A new subfamilial arrangement for the Dromiidae de Haan, 1833, with diagnoses and descriptions of new genera and species (Crustacea, Decapoda, Brachyura)

Danièle GUINOT Département Milieux et Peuplements aquatiques, Muséum national d’Histoire naturelle, 61 rue Button, F-75231 Paris cedex 05 (France) [email protected]

Marcos TAVARES Universidade Santa Úrsula, Instituto de Ciências Biológicas e Ambientáis, Rio de Janeiro, 22231-040 (Brazil) [email protected]

Guinot D. & Tavares M. 2003. — A new subfamilial arrangement for the Dromiidae de Haan, 1833, with diagnoses and descriptions of new genera and species (Crustacea, Decapoda, Brachyura). Zoosystema 25 (1) : 43-129.

ABSTRACT A review of the morphology among members of the Dromiacea (and among other members of the Podotremata) revealed that the present classification of the Dromiidae does not reflect the wide variation of morphological patterns within the family. The Dromiidae are for the first time subdivided into three subfamilies: Dromiinae de Haan, 1833 n. status (type genus: Dromia Weber, 1795): Hypoconchinae n. subfam. (type genus:Hypoconcha G uérin- Méneville, 1854); and Sphaerodromiinae n. subfam. (type genus: Sphaerodromia Alcock, 1899). Thirty-eight dromiid genera are recognized herein: 34 in the Dromiinae n. status, five of which are new:Lamarckdromia n. gen. (type species: Dromia globosa Lamarck, 1818),Lewindromia n. gen. (type species: Cryptodromiopsis unidentata Riippell, 1830), Mclaydromia n. gen. (type species: Mclaydromia colini n. gen., n. sp.),Moreiradromia n. gen. (type species: Dromidia antillensis Stimpson, 1858), andStebbingdromia n. gen. (type species: Dromidiopsis plumosa Lewinsohn, 1984); one in the Hypoconchinae n. subfam.Hypoconcha ( Guérin-Méneville, 1854); and two in the Sphaerodromiinae n. subfam.Sphaerodromia ( , Eodromia McLay, 1993). The diagnoses of the following four dromiine genera are emended: Austrodromidia McLay, 1993; Cryptodromiopsis Borradaile, 1903; Dromidia Stimpson, 1858; andDromidiopsis Borradaile, 1900. The monotypic

ZOOSYSTEMA • 2003 • 25(1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris,www.zoosystema.com 43 Guinot D. & Tavares M.

Platydromia Brocchi, 1877 (type species:Dromia spongiosa Stimpson, 1858) is KEYWORDS Crustacea, resurrected. Seven dromiine genera are discussed in detail: Conchoecetes Podotremata, Stimpson, 1858;Desmodromia McLay, 2001; Epipedodromia André, 1932; Dromiidae, Fultodromia McLay, 1993; Hemisphaerodromia Barnard, 1954; Dromiinae n. status, Hypoconchinae n. subfam., Homalodromia Miers, 1884; and Pseudodromia Stimpson, 1858. Sphaerodromiinae n. subfam., Sphaerodromia , Eodromia, Hypoconcha and the enigmatic Frodromia are ana­ uropod, thoracic sternum, lyzed in detail. Special reference is made to the morphology of the thoracic spermathecae, sternum, spermathecae at the extremity of sternal sutures 7/8, uropods, new subfamilies, vestigial male pleopods on abdominal somites 3-5, coxa of the fifth pereopod new genera, new species, and the penis. A key to the families of the Dromiacea and the subfamilies of phylogeny. the Dromiidae is provided.

RÉSUMÉ Nouvel arrangement subfamilial pour les Dromiidae de Haan, 1833, avec dia­ gnoses et descriptions de nouveaux genres et espèces (Crustacea, Decapoda, Brachyura). Une revue de diverses dispositions morphologiques chez les Dromiacea (et plus largement chez les Podotremata) a révélé que la classification actuelle des Dromiidae ne reflétait pas la richesse des patterns morphologiques dans cette famille. La famille des Dromiidae est ici, pour la première fois, subdivisée en trois sous-familles : Dromiinae de Haan, 1833 n. status (genre type Dromia: Weber, 1795) ; Hypoconchinae n. subfam. (genre typeHypoconcha : Guérin- Méneville, 1854) ; et Sphaerodromiinae n. subfam. (genre type : Sphaerodromia Alcock, 1899). Trente-huit genres de Dromiidae sont recon­ nus : 34 genres de Dromiinae n. status, dont cinq nouveauxLamarckdromia : n. gen. (espèce type : Dromia globosa Lamarck, 1818),Lewindromia n. gen. (espèce type : Cryptodromiopsis unidentata Riippell, 1830), Mclaydromia n. gen. (espèce type : Mclaydromia colini n. gen., n. sp.),Moreiradromia n. gen. (espèce type : Dromidia antillensis Stimpson, 1858) ; etStebbingdromia n. gen. (espèce type : Dromidiopsis plumosa Lewinsohn, 1984) ; un genre d’Hypoconchinae n. subfam. Hypoconcha ( Guérin-Méneville, 1854) ; et deux genres de Sphaerodromiinae n. subfam. Sphaerodromia ( et Eodromia McLay, 1993). La diagnose de quatre genres de Dromiinae n. status Austrodromidia ( McLay, 1993, Cryptodromiopsis Borradaile, 1903, Dromidia Stimpson, 1858 et Dromidiopsis Borradaile, 1900) est émendée. Platydromia Brocchi, 1877 (espèce type : Dromia spongiosa Stimpson, 1858), monotypique, est réhabi­ MOTS CLÉS Crustacea, lité. Sept genres dromiens sont traités de façon détaillée Conchoecetes: Podotremata, Stimpson, 1858,Desmodromia McLay, 2001, Epipedodromia André, 1932, Dromiidae, Fultodromia McLay, 1993, Hemisphaerodromia Barnard, 1954, Dromiinae n. status, Hypoconchinae n. subfam., Homalodromia Miers, 1884 etPseudodromia Stimpson, 1858. Sont étudiés en Sphaerodromiinae n. subfam., détail les genres Hypoconcha, Sphaerodromia , Eodromia et l’énigmatique genre uropode, sternum thoracique, Frodromia. Nous avons privilégié les caractères suivants : organisation du sternum thoracique, morphologie de la spermathèque à l’extrémité de la suture sternale 7/8, de l’uropode, des pléopodes mâles vestigiaux sur les nouveaux genres, nouvelle espèce, somites abdominaux 3-5, de la coxa du cinquième péréiopode et du pénis. phylogénie. Une clé des familles de Dromiacea et des sous-familles de Dromiidae est fournie.

44 ZOOSYSTEMA « 2003 « 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

INTRODUCTION 27) coined the vernacular term “Anomaux” (latinized into Anomalia by Latreille [1817b: 358]; With more than 100 species, the Dromiidae de see McLaughlin & Holthuis 1985) to separate Haan, 1833, or sponge crabs, is the largest family the Hippidae Latreille, 1825, Paguridae Latreille, of primitive brachyuran crabs. Dromiids occur in 1802, and Galatheidae Samouelle, 1819 from the tropical and warm temperate waters worldwide. Macrura Latreille, 1802, he retained Dromia as a Five new genera are described, bringing to 38 the part of the Brachyura. Until H. Milne Edwards total number of dromiid genera recognized in the (1832), the position of the Dromiidae as part of current paper. the Brachyura remained unchallenged (e.g., The Dromiidae is currently regarded as part of Latreille 1802, 1806, 1817a, b, 1825, 1829; the monophyletic Brachyura Latreille, 1802. Desmarest 1825; Guérin-Méneville 1832; de Two major clades are recognized within the Haan 1833). However, H. Milne Edwards Brachyura: 1) the Podotremata Guinot, 1977, (1832: 313, 317, 320, 323, 324) transferred the containing all crabs with paired spermatheca “dromies” (Dromia) to the “Anomoures”, and (Tavares & Secretan 1993; Guinot & Tavares subsequently maintained this same view 2001); and 2) the assemblage Heterotremata (H. Milne Edwards 1834-1837). H. Milne Guinot, 1977-Thoracotremata (Eubrachyura Edwards’ viewpoint was followed by influential de Saint Laurent, 1980), which includes all crabs authors such as Lamarck (1838), Dana (1852, with paired vulvae (Guinot 1977, 1978, 1979a). 1853), Stimpson (1858), and Henderson (1888), We consider here the section Podotremata as although not by Lucas (1840, 1850). Boas containing three subsections: the subsection(1880) transferred the Dromiidae back to the Dromiacea de Haan, 1833, which consists in two Brachyura, and his arguments were adopted by superfamilies: Homolodromioidea Alcock, 1900 Ortmann (1892), Alcock (1900), Borradaile (one family: Homolodromiidae Alcock, 1900) (1903a, b), Ihle (1913), and Bouvier (1896, and Dromioidea de Haan, 1833 (two families: 1940), and have since been followed by many Dromiidae de Haan, 1833 and Dynomenidae carcinologists. For some time, the dromiids and Ortmann, 1892); the subsection Homolidea de related crabs were named “Brachyura Anomala”, Haan, 1839 (one superfamily Homoloidea, with as opposed to the “Brachyura genuina” (Alcock three families: Homolidae de Haan, 1839, 1899; Stebbing 1910). Latreilliidae Stimpson, 1858, and Poupiniidae It should be noted that, in a poorly known paper, Guinot, 1991); the subsection Archaeobrachyura H. Milne Edwards (1832) was the first to recog­ Guinot, 1977 (three families: Cyclodorippidae nize the basic types of female genital orifices in Ortmann, 1892, Phyllotymolinidae Tavares, the Decapoda: 1) gonopore on the coxa of the 1998, Cymonomidae Bouvier, 1897; and thethird pereopod; and 2) gonopore on the thoracic superfamily Raninoidea de Haan, 1839). sternite 6, already named “vulve” by Desmarest (1825: 62, 63, pi. 2, fig. 13) (see Latreille 1829: H is t o r ic a l a c c o u n t 29; H. Milne Edwards 1832: 304; 1837: 168). Before the formal establishment of the Brachyura Because the Dromiidae and related families share by Latreille (1802), the genus Dromia W eber, with the Macrura and Anomura the female gono­ 1795 (type genus of the Dromiidae; type species: pore on the coxa of the third pereopod, H. Milne Cancer personatus Linnaeus, 1758 by subsequent Edwards (1832: 304, 305) decided to remove designation by Holthuis 1962) had appeared in a them from the Brachyura where the female gono­ list along with a number of other crab genera pore opens on the thoracic sternite 6: “La posi­ (Weber 1795: 92). When Latreille (1802: 20, 22; tion des vulves éloigne encore tous les 1803: 155, 188, 382; see Dupuis 1986) estab­ Anomoures, ainsi que les Macroures, du groupe lished the Brachyura, he formally placed Dromia des Brachyures, et ce caractère est si facile à within that group. Even when Latreille (1817a: constater, même sur les individus desséchés et

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conservés dans les collections, qu’il est étonnantdoned (McLay 1993: 111) and more attention que jusqu’ici on y ait fait si peu d’attention”. devoted to characters such as carapace ornamen­ H. Milne Edwards (1832: 327, 331) recognized a tation, arrangement of spines on the legs, spe­ “famille naturelle” containing the genera cially on the two last pairs, number of abdominal “Ranine, Homole, Dromie et Dynomene”. Hesegments, and uropod plates. Altough these was the first to give a good definition and key for criteria are of good use in defining a number of these genera as forming a coherent group but left dromiid genera, a comprehensive view of it unnamed. This large group corresponds to the morphology and systematics of the group should Dromiacea, the “Decapodum Brachygnathorum include a detailed study of the thoracic sternum. Familia T ertia” of de H aan (1833: ix, x; 1839: A new view of the Dromiidae will likely emerge 102). The “tribu des Dromiens” of H. Milne when a clear understanding of the dromiid Edwards (1837: 168) was restricted to the genera thoracic sternum is achieved. Dromia Weber, 1795 andDynomene G uérin- Méneville, 1832, clearly distinct from the “tribu P r e l im in a r y morphological considerations des Homoliens” and “tribu des Raniniens”. The female thoracic sternum changes dramatically H. Milne Edwards (1837: 167) divided the during ontogeny (Gordon 1950: fig. 23), and the Anomura into two groups: the “Ptérygures” most drastic change refers to the development of (abdomen with paired mobile uropods) and the the female sternal sutures 7/8 (the “sternal “Apterures” (abdomen without biramous grooves” or “sternal furrows” of earlier authors). uropods). The true anomurans were placed in the Just because a character or a character complex Ptérygures, whereas the Dromiidae and their rela­ changes during ontogeny is not, in itself, an argu­ tives were arranged in the Apterures. Fransenet ment for not using it to define genera (Manning al. (1997: 159) rightly credited the name & Holthuis 1981: 15-17; McLay 1993: 122, 150; Anomura to H. Milne Edwards, 1832, and not to 2001a: 86); it is evident that morphological com­ MacLeay (1838: 54) who first latinized parisons should only be made between fully devel­ “Anomoures” into “Anomura”. The position ofoped individuals. The generic condition of female the Dromiidae within the Brachyura has been sternal sutures 7/8 does not detract their useful­ recently challenged by studies based on molecular ness in the arrangement and reveals to be a way as data (Spears et al. 1992; Spears & Abele 1996; see fruitful as “overall appearance and camouflage- Martin 2001; Martin & Davis 2001). However, carrying equipment” (McLay 2001a: 86). As a re­ the debate on the relationships of the sponge sult of the extension of sutures 7/8, the thoracic crabs suffers from the lack of accurate under­ sternum of adult females appears very much dis­ standing of dromiid morphology, and from torted, the sternites 7 and 8 occupying much of insufficient molecular data. Despite being the the ventral surface of the céphalothorax (Tavares largest and most diverse family of podotreme 1994: fig. 37C, D). In many dromiids, the female crabs, no subfamilial divisions have yet been sternal surface also becomes more complex, owing attempted for the Dromiidae (Borradaile 1903a, to the fact that, often, the apertures of spermathe­ b; Ihle 1913; Forest 1974; Lewinsohn 1977,cae end apart on (or beneath) two more or less 1979, 1984; McLay 1993, 1998, 2001a, b; prominent tubercules, or more or less together on McLay et al. 2001). (or beneath) a unique tubercle. In many mature McLay’s (1993) recent revision of sponge crabs females a sperm plug, fixed to a more or less large was a major step towards a comprehensive pic­ part of the sternum, has been observed. The pres­ ture of morphology and systematics of the ence of a sperm plug, however, does not imply Dromiidae. Because many genera and species had that the second gonopod of the male has not en­ been poorly defined, McLay’s work focused pri­ tered the aperture of spermatheca to penetrate for marily on dromiid diversity. Emphasis on the a more internal insemination. When surrounding nature of female sternal sutures 7/8 was aban­ the spermathecal apertures, the plaque of hard­

