Two new species and a new record of Perisesarma (Decapoda, Brachyura, Grapsidae, Sesarminae) from Indonesia

Special Issue for Prof Jacques Forest

TWO NEW SPECIES AND A NEW RECORD OF PERISESARMA (DECAPODA, BRACHYURA, GRAPSIDAE, SESARMINAE) FROM INDONESIA BY DWI LISTYO RAHAYU 1·3) and PETER J. F. DAVIE2 ) 1) Research and Development Center for Oceanology, Indonesian Institute of Sciences (LIPI), 2)

JI. Pasir Putih 1, Ancol Timur, Jakarta 14430, Indonesia Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101, Australia

ABSTRACT Two new species and a new distributional record of the sesarmine crabs of the genus Perisesarma are recorded from a mangrove area in southwestern Irian Jaya, Indonesia. Perisesarmaforesti sp. nov. and P. cricotus sp. nov. can be most easily separated from their congeners by the number, shape, and ornamentation of the cheliped dactylar tubercles. Perisesarma cricotus is also recorded from Ambon, eastern Indonesia. Perisesarma semperi (Btirger,. 1894) is a first record for Indonesia and marks a significant southerly range extension.

Deux especes nouvelles sont decrites et une nouvelle reference geographique des crabes Sesarminae Perisesarma est fournie a partir de specimens recoltes sur Ia cote sud-occidentale de I'Irian Jaya, Indonesie. Perisesarma foresti sp. nov. et P. cricotus sp. nov. peuvent etre differenciees de leurs congeneres par le nombre, Ia forme et l'ornementation des tubercules du dactyle du chelipede. Perisesarma cricotus est aussi signa1e d' Ambon, Indonesie orientale. Perisesarma semperi (Btirger, 1894) est trouve pour 1a premiere fois en lndonesie, indiquant une extension significative de sa presence vers 1e sud.

INTRODUCTION

De Man (1895) described the subgenus Perisesarma for two new species, Sesarma (Perisesarma) eumolpe De Man, 1895 and Sesarma (Perisesarma) onychophorum De Man, 1895, and additionally included Sesarma dussumieri A. MilneEdwards, 1853, in the new subgenus. De Man (1888: 175) had earlier included 3) Former address: Timika Environmental Laboratory, PT Freeport Indonesia, P.O. Box 109, Timika 99910, Irian Jaya, Indonesia. © Koninklijke Brill NV, Leiden, 2002 Also available online: www.brill.nl

Crustaceana 75 (3-4): 597-607

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S. bidens De Haan, 1833, S. dussumieri A. Milne-Edwards, 1853, S. guttata A. Milne-Edwards, 1869, S. haswelli De Man, 1887, and S. livida A. MilneEdwards, 1869, in his description of Sesarma "section C", and while failing to directly attribute them to Perisesarma in his later paper, nevertheless the close relationship of all these species was clearly established. Campbell (1967) included 13 species in his key to this group (then referred to as Sesarma (Chiromanthes)). He (Campbell, 1967) included all species earlier mentioned by De Man (1888, 1895), as well as Sesarma semperi Biirger, 1894, Sesarma indiarum Tweedie, 1940, and four new taxa: Sesarma (Chiromanthes) brevicristatum Campbell, 1967, Sesarma (C.) darwinensis Campbell, 1967, Sesarma (C.) semperi longicristatum Campbell, 1967, and Sesarma (C.) messa Campbell, 1967. Chiromanthes Gistel, 1848, was proposed as a replacement name for Pachysoma DeHaan, 1833, and was long considered the appropriate name for the genus here called Perisesarma. Holthuis (1977: 170) pointed out that this was the result of the designation of Grapsus (Pachysoma) bidens DeHaan, 1833, as the type species of Pachysoma by Rathbun (1918: 284), as this species was not originally included in Pachysoma by DeHaan (1833). Chiromanthes is the correct available name for the group previously referred to as Holometopus H. Milne-Edwards, 1853. The genus Perisesarma in Indonesian waters is represented by P. onychophorum De Man, 1895 and P. indiarum (Tweedie, 1940). The present paper further records P. semperi (Btirger, 1894) and describes two new species, P.foresti and P. cricotus from Irian Jaya and Ambon. The specimens for this study were collected during the Ecological Risk Assessment Program of the Environmental Department of PT Freeport, Indonesia, in the mangrove area of Mirnika (04°40'-05°05'S 136°35'-137°20'E), Irian Jaya, Indonesia. Specimens are deposited in the Environmental Department of PT Freeport Indonesia, Irian Jaya (PTFI); Zoological Museum Bogor (MZB) of the Indonesian Institute of Sciences, Zoological Reference Collection (ZRC) of the Raffles Museum, National University of Singapore, and Queensland Museum (QM), Brisbane, Australia. Measurements (in mm) given in the text are of the carapace breadth (measured at the widest point) followed by length, for the biggest and smallest specimen. G 1 = male first gonopod. Perisesarma De Man, 1895 Perisesarma De Man, 1895: 208. Sesarma (Perisesarma): De Man, 1902: 541.- Holthuis, 1977: 170-171. Sesarma (Chiromanthes): Rathbun, 1909: 22; 1910: 309; 1918: 284.- Tesch, 1917: 235, 255.Campbell, 1967: 2.

