GUINOT D. & TAVARES M., 2000. — Conchoedromia alcocki Chopra, 1934: megalopa of Conchoecetes...

Conchoeodromia alcocki Chopra, 1934: Megalopa of Conchoecetes artificiosus (Fabricius, 1798)(Decapoda, Brachyura, Dromiidae)Author(s): Danièle Guinot and Marcos TavaresReviewed work(s):Source: Journal of Crustacean Biology, Vol. 20, No. 2, Special Number Dedicated to Dr.Raymond B. Manning (Jun., 2000), pp. 301-309Published by: The Crustacean SocietyStable URL: http://www.jstor.org/stable/1549508 .Accessed: 08/05/2012 11:17

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

CONCHOEODROMIA ALCOCKI CHOPRA, 1934: MEGALOPA OF CONCHOECETES ARTIFICIOSUS (FABRICIUS, 1798) (DECAPODA,

BRACHYURA, DROMIIDAE)

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])

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 genera Conchoeodromia Chopra, 1934, and Conchoecetes Stimpson, 1858, are synonymized. Illustra- tions of comparative appendages of the megalopal and adult stages are provided. The morphological features of P4 and P5 that allow members of the shell-carrying dromiids Conchoecetes and Hypoconcha Gu6rin-Meneville, 1854, to hold a bivalve shell are compared.

Chopra (1934: 477, pl. 8, figs. 1-6) established Conchoeodromia alcocki as a new genus and species for two small crabs (6.2 x 5.2 and 5.6 x 4.7 mm) from the Bay of Ben- gal at Sandheads, off the mouth of the Hoo- gly 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 assigned to C. alcocki.

Conchoeodromia alcocki is the only species in the genus and has been mentioned in the literature only occasionally. Balss (1957: 1605) referred to Conchoeodromia as an intermediate between the Dromiidae and the Homolodromiidae. Miyake and Takeda (1970: 27) and Takeda (1985: 100) stated that Conchoeodromia share some aberrant characters with the genus Genkaia Miyake and Takeda, 1970 (Tavares, 1993; 1996). T. Sakai (1976: 7) never examined Conchoeodromia alcocki, but the relationship between Con- choeodromia and Genkaia seemed possible to him. McLay (1993: 122, 123) referred to Conchoeodromia as "enigmatic and obscure."

Although the type material of Conchoeo- dromia alcocki, deposited in the Zoological Survey of India, Calcutta, was not available for study, we consider the information available to be sufficient to regard Conchoeodro- mia alcocki as a megalopal stage of Con- choecetes artificiosus (Fabricius, 1798). As a result the genus Conchoeodromia Chopra,

1934, merges into the synonymy of Con- choecetes Stimpson, 1858.

Had the dromiid megalopa and first crab stage been better documented at that time, someone with Chopra's experience would cer- tainly have recognized his material as a megalopal stage. To our knowledge the only dromiid postlarvae known prior to 1934 were those of Austrodromidia octodentata (Haswell, 1888), described by Hale (1925: 406, 407, fig. la) as a brood young of Cryptodromia octodentata, and of Stimdromia lateralis (Gray, 1831) as a brood young of Paradromia lateralis (Hale, 1925: 410, pl. 40, fig. la-f).

Because Conchoecetes and Hypoconcha Guerin-M6neville, 1854, are the only dromiids to carry a bivalve shell, opportunity is taken herein to elaborate on the morphological features of their P4 and P5.

Abbreviations: MNHN (Museum national d'Histoire naturelle, Paris); ZSI (Zoological Survey of India, Indian Museum, 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 specimen figured by Chopra (1934, pl. 8, figs. 1-6) is the holotype.

Morphology Conchoeodromia alcocki presents typical

features of the megalopal stage, such as long

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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.

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GUINOT AND TAVARES: CONCHOEODROMIA ALCOCKI: MEGALOPA OF CONCHOECETES ARTIFICIOSUS

B I'

^^^0,25mm '.cz-\K

D

I 0

0,25mm

I

I 0,1mm

{

C

E

G

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 Dro- miacea 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 Con- choeodromia alcocki commonly occur in the

megalopal stage and, typically, are not retained in adults. These could explain why Chopra (1934: 478) included Conchoeodro- mia alcocki in the Dromiidae but considered that it "possesses a number of characters that are not usually met with in this family." Ad- ditionally, the features used by Chopra to de- fine 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

A

F

0,1 mm

303

I


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 Conchoe- cetes 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 sur- face of the carapace with "well impressed grooves" (Chopra, 1934) and body ornamen- tation 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 Conchoe- cetes 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 Conchoeo- dromia alcocki and in the megalopa of Con- choecetes 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 dis- tinctly 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 Stimp- son, 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 open- ings 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 Conchoeo- dromia alcocki and in the megalopa of Con- choecetes 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 sur- prising that Chopra (1934: 480) found "no distinct platelets" (uropods) in Conchoeo- dromia 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. Nowa- days, 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

304


2,5mm

1.,25 mm

B

D C

mp

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 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, P5 with its upturned dactyl. P4 and P5 drawn at the same magnification.

McLay (1998: 341, 344) the genus Alain- odromia 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). Typ- ically, 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 reported by Wear (1977: 576) ranges from 2.7

305

I


B A

C 1,25 mm

2,5mm

2,5 mm

F

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, 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 Con- choecetes artificiosus is about 3.4 x 2.7 mm. The only known specimens of Conchoeodro- mia alcocki are larger (6.2 x 5.2 mm and 5.6 x 4.7 mm), but their size is far from that at- tained 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

306


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 com- parison, 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 (Lewin- sohn, 1984: 119).

All the characters presented by Conchoe- dromia alcocki agree with those of the megalopa of Conchoecetes artificiosus; the only puzzle is posed by the carapace of Conchoe- dromia alcocki considerably longer than that of Conchoecetes artificiosus. It is worth noting, however, that the sizes of the two indi- viduals of Conchoedromia alcocki signifi- cantly 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 (Al- cock, 1900: 152, 1901: 42; Chopra, 1934: 477). Just before the description of Con- choeodromia alcocki, 17 specimens of Con- choecetes 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

the megalopal stage (Sankolly and Shenoy, 1968; Kircher, 1970; Lang and Young, 1980; see Figs. 1A, B, 2A, E, I, in this report). Bor- radaile (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 quadrangular projection 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 impossi- ble 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 appointment of 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

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Development of Science and Technology, Brasilia) for support in the form of ongoing grant 30.09.15/97-7.

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RECEIVED: 16 March 1999. ACCEPTED: 28 September 1999.

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GUINOT D. & TAVARES M., 2000. — Conchoedromia alcocki Chopra, 1934: megalopa of Conchoecetes...

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