46 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

ened secretions was sometimes pierced by an The correct identification of the sternites may be opening, probably leaving a passageway to the difficult in the Podotremata since the sternal spermathecae. The displacement of female sutures sutures are located only laterally. Because the 7/8 and their terminal apertures to the anterior appendages are articulated on the sternum by part of thoracic sternum (see Gordon 1950: 244- their condyles, thoracic sternites are usually locat­ 247, fig. 23) is not exclusive to the Dromiidae (see ed by finding the corresponding appendages. Key to families of Dromiacea and subfamilies of Sometimes it is easier to locate the gynglymes Dromiidae, included in this study). A similar for­ (H. Milne Edwards 1851: 52). The gynglyme ward movement of sutures 7/8 is also encountered(from the Greek, gym female) is a type of socket in the more advanced podotreme families (Cyclo­ hollowed on the sternal surface, more precisely dorippidae Ortmann, 1892, specially in on the episternite, which receives one of the artic­ Neocorycodus Tavares, 1993, see Tavares 1996: ular condyles (sternal condyle) of the pereopod fig. 23D; Cymonomidae Bouvier, 1897, and (Fig. 16); the other condyle (pleural condyle) is Phyllotymolinidae Tavares, 1998, see Tavares articulated on the pleurite of the same metamere. 1994), but the apertures of spermathecae on the Ongoing studies on the primitive brachyuran anterior part of sternum correspond to a some­crabs have necessitated a review of the morpho­ what different pattern. logical patterns within the Dromiacea (Homo­ In conclusion, all Podotremata are characterized lodromiidae, Dynomenidae, and Dromiidae). As by the presence of paired spermathecae that open far as the Dromiidae are concerned, it was found independently on the line of sutures 7/8, and the that the current classification does not reflect the podotreme female thoracic sternum is markedly wealth of morphological patterns found within modified for a sexual function (for the special the family. In the literature the dromiacean uro- condition of spermathecal apertures in the pod, for example, is always described by the sim­ Raninoidea, see Hartnoll 1979). plistic expression of “vestigial”, without taking Besides a rather similar podotreme groundplan into account whether the uropod shows as a ven­ (with the thoracic sternum essentially exposed by tral plate (Homolodromiidae, Dromiidaepro its somites 4-8), diverse patterns are encountered parte), a dorsal plate (Dynomenidae, Dromiidae in the Dromiidae. Particular attention has been pro parte) (see G uinot & Tavares 2001: fig. 15), paid during this study to arrangement of the or, rarely, is lost. Because the dromiid uropod is anterior thoracic sternites. Because sternites 1-3 so diverse, emphasis is here given to its character are extremely narrow, and located at lower plane, states (see Patterns of uropods and vestigial male they are often concealed. Nevertheless, in some pleopods 3-5, and Table 1), all the more since the dromiids sternites 1 and 2 are discernible, or, at dorsal plate is very often used to hold the imma­ least in males, only sternite 3 may be exposed and ture female and male abdomens, while the ventral clearly visible dorsally. Sternite 4 shows an array plate does not fill any apparent function. of variations in both sexes. When male abdomen The study of the coxa of the fifth pereopod in is flexed against ventral surface, either the tho­ the male, with its gonopore and penis, generally racic sternum is entirely covered by the abdomen, overlooked in the Decapoda, have received par­ or the thoracic sternum is left partly uncovered ticular attention here and led to the recognition anteriorly. In neither case, a part of thoracic ster­ of two main dromiid patterns (see Patterns of num between the abdominal margin and the P5 coxa and penis). The condition of the coxae of the pereopods is exposed at level of ster­ female gonopore on the coxa of third pereopod nites 5 to 8 (episternite 5 excepted) (Guinot & has not been specified. It may show as occluded Tavares 2001). Only some anterior sternal parts, or perforated, probably in relation to the moult especially sternite 4, including its episternite which seems usual before mating in the female (episternite 4), remain uncovered, and, in a few dromiids (see Gordon 1950: 245; Hartnoll 1975: cases, also the small episternite 5. 669).

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N e w p r o p o s a l s — Dromidiopsis globosa (Lamarck, 1818), previ­ A preliminary subfamilial division for the ously Dromia globosa Lamarck, 1818, becomes Dromiidae is proposed herein. Three subfamilies the type species of Lamarckdromia n. gen. are recognized: Dromiinae de Haan, 1833 n. sta­ — Dromidiopsis dubia Lewinsohn, 1984 is herein tus, Hypoconchinae n. subfam., and Sphaero­ placed in Mclaydromia n. gen. A new species, dromiinae n. subfam. The new subfamilial M . colini n. gen., n. sp., is described. system is largely based upon the organization of — Dromidiopsis plumosa Lewinsohn, 1984, origi­ the thoracic sternum, the male abdomen, the nally placed questionably in Dromidiopsis and later morphological relationship between thoracic ster­transferred by McLay (1991) to Dromidia, and num and abdomen, the uropods and their role in then by McLay (1993, 2001a) to Cryptodromiopsis, holding of abdomen, the male coxa of P5 and becomes the type species of Stebbingdromia n. gen. penis, the female thoracic sternal sutures 7/8, the In adopting the same way than McLay (1993) apertures of spermathecae, and the condition (if the generic names are formed by combining the known) of all the abdominal appendages includ­ name of a person withDromia. ing vestigial male pleopods and uropods. All The examination of extensive material, generally descriptions, generic comparisons, and illustra­ including the type species, was necessary to clear tions concentrate on the ventral surface of theup a number of systematic uncertainties. This has body. The tomentum has not been described nor allowed a redefinition of other genera, as follows: figured, and, by stating “thoracic sternite 4 visible Austrodromidia McLay, 1993; Conchoecetes dorsally”, we do not refer to its possible conceal­Stimpson, 1858;E odrom ia McLay, 1993; m ent by setae. Epipedodromia André, 1932; Frodromia McLay, The reappraisal of several species has necessitated 1993; Fultodromia McLay, 1993; H em is­ a strict delineation of a number of genera (theyphaerodromia Barnard, 1954; Homalodromia are indicated “sensu nobis” in the present paper), Miers, 1884; Hypoconcha Guérin-Méneville, and the establishment of five new ones: 1854; Pseudodromia Stimpson, 1858; and 1) D ro m id ia Stimpson, 1858 andCrypto­ Sphaerodromia Alcock, 1899. A few genera and dromiopsis Borradaile, 1903 are redefined on the species are in need of review, for example the basis of their type species, D. hirsutissima large and heterogeneousDromia Weber, 1795, (Lamarck, 1818) and Cryptodromiopsis tridens and some poorly known monotypic, endemic Borradaile, 1903, respectively. As a result, several genera to southern Australia and southern Africa. species formerly described in or transferred to Some synonymies are restricted. A key to separate Dromidia or Cryptodromiopsis, are removed from the families of the Dromiacea and subfamilies of Dromidia sensu nobis and fromCryptodromiopsis the Dromiidae is provided. sensu nobis. They are as follows: — Dromidia spongiosa Stimpson, 1858 is assigned Abbreviations to Platydromia Brocchi, 1877, which is resurrected G l first male pleopod or gonopod; G2 second male pleopod or gonopod; to accomodate it. mxp3 third maxilliped; — Dromidia antillensis Stimpson, 1858 and P1-P5 first to fifth pereopods; D. sarraburei Rathbun, 1910 are transferred to P13-P15 third to fifth pleopods; Moreiradromia n. gen. AM Australian Museum, Sydney; BM The Natural History Museum (ex British — Dromia unidentata Riippell, 1830, transferred Museum [Natural History]), London; by Kossmann (1880) toDromidia and later by EMU Estación Mazatlán, Universidad Nacional McLay (1993) to Cryptodromiopsis, becomes the Autónoma, México; type species of Lewindromia n. gen. M N H N Muséum national d’Histoire naturelle, Paris; QM Queensland Museum, Brisbane; 2) Dromidiopsis Borradaile, 1900 is redefined on RM NH Nationaal Natuurhistorisch Museum, Leiden; the basis of its type species, D. australiensis ZRC National University of Singapore, Raffles (Haswell, 1882). Museum, Zoological Reference Collection.

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Descriptive terminology essentially follows that of Moreiradromia n. gen.; Paradromia Balss, 1921; McLay (1993). Measurements of carapace length Petalomera Stimpson, 1858;Platydromia Brocchi, 1877;Pseudodromia Stimpson, 1858;Speodromia X carapace width are given in millimeters (mm). Barnard, 1947; Stebbingdromia n. gen. (uncertain status); Most material used in this study is deposited at Sternodromia Forest, 1974;Stimdromia McLay, 1993; the M NHN. Other specimens came from various Takedromia McLay, 1993; Tunedromia McLay, 1993. institutions. D e s c r ip t io n Carapace varying from longer than wide, as wide SYSTEMATICS as long to much wider than long; convex, clearly subdivided into anterior and posterior portions; Section PODOTREMATA Guinot,1977 dorsal regions rather well-defined and variously Subsection DROMIACEA de Haan, 1833 ornamented; lateral margins usually rounded. Superfamily D rom ioidea de Haan, 1833 Front variously shaped, often with median ros­ Family D rom iidae de Haan, 1833 trum and two pseudorostral teeth, one at each side of rostrum. Supra- and infraorbital margins Subfamily D rom iinae de Haan, 1833 n. status usually toothed and separated from each other by deep notch. Orbits small, rather circular, orient­ Dromies H. Milne Edwards, 1832: 302, sq. ed more or less horizontally. Ocular stalk rather short, more or less thick, diversely shaped. Dromiacea de Haan, 1833 pro parte, ix, x; 1839 pro parte. 102. Thoracic sternum narrow. Gynglymes of thoracic sternites 1-3 largely spaced from each other, situ­ Dromiens — H. Milne Edwards 1837pro parte. 167, 168. ated at lower plane. Sternite 4 forming plate in contact with bases of the mxp3 or separated from Dromites — Lucas 1840pro parte. 112. mxp3 by sternite 3 when exposed. Episternites Dromiidae — Ortmann 1892: 541, 543. — Barnard 4 and 5 more or less narrow, with gynglymes of 1950: 306. — Holthuis 1962: 56. — Manning & Holthuis 1981: 11 [Name 356 on Official List]. — PI and P2 in almost terminal location. Female McLay 1993 pro parte. 111-251. — Guinot et al. sternal sutures 7/8 long (exceptStebbingdromia 1994: 255, fig. 7. — Hendrickx 1995pro parte. 127. n. gen.), and apertures of spermathecae opening — Ng 1998: 1056, 1063, 1085. — Nget al. 2000 pro into thoracic sternum far beyond level of coxa of parte. 156; 2001 pro parte. 5. — Melo & Campos 1999 pro parte. 274. — Martin & Davis 2001 pro parte. 49, P3 (except Stebbingdromia n. gen.), apart or 74. — Chen & Haibao 2002 proparte\ 73, 541. together, on two tubercules or on central promi­ nence; apertures usually showing as minute pores Type G EN U S. — Dromia Weber, 1795 (type species: Cancer personatus Linnaeus, 1758 by subsequent desig­ at extremities of sutures 7/8, exceptionally as slits nation under the Plenary Powers of the International(.Sternodromia ). Anterior part of thoracic sternite Commission on Zoological Nomenclature, Opinion4 and episternites 4 and 5 either completely cov­ 688, see Holthuis 1962). Name 1568 on Official List. ered or left diversely uncovered by male abdomen GENERA INCLUDED. — Ascidiophilus Richters, 1880; when folded. Deep sterno-coxal depressions usu­ Alainodromia McLay, 1998; Austrodromidia McLay, 1993; Barnardromia McLay, 1993; Conchoecetes ally present. Male abdomen length variable, Stimpson, 1858;Cryptodromia Stimpson, 1858; when flexed attaining the coxa of mxp3 or Cryptodromiopsis Borradaile, 1903; Desmodromia diverse levels of the P 1 coxa. Male abdomen gen­ McLay, 2001; D rom ia Weber, 1795; D rom idia erally narrow, often without pleural parts recog­ Stimpson, 1858;Dromidiopsis Borradaile, 1900; Epigodromia McLay, 1993; Epipedodromia André, nizable, or sometimes broader; all segments free 1932; E udrom idia Barnard, 1947; Exodromidia (exceptionally segment 5 and 6 more or less Stebbing, 1905; Fultodromia McLay, 1993; fused), segment 6 not noticeably extended later­ H aledrom ia McLay, 1993; Hemisphaerodromia ally. Male P13-P15 usually absent, but sometimes Barnard, 1954; Homalodromia Miers, 1884; Lamarckdromia n. gen.; Lauridromia McLay, 1993; vestigial. In males and immature females, Lewindromia n. gen.; Mclaydromia n. gen.; uropods showing as salient calcified dorsal plates,