Type species. - Sesarma dussumieri A. Milne-Edwards, 1853, designated by Campbell (1967: 2); not S. bidens (De Haan, 1833) as designated by Rathbun

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( 1918: 284), as that species was not originally included in Perisesarma by De Man. Not Sesarma (Perisesarma) eumolpe De Man, 1895, as designated by Holthuis (1977: 170) as Campbell's (1967) action has priority. Gender neuter. Perisesarma foresti sp. nov. (figs. 1, 3B, E) Material: - Holotype male (20.7 x 17.5 mm), Kamora, Mimika, Irian Jaya, Indonesia, coll. J. Volosin 3/iv/2000 (MZB). Paratypes: 4 males (23.5 x 20.5-10.0 x 8.3 mm), 3 females (23.5 x 19.022.5 x 18.0 mm), Kamora, Mimika, Irian Jaya, Indonesia, coll. G. Setyadi 4/vii/1999 (MZB); 36 males (24.5 x 20.0-7.3 x 5.9 mm), 29 females (19.0 x 15.6-5.7 x 4.6 mm), 9 ovig. females (18.8 x 15.3-13.1 x 10.4 mm), same locality as preceding, coll. J. Volosin 3/iv/2000 (ZRC, QM). Other material: 6 males (22.0 x 17.5-13.0 x 10.4 mm), Cargo Dock Portsite, Mimika, Irian Jaya, Indonesia, coll. G. Setyadi 29/vii/1999 (PTFI); 4 males (20.0 x 16.5-10.4 x 8.2 mm), 2 females (14.1 x 11.4 and 8.1 x 6.6 mm), same locality as preceding, coll. G. Setyadi 8/xii/1999; 5 males (23.0 x 19.5-17.6 x 14.4 mm), 2 ovig. females (20.5 x 16.5 and 19.5 x 15.5 mm), Minajerwi, Mimika, Irian Jaya, Indonesia, coli. G. Setyadi 5/vii/1999; 17 males (19.4 x 16.3-8.9 x 7.3 mm), 1 female (8.7 x7.1 mm), Ajkwa, Mimika, Irian Jaya, Indonesia, coll. G. Setyadi 18/viii/1999; 9 males (19.1 x 15.8-10.3 x 8.5 mm), 13 females (16.9 x 15.6-6.3 x 5.3 mm), 2 ovig. females (16.0 x 13.1 and 14.2 x 11.5 mm), same locality as preceding, coli. J. Volosin 2/iv/2000. Comparative material. - Perisesarma brevicristatum (Campbell, 1967), 3 males (10.9 x 9.0, 16.6 x 13.4, 20.4 x 17.0 mm), Murray River, Queensland, Australia, coll. P. Davie, 15/v/78, determined by P. Davie (QM W 8269). Perisesarma messa (Campbell, 1967), 1 male, (23.4 x 19.0 mm), Starcke River, N. Queensland, Australia, 14°471611 S 145°001711 E, coll. P. Davie and J. Short, 12/xi/1992, determined by P. Davie (QM W 18203). Perisesarma onychophora (De Man, 1895), 2 males, (20.4 x. 16.1-16.4 x 13.1 mm), Sungei Sendok, Singapore, coll. C. L. Soh (C.10), 9/i/1966, determined by R. Serene (ZRC 1972.3.6.6).

