilen­ 2000: Fig. 5A, I, J). These differences sug­ sis (an obligate symbiont of sea urchins), P. gest that P. mutuensis is not a true Pinnax­ major and P. mutuensis Sakai, 1939 odes. However, to refer P. mutuensis to (known only from bivalves), and comment­ Holothuriophilus as suggested by Takeda & ed that the latter should be transferred to Prince Masahito (2000) is also unsatisfac­ Holothuriophilus. They noted that for P. tory. Species of Holothuriophilus have mutuensis, the "fusion of the ischium-mer- quadrate carapaces which are clearly wider us of the third maxilliped was not men­ than long (rather than round) and the WL tioned in the original description; it is es­ are proportionally shorter (Fig. 7D-F; Man­ sentially complete, and a quite imperfect ning 1993b: figs. 2a, 3). It is also pertinent suture may be traceable only on the external to note that while members of Holothurio­ surface under special light control." (Tak­ philus are symbiotic in holothurians, P. mu­ eda & Prince Masahito 2000:110). This, tuensis is only known from bivalves. For however, is not obvious from their figure the moment, we retain Pinnotheres mutuen­ (Takeda & Prince Masahito 2000: fig. 2G) sis in Pinnaxodes for lack of a better clas­ that does not show any trace of a suture sification. It may need to be referred to its between the ischium and merus, with only own genus at a later date. a suture shown between the ischium and ba­ The MXP3 of P. chilensis (Fig. 7A, B) sis. We compared specimens of all three actually closely resembles that of Holothu­ species, and the MXP3 of P. major (Fig. riophilus (see Manning 1993b: fig. 2b, c; 6B) is essentially similar to that of P. mu­ present Fig. 7C), and interestingly, both tuensis (Fig. 6F), with a distinct suture pre­ taxa are from the eastern Pacific. With re­ sent between the ischium and basis, but the gards to the other supposed differences be­ ischium and merus are completely fused, tween Pinnaxodes and Holothuriophilus with only the faint trace of a suture demar­ mentioned by Manning (1993b), we find VOLUME 1 16, NUMBER 4 917

Fig. 7. A, B, Pinnaxodes chilensis (H. Milne Edwards, 1837), female (14.4 by 12.7 mm) (ZMUC), A, outer surface of right MXP3, setae removed, suture between ischium and merus very faint, not drawn; B, inner surface of right MXP3, setae removed, propodus and dactylus detached. C-F, Holothuriophilus trapeziformis Nauck, 1880, lectotype female (7.7 by 4.8 mm) (SMF); C, right MXP3; D-F WL2-4 respectively. (All structures denuded.) Scales = 1.0 mm. the difference in carapace shape not at all Sinaloa, in Mexico. Nauck (1880:24, 66) obvious, especially between P. major and did not state how many specimens he had H. trapeziformis. The proportions of the or even where it was from, commenting dactyli of the WL are also not that distinct, only that it was a "Parasit in Holothuria with those of Holothuriophilus only slightly maxima Semper" (Nauck 1880:66). Burger shorter than those of the three Pinnaxodes (1895:380) subsequently listed one female species examined here (Fig. 7D-F). (as Pinnotheres trapeziformis) measuring With regard to the specimen of Holothu­ 8.5 by 5.0 mm from Mazatlan, from Hol­ riophilus trapeziformis in the SMF, it car­ othuria maxima Semper. De Man (1887: ries the old catalogue number "ZMG 67/ 722) also measured one specimen from 565a" and was collected from Mazatlan, Nauck, noting it was a female 13.8 by 10.5 918 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