ZOOSYSTEMA • 2003 • 25 (1) 49 Guinot D. & Tavares M.

often playing role in abdominal holding, or asA total of 34 genera are herein included in the ventral plates or lobes (see Patterns of uropods subfamily Dromiinae. The present list of dromiid and vestigial male pleopods 3-5). Mature female genera should be regarded as provisional, and fur­ uropods generally more horizontal, not so salient.ther adjustements and emendations might be Male telson usually short, variously shaped. necessary. For example, the large genus Dromia, Holding of male and immature female abdomens as currently defined (McLay 1993: 149, table 2), variable, usually efficient, a variable number of appears to be composite, and we regard it as pereopods being involved. Chelipeds with or Dromia s.l. We consider Sternodromia valid. without epipod; podobranch absent. P2 and P3 often lobed or nodose; propodus short, lacking distal propodal spine (except Stebbingdromia Genus Austrodromidia McLay, 1993 sensu nobis n. gen.); dactylus curved and armed with spines (Figs 1; 2) on inner border. P4 and P5 reduced, similar in size Dromia — Haswell 1882a pro parte\ 755; 1882bpro and shape (rare exception: Conchoecetes), shorter parte. 139, 140. (Non DromiaSd eber, 1795). than preceding ones, oriented subdorsally (P4) or Dromidiopsis — Ihle 1913 pro parte. 25. (N on dorsally (P5), prehensile; subcheliform apparatus Dromidiopsis Borradaile, 1900). present on propodus and dactylus and formed by Dromidia — Rathbun 1923a: 147. — Griffin 1972: 52. variable number of spines, varying from multiple (NonDromidia Stimpson, 1858). to only one, sometimes without opposing propo­ Cryptodromia — Rathbun 1923a: 151. — Hale 1925: dal spines. Male P5 coxa unmodified; penis 406; 1927: 107. — Griffin 1972: 53. (Non emerging as long and mobile calcified tube Cryptodromia Stimpson, 1858). (“penial tube”) (see Patterns of P5 coxa and Austrodromidia McLay, 1993 pro parte. 125, 185, penis). G2 long, with long, styliform and needle­ 186, table 6. — McLayet al. 2001 pro parte. 733, 740, like flagellum (except Stebbingdromia n. gen.), 743, table 2. without exopod. T ype SPECIES. — Dromidia australis Rathbun, 1923 by original designation (McLay 1993: 185). Gender: Carrying behaviour feminine. Sponges, compound or solitary ascidians, soft SPECIES INCLUDED. — Dromidia australis Rathbun, coral or actinians occasionally, bivalve shells 1923; Dromia octodentata Haswell, 1882. Cryptodromia incisa Henderson, 1888, from Australia rarely (see discussion under Shell-carrying behav­ and Japan, andDromidia insignis Rathbun, 1923, from iour), fragments of weed. Some dromiine mem­ Australia, assigned to Austrodromidia by McLay (1993: bers (Epigodromia, Takedromia ), with small last 185), were not available for study. Whether or not they pereopods, are not known to carry any camou­belong toAustrodromidia deserves further investigation. flage (Wicksten 1986a: 364; McLay 1993: 213, D istribution . — Australia. 216, 219, 224; 2001b: 7; Guinot et al. 1995: 385, 401; Ng et al. 2000: 157). D e s c r ip t io n Carapace slightly wider than long, convex; dorsal R em a r k s surface with regions not well-defined; branchial The following four dromiid genera have not beengroove marked. Anterolateral margin beginning included in the Dromiinae n. status:Hypoconcha at orbital level, armed with several teeth; postero­ Guérin-Méneville, 1854 referred to Hypoconchi­lateral margins toothed, not markedly convergent nae n. subfam. (see Hypoconchinae n. subfam.); posteriorly. Front narrow, appearing tridentate, and Eodromia McLay, 1993 and Sphaerodromia rostral tooth deflexed, two pseudorostral teeth; Alcock, 1899, referred to Sphaerodromiinae supraorbital, suborbital and exorbital teeth well- n. subfam. (see Sphaerodromiinae n. subfam.); developed. Exopod of antennal basal article the subfamilial status of Frodromia McLay, 1993 thickly developed, directed downwards, internal needs a re-appreciation. corner strongly produced. Mxp3: coxae with

50 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

1-2 mxp3

mxp3 1-2

cx1 cx1

cx2

cx2

3 5( e5

cx3

cx2 r 4/5

cx3

cx4

cx5

Fig. 1. — Austrodromidia australis (Rathbun, 1923), New South Wales, off Eden, Goonambee, 1922, S 29.5 x 33 mm (AM P5777); A, thoracic sternum, two front views; B, abdomen, dorsal view;C, abdomen, ventral view; note uropods showing as very narrow ventral plates. Abbreviations:a6, abdominal segment 6;cx1-cx5, coxae of P1-P5;e4, e5, épisternites 4, 5; mxp3, external maxil- liped; pr, holding prominence;pt, penial tube; t, telson; u, uropod; 1-2, sternites 1-2; 3-8, sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-778. Dotted line indicates difference in level. Scale bar: 5 mm.

ZOOSYSTEMA • 2003 • 25 (1) 51 Guinot D. & Tavares M.

1-2 mxp3

cx1

cx2

5/6

e5

cx3

mpx3 'I 1-2 cx2

cx1 cx3 5/6 sp

cx2

-4/5- - - h

cx4

/c x 3

cx5

Fig. 2. — Austrodromidia octodentata (Haswell, 1882), South Australia, Kangaroo Island,Endeavour, Rathbun det. Cryptodromia octodentata; A, B, S 39 x 42 mm (AM E. 801); A, thoracic sternum, two front views;B, abdomen, ventral view; note uropods reduced to nearly obsolete buds;C, 9 44 x 46 mm (AM P. P2311), thoracic sternum and apertures of spermathecae, covered with spermatic plug. Abbreviations: a6, abdominal segment 6;cx1-cx5, coxae of P1-P5; e4, e5, episternites 4, 5; mxp3, external maxilliped; o, female gonopore;pr, holding prominence;pt, penial tube; sp, spermatic plug; t, telson;u, uropod; 1-2, sternites 1-2; 3-8,stern­ ites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 5 mm.

52 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

(A. australis) or without (A. octodentata) narrow female sternal sutures 7/8 as ending together gap between them. between P2 (McLay 1993: 185, 186, table 6). Thoracic sternites 1-3 clearly (A. australis, These structures, however, have not been previ­ Fig. IA, B) or weakly (A. octodentata, Fig. 2A, C) ously described or figured in detail, even in the visible. Male thoracic sternite 4 markedly nar­ type species A. australis (Rathbun 1923a: 147) rowing anteriorly, with anterior margin truncate.(see Hale 1927: 106; Griffin 1972: 52 as Female sternal sutures 7/8 rather long, ending at Dromidia australis). The examination of a male of level of P2; apertures of spermathecae wide apart A. australis (AM P. 5777) revealed two very small between P2, beneath very prominent epister­ but distinct ventral plates hidden under the setae nites 4 (Fig. 2C). Sternite 3 remaining exposed that cover the foliaceous posterior part of ventral when male abdomen is applied against ventral surface of segment 6, and not visible dorsally surface; sternite 4 almost completely covered; (Fig. IB, C). In A. octodentata (Fig. 2B, C) the episternite 4 and episternite 5 exposed. male uropods are even more rudimentary, Male abdomen with all segments free, completely reduced to nearly obsolete buds. These buds are covering most of sterno-abdominal depression;easily overlooked and have been interpreted as telson bluntly triangular or rounded, almost “absent” (McLay 1993). In A. octodentata, how­ reaching mxp3, its base enlarged; pleural parts ever, the bud is found exactly in the same loca­ visible. Male segment 6 with external borders deeply tion where an uropod should be. InA. australis hollowed (but not thickened) in anterior part, andthe uropods are reduced to small plates. The broadly expanded in posterior part. Males without uropods are indistinct in the females ofA. octo­ vestigial pleopods. Male uropods as ventral lobes, dentata. We had no access to females ofA. aus­ small and narrow(A. australis), or inconspicuous, tralis and therefore its uropod condition remains or nearly obsolete(A. octodentata). Uropods not unconfirmed. Rathbun’s (1923a: 148) descrip­ involved in holding of abdomen. Abdomen holdtions of the sternal sutures 7/8 of A.australis (as folded by sharp or cristiform prominence on P2 Dromidia australis) are also not accurate. We sus­ coxa. Female uropods indistinct. pect that A. australis and A. octodentata (Fig. 2C) Chelipeds without epipod. P2 and P3 short, not share a similar condition in this regard, i.e. knobbed; propodus without distal spine; inner sutures 7/8 end apart instead of ending together. margin of dactylus armed with spines. P4 and P5 Austrodromidia octodentata, one of the largest reduced; P5 longer and slender, propodus short Australian sponge crabs, is known to have direct and broad; P4 carrying terminal apparatus development and brood its young, the abdomen formed by two distal propodal spines opposing forming a pouch for the young crabs (Hale 1927: curved dactylus; P5 terminal apparatus formed 109, figs 104, 105). A colour photograph of a by one distal propodal spine opposing dactylus; a specimen from South Australia carrying a colour­ long spine on outer margin of dactylus. ful colonial ascidian is given by Debelius (1999: Male P5 coxa with mobile penial tube (Figs IA; 2A). 249 as Cryptodromia octodentata). The status of Cancer aegagropila Fabricius, 1787, Carrying behaviour described supposedly from Australia and syn­ Sponges, ascidians, “marine growths, including onym o fDromia australiasae Weber, 1795 two species of plants”, “fragments of weed or {nomen nudum) (see Holthuis 1962), needs fur­ large shell” (Hale 1925: 406, pi. 40A; 1927: 108, ther investigation. 109); see discussion under Discussion, Shell-carrying behaviour. Genus Conchoecetes Stimpson, 1858 R em a r k s (Fig. 3) In Austrodromidia the uropods have been report­ Dromia — Fabricius 1798pro parte. 360. — Haswell ed as “reduced and concealed or absent” and the 1882bpro parte. 141. (NonD rom ia^!eber, 1795).

ZOOSYSTEMA • 2003 • 25 (1) 53 Guinot D. & Tavares M.

1-2 mxp3 1-2 mxp3

e4

cx2

e5

sp cx3 cx2

cx4 cx3

cx2

cx3

a6

cx4

cx5

Fig. 3. — A, Conchoecetes artificiosus (Fabricius, 1798), Madagascar, Nosy-Bé, 9 23 x 22 mm (MNFIN-B 6890), thoracic sternum, two front views, and spermathecae;B, C, Conchoecetes intermedius Lewinsohn, 1984;B, Madagascar, S 1 6 x 1 7 mm, holotype (MNFIN-B 6891), thoracic sternum with abdomen;C, Moluccas, S 9 x 9 mm (MNHN-B 15890), abdomen, ventral view; note absence of vestigial pleopods. Abbreviations:a3-a6, abdominal segments 3-6;cx1-cx5, coxae of P1-P5;e4, e5, episternites 4, 5; mxp3, external maxilliped (coxae separated for clarity); n, notch on margin of abdominal segmento, 5; female gonopore;pr, holding prominence;s, aperture of spermatheca; sp, sperm plug; t, telson; u, uropod; 1-2, sternites 1-2 (normally covered by mxp3 coxae, closely approximated); 3-8,sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 1 mm.

