Conchoeodromia alcocki Chopra, 1934: Megalopa of Conchoecetes artificiosus (Fabricius, 1798) (Decapoda, Brachyura, Dromiidae) Author(s): Danièle Guinot and Marcos Tavares Reviewed work(s): Source: Journal of Crustacean Biology, Vol. 20, No. 2, Special Number Dedicated to Dr. Raymond B. Manning (Jun., 2000), pp. 301-309 Published by: The Crustacean Society Stable URL: http://www.jstor.org/stable/1549508 . Accessed: 08/05/2012 11:17 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. The Crustacean Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Crustacean Biology. http://www.jstor.org JOURNALOF CRUSTACEANBIOLOGY,20, SPECIALNUMBER2: 301-309, 2000 CONCHOEODROMIAALCOCKICHOPRA, 1934: MEGALOPA OF CONCHOECETESARTIFICIOSUS(FABRICIUS, 1798) (DECAPODA, BRACHYURA, DROMIIDAE) Daniele Guinotand Marcos Tavares (DG) Museum nationald'Histoirenaturelle,Laboratoirede Zoologie (Arthropodes),61 rue Buffon, 75005 Paris,France(e-mail: guinot@mnhn.fr);(MT) UniversidadeSantaUrsula, Institutode Ciencias Biol6gicas e Ambientais,Rio de Janeiro22231-040, Brazil (e-mail: mtavares@ax.apc.org) ABSTRACT alcockiChopra,1934, knownfrom a single localityin the Bay of Bengaland Conchoeodromia fromonly two small specimens(6.2 x 5.2 and5.6 x 4.7 mm), is shownto be a megalopaof Conchoecetes artificiosus (Fabricius, 1798). Conchoeodromia alcocki presents several characteristics of the megalopalstage, such as long feelerson the tip of the fifthpereiopod.As a result,the genera Conchoeodromia Chopra,1934, and ConchoecetesStimpson,1858, are synonymized.Illustrations of comparativeappendagesof the megalopaland adultstagesare provided.The morphological featuresof P4 and P5 that allow membersof the shell-carryingdromiidsConchoecetesand 1854, to hold a bivalveshell are compared. HypoconchaGu6rin-Meneville, Chopra (1934: 477, pl. 8, figs. 1-6) established Conchoeodromia alcocki as a new 1934, merges into the synonymy of Conchoecetes Stimpson,1858. Had the dromiidmegalopa and first crab stage been better documentedat that time, someonewithChopra'sexperiencewouldcertainlyhaverecognizedhis materialas a megalopalstage.Toourknowledgethe onlydromiid postlarvaeknownpriorto 1934 were those of genus and species for two small crabs(6.2 x 5.2 and 5.6 x 4.7 mm) from the Bay of Bengal at Sandheads,off the mouthof the Hoogly River. His material came from about 37-m depth, in soft ooze-like mud with patches of sand and shells. No more specimens have been assignedto C. alcocki. Austrodromidia octodentata (Haswell, 1888), Conchoeodromia alcocki is the only describedby Hale (1925: 406, 407, fig. la) as species in the genus andhas been mentioned a brood young of Cryptodromia octodentata, in the literature only occasionally. Balss and of Stimdromia lateralis (Gray, 1831) as a (1957: 1605) referred to Conchoeodromia as brood young of Paradromia lateralis (Hale, choeodromia and Genkaia seemed possible to Guerin-M6neville,1854, are the only dromiids to carry a bivalve shell, opportunityis takenhereinto elaborateon the morphological featuresof theirP4 and P5. Abbreviations:MNHN (Museumnational d'Histoirenaturelle,Paris);ZSI (Zoological Survey of India, IndianMuseum,Calcutta); Mxp3, third maxilliped; P2-P5, second to fifth pereiopods; P12-P15, second to fifth pleopods. Measurements of the carapace (length x width) are given in millimeters (mm). The type specimenfiguredby Chopra (1934, pl. 8, figs. 1-6) is the holotype. an intermediatebetween the Dromiidaeand the Homolodromiidae.Miyake and Takeda (1970: 27) andTakeda(1985: 100) statedthat Conchoeodromiashare some aberrantcharacters with the genus Genkaia Miyake and Takeda,1970 (Tavares,1993; 1996). T. Sakai (1976: 7) never examined Conchoeodromia