mm. The size discrepancy between Burger's eres niaculcitns Say, 1818 (Brachyura: Pinnoth- and De Man's specimens is difficult to rec­ eridae).—Journal of Crustacean Biology 9:672- oncile and suggests that Nauck may have 679. —. 2002. Two new genera of pinnotherid crabs had more than one specimen. As such, we from the tropical eastern Pacific (Decapoda: prefer to regard the present SMF specimen Brachyura: Pinnotheridae).—Journal of Crusta­ as a syntype and we here designate it as the cean Biology 22:328-336. lectotype of Holothuriophilus trapeziifor- -, & R. B. Manning. 2000. The identities of Pin­ mis Nauck, 1880. The present female spec­ notheres luiclus Holmes, 1895, and P. niidiis •sensu Weymouth. 1910 (Crustacea: Decapoda: imen in the SMF (Fig. 7C-F) measures ap­ Pinnotheridae).—Proceedings of the Biological proximately 7.7 by 4.8 mm, closer to Bur­ Society of Washington I 13:799-805. ger's data. This specimen is in poor condi­ -, & . 2000. Authorship and diagnosis of tion, with the carapace removed, very soft the genus Arcotheres Manning, 1993 (Crusta­ and somewhat deformed (making precise cea: Brachyura: Pinnotheridae).—Raffles Bul­ measurements difficult). letin of Zoology 49:167-170. -, V. Dfaz, & J. A. Gamboa-Contreras. 1998. Notes on distribution and taxonomy of five Acknowledgements poorly known species of pinnotherid crabs from the eastern Pacific (Crustacea: Brachyura: Pin­ The authors are most grateful to Michael notheridae).—Proceedings of the Biological So­ Tiirkay and Andreas Allspach (SMF) for ciety of Washington 1 1 1:372-381. their help with the Gottingen specimens, Chopra, B. 1931. Further notes on Crustacea Decapoda and Oliver Coleman (Berlin Museum) for in the Indian Museum. II. On some decapod Crustacea found in the cloaca of holothurians.— their help in locating specimens. We are Records of the Indian Museum 33(3):303-324. grateful to Ernesto Campos, and Alain Chuang, S. H. 1961. On Malayan Shores. Muwu Sho- Crosnier, for their very helpful comments sa, Singapore, 225 pp. that have strengthened the discussion. The Doflein, F. 1904. Brachyura. Wissenschaften Ergeben- study has been partially supported by a re­ isse Deutschen Teifsee 'Valdivia' Expedition 6: search grant to the first author from the Na­ i-xiv, 1-314, figs. 1-68, pis. 1-57. Garth, J. S. 1957. The Crustacea Decapoda of Chile. tional University of Singapore. The second Reports of the Lund University Chile Expedi­ author's studies of pinnotherid crabs were tion 1948-49, 29.—Lunds Universitets Arssk- supported by the Smithsonian Marine Sta­ rift, new series (2)53(7): 1-130. tion at Fort Pierce, Florida. This is contri­ Green, T. M. 1992. Pinnaxodes gigas, a new species bution no. 486 from that program. Several of pinnotherid crab from the Gulf of California of the figures of Holotheres flavus and (Decapoda: Brachyura: Pinnotheridae).—Pro­ ceedings of the Biological Society of Washing­ Buergeres holothuriae were prepared by ton 105:775-779. Lilly King Manning. Hamel, J.-E, R K. L. Ng, & A. Mercier. 1999. Life cycle of the pea crab Pinnotheres halingi sp. Literature Cited nov., an obligate symbiont of the sea cucumber Holothuria scahra Jaeger.—Ophelia 50(3): 149- Baker, W. H. 1907. Notes on South Australian Deca­ 175. poda Crustacea. Part V.—Transactions, Pro­ Heller, C. 1865. Crustaceen. In Reise der osterrei- ceedings and Report of the Royal Society of chischen Fregatte 'Novara' um die Erdre, in den South Australia 31:173-191, pis. 23-25. Jahren 1857, 1858, 1859, unter den Befehlen Bosc, L. A. G. 1802. Histoire naturelle des Crustaces, des Commodore B. von Wullerstorf-Urbair. contenant leur description et leurs moeurs, avec Zoologische Theil 2(3): 1-280, pis. 1-25. figures dessinees d'apres nature. Deterville, Par­ Lanchester, W. F. 1900. On a collection of Crustacea is, 1:1-258, pis. 1-8; 2:1-296, pis. 9-18. made at Singapore and Malacca.—Part I. Crus­ Burger, O. 1895. Ein Beitrag zur kenntniss der Pin- tacea Brachyura.—Proceedings of the Zoologi­ notherinen.—Zoologische Jahrbiicher, Abthei- cal Society of London 1900:719-770, pis. 44- lung fiir Systematik, Geographic und Biologic 47. der Thiere 8:361-390, pis. 9. 10. Man, J. G. De. 1887. Uebersicht der Indo-pacifischen Campos, E. 1989. Tumidotheres, a new genus for Pin­ Arten der Gattung Sesarma Say nebst einer Kri- notheres margarita Smith, 1869, and Pinnoth­ tik der von W Hess und E. Nauck in den Jahren VOLUME 1 16. NUMBER 4 919