54 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

Conchoecetes Stimpson, 1858: 266; 1907: 180. —applied against ventral surface. Posterior sternites Henderson 1893: 407. — Alcock 1900: 150; 1901: tilted backwards. 40. — Borradaile 1903a: 301. — Ihle 1913: 50. — Chopra 1934: 478. — T. Sakai 1936: 41; 1976: 26. Male abdomen with all segments free, not — Barnard 1950: 308. — Lewinsohn 1984: 119. —completely covering sterno-abdominal depression; Dai & Yang 1991: 30. — Tirmizi & Kazmi 1991: 14. telson rounded at tip. Male segment 6 with sub­ — McLay 1993: 123, 174, table 5; 2001b: 8. — parallel external borders. Vestigial pleopods absent Guinot & Bouchard 1998: 621, 622. — Guinot & Tavares 2000: 301. — Ng et al. 2000: 156-159. — in males (papillae on segment 3 maybe present, to be McLay et al. 2001: 743, table 2. — Chen & Haibao verified) (Fig. 3C). Uropods (Fig. 3B, C) showing 2002: 73, 97, 540. as salient dorsal plates, involved in holding of ab­ Conchoedromia Chopra, 1934: 477 (type species: domen, and acting together with strong and orna­ Conchoedromia alcocki Chopra, 1934 by original desig­ mented prominence on P2 coxa; episternite 5 with nation; gender: feminine). some granules; P3 coxa with rounded tubercle T ype SPECIES. — Dromia artificiosa Fabricius, 1798 by matching notch on lateral margin of abdominal monotypy. Gender: masculine. segment 5; granule on P4 coxae similar to that on SPECIES INCLUDED. — Conchoecetes andamanicus P3, perhaps too small to be efficient. Alcock, 1900; Dromia artificiosa Fabricius, 1798; Chelipeds with epipod; P1-P3 with some nodosi­ Conchoecetes intermedius Lewinsohn, 1984. ties or tubercles. Propodus of P2 and P3 without Conchoecetes canaliculatus Yang & Dai, 1994 was regarded as a probable junior synonym ofC. inter- distal spine. P4 and P5 very dissimilar in posi­ medius by Ng et al. (2000). tion, size and shape, and with grasping system to

D istribution . — Indo-West Pacific. hold bivalve shell. P4 noticeably heavy, ending in thick propodus and in long curved dactylus; pos­ D e s c r ip t io n terior border of propodus bearing hollow, socket- Carapace as long as wide or slightly wider than like projection, with mobile process. P5 small long, subpentagonal. Dorsal surface flattened, and ending in simple, upturned dactylus. sometimes partly membranous, with some Male P5 coxa with mobile penial tube. regions defined; cervical and branchial grooves distinct. Anterolateral margin long, unarmed or Carrying behaviour with small teeth only; posterolateral margins with Exclusively involving bivalve shells (Nishimura or without tooth, straight or convergent posteri­1987: pi. M4; Nget al. 2000: fig. lb). See under orly. Front narrow, with three subequal teeth, the Discussion, Shell-carrying behaviour. deflexed rostral tooth and two pseudorostral teeth; supraorbital tooth small; suborbital and R em a r k s exorbital teeth not marked. Antenna: urinal article The case of Cancer mutus Linnaeus, 1758 relatively narrow, widening towards beak-like (p. 625), described with a smooth and anteriorly part; anterior part of beak narrow, acute, poste­ truncated carapace and with a brown anterior rior part rounded; exopod of basal article well- border, and for which the indication “? Medi­ developed, with internal corner acutely produced terranea” could be erroneous, is interesting. The and curved. Orbits rounded; ocular stalks short, type specimen(s) is (are) no longer extant. The eyes large. Mxp3: coxae closely approximated. name was subsequently used by Herbst (1783: Thoracic sternite 3 partly visible dorsally in both 116), who listed the species, but without any sexes (sternites 1-2 concealed) (Fig. 3A, B). figure. The species was then forgotten until the Thoracic sternite 4 broad, with anterior margin name was used again by K. Sakai (1999), this truncate. Female sternal sutures 7/8 separated time for a dromiid. Sakai (1999: pi. 4, fig. F) wide apart; apertures of spermathecae ending figured a specimen found in the Berlin Museum apart on two raised tubercles placed between P2 which had been identified and labelled by Herbst (Fig. 3A). In males, sternite 4 and episternites 4 as “Conchoecetes mutus Linnaeus, 1758”. The and 5 remaining exposed when male abdomen specimen figured by Sakai corresponds to a

ZOOSYSTEMA • 2003 • 25 (1) 55 Guinot D. & Tavares M.

Conchoecetes species (C. intermedius), whereas the tooth, which is wide apart, uninterrupted, eave- characters noted by Linnaeus are that of a trape- like; suborbital and exorbital teeth well-devel­ ziid crab, probably Tetralia Dana, 1851 (or oped. Proepistome developed, raised. Subhepatic Tetraloides Galil, 1986) (P. Castro and P. K. L. area convex, prominent dorsally. Antenna: urinal Ngpers. comm.). article upturned, with urinal opening placed McLay (1993, 2001b) and McLay et al. (2001) above axis of urinal article, and very narrow, argued for the close relationships between widening towards beak-like part; basal article Conchoecetes and Hypoconcha, mostly based on much enlarged and touching front on both cor­ the special condition of P4 and P5, the obligateners; exopod very developed and with internal shell-carrying behaviour, and the similarities in corner so produced that article 4 is completely or larval and postlarval development. For a compar­ almost included. Mxp3: coxae separated by gap. ison of the specialized morphological features Thoracic sternite 1-2 discernible; sternite 3 visi­ that allow the two genera to grasp a shell, see ble dorsally as a short triangular plate. Male tho­ G uinot & Tavares (2000: figs 4, 5). The system­ racic sternite 4 with external borders oblique. atic position ofConchoecetes is discussed under Female sternal sutures 7/8 long, separated wide Hypoconchinae n. subfam. apart proximally, but abruptly joined at level of The precise condition of the vestigial pleopods, P2; apertures of spermathecae ending together on suspected to be absent, needs to be verified. slight tubercle between chelipeds. When male The development ofConchoecetes only includes two abdomen applied against ventral surface, sternites zoeal stages and the megalopa (Sankolli & Shenoy 1-3 and extreme anterior part of sternite 4 (with 1968; see McLayet al. 2001: 740, 744, table 1). episternite 4) remaining visible. Male abdomen with all segments free, not completely covering sterno-abdominal depres­ Genus Cryptodromiopsis Borradaile, 1903 sensu nobis sion; no pleural parts discernible; telson broadly (Fig. 4) triangular. Male segment 6 with external borders Cryptodromiopsis Borradaile, 1903a: 299, 302; 1903b: subparallel on anterior half. No vestigial pleopods 578. — Tweedie 1950: 106. — McLay 1991pro in males. Uropods showing as salient dorsal parte: 467, 469; 1993pro parte: 187, table 6; 2001a plates, obliquely oriented, very mobile. Abdo­ pro parte. 84. minal holding by granulous crest on P2 coxae, far D rom idia — Lewinsohn 1979pro parte\ 3. (Non from uropods (Fig. 4A, B). Dromidia Stimpson, 1858). Chelipeds without epipod, and of moderate size, T ype SPECIES. — Cryptodromiopsis tridens Borradaile, knobbed; fingers gaping, prehensile margin of 1903 by monotypy (senior synonym ofDromidia fixed finger markedly concave; both cutting mar­ fenestrata Lewinsohn, 1979). Gender: feminine. gins without proximal teeth, only armed with SPECIES INCLUDED. — Cryptodromiopsis tridens interlocking distal teeth. P2 and P3 short, Borradaile, 1903. knobbed; propodus without distal spine; inner D istribution . — Indo-West Pacific. margin of dactylus armed with spines. P4 and P5 reduced, P5 longer and more slender. P4 and P5 D e s c r ip t io n carrying terminal apparatus formed mostly by Carapace distinctly wider than long, convex; one distal propodal spine opposing curved dacty­ dorsal surface with regions not well-defined; lus which ends in horny spine; a stout outer branchial groove quite distinct. Anterolateral propodal spine. P5 dactylus with spine on outer margin toothed; posterolateral margins without margin. tooth, and not markedly convergent posteriorly. G1 with setose apex, armed with sharp tubercle. Front narrow, appearing tridentate, rostral toothG2 with thin, needle-like, long flagellum, not long and deflexed, two pseudorostral teeth weak­ overreaching sterno-abdominal depression. er; border joining rostrum and supraorbital Male P5 coxa with mobile penial tube (Fig. 4B).

56 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

1-3 mxp3

cx1 mxp3 1-3

cx1 cx2

e5 M e4

cx2 cx3

e5 cx4 cx3 cx5

Fig. 4. — Cryptodromiopsis tridens Borradaile, 1903, Polynesia, Moorea, 1979,C. Bouchon coll.,â 6.2 x 7.5mm (MNHN-B 22579); thoracic sternum with(A) and without (B) abdomen. Abbreviations:a6, abdominal segment 6;cx1-cx5, coxae of P1-P5;e4, e5, epis­ ternites 4, 5; mxp3,external maxilliped; pr, holding prominence;pt, penial tube; t, telson;u, uropod; 1-3, sternites 1-3; 4-8,sternites 4-8; 6/7-7/8,thoracic sternal sutures 6/7-7/8. Scale bar: 1 mm.

Carrying behaviour genus contains six species for certain”, which is Sponges, compound ascidians. the number of species included in his (McLay 1993: 187, table 6) previous definition ofCrypto­ R e m a r k s dromiopsis. Borradaile (1903a: 299, 302; see 1903b: 578) The examination of several species presently erected Cryptodromiopsis mostly to separate from included in Cryptodromiopsis led to the conclu­ Cryptodromia Stimpson, 1858 (type species: sion that the genus is indeed a composite one. C. coronata Stimpson, 1858 by original designa­The following species are now being removed tion) those species with female sternal sutures 7/8 from Cryptodromiopsis sensu nobis: 1)C. uniden­ ending together (i.e. converging), in contrast totata (Ruppeli, 1830) transferred to Lewindromia wide apart in Cryptodromia. n. gen.; 2) C. antillensis (Stimpson, 1858) and While studying specimens of Cryptodromiopsis C. sarraburei (Rathbun, 1910) transferred to tridens from French Polynesia, McLay (1991: Moreiradromia n. gen.; and 3) C. plum osa 467-470, fig. 5) remarked that at that time the (Lewinsohn, 1984) transferred toStebbingdromia species included in Cryptodromiopsis did “not n. gen. make a natural group, having very little in For the time being,Dromia (Cryptodromia :) bul­ common with each other”. More recently, how­lifera Alcock, 1900 is perhaps better placed in ever, McLay (2001a: 84) maintained that “the Cryptodromia.

ZOOSYSTEMA • 2003 • 25 (1) 57 Guinot D. & Tavares M.

The two Chinese species Cryptodromiopsis dubia Genus Desmodromia McLay, 2001 Dai, Yang, Song & Chen, 1981 andC. planaria Dai, Yang, Song & Chen, 1981 were not exam­Desmodromia McLay, 2001b: 1-8. ined. Their status was suggested as doubtful by T ype SPECIES. — Desmodromia griffini McLay, 2001 McLay (1993: 187). The male abdomen of by original designation. Gender: feminine. C. dubia figured by Chen & Haibao (2002: SPECIES INCLUDED. — Desmodromia griffini McLay, fig. 42.4) seems rather similar to that of C. tridens 2001; D. tranterae McLay, 2001. (Fig. 4A). D istribution . — Australia. Thus, Cryptodromiopsis is herein reduced to the type species, C. tridens. Because of similarities D e s c r ip t io n between C. tridens and C. coronata Stimpson, See McLay 2001b: 1-8, figs 1-3, table 1. 1858, the type species ofCryptodromia, McLay (1991: 470; 1993: 188) argued that Crypto­ R em a rks dromiopsis is perhaps no longer necessary.Crypto­ The inclusion in the Dromiinae n. status of the dromiopsis tridens has been well illustrated by presumably shell-carrying species of the genus Lewinsohn (1979: fig. 1, asDromidia fenestrata) Desmodromia (regarded as an “unusual dromiid and by McLay (1991: fig. 5). Its main resem­ crab” by McLay 2001b: 1) is puzzling. The males blances with Cryptodromia coronata are as of both species are unknown and the description follows: 1) thoracic sternite 3 visible (Fig. 4); of the females is incomplete. Important informa­ 2) episternite 4 uncovered when male abdomen is tion on the two smallDesmodromia species (not folded; 3) general shape of male abdomen and available during this study) is missing, such as: similar holding of abdomen; 4) G2 needle-like 1) organization of thoracic sternum; 2) shape of but not much longer than Gl; and 5) similar male abdomen; 3) male uropods (showing as shape of urinal article of antenna. well-developed dorsal plates in an immature The differences that enable to distinguish female ofD. tranterae)-, 4) complete male pleopo- Cryptodromiopsis from Cryptodromia include: dal formula; and 5) morphology of male P5 coxa 1) apertures of spermathecae approximated on a (presence or not of a differentiated mobile penial median tubercle (wide apart in Cryptodromia)-, tube). Besides, it remains unclear whether the last 2) front narrow, with weak pseudorostral teeth, legs are really dissimilar in size and shape (also and presence of a frontal eave (pseudorostral their terminal grasping apparatus have not been teeth developed and no eave in Cryptodromia)-, figured in detail) and whether the female thoracic 3) antennal basal article broad and long, reaching sternites 7 and 8 are tilted. The female sternal front (not so developed inCryptodromia)-, and sutures 7/8 ending apart between P2 (McLay 4) male thoracic sternite 4 with external borders 2001b: 2) are typically dromiine. oblique and anterior border somewhat triangular Despite some resemblance of the short and (external borders subparallel and anterior border upturned dactyli of P4 and P5 to that of the squarely truncate in Cryptodromia). shell-carrying Hypoconcha, M cLay (2001b: 8) Even if these differences will in the future prove separated Desmodromia in his key from the cou­ to be within the range of variation of the largeplet Conchoecetes! Hypoconcha. genus Cryptodromia, one will have to consider As far as we can tell, Desmodromia and Conchoecetes that Cryptodromia s.l., as currently defined with (Fig. 3) share the following characters: 1) female some 20 species, is an heterogeneous assemblage. sternal sutures 7/8 ending between P2; and 2) ab­ Cryptodromiopsis tridens is distinguished from all domen hold folded by dorsal uropods and, most­ other dromiids by presence of two conspicuous ly, by a structure on P2 coxae. Besides their smooth, naked areas at posterior angles of cara­different carapaces, Desmodromia and Conchoe­ pace (see Lewinsohn 1979: fig. 1A, asDromidia cetes differ from each other in: 1) epipod absent on fenestrata). PI in Desmodromia (present inConchoecetes)-, 2) P4

