JOURNAL OF BIOLOGY, 20, SPECIAL NUMBER 2: 301-309, 2000

CONCHOEODROMIA ALCOCK/ CHOPRA, 1934: MEGALOPA OF CONCHOECETES ARTIFICIOSUS (FABRICIUS, 1798) (, BRACHYURA, )

Daniele Guinot and Marcos Tavares

(DG) Museum national d'Histoire naturelle, Laboratoire de Zoologie (Arthropodes), 61 rue Buffon, 75005 Paris, France (e-mail: [email protected]); (MT) Universidade Santa Ursula, Instituto de Ciencias

Biol6gicas e Ambientais, Rio de Janeiro 22231-040, Brazil (e-mail: [email protected]) Downloaded from https://academic.oup.com/jcb/article/20/5/301/2419523 by guest on 01 October 2021

ABSTRACT Conchoeodromia alcocki Chopra, 1934, known from a single locality in the Bay of Bengal and from only two small specimens (6.2 x 5.2 and 5.6 x 4.7 mm), is shown to be a megalopa of Con· choecetes artificiosus (Fabricius, 1798). Conchoeodromia alcocki presents several characteristics of the megalopal stage, such as long feelers on the tip of the fifth pereiopod. As a result, the gen­ era Conchoeodromia Chopra, 1934, and Conchoecetes Stimpson, 1858, are synonymized. Illustra­ tions of comparative appendages of the megalopal and adult stages are provided. The morpholog­ ical features of P4 and P5 that allow members of the shell-carrying dromiids Conchoecetes and Hypoconcha Guerin-Meneville, 1854, to hold a bivalve shell are compared.

Chopra (1934: 477, pl. 8, figs. 1-6) estab­ 1934, merges into the synonymy of Con­ lished Conchoeodromia alcocki as a new choecetes Stimpson, 1858. and species for two small (6.2 x Had the dromiid megalopa and first 5.2 and 5.6 x 4.7 mm) from the Bay of Ben­ stage been better documented at that time, gal at Sandheads, off the mouth of the Hoo­ someone with Chopra's experience would cer­ gly River. His material came from about tainly have recognized his material as a mega­ 37-m depth, in soft ooze-like mud with lopal stage. To our knowledge the only drorniid patches of sand and shells. No more speci­ postlarvae known prior to 1934 were those of mens have been assigned to C. alcocki. Austrodromidia octodentata (Haswell, 1888), Conchoeodromia alcocki is the only described by Hale (1925: 406, 407, fig. 1a) as species in the genus and has 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, an intermediate between the Dromiidae and 1925: 410, pl. 40, fig. 1a-f). the Homolodromiidae. Miyake and Takeda Because Conchoecetes and Hypoconcha (1970: 27) and Takeda (1985: 100) stated that Guerin-Meneville, 1854, are the only drorni­ Conchoeodromia share some aberrant char­ ids to carry a bivalve shell, opportunity is acters with the genus Genkaia Miyake and taken herein to elaborate on the morpholog­ Takeda, 1970 (Tavares, 1993; 1996). T. Sakai ical features of their P4 and P5. (1976: 7) never examined Conchoeodromia Abbreviations: MNHN (Museum national alcocki, but the relationship between Con­ d'Histoire naturelle, Paris); ZSI (Zoological choeodromia and Genkaia seemed possible to Survey of India, Indian Museum, Calcutta); him. McLay (1993: 122, 123) referred to Mxp3, third maxilliped; P2-P5, second to Conchoeodromia as "enigmatic and obscure." fifth pereiopods; Pl2-Pl5, second to fifth Although the type material of Conchoeo­ pleopods. Measurements of the carapace alcocki, deposited in the Zoological (length x width) are given in millimeters Survey of India, Calcutta, was not available (mm). The type specimen figured by Chopra for study, we consider the information avail­ (1934, pl. 8, figs. 1-6) is the holotype. able to be sufficient to regard Conchoeodro­ mia alcocki as a megalopal stage of Con­ Morphology choecetes artificiosus (Fabricius, 1798). As Conchoeodromia alcocki presents typical a result the genus Conchoeodromia Chopra, features of the megalopal stage, such as long

301 302 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000 Downloaded from https://academic.oup.com/jcb/article/20/5/301/2419523 by guest on 01 October 2021