1865 und 1880 beschriebenen Decapoden.— nota descripsit.—Archiv fiir Naturgeschichte Zoologische Jahrbiicher, Abtheilimg fiir Syste- 2(1): 134-144, pi. 4. matik. Geographic und Biologic der Thiere 2: Pregenzer, C. 1988. A redescription of Pinnotheres 639-689, pi. 1. hoh)thiiriensis Baker, 1907 and Pinnotheres Manning, R. B. 1993a. West African pinnotherid crabs subglohosus Baker, 1907 with a reassignment to subfamily Pinnotherinae (Crustacea, Decapoda, the genus Ostracotheres (Decapoda, Brachyu­ Brachyura).—Bulletin du Museum national ra).—Crustaceana 55:17-28. d'Histoire naturelle, Paris (4)!5A(]-4):125- Rathbun, M. J. 1893. Scientific results of explorations 177. by the U.S. Commission Steamer 'Albatross'. . 1993b. Three genera removed from the syn­ XXIV. Description of new genera and species onymy of Pinnotheres Bosc, 1802 (Brachyura: of crabs from the west coast of North America and the Sandwich Islands.—Proceedings of the Pinnotheridae).—Proceedings of the Biological United States National Museum 16:223-260. Society of Washington 106:523-531. . 1900. The catametopous or grapsoid crabs of . 1998. A new genus and species of pinnotherid North America.—American Naturalist 34:583- crab (Crustacea, Decapoda, Brachyura) from In­ 592. donesia.—Zoosystema, Paris 20(2):357-362. . 1918. The grapsoid crabs of America.—Bul­ Milne Edwards, H. 1837. Histoire naturelle des Crus- letin of the United States National Museum 97: taces, comprenant I'anatomic, la physiologic et 1-461. la classification de ces animaux. Atlas [1834, Sakai, T 1939. Studies on the crabs of Japan. IV. 1837, 1840]: 1-32, plates 1-14, 14 bis, 15-25, Brachygnatha, Brachyrhyncha. Tokyo, pp. 365- 25 bis, 26-42. Librairie Encyclopedique de 741, pis. 42-111. Roret, Paris. . 1976. Crabs of Japan and the adjacent seas. Nauck, E. 1880. Das Kaugerust der Brachyuren (mit In three volumes; English text, pp. beschreibung neur gattungen und arten, Z. T. xxix+773pp., Japanese text, pp. 1-461, Plates von C. Semper).—Zeitscrift wissenschaften volume, pp. 1-16, pis. 1-251. Kodansha Ltd., Zoologische Leipzig 34:1-69, pi. 1. Tokyo. Nobili, G. 1901. Decapodi raccoiti dal Dr. Filippo Sil- Schmitt, W. L., J. C. McCain, & E. Davidson. 1973. vestri nell'America meridonale.—Bolletino dei Decapoda I. Brachyura I. Fam. Pinnotheridae. Museo di Zoologia ed Anatomia comparata del- Crustaceorum Catalogus, Dr. W. Junk, Den la R. Universita di Torino I6(402):l-16. Haag 3:1-160. . 1905. Decapodes nouveaux des cotes Semper, K. 1880. Natiirlichen existenzbedingungen de d'Arabic et du Golfe Persique. (Diagnoses pre- Thiere. Erster theil. F. A. Brockhaus, Leipzig. liminaires).—Bulletin du Museum national Pp. 98-100. d'Histoire naturelle, Paris 11(3):158-164. Takeda, M., & Prince Masahito. 2000. Systematic . 1906. Faune carcinologique de la Mer Rouge. notes on the pinnotherid crabs of the genus Pin- Decapodes et stomatopodes.—Annales de Sci­ naxodes (Crustacea: Decapoda: Brachyura).— Bulletin of the National Science Museum, Se­ ences Naturelles Zoologie, Paris (9)4(1-3): 1- ries A (Zoology) 20:99-112. 347, pis. 1-11. Tesch, J. J. 1918. Decapoda Brachyura II. Goneplaci- Ortmann, A. 1894. Die Decapoden—Krebse des dae and Pinnotheridae.—Siboga Expeditie Strassburger Museums, mit besonderer Beriick- Monographite 39cl: 149-295, pis. 7-18. sichtigung der van Herrn Dr. Doderlein bei Ja­ Van den Spiegel, D., & M. Jangoux. 1989. Sur la sym- pan und bei den Liu-Kiu-Insein gesammelten biose entre le Pinnotheride Pinnotheres villos- und zur Zeit im Strassburger Museum aufbe- issimus (Crustacea, Decapoda) et I'Holothurie wahrten Formen. VIIL Theil. Abtheilung: Actinopyga mauritiana (Echinodermata).—Vie Brachyura (Brachyura genuine Boas) IIL Un- Marine, Marseille 10:205-213. terabtheilung: Cancroidea, 2. .section: Cancri- Wells, H. W, & M. J. Wells. 1961. Observations on nea, 2. Gruppe: Catametopa.—Zoologische Pinnaxodes floridensis, a new species of pin­ Jahrbiicher, Abtheilung fiir Systematik, Geogra­ notherid crustacean in holothurians.—Bulletin phic und Biologic der Thiere 7:683-772, pi. 23. of Marine Science of the Gulf and Caribbean Poeppig, E. 1836. Crustacea chilensia nova aut minus 11:267-279.