58 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

and P5 more or less similar, with same dactylus Dromidia Stimpson, 1858pro parte-. 225; 1907 pro which is simply upturned and without opposing parte-. 170. — Borradaile 1903a pro parte-. 299. — Stebbing 1910pro parte-. 342. — Barnard 1950 pro propodal spines (P4 and P5 markedly dissimilar in parte. 319. — Tirmizi & Kazmi 1991 pro parte-. 27. — size and shape in Conchoecetes, with P4 very stout McLay 1993 pro parte-. 125, 183, table 5. — McLay et and specialized subcheliform apparatus, P5 fili­ al. 2001 pro parte-. 740, table 3. form and ending in small upturned dactylus; see N onPlatydromia Brocchi, 1877: 54 (type species: Guinot & Tavares 2000: fig. 4); and 3) abdominal P. depressa Brocchi, 1877 by monotypy). holding by structure on P2 coxa only (additional NonParasphaerodromia Spiridonov, 1992: 69 (type efficient structure on P3 coxae and inefficient species: P. subglobosa Spiridonov, 1992 by original structure on P4 coxae inConchoecetes, Fig. 3B; see designation). Guinot & Bouchard 1998: 622, fig. 4A). T ype SPECIES. — Dromia hirsutissima Lamarck, 1818 The carapace of Desmodromia, although not by original designation (Stimpson 1858: 225). membranous on posterior half as inHypoconcha, Gender: feminine. Lamarck (1818: 264) stated that his Dromia hirsutissi­ may be poorly calcified and flattened, and the ma came from Cape of Good Hope (“Cap de Bonne- eaves overhang eyes. The differences between Espérance”) and that it was deposited in the Paris Desmodromia and Hypoconcha consist of: 1) epi- Museum. H. Milne Edwards (1837: 176 as Dromia hirtissima, sic) seems to have examined the same mate­ pod absent on P1 inDesmodromia (present in rial and also mentioned that it was housed in the Paris Hypoconcha)-, 2) uropods as dorsal plates (as Museum. The only specimen ofDromidia hirsutissima minute ventral plates inHypoconcha, Fig. 19B, (Lamarck, 1818) in the M N H N collections is a male C); 3) female sternal sutures 7/8 ending between (26.6 X 31 mm) labeled “Cap de Bonne-Espérance” (MNHN-B 22034), and without details on the label. P2 (more posteriorly inHypoconcha, Fig. 19A); This specimen, presently preserved in alcohol (at one 4) P2 coxae and uropods functioning in abdomi­time dry, and perhaps rehydrated), is presumed to be nal holding (inHypoconcha either a differentia­ the type of Dromia hirsutissima and is selected here as tion on P1 coxa without involvement of uropodsthe lectotype of the species. or only provided by strong natural flexion of SPECIES INCLUDED. — Dromia hirsutissima Lamarck, abdomen, see Guinot & Bouchard 1998: fig. 1C, 1818. Whether Dromidia aegibotus Barnard, 1947, Dromidia D); and 5) P4 and P5 simply ending in upturned dissothrix Barnard, 1947 and Dromidiopsis cornuta dactylus (with diverse specialized features in Barnard, 1947 belong toD romidia, as stated by Hypoconcha, see under Hypoconchinae n. sub­ McLay (1993), deserves further investigation. fam., and Guinot & Tavares 2000: fig. 5). Dromidia spongiosa Stimpson, 1858 is herein trans­ ferred to Platydromia Brocchi, 1877. For a comparison ofDesmodromia w ith Epipe­ dodromia (Fig. 7A) and Homalodromia (Fig. 9), D istribution . — South Africa. in which the eyes and cephalic appendages are also ventrally located, see discussion of D e s c r ip t io n Homalodromia. Carapace wider than long, convex; dorsal surface only with low gibbosities; only branchial groove Carrying behaviour marked. Anterolateral margin long, with one Unknown, but supposedly bivalve shells (McLay tooth separated from exorbital angle by a deep 2001b: 7, 8). For comments on shell-carrying inconcavity, and followed by a prominence; a blunt Dromiidae, see under Discussion, Shell-carrying tooth behind branchial groove; posterolateral behaviour. margins very short, markedly convergent posteri­ orly. Front inclined, narrow, with median rostral tooth directed downwards, and two pseudorostral Genus Dromidia Stimpson, 1858 sensu nobis teeth; supraorbital, suborbital and exorbital teeth (Fig. 5) present. Antenna: basal article with exopod more Dromia — Lamarck 1818pro parte-. 264. — Lucas developed than internal angle which is hardly pro­ 1840 pro parte-. 113. (Non Dromia Weber, 1795). duced. Mxp3: coxae separated by narrow gap.

ZOOSYSTEMA • 2003 • 25 (1) 59 Guinot D. & Tavares M.

cx1 cx1

Mr

cx2 cx2 a6

cx3 a5

cx4

cx5

cx3 5/6

cx4

cx5

Fig. 5. — Dromidia hirsutissima (Lamarck, 1818), “Cap de Bonne-Espérance”,S 26.6 x 31 mm, lectotype (MNHN-B 22034); A, tho­ racic sternum, two front views;B, C, abdomen, dorsal and ventral views; note uropods showing as ventral plates, hardly visible dor­ sally. Abbreviations: a5, a6, abdominal segm ents 5, 6;cx1-cx5, coxae of P1-P5; e4, e5, episternites 4, 5; pr, spiniform holding prominence;pt, penial tube; v4, v5, elongated vestigial pleopods 4 and 5 (pleopods present also on segmentt, 3); telson; u, uropod; 4-6, sternites 4-6; 5/6-7/8, thoracic sternal sutures 5/6-7/8. Dotted line indicates difference in level. Scale bars: 2 mm.

Thoracic sternite 3 hardly visible dorsally; ster­ part of sternite 4 and episternite 4 remaining nite 4 forming plate weakly hollowed medially, exposed. with lateral borders oblique and anterior margin Male abdomen long, wide, with distinct spaced gently rounded (Fig. 5A, B). Female sternal pleural parts, and with all segments free; telson sutures 7/8 long, with apertures of sperma­ broadly triangular, ending in acute tip (Fig. 5B, thecae ending together on slight prominenceC). Male segment 6 with external borders paral­ between chelipeds. When male abdomen is lel. Vestigial pleopods present in males, as elon­ applied against ventral surface, only a small gated lobes on segments 3-5 (Fig. 5C). Uropods

60 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

showing as narrow ventral plates, deeply inserted ending in acute tip, leaving exposed anterior part ventrally, oblique, hardly visible dorsally. of sternite 4 and episternite 4 (telson bluntly Uropods not involved in holding of abdomen.rounded at tip, almost completely recovering On P2 coxa a strong spine, directed backwards, sternite 4, but episternites 4 and 5 exposed, in which may maintain abdominal segment 5 only Austrodromidia)-, and 6) abdomen maintained when P2 are moved backwards. folded by a strong P2 coxal spine, directed back­ Chelipeds long, without epipod. PI, P2 and P3 wards (a sharp prominence on P2 coxa in thick, not nodose or ridged; propodus of P2 and Austrodromidia). P3 without distal spine; inner margin of dactylus The presence of uropods is not mentioned by armed with spines. P4 and P5 reduced, P5 being Barnard (1947, 1950) inDromidia aegibotus and more slender. Propodus of P4 and P5 with distal Dromidiopsis cornuta, meaning perhaps that they spine opposing dactylus, which is very long and are ventral. In both species, the apertures of sper­ ends in horny spine; a spine on external border ofmathecae end on prominence between PI. In P5 propodus. Dromidiopsis aegibotus telson ends in sharp point Male P5 coxa with mobile penial tube (Fig. 5A). and P2 coxa bears a sharp spine directed back­ wards, as in Dromidia hirsutissima. The generic Carrying behaviour status of these two species and ofD. dissothrix, Individuals carry a compound ascidian. only known by a female, remains uncertain.

R em a r k s McLay (1993: 183) stressed the confusion Genus Dromidiopsis Borradaile, 1900 sensu nobis regarding the concept ofDromidia, for which (Fig. 6) “no fewer than eight definitions have been pub­ lished”. The present new restricted description is Dromia — H. Milne Edwards 1837 pro parte-. 178. — Haswell 1882a pro parte-. 755; 1882bpro parte-. 139. based on the type species of the genus, D. hir­ (NonDromiaSd eber, 1795). sutissima, previously poorly known and endemic Dromidiopsis Borradaile, 1900: 572; 1903a: 298; to South Africa (see Barnard 1950: 320, fig. 6la- 1903b pro parte-. 576. — Holthuis 1962: 56. — c). Stimpson (1858: 225) clearly defined the Lewinsohn 1984pro parte-. 95, 97-103. — Forest genus by the female sternal sutures 7/8 (“sulci”) 1974pro parte. 72, 74, 102, 103. — McLay 1993 pro ending together between chelipeds, “in tubercu­ parte: 135-137, table 2; 2001a pro parte-. 79, 80. — McLay et al. 2001 pro parte-. 733, 742, tables 2, 3. — lum approximati”. For Rathbun (1923b: 67, 68) Chen & Haibao 2002 pro parte-. 102, 541. the Hawaiian Dromia hirsutissima o f D an a T ype SPECIES. — Dromia australiensis Haswell, 1882 (1852) and Edmondson (1922) belong to by monotypy and subsequent designation (see Dromia dormia (Linnaeus, 1763). Holthuis 1962: 56; McLay 1993: 135, 136, and The differences that enable to distinguish below). Gender: feminine. Dromidia from Austrodromidia (Figs 1; 2), both SPECIES INCLUDED. — Dromia australiensis Haswell, sharing ventral uropods, include: 1) uropods nar­ 1882; Dromidiopsis edwardsi Rathbun, 1919; and row, oblique, deeply inserted but developed and Dromidiopsis tridentata Borradaile, 1903. Probably also Sphaerodromia lethrinusae Takeda & Kurata, 1976 only slightly visible dorsally in Dromidia (very included in Dromidiopsis by McLay (1993: 135, 139), small plates, may be indistinct in A u stro ­ see below. dromidia)-, 2) vestigial pleopods P13-P15 present Two species assigned to Dromidiopsis by McLay (absent inAustrodromidia ); 3) apertures of sper­ (1993, 2001), D. globosa (Lamarck, 1818) and D. dubia Lewinsohn, 1984, are herein excluded from mathecae ending together on slight prominenceDromidiopsis sensu nobis:D. globosa becomes the type between PI (ending wide apart between P2 in species of Lamarckdromia n. gen., whileD. dubia is Austrodromidia ); 4) male segment 6 with external included in Mclaydromia n. gen. The generic status of borders parallel (anteriorly concave and posteri­ Dromidiopsis richeri McLay, 2001 remains doubtful. orly expanded in Austrodromidia)-, 5) male telson D istribution . — Indo-West Pacific.

ZOOSYSTEMA • 2003 • 25 (1) 61 Guinot D. & Tavares M.

mxp3

cx1 cx1

cx2

cx3

a5 cx3 cx4

cx5

Fig. 6. — A, Dromidiopsis tridentata Borradaile, 1903, New Caledonia, LAGON, stn DW 554, ovigerous 9 12.7x 12 mm (MNHN-B 22550), thoracic sternum and spermathecae; B, D. australiensis (Haswell, 1882), Australia, Queensland, Cape Bedford, D. J. G. Griffin det., ô 15.4 x 14 mm (AM P16598), thoracic sternum with abdomen. Abbreviations:a5, a6, abdominal segm ents 5, 6;cx1- cx5, coxae of P1-P5;e4, episternite 4; mxp3, external maxilliped; o, female gonopore;pr, holding prominence;s, aperture of sper- matheca; t, telson (setae covering sternite 4 not shown);u, uropod; 4-8, sternites 4-8; 4/5-7/8, thoracic sternal sutures 4/5-718. Dotted line indicates difference in level. Scale bars: 2 mm.