alcocki, but the relationshipbetween Conhim. McLay (1993: 122, 123) referred to as "enigmaticandobscure." Conchoeodromia Althoughthe type materialof Conchoeodromiaalcocki, deposited in the Zoological Survey of India, Calcutta,was not available for study,we considerthe informationavailable to be sufficientto regardConchoeodromia alcocki as a megalopal stage of Con- 1925: 410, pl. 40, fig. la-f). choecetes artificiosus (Fabricius, 1798). As Because Conchoecetes and Hypoconcha Morphology Conchoeodromia alcocki presents typical a result the genus ConchoeodromiaChopra, featuresof the megalopalstage, such as long 301 302 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000 Fig. 1. A-F, reproduction of Chopra's (1934: pl. 8, figs. 1-6) original photographs of the male holotype (6.2 x 5.2 mm) of Conchoeodromia alcocki (ZSI-C1689/1), actually the megalopa of Conchoecetes artificiosus (Fabricius). A, dorsal view; B, ventral view; C, detail of anterior view of ventral parts; D, Mxp3; E, external view of the left cheliped; F, P5. Notice the long feelers (f) at the dactyl tip. GUINOT AND TAVARES: CONCHOEODROMIA ALCOCKI: MEGALOPA OF CONCHOECETES ARTIFICIOSUS B A I 0,1mm 303 C I' ^^^0,25mm '.cz-\K { D E 0 I 0,25mm I I G F 0,1 mm 0,25 mm Fig. 2. The megalopa of Conchoecetes artificiosus (Fabricius), from Bombay, after Sankolli and Shenoy (1968: figs. 8, 9). A, dorsal view; B, cheliped; C, Mxp3; D, P3; E, P5, notice the long feelers (f); F, telson with its posterior border deeply notched and fringed with long setae; G, uropod; H, pleopod; I, P4, notice the toothed process (p) on the propodus. feelers at the tip of P5 (Fig. 1A, F). Lebour (1928) used the term feelers to describe the special setae found at the tip of P5 of the megalopa of brachyurans(Gumey, 1942: 276, fig. 114 F). Williamson (1976: 405) believed that the feelers of the megalopa of the Dromiacea indicated a close relationship between Dromiacea and Brachyura, rather than between Dromiacea and Anomura. Felder et al. (1985) refers to these setae as "brachyuran feelers." Actually, feelers are present in many brachyuran and anomuran megalopae (Rice, 1981). To our knowledge, the feelers first appear in the megalopa and are not retained in the first crab stage. Several other characters shown by Conchoeodromia alcocki commonly occur in the megalopal stage and, typically, are not retained in adults. These could explain why Chopra (1934: 478) included Conchoeodromia alcocki in the Dromiidae but considered that it "possesses a number of characters that are not usually met with in this family." Additionally, the features used by Chopra to define the genus Conchoeodromia are also found in Conchoecetes, most of them already in the megalopal stage. Thus, as in any dromiid megalopa, in Conchoeodromia alcocki (Fig. 1A) the body is markedly longer than wide and with subparallel borders (Rice and Provenzano, 1966; Sankolly and Shenoy, 1968; Kircher, 1970; Rice et al., 1970; Lang and Young, 1980; Wear, 1977). Even in the first crab of Austrodromidia octodentata and 304 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000 Fig. 3. Conchoecetes artificiosus (Fabricius), adult female, 23 x 22 mm, Nosy B6 (MNHN-B 6890). Notice the strong P4 and the filiform P5. Stimdromia lateralis the carapace remains longer than wide (Hale, 1925: 410, pl. 40, fig. la-f). Typically, in adult dromiid crabs the carapace is as wide as long or even wider than long; only a few representatives of the family, such as Ascidiophilus caphyraeformis Richters, 1880, and Epigodromia rotonda McLay, 1993, present an elongated body. Chopra (1934: 478) referred to the body shape of Conchoeodromia alcocki as "roughly