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 che­ liped; F, P5. Notice the long feelers (f) at the dactyl tip. GUINOT AND TAVARES: CONCHOEODROMIA ALCOCK/: MEGALOPA OF CONCHOECETES ARTIF/C/OSUS 303

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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 poste­ rior 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 megalopal stage and, typically, are not re­ (1928) used the term feelers to describe the tained in adults. These could explain why special setae found at the tip of P5 of the Chopra (1934: 478) included Conchoeodro­ megalopa of brachyurans (Gurney, 1942: 276, mia alcocki in the Dromiidae but considered fig. 114 F). Williamson (1976: 405) believed that it "possesses a number of characters that that the feelers of the megalopa of the Dro­ are not usually met with in this family." Ad­ rniacea indicated a close relationship between ditionally, the features used by Chopra to de­ and Brachyura, rather than be­ fine the genus Conchoeodromia are also tween Dromiacea and Anomura. Felder et al. found in Conchoecetes, most of them already (1985) refers to these setae as "brachyuran in the megalopal stage. Thus, as in any feelers." Actually, feelers are present in many dromiid megalopa, in Conchoeodromia al­ brachyuran and anomuran megalopae (Rice, cocki (Fig. lA) the body is markedly longer 1981). To our knowledge, the feelers first ap­ than wide and with subparallel borders (Rice pear in the megalopa and are not retained in and Provenzano, 1966; Sankolly and Shenoy, the first crab stage. 1968; Kircher, 1970; Rice et al., 1970; Lang Several other characters shown by Con­ and Young, 1980; Wear, 1977). Even in the choeodromia alcocki commonly occur in the first crab of Austrodromidia octodentata and 304 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000

dus and a long, curved dactyl. In Conchoeo­ dromia alcocki and in the megalopa of Con­ choecetes artificiosus, the propodus of P4 bears a distinct toothed projection (Figs. l A, B, 2A, 1). In the adult of Conchoecetes arti­ ficiosus (Figs. 3, SA, C-E) this projection is quadrangular and hollowed in the middle as a socket, into which a mobile process end­ ing in a corneous tip can sink. Additionally, Conchoeodromia alcocki (Fig. IF) and the megalopa of Conchoecetes artijiciosus (Fig. Downloaded from https://academic.oup.com/jcb/article/20/5/301/2419523 by guest on 01 October 2021 2A, E) share a subdorsal P5 ending in a minute dactyl. This combination of characters Fig. 3. Conchoecetes artijiciosus (Fabricius), adult fe­ male, 23 x 22 mm, Nosy Be (MNHN-B 6890). Notice of P4 and P5 is unique among the dromiids. the strong P4 and the filiform PS. In the photographs given by Chopra, the abdomen of Conchoeodromia alcocki is dis­ tinctly convex and partially folded in a hol­ Stimdromia latera/is the carapace remains low between the pereiopods (Fig. lA, B). The longer than wide (Hale, 1925: 410, pl. 40, fig. extended abdomen of Conchoecetes artifi­ la-f). Typically, in adult dromiid crabs the ciosus (Fig. 2A), as depicted by Sankolly and carapace is as wide as long or even wider than Shenoy (1968), may be either a natural state long; only a few representatives of the fam­ or a mere artefact and is not inconsistent with ily, such as Ascidiophilus caphyraeformis our hypothesis. It is worth noting that in Evius Richters, 1880, and Epigodromia rotonda ruber Moreira, 1912, actually a megalopal McLay, 1993, present an elongated body. stage of Cryptodromiopsis antillensis Stimp­ Chopra (1934: 478) referred to the body son, 1858 (Franco, 1998; Rathbun, 1937: 31, shape of Conchoeodromia alcocki as pl. 8, figs. 1, 2, as Dromia erythropus (George "roughly pentagonal." The carapace closely Edwards, 1771)), the abdominal segments are resembles that of the megalopa of Conchoe­ not completely extended either (Moreira, cetes artificiosus (Fig. 2A), which shows a 1912: 322, fig. 1). "tendency towards the pentagonal shape of Neither the gonopods nor the sexual open­ the adult" (Sankolli and Shenoy, 1968: 103). ings were mentioned in Chopra's description The conspicuous lobulation of the dorsal sur­ of Conchoedromia alcocki, although he said face of the carapace with "well impressed that the two specimens were males. Pleopods grooves" (Chopra, 1934) and body ornamen­ 2-5 are well developed in the megalopa of tation consisting of minute serrules, granules, Conchoecetes artificiosus (see Sankolli and and teeth are unusual for an adult dromiid. Shenoy, 1968: 108) (Fig. 2H). In Conchoeo­ Rather, it resembles the megalopal stage of dromia alcocki and in the megalopa of Con­ Conchoecetes artificiosus which is heavily choecetes artijiciosus (Sankolli and Shenoy, built, with a globose and grooved carapace 1968: fig. 2) the telson is medially concave, and relatively stout pereiopods. Typically, in rather long, and profusely hairy (Fig. 2A, F), adult dromiids the Mxp3 are operculiform, as in many brachyuran megalopae. It is sur­ whereas in Conchoeodromia alcocki and in prising that Chopra (1934: 480) found "no the megalopa of Conchoecetes artificiosus the distinct platelets" (uropods) in Conchoeo­ Mxp3 are pediform and separated by a gap dromia alcocki. It may well be possible that (Figs. lD, 2C). In Conchoeodromia alcocki the uropods are ventrally situated, as in the the trigonal shape of the chela (Fig. IE) and megalopa of Conchoecetes artificiosus (Fig. the serrate borders of P2 and P3 are more 2A), and went unnoticed by Chopra. Nowa­ similar to that of the megalopa of Conchoe­ days, it is well established that uropods (as cetes artijiciosus (Fig. 2B, D) than to the adult dorsal or, more rarely, ventral platelets) are (Fig. 3). present in all known adult dromiids, with rare Conchoeodromia alcocki and all known exceptions such as Ascidiophilus caphyrae­ species of Conchoecetes are the only dromi­ formis Richters, 1880 (Guinot, 1995; Guinot ids with P4 stronger than P5 and much stouter and Bouchard, 1998) and Austrodromidia oc­ than P2 and P3; P4 ends in a heavy propo- todentata (fide McLay, 1993). According to GUINOT AND TAVARES: CONCHOEODROMIA ALCOCK!: MEGALOPA OF CONCHOECETES ARTIFICJOSUS 305