D e s c r ip t io n gressively close to each other; apertures of sper­ Carapace longer than wide, strongly convex; dor­ mathecae ending between PI or just behind sal surface smooth, with regions not defined; them, together on central prominence. When branchial groove poorly defined. Anterolateral male abdomen is applied against ventral surface, margin entire or armed with several blunt (vari­ no part or only extreme anterior part of sternite able) teeth; only a small tooth behind branchial 4, with small episternite 4, remaining visible groove; posterolateral margin about as long as(setae of telson anteriorly covering sternite 4); anterolateral margin. Front appearing almostepisternite 5 not visible. entire, only bilobed or with two rounded very Male abdomen with segments 5-6 fused (sutures low pseudorostral teeth; rostrum strongly may be partly visible); telson long, ovate and deflexed, not or hardly visible dorsally; no supra­ rounded at tip. Male segment 5 with posterior orbital and exorbital teeth; suborbital tooth faint­ angles expanded; segment 6 with external borders ly indicated. Antenna: urinal article thick, slightly oblique and slightly thickened on edge. No ves­ wider than long, and with anterior part of beak tigial pleopods in males. Uropods as dorsal plates very narrow; basal article with exopod well-devel­ narrow and oriented vertically in males, as hori­ oped and internal corner acutely produced. zontal plates in females. Uropods involved in Mxp3: coxae not completely approximated. holding of abdomen, provided by strong dentate Thoracic sternite 3 very slightly visible. Thoracic crest on coxa of P2; in addition, presence of a sternite 4 narrow, with anterior margin bluntly granular prominence on PI coxa, in contact with truncate. Female sternal sutures 7/8 getting pro­ telson.

62 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

Chelipeds with epipod. P2 and P3 short and figs A, B) compared D. australiensis with D. tri­ stout, not knobbed or nodose; propodus of P2dentata and cleared up the synonymy of these and P3 without distal spine; inner margin of two species, many important morphological dactylus armed with spines. P4 and P5 reduced, characters of D. australiensis remained obscure. but P5 relatively long and, when extended for­ This situation led McLay (1993: 135, table 2; ward, reaching as far as outer orbital angle. 2001a: 79) to propose a compositeDromidiopsis. Propodus of P4 only slightly longer than wide. According to McLay (2001: 80, key), the follow­ P5 much longer than P4, merus and carpus elon­ ing seven species belong toDromidiopsis-. D. aus­ gated; propodus, clearly longer than wide, being traliensis, D. dubia, D. edwardsi, D. globosa, noticeably longer than that of P4. Subcheliform D. lethrinusae, D. tridentata and D. richeri. apparatus formed by one distal spine opposing Dromidiopsis sensu nobis is herein restricted and dactylus; on P5 one outer propodal spine and onenow includes onlyD. australiensis, D. edwardsi outer dactylus spine. and D. tridentata, and perhaps D. lethrinusae. Its P5 coxa with mobile penial tube. main diagnostic features are as follows: 1) front Male G2 with needle-like flagellum almost reach­ and orbital border entire or without marked ing anterior margin of sternite 4 and completely teeth; 2) male abdomen with segments 5-6 fused; covered by abdomen. 3) telson long; 4) male uropods showing as dorsal plates vertically oriented (Fig. 6B); and 5) female Carrying behaviour sternal sutures 7/8 long, and apertures of sper­ Sponges, compound or solitary ascidians. mathecae ending between PI or just behind them, together on central prominence (Fig. 6A). R em a r k s Dromidiopsis edwardsi, a new name given by Dromia australiensis (Haswell, 1882) is the type Rathbun (1919: 197) to the Indo-West Pacific species of Dromidiopsis (Borradaile 1900: 572) by crab identified as “Dromia caputmortuum” by monotypy (Holthuis 1962: 56). However, H. Milne Edwards (1837: 178, “from Indian McLay (1993: 135) indicated Dromia australien­ Ocean”) (nonCancer caputmortuum Linnaeus, sis as type species “by present designation”. On 1766), remains inDromidiopsis, sensu nobis. We account of the fact that “most records ofD. aus­ examined the material labeled Dromia caputmor­ traliensis in the literature are more likely to repre­ tuum by H. Milne Edwards (1837), two dry spec­ sen t D. tridentata th a n D. australiensis”, imens without locality and in poor condition including those of Borradaile (1900) and (MNHN-B 1 and 2). These specimens constitute Lewinsohn (1984: 100), McLay concluded that the syntypes of Dromidiopsis edwardsi. Because a “the name of the genus Dromidiopsis[. ], definition holotype has not been designated, the female of the genus, and description of the type species specimen MNHN-B 2, with the mention “Exp. all occured at different times”. Borradaile (1903a, b) de VAstrolabe”, is now selected as the lectotype, “gave a definition ofDromidiopsis which clearly and the remaining individual is the paralectotype. included D. tridentata but not necessarilyDromia McLay (1993: 137) remarks that “there is a need australiensis” (McLay 1993: 135, 136), hence the to clarify the validity of the records outside necessity to clearly designate here the type species Australia” for D. edwardsi (see Rathbun 1923a: ofDromidiopsis. 145). The records from Indian Ocean and Dromidiopsis australiensis is a poorly know n Indonesia may belong to another species, perhaps species, only briefly described and not figured by D. tridentata. The sperm of D. edwardsi has been Haswell (1882a: 755; 1882b: 139). Unfortu­described by Jamieson et al. (1993). nately, in recent years not enough attention hasSphaerodromia lethrinusae Takeda & Kurata, been paid to this species (McLay 1993: 136, 137 1976, described on basis of small specimens, was in key; 2001a: 79, 80 in key). Although assigned to Dromidiopsis by McLay (1993: 135, Lewinsohn (1984: 99-101, pi. 3, fig. A, pi. 4, 139) but left in Sphaerodromia Alcock, 1899 by

ZOOSYSTEMA • 2003 • 25 (1) 63 Guinot D. & Tavares M.

Chen & Haibao (2002: 76, fig. 29). The original sternal sutures 7/8 ending at level of the che­ immature female was shown with incompletely lipeds. The differences between Dromidiopsis and developed sternal sutures 7/8 (Takeda & Kurata Lauridromia are very few (see McLay 1993: 135, 1976: fig. 1.4), but mature females examined by 145, table 2): small size of individuals in McLay (1993: 140) has long sutures 7/8 which Dromidiopsis (large size in Lauridromia)-, aper­ end between chelipeds, and also other characters tures of spermathecae located together on central indicating that S. lethrinusae should be placed in prominence (Fig. 6A) (wide apart on long and Dromidiopsis. strong tubercles in typical Lauridromia). In Dromidiopsis globosa and D. dubia should no L. indica (Gray, 1831), however, the female ster­ longer be referred toDromidiopsis sensu nobis. nal sutures 7/8 end together at summit of two For D. globosa we establish Lamarckdromia coalescent tubercles; in these respects,L. indica n. gen. (Fig. 10), whose main characteristics are: seems to be close toDromidiopsis. 1) all abdominal segments free and presence of pleural parts; 2) uropods showing as completely concealed ventral plates; 3) apertures of sper­ Genus Epipedodromia André, 1932 mathecae ending between P2; and 4) uropods (Fig. 7A) not used to maintain the abdomen when folded. Platydromia Fulton & Grant, 1902a: 57 (pre-occupied Dromidiopsis dubia, also excluded from D rom i­ by Platydromia Brocchi, 1877, type species: diopsis sensu nobis, is herein attributed toMclay­ Platydromia depressa Brocchi, 1877, junior synonym of dromia n. gen., whose male uropods show as salient Dromidia spongiosa Stimpson, 1858, see under Platydromia spongiosa (Stimpson, 1858)). — Fulton & dorsal plates that are obliquely oriented (Fig. 12A, Grant 1906a: 11; 1906b: 20. — Hale 1925: 412; M . colini n. sp.). A male abdomen with all seg­ 1927: 105. — Griffin 1972: 52. ments free is another difference betweenM clay­ Epipedodromia André, 1932: 180 (replacement name, dromia n. gen. andDromidiopsis sensu nobis. with same type species: Epipedodromia thomsoni). — The generic status ofDromidiopsis richeri remains McLay 1993: 224, 225, table 8; 2001b: 2, 7, table 1. uncertain. McLay (2001a: 82, figs 1, 4A) indicat­ — McLay et al. 2001 pro parte-. 743. ed in his key that D. richeri belonged to the third T ype SPECIES. — Platydromia thomsoni Fulton & couplet, containingD. globosa. Dromidiopsis richeri Grant, 1902 by monotypy. Gender: feminine. is only known from two immature females (perhaps SPECIES INCLUDED. — Epipedodromia thomsoni parasitized), in which the apertures of spermathe­ (Fulton & Grant, 1902). cae lie between P3 (probably more forward in D istribution . — Australia. sexually mature females), the uropods (present in the smallest specimen, absent in the largest) show D e s c r ip t io n as dorsal plates, vertically oriented, and the Males not examined by the authors. abdominal holding is still effective.D. globosa, on Carapace wider than long, subpentagonal; dorsal the other hand, is the type species of Lamarck­ surface flattened, not membranous, with regions drom ia n. gen. (Fig. 10), characterized by not defined; branchial groove indistinct, not uropods showing as ventral plates and by antero­ marked by tooth. Prominent ledge present poste­ lateral border of carapace having a single tooth. rior to front, and limiting anterior margin of Dromidiopsis globosa and D. richeri cannot be carapace, front and cephalic parts at lower plane placed in the same genus. and showing as “false front”, marked by hairy Dromidiopsis sensu nobis andLauridromia ridge and divided into four parts by deep grooves. McLay, 1993, which consist of large-sized When viewed from above, carapace quadrate. species, share many features: shape of male Lateral margin formed by anterior part (slightly abdomen (with segments 5 and 6 partly fused), convex and corresponding to lateral border of uropods narrow and vertically oriented, holding “false front”), by straight medial part (anteriorly prominences on P2 and also on PI coxae, anddelimitated by notch), and by concave border

64 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

mxp3 1-3

cx1 cx1

4/5 cx2

4/5 cx2 5/a cx3

cx3

cx5

Fig. 7. — A, Epipedodromia thomsoni (Fulton & Grant, 1902), Australia, Victoria, Port Phillip, 9 9x11mm (BM 1884-39), thoracic ster­ num and spermathecae; B, Hemisphaerodromia monodus (Stebbing, 1918), Madagascar, Fort-Dauphin, Decary coll., det.H. abellana Barnard, 1954, 9 16x18 mm (MNFIN-B 12725), thoracic sternum and spermathecae. Abbreviations:cx1-cx5, coxae of P1-P5;e4, e5, episternites 4, 5; mxp3,external maxilliped; o, female gonopore;r, medial ridge; s, aperture of spermatheca; 1-3, sternites 1-3; 4-8,sternites 4-8;4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 2.5 mm.

corresponding to posterolateral margin. Frontat tip. No vestigial pleopods in males (to be partly visible dorsally, entire; rostrum truncate verified). Uropods absent (verified in ovigerous medially; pseudorostral teeth eave-like, turned females only). Holding of abdomen by raised upwards. Proepistome small but marked by knob on P2 coxae. Female abdomen well- raised ridge. Orbits deep; eyes small. No supra­ developed, first three segments positioned dor­ orbital and exorbital teeth; suborbital tooth form­ sally. ing thickening. Antenna: urinal article withChelipeds without epipod. P2 and P3 longer anterior part of beak very narrow and posterior than chelipeds, smooth. P4 and P5 reduced; P5 part shorter and rounded; basal article with exo­ longer, its curved merus almost as long as lateral pod and internal corner similarly developed and margin of carapace. Subcheliform apparatus produced. Mxp3: coxae separated by gap. formed by single distal propodal spine opposing Pterygostomial region soft. dactylus. Thoracic sternite 3 distinctly developed and com­ pletely visible. Thoracic sternite 4 forming raised Carrying behaviour piece, with medial part triangular and lateral “Unknown, but probably sponge” (McLay parts largely expanded. In females, posterior ster­ 2001b: 2, table 1). nites sharply and vertically tilted to form brood chamber; female sternal sutures 7/8 ending apart; R em a r k s apertures of spermathecae between PI and P2, at Epipedodromia thomsoni has the front and cepha­ level of episternite 4, and beneath raised medial lic parts being at lower plane, that provides a ridge (Fig. 7A). somewhat similar appearance than in Male abdomen with all segments free, acutely Desmodromia and Homalodromia and also in triangular in shape but telson rounded, obtuse Hypoconcha, see under Homalodromia.

ZOOSYSTEMA • 2003 • 25 (1) 63 Guinot D. & Tavares M.