pentagonal." The carapace closely resembles that of the megalopa of Conchoecetes artificiosus (Fig. 2A), which shows a "tendency towards the pentagonal shape of the adult" (Sankolli and Shenoy, 1968: 103). The conspicuous lobulation of the dorsal surface of the carapace with "well impressed grooves" (Chopra, 1934) and body ornamentation consisting of minute serrules, granules, and teeth are unusual for an adult dromiid. Rather, it resembles the megalopal stage of Conchoecetes artificiosus which is heavily built, with a globose and grooved carapace and relatively stout pereiopods. Typically, in adult dromiids the Mxp3 are operculiform, whereas in Conchoeodromia alcocki and in the megalopa of Conchoecetes artificiosus the Mxp3 are pediform and separated by a gap (Figs. ID, 2C). In Conchoeodromia alcocki the trigonal shape of the chela (Fig. IE) and the serrate borders of P2 and P3 are more similar to that of the megalopa of Conchoecetes artificiosus (Fig. 2B, D) than to the adult (Fig. 3). Conchoeodromia alcocki and all known species of Conchoecetes are the only dromiids with P4 stronger than P5 and much stouter than P2 and P3; P4 ends in a heavy propo- dus and a long, curved dactyl. In Conchoeodromia alcocki and in the megalopa of Conchoecetes artificiosus, the propodus of P4 bears a distinct toothed projection (Figs. 1A, B, 2A, I). In the adult of Conchoecetes artificiosus (Figs. 3, 5A, C-E) this projection is quadrangular and hollowed in the middle as a socket, into which a mobile process ending in a corneous tip can sink. Additionally, Conchoeodromia alcocki (Fig. IF) and the megalopa of Conchoecetes artificiosus (Fig. 2A, E) share a subdorsal P5 ending in a minute dactyl. This combination of characters of P4 and P5 is unique among the dromiids. In the photographs given by Chopra, the abdomen of Conchoeodromia alcocki is distinctly convex and partially folded in a hollow between the pereiopods (Fig. 1A, B). The extended abdomen of Conchoecetes artificiosus (Fig. 2A), as depicted by Sankolly and Shenoy (1968), may be either a natural state or a mere artefact and is not inconsistent with our hypothesis. It is worth noting that in Evius ruber Moreira, 1912, actually a megalopal stage of Cryptodromiopsis antillensis Stimpson, 1858 (Franco, 1998; Rathbun, 1937: 31, pl. 8, figs. 1, 2, as Dromia erythropus (George Edwards, 1771)), the abdominal segments are not completely extended either (Moreira, 1912: 322, fig. 1). Neither the gonopods nor the sexual openings were mentioned in Chopra's description of Conchoedromia alcocki, although he said that the two specimens were males. Pleopods 2-5 are well developed in the megalopa of Conchoecetes artificiosus (see Sankolli and Shenoy, 1968: 108) (Fig. 2H). In Conchoeodromia alcocki and in the megalopa of Conchoecetes artificiosus (Sankolli and Shenoy, 1968: fig. 2) the telson is medially concave, rather long, and profusely hairy (Fig. 2A, F), as in many brachyuran megalopae. It is surprising that Chopra (1934: 480) found "no distinct platelets" (uropods) in Conchoeodromia alcocki. It may well be possible that the uropods are ventrally situated, as in the megalopa of Conchoecetes artificiosus (Fig. 2A), and went unnoticed by Chopra. Nowadays, it is well established that uropods (as dorsal or, more rarely, ventral platelets) are present in all known adult dromiids, with rare exceptions such as Ascidiophilus caphyraeformis Richters, 1880 (Guinot, 1995; Guinot and Bouchard, 1998) and Austrodromidia octodentata (fide McLay, 1993). According to GUINOT AND TAVARES: CONCHOEODROMIA ALCOCKI: MEGALOPA OF CONCHOECETES ARTIFICIOSUS 305 B 2,5mm C D mp I 1.,25 mm 1,25 mm F G 2,5 mm Fig. 4. A-D, Conchoecetes