2,5mm B

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1,25 mm G

1,25mm

2,5mm

Fig. 4. A-D, Conchoecetes artificiosus, female, Nosy Be (MNHN-B 6890). E-0, C. intermedius, holotype male, Madagascar (MNHN-B 6891); A, E, P4 with a quadrangular projection (p) on the propodus and the mobile process (mp); C, D, G, detail of the quadrangular projection (p) of the propodus of P4 and of the mobile process (mp); B, F, PS with its upturned dactyl. P4 and P5 drawn at the same magnification.

McLay (1998: 341, 344) the genus Alain­ the species with one or two zoea stages). Typ­ odromia McLay, 1998, lacks uropod plates. ically, the uropod is ventrally situated, well The reexamination (this report) of the type developed, and biramous in the megalopa and material of the sole species in the genus, A. becomes uniramous and strongly reduced in timorensis McLay, 1998, revealed, however, subsequent stages. the presence of dorsal uropod plates (see also McLay, 1998, fig. 3). According to Franco Size (1998) the uropods appear in the third zoeal The megalopae of dromiids are relatively stage (in the dromiid species with three to large. The average carapace length and width six zoea stages) or in the megalopal stage (in reported by Wear ( 1977: 576) ranges from 2. 7 306 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2, 2000

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Fig. 5. A, P5 of Hypoconcha arcuata Stimpson, male, Sombrero (MNHN- B 22065); B- F, P4 and P5 of H. pana­ mensis Smith, female, Lower California (MNHN- B 20865). A, F, P5 with its short propodus and dactyl; B, C, de­ tail 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 respec­ have been reported from several localities: tively. From Sankolli and Shenoy's (1968) Madras, 23 x 24 mm (Henderson, 1893: 408); figure, the carapace of the megalopa of Con­ the Persian Gulf, 34 x 36 mm (Nobili, 1906: choecetes arti.ficiosus is about 3.4 x 2.7 mm. 94); Bombay, 15 x 16 mm (Chhapgar, 1969: The only known specimens of Conchoeodro­ 609); and Nosy Be, 23 x 22 mm (this report, mia alcocki are larger (6.2 x 5.2 mm and 5.6 Figs. 3, 4A-D). In the Herbst's Collection (K. x 4.7 mm), but their size is far from that at­ Sakai, 1999: 14, pl. 4E) there is a female 26.5 tained by the adults of Conchoecetes arti.fi­ x 28.5 mm labelled "Indian Ocean." Tirmizi ciosus. Adults of Conchoecetes artificiosus and Kazmi (1991: 15) reported a male 20.5 GUINOT AND TAVARES: CONCHOEODROMIA ALCOCKJ: MEGALOPA OF CONCHOECETES ARTIFIC/OSUS 307 x 22.5 mm from Karachi, Pakistan. The the megalopal stage (Sankolly and Shenoy, smallest Conchoecetes artificiosus examined 1968; Kircher, 1970; Lang and Young, 1980; by Barnard (1950: 309) had a carapace length see Figs. 1A, B, 2A, E, I, in this report). Bor­ of 7 mm, and the largest was 27 mm. In com­ radaile (1903) suggested that the two genera parison, Conchoecetes andamanicus Alcock, (and Sphaerodromia) were closely related. 1900, is a much smaller species. Ihle (1913: However, according to McLay (1993: 229), 50) recorded an ovigerous female from the "While Conchoecetes probably belongs in the west coast of New Guinea with a carapace Dromiidae, the placement of Hypoconcha is length of 5 mm; the specimen reported by doubtful." Therefore, we compare P4 and P5 Laurie (1906: 353) from the Gulf of Manaar from both genera here. had a carapace length of 10.2 mm and was On close scrutiny the resemblance between Downloaded from https://academic.oup.com/jcb/article/20/5/301/2419523 by guest on 01 October 2021 probably an adult male. The largest (7 .5 x 7.0 Conchoecetes and Hypoconcha proved to be mm) C. andamanicus examined by Alcock superficial. In Conchoecetes, P4 and P5 