4 Fultodromia McLay, 1993: 124, 162, table 3.

T ype species. — Dromia nodipes Guérin-Méneville, 1832 by original designation (McLay 1993: 162). cx1 (.Dromia nodipes Lamarck, 1818: 264 is a nomen nudum). Gender: feminine. A specimen ofDromia nodipes (female 22.5 X 23 mm, MNHN-B 15, rehydrated and now in alcohol) is regarded as the presumed type and selected here as the lectotype. It is not accompanied by any original label cx2 indicating the country of origin. This agrees to the question mark in the caption of the figure by Guérin- Méneville (1832: 11) and in the text of H. Milne a6 Edwards (1837: 170). The mention “Cap de Bonne- Espérance” in the MNHN inventory register most probably results from a mistake of a subsequent tran­ scription: it is perhaps useless to assign the reference for Port Esperance or Esperance Bay in South cx3 Australia (McLay 1993: 162). Therefore, the origin of the lectotype remains unknown. However, Fultodromia nodipes is most probably an Australian species, where it has been found by Baker (1907: 180, pi. 25, fig. 1, as Cryptodromia depressa), Hale (1927: 112, fig. 110, as Petalomera depressa) and Rathbun Fig. 8. — Fultodromia spinifera (Montgomery, 1931), Moluccas, (1923a: 154, as P. depressa). Pele, stn AN 1/1, S 17.5x 17.5 mm, abdomen (after Bouchard S p e c ie s i n c l u d e d . — Dromia nodipes G uérin- 2000: fig. 18E). Abbreviations:a6, abdominal segment 6;cx1-cx3, coxae of P1-P3; pr, holding prominence;t, telson;u, uropod; Méneville, 1832 (senior synonym ofCryptodromia 4, sternite 4. Scale bar: 1 mm. tumida Stimpson, 1858;Petalomera depressa Baker, 1907, and Dromidiopsis michaelseni Balss, 1935); Cryptodromia tumida var. spinifera Montgomery, The examination of male ofEpipedodromia thom­ 1931. soni will show whether the uropods are missing D istribution . — Australia. (McLay 1993: 225, table 8; 2001b: 2, table 1), or are simply reduced to small ventral plates. D e s c r i p t i o n The species is characterized by the development Carapace almost as long as wide, convex; dorsal of a brood chamber and by direct development. surface with regions not well-defined; branchial As Hale (1925: 412), we found very few large groove indistinct but a tooth just behind it. eggs in the pouch. Anterolateral margin joining exorbital angle and armed with several developed blunt teeth; pos­ terolateral margins nearly straight. Front with Genus Fultodromia McLay, 1993 deflexed rostral tooth and two prominent (Fig. 8) pseudorostral teeth; supraorbital and exorbital teeth prominent; suborbital tooth may be devel­ Dromia — Guérin-Méneville 1832: pi. 14, fig. 1. — oped. Antenna: urinal article with anterior part of H. Milne Edwards 1837 pro parte\ 170, 177. (Non DromiaW e ber, 1795). beak longer and more acute than posterior ones; basal article with exopod very long and thickened Cryptodromia — Stimpson 1858pro parte'. 255; 1907 pro parte. 172. — Baker 1907: 180. — Ihle 1913pro and with internal corner produced, both enclos­ parte\ 32. — Montgomery 1931: 413. (Non ing two following articles. Mxp3: coxae closely Cryptodromia Stimpson, 1858). approximated. Petalomera — Rathbun 1923a pro parte. 154. — Hale 1927 Thoracic sternite 3 not visible. Male thoracic proparte\ I l l , 112. (Non Petalomera Stimpson, 1858). sternite 4 with anterior margin bluntly triangular; Dromidiopsis — Balss 1935: 113. (Non Dromidiopsis episternites 4 and 5 narrow. Female sternal Borradaile, 1900). sutures 7/8 reaching forward to between coxae of

66 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

PI; apertures of spermathecae apart, but not sep­ T ype SPECIES. — Cryptodromia monodus Stebbing, arated by wide space. When male abdomen is 1918 by monotypy (senior synonym ofH em i- sphaerodromia abellana Barnard, 1954). Gender: femi­ applied against ventral surface, only a small part nine. of sternite 4 exposed; episternite 4 not visible or SPECIES INCLUDED. — Hemisphaerodromia monodus only a minute part discernible; episternite 5 not (Stebbing, 1918). visible. Male abdomen with all segments free, wide, and D istribution . — Indian Ocean. with characteristic blunt but developed promi­ nences in latero-posterior angles; telson with baseD e s c r i p t i o n very enlarged and may be markedly concave at Carapace wider than long, rounded/pentagonal, tip. Male segment 6 with external borders thick­ strongly convex; dorsal surface smooth, with ened on anterior half. No vestigial pleopods inregions not well-defined; branchial groove dis­ males. Male uropods showing as much salient tinct. Anterolateral margin with only blunt small and mobile dorsal plates, with petaloid expan­ teeth; posterolateral margins straight, with blunt sions similar to those of posterior angles oftooth. Front entire and continuous to orbital abdominal segments (Fig. 8). Holding ofmargin, a small rostral tooth and two eave-like abdomen consisting of a strong serrated promi­pseudorostral teeth; no suborbital nor exorbital nence on P2 coxa, far from uropods. teeth. Antenna: urinal article developed, with Chelipeds with epipod. All pereopods short and only anterior part of beak acute, posterior part thick. P2 and P3 nodular; propodus without dis­ being wide; basal article with exopod well devel­ tal spine; inner margin of dactylus with spines. oped and internal corner thickly produced, both P4 and P5 reduced, with terminal apparatus enclosing two following articles. Mxp3: coxae formed by up to two distal propodal spines closely approximated. opposing dactylus; two or three other spines onThoracic sternite 3 not visible. Male thoracic outer propodal margin. sternite 4 with anterior margin truncate. Female Male P5 coxa with long mobile penial tube. sternal sutures 7/8 ending apart behind P2; aper­ tures of spermathecae wide apart on tubercule, at Carrying behaviour level of episternite 5 (Fig. 7B). When male Sponges, compound ascidians. abdomen is applied against ventral surface, only anterior part of sternite 4 and narrow episternite R em a r k s 4 exposed. The female sternal sutures 7/8 end wide apart “in Male abdomen with all segments free, wide; tel­ a small mound” inFultodromia nodipes, and “in a son with base enlarged and bluntly rounded at transverse ridge” in F. spinifera (Montgomery tip. Male segment 6 with external borders deeply 1931: 414, pi. 29, fig. 3a). hollowed and much thickened in anterior part. No vestigial pleopods in males. Male uropods showing as markedly salient and mobile dorsal Genus Hemisphaerodromia Barnard, 1954 plates. Uropods involved in holding of abdomen, (Fig. 7B) which is particularly efficient, being provided with whole base of P2 coxa which bears a serrated Cryptodromia — Stebbing 1918: 56. — Barnard 1950 pro parte\ 307, 328. (Non Cryptodromia Stimpson, salient ridge, tightly encircled by depression on 1858). border of segment 6. Female uropods visible dor­ HemisphaerodromiaV>a.’C'aa.ïA, 1954: 100.— Lewinsohn sally. 1979: 10; 1984: 117. — McLay 1993: 124, 159, Chelipeds with an epipod. All pereopods short table 3- — Guinot & Bouchard 1998: 624, 628. and stout. P2 and P3 lobed; propodus without Petalomera — Kensley 1970 pro parte: 110. (Non distal spine; inner margin of dactylus with few Petalomera Stimpson, 1858). tiny spines. P4 and P5 reduced, with terminal

ZOOSYSTEMA • 2003 • 25 (1) 67 Guinot D. & Tavares M.

apparatus formed by one distal propodal spine resemblance and possible confusion withHomolo­ opposing dactylus; only one another very small dromia A. Milne Edwards, 1880 (Homolodromiidae) was unnecessary (ICZN 1999: article 56.2). spine on outer propodal margin. Male P5 coxa with long mobile penial tube. SPECIES INCLUDED. — Homalodromia coppingeri Miers, 1884, and perhaps another species,Homa­ lodromia unidentata (Ihle, 1913) (see Takeda 1977: Carrying behaviour 73; McLay 1993:227). Compound ascidians. D istribution . — Indo-West Pacific.

R e m a r k s As pointed out by Barnard (1954) and McLay D e s c r ip t io n (1993), Hemisphaerodromia resembles Sphaero­ Carapace longer than wide, whole anterior part dromia only by the shape of carapace, particularly strongly deflected; rest of dorsal surface flattened, the front. Nevertheless, Hemisphaerodromia is a with regions not defined; branchial groove dis­ typical dromiine, with salient dorsal uropods, tinct, may be sometimes marked by tooth. male P5 coxa bearing a long penial tube, female Subhepatic area inflated. Eyes and cephalic sternal sutures 7/8 ending behind P2 and aper­ appendages not visible dorsally. Anterolateral tures of spermathecae rather far from female margin convex, may be sometimes with single gonopores on P3 (Fig. 7B). small tooth oriented ventrally; posterolateral mar­ Fultodromia (Fig. 8) and Hemisphaerodromia are gins convergent. Front particular, at lower plane close and characterized by uropods showing as and appearing quadridentate in counting salient and mobile dorsal plates. The uropod and pseudorostral and supraorbital teeth; rostrum the coxal prominence on P2 are near each other markedly deflexed, without tooth; each promi­ in Hemisphaerodromia (Guinot & Bouchard nent acute pseudorostral tooth fused with simi­ 1998: fig. 3C, D), not inFultodromia (Bouchard larly shaped supraorbital tooth, together forming 2000: 82, figs 18E, 20D); the abdominal seg­ broad, concave eave. Suborbital tooth acute, ment 6 is markedly modified on both. In long, visible dorsally; exorbital tooth marked. Hemisphaerodromia (Fig. 7B) the apertures of Proepistome small but marked by raised ridge. spermathecae are located at level of episternites 5 Antenna: basal article much longer than wide, between P2, while in Fultodromia they reach for­ with enlarged exopod and internal corner similar­ ward between coxae of P1. The terminal carrying ly strongly produced; following articles well apparatus on P4 and P5 also distinguishes the developed. Mxp3: coxae closely approximated. two genera. Pterygostomial region soft. Thoracic sternite 3 not visible dorsally. Thoracic sternite 4 with anterior margin bluntly truncate, Genus Homalodromia Miers, 1884 in contact with mxp3 coxae. When male (Fig. 9) abdomen is applied against ventral surface, ante­ rior part of sternite 4 and epsisternite 4 remain­ Homalodromia Miers, 1884: 553. — McLay 1993: ing visible. In females, posterior sternites 125, 225, table 8; 2001b: 2, 7, table 1. — McLay et al. 2001 pro parte: 740, table 3. obliquely tilted; female sternal sutures 7/8 ending apart, just at level of PI; apertures of spermathe­ Pseudodromia — Alcock 1900pro parte\ 149. (Non Pseudodromia Stimpson, 1858). cae at each extremity of tubular curved promi­ nence forming bridge between PI coxae Lasiodromia Alcock, 1901: 56 (type species: Homalo­ (Fig. 9B). dromia coppingeri Miers, 1884). — Ihle 1913: 51. Male abdomen with all segments free, triangular T y p e SPECIES. — Homalodromia coppingeri M iers, in shape but with telson very long and obtuse at 1884 by monotypy. The replacement nameLasiodromia created by Alcock tip. No vestigial pleopods in males. Male uropods (1901) forHomalodromia Miers, 1884, because of the as elongated and mobile dorsal plates, vertically

68 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

cx1 sp mxp3

cx2

cx1 / ^ I cx3

cx2

cx4

Fig. 9. — A, Homalodromia coppingeri Miers, 1884, Seychelles, S 5.2 x 5.3 mm (MNHN-B 26130), thoracic sternum with abdomen; B, Homalodromia? coppingeri Miers, 1894, New Caledonia, LAGON, stn 556, 9 7.8x 9 mm (MNHN-B 22528), thoracic sternum and spermathecae; note apertures of spermathecae at level of P1, concealed by sperm plug, and vestigial holding prominence on P2 coxa of female. Abbreviations:a6, abdominal segment 6;b, tubular bridge between apertures of spermathecae; cx1-cx4,coxae of P1-P4; e4, episternite 4; mxp3,external maxilliped; o, female gonopore;pr, holding prominence;sp, sperm plug; t, telson;u, uropod; V, vestige of the holding prominence in the female;4-8, sternites 4-8;4/5-778, thoracic sternal sutures4/5-718. Scale bar: 1 mm. oriented, involved in holding of abdomen;assertion thatHomalodromia is most closely relat­ uropods remaining rather developed in females. ed to Epipedodromia (McLay 1993: 225), several Abdominal holding provided by particularly characters of the ventral surface of body separate high, cupuliform and denticulated prominence the two genera, notably: 1) sternite 3 not visible on coxa of P2 (Fig. 9A), which remains as a in Homalodromia (Fig. 9) (well-developed and vestige in mature females (Fig. 9B); on PI coxa, a completely visible in Epipedodromia , Fig. 7A); small tuberculate prominence. 2) uropods as elongated dorsal plates vertically Chelipeds without epipod, slighdy more massive oriented, constituting a full-lock system with than P2 and P3, none of these verrucose or dilat­ uropods completely involved in abdominal hold­ ed. P2 and P3 propodus without distal spine, and ing (see Guinot & Bouchard 1998: fig. 3B) inner margin of dactylus armed with several small (uropods absent, at least in ovigerous females, in spines. P4 and P5 reduced but unequal; P5 Epipedodromia ; their absence is to be verified in almost as long as P2, merus not so long as lateralmales); 3) apertures of spermathecae apart, each margin of carapace, however. Subcheliform appa­ located at the extremity of tubular prominence ratus formed by single distal propodal spine forming a bridge just behind PI coxae (Fig. 9B) opposing the dactylus. (apart, beneath raised medial thickening between Male P5 coxa with mobile penial tube. PI coxae in Epipedodromia , Fig. 7A); and 4) female abdomen normally widened (largely Carrying behaviour expanded and forming a brood chamber in Sponges. Epipedodromia ). Homalodromia resembles Dromidiopsis sensu R e m a r k s nobis (Fig. 6) by the shape of their male Despite an overall resemblance of the carapace abdomen, presence of uropods shaped as elongat­ (in particular the front), which permitted the ed dorsal plates, that are vertically oriented, and