artificiosus, female, Nosy B6 (MNHN-B 6890). E-G, C. intermedius, holotype male, Madagascar (MNHN-B 6891); A, E, P4 with a quadrangularprojection (p) on the propodus and the mobile process (mp); C, D, G, detail of the quadrangularprojection (p) of the propodus of P4 and of the mobile process (mp); B, F, P5 with its upturned dactyl. P4 and P5 drawn at the same magnification. McLay (1998: 341, 344) the genus Alainodromia McLay, 1998, lacks uropod plates. The reexamination (this report) of the type material of the sole species in the genus, A. timorensis McLay, 1998, revealed, however, the presence of dorsal uropod plates (see also McLay, 1998, fig. 3). According to Franco (1998) the uropods appear in the third zoeal stage (in the dromiid species with three to six zoea stages) or in the megalopal stage (in the species with one or two zoea stages). Typically, the uropod is ventrally situated, well developed, and biramous in the megalopa and becomes uniramous and strongly reduced in subsequent stages. Size The megalopae of dromiids are relatively large. The average carapace length and width reportedby Wear (1977: 576) ranges from 2.7 306 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000 B A C 1,25 mm 2,5mm F 2,5 mm Fig. 5. A, P5 of Hypoconcha arcuata Stimpson, male, Sombrero (MNHN-B 22065); B-F, P4 and P5 of H. panamensis Smith, female, Lower California (MNHN-B 20865). A, F, P5 with its short propodus and dactyl; B, C, detail of the P5 upturned dactyl; D, detail of the dactyl of P4; E, stout P4. P4 and P5 drawn at the same magnification. Arrow points to prop-up plate. to 3.6 mm and from 2.2 to 3.1 mm respectively. From Sankolli and Shenoy's (1968) figure, the carapace of the megalopa of Conchoecetes artificiosus is about 3.4 x 2.7 mm. The only known specimens of Conchoeodromia alcocki are larger (6.2 x 5.2 mm and 5.6 x 4.7 mm), but their size is far from that attained by the adults of Conchoecetes artificiosus. Adults of Conchoecetes artificiosus have been reported from several localities: Madras, 23 x 24 mm (Henderson, 1893: 408); the Persian Gulf, 34 x 36 mm (Nobili, 1906: 94); Bombay, 15 x 16 mm (Chhapgar, 1969: 609); and Nosy Be, 23 x 22 mm (this report, Figs. 3, 4A-D). In the Herbst's Collection (K. Sakai, 1999: 14, pl. 4E) there is a female 26.5 x 28.5 mm labelled "Indian Ocean." Tirmizi and Kazmi (1991: 15) reported a male 20.5 GUINOTAND TAVARES:CONCHOEODROMIA ALCOCKI:MEGALOPA OF CONCHOECETES ARTIFICIOSUS x 22.5 mm from Karachi, Pakistan. The smallest Conchoecetes artificiosus examined by Barnard(1950: 309) had a carapace length of 7 mm, and the largest was 27 mm. In comparison, Conchoecetes andamanicus Alcock, 1900, is a much smaller species. Ihle (1913: 50) recorded an ovigerous female from the west coast of New Guinea with a carapace length of 5 mm; the specimen reported by Laurie (1906: 353) from the Gulf of Manaar had a carapace length of 10.2 mm and was probably an adult male. The largest (7.5 x 7.0 mm) C. andamanicus examined by Alcock (1901: 43) comes from Andamans. The third species in the genus, Conchoecetes intermedius Lewinsohn, 1984, seems to have an intermediate size between Conchoecetes artificiosus and C. andamanicus; the male holotype of Conchoecetes andamanicus from Madagascar measures 16 x 17 mm (Lewinsohn, 1984: 119). All the characters presented by Conchoedromia alcocki agree with those of the megalopa of Conchoecetes artificiosus; the only puzzle is posed by the carapace of Conchoedromia alcocki considerably longer than that of Conchoecetes artificiosus. It is worth noting, however, that the sizes of the two individuals of Conchoedromia alcocki significantly differ from one another, 6.2 x 5.2 