Etymology. - This species is dedicated to Professor Jacques Forest to honour his enormous contribution to crustacean systematics. Description. - Carapace rectangular; greatest width between exorbital angles c. 1.22 (1.15-1.26) times broader than long. Carapace surface smooth, shining, punctate; with sparse transverse rows or tufts of minute setae. Front c. 0.54 times carapace width, moderately defiexed, with broad median emargination. Post-frontal lobes prominent, median lobes broader than laterals, separated by furrow. Dorsal carapace regions moderately well indicated, with gastric region most strongly demarcated; lateral branchial ridges prominent. Upper orbital border smooth, lower orbital border finely granulate. Anterolateral margin with sharp exorbital angle and single prominent epibranchial tooth; lateral margin slightly concave, edged with row of short setae. Chelipeds equal, large, robust. Merus with posterior border minutely granulate; distinct subdistal spine; lower border granulate; anterior border granulate with distinct large subdistal spine. Carpus with inner angle not produced, inner margin granular; inner face with longitudinal row .of minute granules; outer surface finely

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e -b,c

--a ----d,e Fig. 1. Perisesarma foresti sp. nov., male, holotype (20.7 mm c.b.) (MZB). a, carapace; b, c, right cheliped; d, male abdomen; e, male first pleopod. Scales: 1 mm.

granular. Upper surface of palm with 2 tranverse pectinated crests. First crest composed of 11-17 tall, broad teeth (14 and 17 on holotype). Second crest well developed, shorter than primary, with 5-9 small teeth (5 and 6 on holotype). Primary and secondary pectinated crests followed by several blunt tubercles; row of granules proximal to second crest. Upper margin of palm with strong blunt tubercles. Outer surface of palm granular, with indistinct median longitudinal ridge, naked except for tuft of short setae in front of first pectinated crest; inner surface of palm with smaller granules. Distal two-thirds of surface of fixed finger almost smooth or very scarcely granular, cutting edge of both fingers with

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smaller and larger, triangular teeth; length of cutting edge 0.45 times length of propodus; ventral border of chela concave at base of fixed finger, minutely granular. Dorsal surface of dactyl bearing 11-12 low tubercles; first 3 tubercles small and oval, followed by more rounded tubercles, larger and well separated. First to ninth tubercles distinct, becoming progressively smaller and almost indiscernible towards tip; each tubercle finely striated longitudinally with shallow, indistinct median groove. Row of sharp, distinct tubercles on proximal two-thirds of inner edge of dorsal surface. Fingers with tips chitinous, intermeshing; adult males with wide gape left when fingers closed. Second pair of walking legs longest, c. 1. 7 times carapace width. Merus of third leg twice as long as wide; carpus twice as long as wide, propodus c. 2.75 times as long as wide, dactyl c. 0.9 times length of propodus. Male abdomen with telson as long as broad at base, evenly rounded, as long as preceding somite; penultimate somite twice as long as wide. Second somite with median length as long as or very slightly longer than lateral edge. G 1 relatively slender, moderately curved; apical process bent, strongly produced, corneous. Setae long, simple, originating at base of apical process and on palp. Female with chelipeds smaller; dactyl slightly curved. Row of sharp, distinct tubercles on proximal two-thirds of inner edge of dorsal surface of dactyl more prominent than dactylar tubercles. Colour. -Carapace black or dark brown with yellowish-green marking. Ocular peduncles orange with brown spot, corneas black. Front and epistome yellowishgreen. Pterygostomial region black or dark purple, gradually lighter purple on branchiostegal region; suborbital region light blue. Ischium of maxilliped light blue; merus dark brown to dark purple. Merus of cheliped brown to dark red, carpus bright to dark red; palm bright red with dark red on upper margin and _lighter red on lower margin; dactyl bright red. First, second, and third somites of abdomen purple; fourth, fifth, and sixth somites brownish-white; telson white; thoracic sternites brown to dark brown. Ambulatory legs with merus mottled blue and purple; carpus, propodus, and dactylus purple; inner surface of merus blue in small individuals. Remarks. ~ Perisesarma foresti sp. nov. is similar to P. brevicristatum (Campred and purbell, 1967) in having 11. dactylar tubercles, and in colouration (dark , pie). But P. brevicristatum is easily distinguished by having dactylar tubercles that are very prominent, subcircular, and dome-shaped, and each having a smooth raised stripe at right-angles to the axis of the dactyl. P. foresti sp. nov. resembles P. messa (Campbell, 1967) in having low dactylar tubercles becoming barely discernable distally, but P. messa has 14-16 dactylar tubercles, while P. foresti has only 11-12 dactylar tubercles that are more prominent. Finally, P. foresti is superficially similar toP. onychophorum (De Man, 1895), in haying the dactylus of