are (1901: 43) comes from Andamans. The third markedly dissimilar in shape, size, and posi­ species in the genus, Conchoecetes inter­ tion. These dissimilarities are already present medius Lewinsohn, 1984, seems to have an in the megalopal stage (e.g., Conchoecetes ar­ intermediate size between Conchoecetes ar­ tificiosus). In adult Conchoecetes artificiosus, tificiosus and C. andamanicus; the male bolo­ P4 is much stronger than P5 and much stouter type of Conchoecetes andamanicus from than P2 and P3, ending in a heavy propodus Madagascar measures 16 x 17 mm (Lewin­ and a long, curved dactyl; the posterior bor­ sohn, 1984: 119). der of the propodus bears a quadrangular pro­ All the characters presented by Conchoe­ jection with a socket into which fits a mo­ dromia alcocki agree with those of the mega­ bile process ending in a corneous tip (Fig. 4C, lopa of Conchoecetes artificiosus; the only D); this mobile process can sink and spring puzzle is posed by the carapace of Conchoe­ back in its socket, and as far as we know it dromia alcocki considerably longer than that is unique among the Brachyura. The bivalve of Conchoecetes artificiosus. It is worth not­ shell is held by the mobile process and the ing, however, that the sizes of the two indi­ dactyl. The P5 (Fig. 4B) is filiform, shorter viduals of Conchoedromia alcocki signifi­ than P4, ending in a simple upturned dactyl. cantly differ from one another, 6.2 x 5.2 mm Conchoecetes intermedius possesses a simi­ and 5.6 x 4.7 mm. lar apparatus (Fig. 4F). In Hypoconcha, P4 is more robust and Geographical Range shorter than P5. As in Conchoecetes, only P5 Conchoecetes artificiosus is widespread is subdorsal in position. The P4 (Fig. 5D, E) throughout the Indo-West Pacific and is com­ and P5 dactyli (Fig. SA-C, F) are short, up­ mon near the type locality of C. alcocki (Al­ turned and can fit in a hollow excavated on cock, 1900: 152, 1901: 42; Chopra, 1934: the distal portion of the propodus. The bivalve 477). Just before the description of Con­ shell, often a clamshell, is held by the crab choeodromia alcocki, 17 specimens of Con­ with its posterior legs (Rathbun, 1937: 44, pl. choecetes artificiosus were recorded from the 9, figs. 4, 5; Brusca, 1980: 318, fig. 20.47b; type locality, the Hoogly River, by Chopra Hendrickx, 1997: 29). According to Williams (1933: 28; 1934: 477, footnote) himself. (1984: 257), H. arcuata clings so tightly to its shell that its removal is almost impossi­ Shell-carrying Dromiids ble without damaging the crab. Species of Conchoecetes and Hypoconcha Guerin-Meneville, 1854; are the only dromi­ ACKNOWLEDGEMENTS ids that always carry a bivalve shell (note, This paper is dedicated to Raymond B. Manning's car­ however, that Hale, 1925: 406, pl. 40 A, re­ cinological oeuvre. We thank Ray for his constant sup­ port and encouragement. Final stages of this paper were ported that Austrodromidia octodentata done while DG held an appointment of Invited Researcher (Haswell, 1888) can also shelter under a bi­ at Universidade Santa Ursula, Rio de Janeiro. We thank valve). The morphological features of P4 and Gary C. B. Poore (Museum Victoria, Melbourne) for his P5 that allow the crab to hold a bivalve shell comments on the manuscript (Gary also kindly checked the English text). The drawings and photographs are by are clearly distinct in Conchoecetes and Michele Bertoncini and Jacques Rebiere respectively Hypoconcha. In both genera the specialized (both from the Museum national d'Histoire naturelle, features of P4 and P5 are already present at Paris). MT thanks the CNPq (National Council for the 308 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 20, SPECIAL NO. 2. 2000