mm and 5.6 x 4.7 mm. Geographical Range Conchoecetes artificiosus is widespread throughout the Indo-West Pacific and is common near the type locality of C. alcocki (Alcock, 1900: 152, 1901: 42; Chopra, 1934: 477). Just before the description of Conchoeodromia alcocki, 17 specimens of Conchoecetes artificiosus were recorded from the type locality, the Hoogly River, by Chopra (1933: 28; 1934: 477, footnote) himself. Shell-carrying Dromiids Species of Conchoecetes and Hypoconcha Guerin-Meneville, 1854, are the only dromiids that always carry a bivalve shell (note, however, that Hale, 1925: 406, pl. 40 A, reported that Austrodromidia octodentata (Haswell, 1888) can also shelter under a bivalve). The morphological features of P4 and P5 that allow the crab to hold a bivalve shell are clearly distinct in Conchoecetes and Hypoconcha. In both genera the specialized features of P4 and P5 are already present at 307 the megalopal stage (Sankolly and Shenoy, 1968; Kircher, 1970; Lang and Young, 1980; see Figs. 1A, B, 2A, E, I, in this report). Borradaile (1903) suggested that the two genera (and Sphaerodromia) were closely related. However, according to McLay (1993: 229), "While Conchoecetes probably belongs in the Dromiidae, the placement of Hypoconcha is doubtful." Therefore, we compare P4 and P5 from both genera here. On close scrutiny the resemblance between Conchoecetes and Hypoconcha proved to be superficial. In Conchoecetes, P4 and P5 are markedly dissimilar in shape, size, and position. These dissimilarities are already present in the megalopal stage (e.g., Conchoecetes artificiosus). In adult Conchoecetes artificiosus, P4 is much stronger than P5 and much stouter than P2 and P3, ending in a heavy propodus and a long, curved dactyl; the posterior border of the propodus bears a quadrangularprojection with a socket into which fits a mobile process ending in a corneous tip (Fig. 4C, D); this mobile process can sink and spring back in its socket, and as far as we know it is unique among the Brachyura. The bivalve shell is held by the mobile process and the dactyl. The P5 (Fig. 4B) is filiform, shorter than P4, ending in a simple upturned dactyl. Conchoecetes intermedius possesses a similar apparatus (Fig. 4F). In Hypoconcha, P4 is more robust and shorter than P5. As in Conchoecetes, only P5 is subdorsal in position. The P4 (Fig. 5D, E) and P5 dactyli (Fig. 5A-C, F) are short, upturned and can fit in a hollow excavated on the distal portion of the propodus. The bivalve shell, often a clamshell, is held by the crab with its posterior legs (Rathbun, 1937: 44, pl. 9, figs. 4, 5; Brusca, 1980: 318, fig. 20.47b; Hendrickx, 1997: 29). According to Williams (1984: 257), H. arcuata clings so tightly to its shell that its removal is almost impossible without damaging the crab. ACKNOWLEDGEMENTS This paper is dedicated to Raymond B. Manning's carcinological oeuvre. We thank Ray for his constant support and encouragement. Final stages of this paper were done while DG held an appointmentof Invited Researcher at Universidade Santa Ursula, Rio de Janeiro. We thank Gary C. B. Poore (Museum Victoria, Melbourne) for his comments on the manuscript (Gary also kindly checked the English text). The drawings and photographs are by Michele Bertoncini and Jacques Rebiere respectively (both from the Musdum national d'Histoire naturelle, Paris). MT thanks the CNPq (National Council for the 308 JOURNALOF CRUSTACEANBIOLOGY,VOL. 20, SPECIALNO. 2, 2000 Development of Science and Technology, Brasilia) for support in the form of ongoing grant 30.09.15/97-7. LITERATURECITED Alcock, A. 1900. Materials for a carcinological fauna of India. N? 5. The Brachyura Primigenia or Dromiacea.