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the adult male cheliped strongly curved, leaving a wide gape between the fingers when closed. P. onychophorum differs from our new species in possessing only 7-9 dactylar tubercles with the most distal being greatly elongated and occupying almost one-quarter the length of the dorsal surface of the dactyL Habitat. - The specimens of the new species were collected on mud or sandy mud substrates in the estuary or in dense mangrove forest. When disturbed, the crab took shelter under tree trunks, in shallow burrows, or burrowed into the muddy bottom of shallow pools. Perisesarma cricotus sp. nov. (figs. 2, 3A, C, D) Material.- Holotype male (16.0 x 13.4 mm), Ajkwa, Mimika, Irian Jay a, Indonesia, 31/iii/2000 (MZB). Paratypes: 23 males (19.3 x 15.5-8.1 x 6.8 mm), B females (18.3 x 14.5-8.2 x 6.8 mm) (MZB); 2 males (15.9 x 13.0 and 15.7 x 12.7 mm), 2 females (16.2 x 13.0 and 13.3 x 10.7 mm) (QM), Ajkwa, Mimika, Irian Jaya, Indonesia, 18/viii/1999; 1 male (15.5 x 12.6 mm), 1 female (13.2 x 11.0 mm) (QM), Waleila River mouth, Ambon, low tide, coli. R. Serene 22/i:x/1970; 2 males (15.0 x 12.1 and 13.7 x 11.2 mm), Kamora, Mimika, Irian Jaya, Indonesia, 3/iv/2000 (ZRC). Other material: 2 males (17.1 x 13.8 and 16.9 x 13.5 mm), Ajkwa, Mimika, lrianJaya, Indonesia, coli. G. Setyadi 11/i/2000 (PTFI); 1 male (12.0 x 9.6 mm), 1 ovig. female (15.0 x 11.5 mm), Sungai Tipoeka, Mimika, Irian Jaya, Indonesia, coli. D. L. Rabayu 10/:x/1999 (PTFI); 8 males (18 .6 x 14.512.6 x 9.6 mm), 7 females, (18.0 x 14.5-7.7 x 6.3 mm), 3 ovig. females (18.3 x 15.2 mm), Kamora, Mimika, Irian Jaya, Indonesia, coiL J. Volosin 3/iv/2000 (PTFI). Comparative material. - Perisesanna indiarum (Tweedie, 1940) 1 male (24.5 x 20.2 mm), Labuan, Malaysia, coli. Tweedie, 1950, determination by Tweedie (ZRC 1964 928/25).

Etymology. - The specific name cricotus is latin meaning to furnish with rings, and refers to the characteristically ringed dorsal dactylar tubercles of the male chela. It is used as a noun in apposition. Description. - Carapace rectangular, greatest width between orbital angles c. 1.22 (1.19-1.26) times broader than long. Carapace surface smooth, shining, punctate; with sparse transverse rows or tufts of minute setae. Front c. 0.56 times carapace width, sinuous medially. Median post-frontal lobes prominent, separated by deep median furrow, broader than laterals; lateral lobes less prominent, separated from medians by shallow furrow. Dorsal carapace regions moderately well indicated, with gastric region most strongly demarcated; lateral branchial ridges prominent. Anterolateral margin with large triangular, sharp exorbital angle, epibranchial tooth smaller; lateral margin slightly concave, lined with row of short setae. Chelipeds equal, large, robust. Merus with posterior border granulate; distinct subdistal spine; lower border granulate; anterior border tuberculate with strong, broad, subdistal spine; outer surface minutely tuberculate. Carpus with inner angle not produced; inner margin tuberculate, inner surface with longitudinal row of

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-a,b,c _ _ ___,e Fig. 2. Perisesarma cricotus sp. nov., male, holotype (16.0 mm c. b.) (MZB). a, carapace; b, c, right cheliped; d, male abdomen; e, male first pleopod. Scales: 1 mm.