Development of Science and Technology, Brasilia) for Pacifico Mexicano. Comision Nacional para el support in the form of ongoing grant 30.09.15/97-7. Conocimiento y uso de Ia Biodiversidad e Instituto de Ciencias del Mar y Limnologfa, Univeridad Nacional Aut6noma de Mexico. 178 pp. LITERATURE CITED Ihle, J. E. W. 1913. Die Decapoda Brachyura der Siboga­ Alcock, A. 1900. Materials for a carcinological fauna Expeditien.l. Dromiacea.-Siboga-Expeditie 39b: 1-96. of India. No 5. The Brachyura Primigenia or Dromi­ Kircher, A. B. 1970. The zoeal stages and glaucothoe acea.-Journal of the Asiatic Society of Bengal 68, Part of Hypoconcha arcuata Stimpson (Decapoda: Dromi­ II. Natural Science. W 3 1899 (1900): 123-169. idae) reared in the laboratory.-Bulletin of Marine Sci­ ---. 1901. Catalogue of the Indian Decapod Crus­ ence 20: 769-792. tacea in the collection of the Indian Museum. Part I. Lang, W. H., and A. M. Young. 1980. Larval develop­ Brachyura. Fasc. 1. Introduction and Dromides or Dro­ ment of Hypoconcha sabulosa (Decapoda: Dromi­

miacean (Brachyura Primigenia). Calcutta, India. ix + idae).-Fishery Bulletin 77: 851-864. Downloaded from https://academic.oup.com/jcb/article/20/5/301/2419523 by guest on 01 October 2021 80 pp. Laurie, R. D. 1906. Report on the Brachyura collected Balss, H. 1957. Decapoda. VIII. Systematik. Pp. by Professor Herdman, at Ceylon, in 1902. Pp. 349-432 1505-1672 in H. G. Bronns, ed. Klassen und Ordnun­ in W. A. Herdman, ed. Report to the Government of gen des Tierreichs. Fiinfter Band, I. Abteilung, 7. Buch, Cey Ion on the pearl oyster fisheries of the Gulf of Man­ 12. Lief. Leipzig und Heidelberg, Germany. aar. Part 5. Suppl. Rep. 40. Barnard, K. H. 1950. Descriptive catalogue of South Lebour, M. V. 1928. The larval stages of the Plymouth African Decapod Crustacea.-Annals of the South Brachyura.-Proceedings of the Zoological Society of African Museum 38: 1-837. London (2): 473-560. Borradaile, L. A. 1903. On the genera of the Dromi­ Lewinsohn, C. 1984. 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