-Journal of the Asiatic Society of Bengal 68, Part II. Natural Science. N? 3 1899 (1900): 123-169. . 1901. Catalogue of the Indian Decapod Crustacea in the collection of the Indian Museum. Part I. Brachyura.Fasc. 1. Introductionand Dromides or Dromiacean (Brachyura Primigenia). Calcutta, India. ix + 80 pp. Balss, H. 1957. Decapoda. VIII. Systematik. Pp. 1505-1672 in H. G. Bronns, ed. Klassen und Ordnungen des Tierreichs. FiinfterBand, I. Abteilung, 7. Buch, 12. Lief. Leipzig und Heidelberg, Germany. Barnard, K. H. 1950. Descriptive catalogue of South African Decapod Crustacea.-Annals of the South African Museum 38: 1-837. Borradaile, L. A. 1903. On the genera of the Dromiidae.-Annals and Magazine of Natural History (7) 11: 297-303. Brusca, R. C. 1980. Common intertidal invertebrates of the Gulf of California. University of Arizona Press, Tucson, Arizona, 2nd ed. 513 pp. Chhapgar,B. F. 1969. More additions to the crab fauna of Bombay State.-Journal of the Bombay NaturalHistory Society 65 (1) 1968 (1969): 608-617. Chopra, B. N. 1933. Further notes on Crustacea Decapoda in the Indian Museum. VI. On the Crustacea Decapoda collected by the Bengal Pilot Service from the mouth of the River Hugly.-Records of the Indian Museum 35: 25-52. 1934. Further notes on Crustacea Decapoda in the Indian Museum. III. On a new dromiid and a rare oxystomous crab from the Sandheads, off the mouth of the Hoogly River.-Records of the Indian Museum 36: 477-481. Felder, D. L., J. W. Martin, and J. W. Goy. 1985. Patterns in early postlarval development of decapods. Pp. 163-225 in A. M. Wenner, ed. Larval growth. Crustacean Issues 2. A. A. Balkema. Rotterdam/Boston. Franco, G. M. 0. 1998. Zo6s, megalopa, e estagios juvenis de Cryptodromiopsis antillensis (Stimpson, 1858): implicacoes sobre a monofilia dos Dromiacea (Crustacea: Decapoda: Brachyura). Master's Thesis. Universidade Federal do Rio de Janeiro. 135 pp. Guinot, D. 1995. Crustacea Decapoda Brachyura:R6vision de la famille des Homolodromiidae Alcock, 1900. Pp. 155-282 in A. Crosnier, ed. R6sultats des Campagnes Musorstom, Volume 13. Memoires du Museum national d'Histoire naturelle 163. , and J.-M. Bouchard. 1998. Evolution of the abdominalholding systems of brachyurancrabs (Crustacea, Decapoda, Brachyura).-Zoosystema 20: 613-694. Gurney,R. 1942. Larvae of decapod Crustacea.The Ray Society, London. 306 pp. Hale, H. M. 1925. The development of two Australian sponge crabs.-Proceedings of the Linnean Society of New South Wales 50: 405-413. Henderson, J. R. 1893. A contribution to Indian Carcinology.-Transactions of the Linnean Society of London, 2nd series, 5: 325-458. Hendrickx, M. E. 1997. Los cangrejos braquiuros(Crustacea: Brachyura: Dromiidae, hasta Leucosiidae) del Pacifico Mexicano. Comision Nacional para el Conocimiento y uso de la Biodiversidad e Instituto de Ciencias del Mar y Limnologia, Univeridad Nacional Aut6noma de M6xico. 178 pp. Ihle, J. E. W. 1913. Die Decapoda Brachyurader SibogaExpeditien.I. Dromiacea.-Siboga-Expeditie 39b: 1-96. Kircher, A. B. 1970. The zoeal stages and glaucothoe of Hypoconcha arcuata Stimpson (Decapoda: Dromiidae) rearedin the laboratory.-Bulletin of Marine Science 20: 769-792. Lang, W. H., and A. M. Young. 1980. Larval development of Hypoconcha sabulosa (Decapoda: Dromiidae).-Fishery Bulletin 77: 851-864. Laurie, R. D. 1906. Report on the Brachyura collected by Professor Herdman,at Ceylon, in 1902. Pp. 349-432 in W. A. Herdman, ed. Report to the Government of Ceylon on the pearl oyster fisheries of the Gulf of Manaar. Part 5. Suppl. Rep. 40. Lebour, M. V. 1928. The larval stages of the Plymouth Brachyura.