minute tubercles; outer margin and outer surface minutely tuberculate. Upper surface of palm with 2 pectinated crests. First pectinated crest composed of 1422 tall teeth (holotype with 16 and 17 teeth). Second crest well developed, almost as long as first pectinated crest with 9-17 short teeth (holotype with 9 and 12 teeth); .row of blunt tubercles proximal to second crest. Upper margin of palm with strong blunt tubercles. Outer surface of palm tuberculate, with indistinct median longitudinal ridge; naked except for tuft of short setae in front of first pectinated crest; inner surface of palm with scattered smaller tubercles. Distal two-thirds of surface of fixed finger almost smooth or very scarcely granular with slight longitudinal ridge ventrally; cutting edge of both fingers with smaller large

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Fig. 3. A, C, D, Perisesarma cricotus sp. nov., male paratype (15.7 mm c. b.) (QM); A, dorsal view; C, outer face of chela; D, distal most dorsal dactylar tubercles; B, E, P. foresti sp. nov.; B, dorsal view of male paratype (26.1 mm c. b.) (QM); E, outer face of chela of male paratype (20.3 mm c. b.) (QM).

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triangular teeth; length of cutting edge 0.4 times length of propodus; ventral border of chelae concave at base of fixed finger, coarsely granular. Dactyl with dorsal surface bearing 11-12 tubercles, first tubercle small, third and fourth tubercles largest, decreasing in size distally; all tubercles prominent, subcircular, ringed with fine circular lines; row of sharp tubercles on proximal two-thirds of inner dorsal margin. Fingers with tips chitinous, intermeshing; narrow gape left when fingers closed. I Second and third pairs of walking legs longer than first and fourth pairs, c. 1.4 times carapace width. Merus of third leg twice as long as wide; carpus twice as long as wide, propodus c. 2.9 times longer than wide, dactyl c. 0.75 times length of propodus. Male abdomen with telson as long as broad at base, evenly rounded, as long as preceding somite; penultimate somite twice as long as wide. G 1 relatively slender, moderately curved; apical process corneous, strongly produced, bent. Setae long, simple, originating at base of apical process and on palp. In female, primary pectinated crest on palm lower and less distinct, second pectinated crest may be reduced to row of tubercles; dactylar tubercles also smaller but always with fine circular rings. Colour (recently preserved). - Carapace greenish brown with dark brown marking; cheliped orange; walking legs with merus light brown with dark brown marking; carpus, propodus, and dactylus light brown with purple median band. Remarks. - Perisesanna cricotus bears a very close resemblance to P brevicristatum (Campbell, 1967). However, in P brevicristatum there are 10-11 dactylar tubercles versus 11-12 in P cricotus. Also the dactylar tubercles are higher and more prominent, more widely spaced, and each is topped with a smooth, sharp, transverse keel that is lacking in P cricotus. There is perhaps an even greater similarity to Perisesanna indiarum Tweedie, 1950, but in that species the dactylar tubercles are larger and lower, especially over the proximal third; also each tubercle is crowned with an obvious narrow transverse chiton-like stripe, and this and the rest of the tubercle are marked by short longitudinal fine lines, not rings as in P. cricotus. Habitat. - The specimens were found amongst mangroves, in the same area as P foresti sp. nov. Distribution.- Mimika region, Irian Jay a, and Ambon, eastern Indonesia. Perisesarma semperi (Bi.irger, 1894) Sesarma semperi Biirger, 1894: 630, pl. 21 fig. I . - Tweedie, 1950: 342, fig. le. Sesarma (Perisesarma) semperi: De Man, 1902: 542. Sesarma (Chiromanthes) semperi: Tesch, 1917: 198. Sesarma (Chiromanthes) semperi semperi: Campbell, 1967: 4 (in key).

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Material.- 2 males (11.2 x 8.5 and 8.9 x 7.1 mm), Portsite, Mimika, Irian Jaya, Indonesia, coli. G. Setyadi 27/vii/1999 (MZB); 2 males (12.8 x 10.3 and 12.2 x 9.8 mm), same locality as preceding, coli. D. L. Rahayu 20/xii/1999 (ZRC); 25 males (15.3 x 11.5-5.1 x 4.3 mm), 17 females (15.8 x 13.06.2 x 4.9 mm), Ajkwa, Mimika, Irian Jaya, Indonesia, coli. J. Volosin, 21/vi/2000 (PTFI & MZB); 2 males (11.5 x 9.7 and 11.2 x 9.5 mm), 1 female (12.0 x 10.2 mm), same data as preceding (QM).