-Proceedings of the Zoological Society of London (2): 473-560. Lewinsohn, C. 1984. Dromiidae (Crustacea, Decapoda, Brachyura)from Madagascarand the Seychelles.-Memoires du Museum national d'Histoire naturelle (4) 6 sect. A (1): 89-129. McLay, C. L. 1993. The sponge crabs (Dromiidae) of New Caledonia and the Philippines with a review of the genera. Pp. 111-251 in A. Crosnier, ed. R6sultats des Campagnes Musorstom, Volume 10. Memoires du Museum national d'Histoire naturelle (A) 156. . 1998. A new genus and species of dromiid crab (Brachyura, Dromiidae) from the Timor Sea, NorthWest Australia with records of other species from the China Sea.-Zoosystema 20: 339-350. Miyake, S., and M. Takeda. 1970. A remarkablespecies of the Dromiacea (Crustacea Decapoda) from the Tsushima Islands, Japan.-Occasional Papers of Zoological Laboratory,Kyushu University, Japan3: 19-28. Moreira, C. 1912. Un crustace nouveau du Br6sil.-Bulletin de la Societe entomologique de France (15): 322-324. Nobili, G. 1906. Crustac6sDecapodes et Stomatopodes, Pp. 13-159 in Mission J. Bonnier et Ch. Perez (Golfe Persique 1901). Bulletin scientifique de la France et de la Belgique 40. Rathbun,M. J. 1937. The oxystomatous and allied crabs of America.-Bulletin of the United States National Museum 166: i-vi, 1-278. Rice, A. L. 1981. The megalopa stage in brachyuran crabs. The Podotremata Guinot.-Journal of Natural History 15: 1003-1011. , R. W. Ingle, and E. Allen. 1970. The larval development of the sponge crab, Dromia personata (L.) (Crustacea, Decapoda, Dromiidea), reared in the laboratory.-Vie et Milieu A, 21 (1A): 223-240. , and A. J. Provenzano. 1966. The larval development of the West Indian sponge crab Dromidia antillensis (Decapoda: Dromiidae).-Journal of Zoology, London 149: 297-319, figs. 1-15, tabl. 1-5. Sakai, T. 1976. Crabsof Japanand the adjacentseas. Kodansha Ldt., Tokyo, in 3 vol., xxix + 773 p. + 461 p. + 16 p., 251 pls. Sakai, K. 1999. J. F. W. Herbst-Collection of Decapod Crustacea of the Berlin Zoological Museum, with remarks on certain species.-Naturalists, Tokushima Biological Laboratory,Shikoku University 5: 1-45, 1 fig., 21 pls., 1 tabl. GUINOTAND TAVARES:CONCHOEODROMIA ALCOCKI:MEGALOPA OF CONCHOECETES ARTIFICIOSUS 309 Sankolly,K. N., and S. Shenoy. 1968. Larvaldevelop- Tirmizi, N. M., and Q. B. Kazmi. 1991. Crustacea: ment of a dromiid crab, Conchoecetes artificiosus Brachyura(Dromiacea,Archaeobrachyura, OxystoIn:MarineFaunaof Pakistan:4.mata,Oxyrhyncha). (Fabr.)(Decapoda,Crustacea)in the laboratory.-Journal of the MarineBiologicalAssociationof India9 (1) Publ. 1. BCCI FoundationChair,Instituteof Marine 1967 (1968): 96-110. Sciences,Universityof Karachi.246 pp., 65 figs. to PetalomTakeda,M. 1985. Recordof a male Genkaiagordonae Wear,R. G. 1977. A largemegalopaattributed erawilsoni(FultonandGrant,1902)(Decapoda,DromiMiyakeandTakedafromJapan(Crustacea: Decapoda: idae).-Bulletin of MarineScience27: 572-573. Brachyura).-SpecialPublicationof the Mukaishima MarineBiologicalStation:97-100. Williams,A. B. 1984. Shrimps,lobsters,andcrabsof the Atlanticcoast of the easternUnited States,Maineto Tavares,M. 1993. CrustaceaDecapoda:Les CyclodoFlorida. SmithsonianInstitutionPress. Washington, a rippidaeet Cymonomidaede l'Indo-Ouest-Pacifique l'exclusiondu genre Cymonomus. D.C. 550 pp. Pp. 253-313 in A. Crosnier,ed. R6sultatsdes CampagnesMusorstom,10. Williamson,D. I. 1976. Larvalcharactersand the oriMemoiresdu Museumnationald'Histoirenaturelle gin of crabs(Crustacea,Decapoda,Brachyura).-Tha156. lassiaJugoslavica10 1974 (1976):401-414. . 1996. Phyllotymolinidae,nouvelle famille de 16 March 1999. BrachyouresPodotremata(Crustacea,Decapoda).- RECEIVED: ACCEPTED:28 September 1999. Zoosystema20: 109-122.