1

Colour. - Carapace mottled brown and yellowish green or light blue. Pterygostomial region dark brown with blue spot; ischium of maxilliped brown with mesial part blue; merus blue. Cheliped with merus and carpus dark brown, upper margin of palm brown; fixed finger and upper surface of propodus red; dactyl yellowish orange. Remarks. - The species is readily distinguished from its congeners by having only seven dactylar tubercles which are circular, prominent, and well spaced. The specimens were collected from mud substrate in mangroves. Distribution. -Philippines, Labuan, Malaysia and herein recorded from southwestern Irian J aya, Indonesia.

ACKNOWLEDGMENTS

The first author is grateful to Drs Peter K. L. Ng and Christoph Schubart for their kind help in the preliminary identification of the new species. We are grateful to the Zoological Reference Collection (ZRC), National University of Singapore, for allowing us to examine specimens of Perisesarma indiarum in their care. We are also pleased to thank Mr. G. Setyadi and Joe Volosin for collecting the specimens.

REFERENCES BURGER, 0., 1894. Beitdige zur Kenntniss der Gattung Sesarma. Zool. Jahrb., (System.) 7: 613-631. CAMPBELL, B. M., 1967. The Australian Sesarminae (Crustacea: Brachyura). Five species of Sesarma (Chiromantes). Mem. Queensland Mus., 15 (1): 1-19. CROSNIER, A., 1965. Crustaces Decapodes Grapsidae et Ocypodidae. Faune de Madagascar, 18: 1-143. HAAN, W. DE, 1833. Crustacea. In: P. F. VON SIEBOLD, Fauna Japonica sive descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava imperium tenent, suscepto, annis 1823-1830 collegit, notis, observationibus et adumbrationibus illustravit: i-xvii, i-xxxi, ix-xvi, 1-243, pis. A-J. [Published from 1833-1850]. (LugduniBatavorum [= Leiden]). HOLTHUIS, L. B., 1977. The Grapsidae, Gecarcinidae and Palicidae (Crustacea: Decapoda: Brachyura) of the Red Sea. Israel Joum. Zool., 26: 141-192. MAN, J. G. DE, 1888. Report on the podophthalmous Crustacea of the Mergui Archipelago, collected for the trustees of the Indian Museum, Calcutta. Journ. Linn. Society, London, 22: 1-312. - - , 1895-1898. Bericht tiber die von Herm Schiffscapitan Storm zu Atjeh, an den westlichen Ktisten von Malakka, Borneo und Celebes sowie in der Java-See gesamrnelten Decapoden und Stomatopoden. Zool. Jahrb., (System.) 9: 75-218; 10, pis. 9-15.

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- - , 1902. Die von Herro Professor Kiikenthal im Indischen Archipel gesammelten Dekapoden und Stomatopoden. In: W. KDKENTHAL, Ergebnisse einer zoologischen Forschungsreise in den Molukken und Borneo. Abh. Senckenberg naturforsch. Ges., 25: 467-929, pis. 19-27. RATHBUN, M. J., 1909. New crabs from the Gulf of Siam. Proc. bioi. Soc. Washington, 22: 107-114. - - , 1910. The Danish Expedition to Siam 1899-1900. V. Brachyura. K. dansk. Vidensk. Selsk. Skr., 7 (4): 301-368 (1-68), text-figs. 1-44, pis. 1-2, 1 map. - - , 1918. The grapsoid crabs of America. Bull. U.S. natl. Mus., 97: i-xxii, 1-461, text-figs. 1-172, pis. 1-161. TESCH, J. 1., 1917. Synopsis of the genera Sesarma, Metasesarma, Sarmatium and Clistocoeloma with a key to determination of the Indo-Pacific species. Zoo!. Meded., Leiden, 3 (2-3): 127-260. TwEEDIE, M . W. F., 1940. New and interesting Malaysian species of Sesarma and Utica (Crustacea Brachyura). Bull. Raffles Mus., 16: 88-113. - - , 1950. Grapsoid crabs from Labuan and Sarawak. Sarawak Mus. Joum., 5 (2): 338-369.

First received 11 May 2001. Final version accepted 25 January 2002.