ZOOSYSTEMA • 2003 • 25 (1) 69 Guinot D. & Tavares M.

abdominal holding provided by crest on P2 footnote. — Borradaile 1900: 571. (NonDromidia coxae. It differs from it by several features: Stimpson, 1858). 1) male abdomen with all segments free (seg­ Dromidiopsis — Rathbun 1923a: 146. — Hale 1927: ments 5-6 fused inDromidiopsis); and 2) aper­ 110; 1941: 281. — Griffin 1972: 53 Dromiopsis, ( sic). — Forest 1974pro parte. 103. — McLay 1993 pro tures of spermathecae apart and located at each parte'. 120, 135, 137; 2001a pro parte'. 79, 80. — extremity of tubular bridge just behind PI coxae M cLay et al. 2001 pro parte'. 733, 742. (Non (ending between PI, together on central promi­Dromidiopsis Borradaile, 1900 sensu nobis). nence, inDromidiopsis). T ype SPECIES. — Dromia globosa Lamarck, 1818 by Several dromiid genera share with Homalodromia present designation. the ventral location of eyes and cephalic ap­ SPECIES INCLUDED. — Dromia globosa Lamarck, 1818 pendages: Epipedodromia, Desmodromia and (senior synonym ofDromidia excavata Stimpson, Hypoconcha. McLay (2001b: 2, table 1) compared 1858). the first three genera, but the morphology of tho­ A damaged, dry specimen of a male, identified as Dromia globosa and without locality, deposited in the racic sternum was not taken into account.Historical Reference Collection (MNHN-B 22033), Differences include: 1) the anterior sternites 3 and bearing the label “it is probably the material studied by 4 (in Epipedodromia large sternite 3 present, stern­ H. Milne Edwards (1837: 177) and sent to M. de Man (1888: 396, footnote, pi. 18, fig. 1)”, is to be ite 4 raised and triangular, at least in females, considered the type of the species and selected here as Fig. 7A; sternite 3 not exposed and sternite 4 the lectotype. bluntly truncate inHomalodromia, Fig. 9); and ETYMOLOGY. — The genus Lamarckdromia n. gen. is 2) the shape of uropods (narrow dorsal plates ver­ dedicated to the eminent French naturalist Jean- tically oriented inHomalodromia, Fig. 9A, said to Baptiste Pierre de Monet, chevalier de Lamarck (1744- be absent inEpipedodromia, described as dorsal in 1829), who described several dromiid species. Gender: feminine. immature females of Desmodromia). In addition to the features of carapace and last pairs of pere- D istribution . — Australia. opods, Hypoconcha is characterized by a peculiar thoracic sternum (Fig. 19A), the presence of male D e s c r i p t i o n vestigial pleopods (Fig. 19B), the uropods show­ Carapace about as long as wide, convex; dorsal ing only as minute ventral lobes (Fig. 19B, C), a surface smooth, with regions not defined; sub- male abdomen that is short and usually flexed at hepatic region hollowed; branchial groove right angles in the middle, sternal female sutures marked and deeply notching external border. 7/8 which are relatively short and located on tilted Anterolateral margin not joining orbit but reach­ surface of posterior part of sternum (Fig. 19A), ing middle of pterygostomial border, and armed and by the presence of an epipod on PI. All of with only one blunt tooth; no tooth behind level these differences support the inclusion of of branchial groove; posterolateral margin short. Homalodromia, Epipedodromia and Desmodromia Front narrow, appearing tridentate, with strongly in the Dromiinae n. status, and the separation ofdeflexed rostrum and developed pseudorostral Hypoconcha in the Hypoconchinae n. subfam. teeth; supraorbital tooth present; no suborbital and exorbital teeth. Orbits deep; eyes small. Antenna: urinal article with anterior part of beak Genus Lamarckdromia n. gen. raised; basal article with exopod developed and (Fig. 10) internal corner produced. Anterior margin of buccal frame formed by raised median wall and Dromia — Lamarck 1818pro parte'. 264. — H. Milne Edwards 1837 pro parte'. 177. — Haswell 1882bpro two acute lateral teeth. Mxp3: coxae separated by parte'. 140. — Henderson 1888pro parte'. 3. — Ihle small gap. 1913 pro parte: 89. (NonD rom ia^!eber, 1795). Thoracic sternite 3 remaining exposed and visible Dromidia — Stimpson 1858pro parte'. 225, 239. — dorsally, specially in males (sternites 1-2 at lower Henderson 1888: 5. — de Man 1888pro parte'. 396, plane) (Fig. 10A, D). Thoracic sternite 4 narrow,

70 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

cx1

e4 cx2

1-2 mxp3

cx1

cx2

cx3

a3 cx4

Fig. 10. — Lamarckdromia globosa (Lamarck, 1818) n. comb., New South Wales, Port Jackson, det.Dromidia excavata Stimpson, McLay redet. Dromidiopsis globosa (MNHN-B 22041); A, 9 25 x 25 mm, thoracic sternum and spermathecae; B-D, S 24 x 25 mm, extremity of abdomen in dorsal view, abdomen in ventral view, and thoracic sternum without abdomen. Abbreviations:a3, a6, abdominal segm ents 3, 6; cx1-cx5,coxae of P1-P5; e4, e5, episternites 4, 5; mxp3, external maxilliped; o, female gonopore;pr, holding prominence;pt, penial tube; s, aperture of spermatheca; t, telson;u, uropod; 1-2, sternites 1-2; 3-8,sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 2.5 mm.

ZOOSYSTEMA • 2003 • 25 (1) 71 Guinot D. & Tavares M.

with triangular anterior margin. Female sternal segments free (segments 5-6 fused inD rom i­ sutures 7/8 long, with apertures of spermathecae diopsis)-, 4) male telson is short and wide, bluntly ending between P2, together on central promi­rounded at tip (longer than wide, and more trian­ nence (Fig. 10A). When male abdomen is gular in Dromidiopsis)-, 5) abdominal holding is applied against ventral surface, extreme anterior provided by tuberculate sharp prominence on P2, part of sternite 4 and small part of episternite without involvement of the uropods (presence of 4 remaining visible. a dentate crest on P2 and uropods involved in Male abdomen with all segments free; presence of abdominal holding inDromidiopsis)-, and 6) there extended pleural parts; telson much broader than are differences in overall shape of carapace. long, rounded at tip. Male segment 6 slightly Since the uropods show as ventral plates in constricted in posterior part and rest of external Dromidiopsis globosa, the species is excluded from borders subparallel. Vestigial pleopods absent in the existing dromiine genera in which uropods males. Uropods showing as ventral rounded/ show as dorsal plates (see Table 1). The uropods ovate plates, well inserted, relatively developed have been described as “small”, “concealed” or but completely concealed (Fig. 10B, C). Uropods “absent” in about 10 dromiine genera. Despite not involved in holding of abdomen. Abdominalthe vagueness and imprécisions in some descrip­ holding provided by very sharp tuberculate tions, D. globosa differs from those 10 genera in prominence on P2 coxa, overhanging base of seg­the details of the carapace, pereopods, thoracic m ent 6 (Fig. 10B, D). sternum, and abdomen. Alone, the features of the Chelipeds not knobbed. P2 and P3 not knobbed carapace would be sufficient to place D. globosa nor nodose; propodus of P2 and P3 without dis­ in its own genus,Lamarckdromia n. gen. tal spine; inner margin of dactylus armed with Additional characters mentioned as follows do spines. P4 and P5 reduced, but P5 much longer not support the inclusion ofD. globosa in the than P4 and, when extended forward, reaching as following genera: 1)Ascidiophilus Richters, 1880; far as the anterolateral tooth; merus and carpus 2) Austrodromidia McLay, 1993 sensu nobis; being noticeably much longer than that of P4. P4 3) Barnardromia McLay, 1993; 4) D rom idia very short and stout, propodus wider than long. Stimpson, 1858 sensu nobis; Epipedodromia 5) Subcheliform apparatus of P4 and P5 formed by André, 1932; 6) Eudrom idia Barnard, 1947; one distal spine opposing the curved dactylus; 7) Exodromidia Stebbing, 1905; 8)Haledromia three developed and subequal outer propodal McLay, 1993; 9) Platydromia Brocchi, 1877; spines on P4; on P5, only one long outer propo­10) Pseudodromia Stimpson, 1858; 11)Speodromia dal spine. Barnard, 1947; and 12) Tunedromia McLay, Male P5 coxa with mobile penial tube (Fig. 10D). 1993. 1) Ascidiophilus, (type species: Ascidiophilus Carrying behaviour caphyraeformis Richters, 1880 by monotypy) and Sponges, compound ascidians. 10) Pseudodromia (type species: Pseudodromia latens Stimpson, 1858 by original designation). R e m a r k s In Dromidiopsis globosa (Fig. 10): thoracic ster­ Dromidiopsis globosa (Lamarck, 1818) (see McLay num “normal” (very narrow, specially sternite 4 1993: 135, 137), from Australia, is excluded showing as small piece inAscidiophilus and from Dromidiopsis sensu nobis (Fig. 6) since: Pseudodromia)-, male telson wider than long and 1) uropods occur as completely concealed ventral rounded at tip (longer than wide and tip pointed plates (dorsal plates vertically oriented and well in Ascidiophilus and Pseudodromia)-, holding of visible dorsally in Dromidiopsis ); 2) females abdomen present, provided by sharp prominence sutures 7/8 only reach bases of P2 (see Forest on P2 coxa (absent inAscidiophilus and 1974: pi. 6, fig. 4 as D. excavata) (reaching bases Pseudodromia)-, when extended forward, P5 of PI inDromidiopsis ); 3) male abdomen has all reaching anterolateral tooth (reaching as far as

72 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

orbital angle inAscidiophilus, or even more elon­ D. globosa-, uropods present, showing as ventral gated in Pseudodromia); and P4 and P5 propodus plates in both sexes (absent in ovigerous females with distal spine opposing dactylus (no distal in Epipedodromia-, absence to be verified in propodal spine opposing dactylus inAscidiophilus males); sternite 3 discernible (Fig. 10A, D) (ster­ and Pseudodromia). Another main difference is nite 3 developed and completely visible in that the uropods show as ventral plates in Epipedodromia, Fig. 7A); in females, posterior Lamarckdromia n. gen. such as inPseudodromia, sternites gently tilted (sharply and vertically tilt­ while they are completely absent in both sexes in ed, with formation of brood chamber in Ascidiophilus. Epipedodromia); and apertures of spermathecae 2) Austrodromidia sensu nobis (Figs 1; 2). In between P2, together on central prominence D. globosa-, male abdominal segment 6 with exter­ (apart, beneath thick medial bridge just behind nal borders subparallel (deeply hollowed anterior­ P I, in Epipedodromia, Fig. 7A). ly and expanded posteriorly in Austrodromidia); 6) Eudromidia (type species: Eudromia frontalis telson with its base not particularly enlarged Henderson, 1888 by monotypy). InD. globosa-. (enlarged in Austrodromidia); and female sternal apertures of spermathecae ending between P2, sutures 7/8 ending together on central promi­together on central prominence (between PI, on nence between P2 (ending wide apart at level of a tubercle in Eudromidia)-, and P4 and P5 P2 in Austrodromidia). reduced, but P5 reaching anterolateral tooth (P4 3) Barnardromia (type species: Cryptodromia hir­ and P5 smaller, P5 being filiform inEudromidia, sutimana Kensley & Buxton, 1984 by original at least in E. frontalis). In E u drom idia the designation), in which the sternal parts and uropods, which have not been figured, are poorly abdomen are not figured (Kensley & Buxton known; McLay (1993: 179) refers to the uropods 1984: 193, fig. 4). In D. globosa-, apertures of as “very small, concealed”. spermathecae ending together on central promi­7) Exodromidia (type species: Dromidia spinosa nence between P2 (behind bases of PI in Studer, 1883 by monotypy). In D. globosa-, male Barnardromia)-, and P4 and P5 propodus with telson rounded at tip (ending in spine in one distal spine opposing dactylus and three Exodromidia)-, vestigial pleopods absent in males outer propodal spines on P4 and one outer(P13-P15 present inExodromidia spinosa (Studer, propodal spine on P5 (dactyli of P4 and P5 1883), or only P15 present inE. bicornis (Studer, opposed by single distal propodal spine in 1883) and E. spinosissima (Kensley, 1977)); tho­ Barnardromia, in McLay 1993: 180, table 5). racic sternite 3 dorsally visible, only by very small 4) D rom idia sensu nobis. InD. globosa-, male part, specially in males (largely exposed in uropods showing as ventral plates totally Exodromidia)-, and male sternite 4 not hollowed concealed (Fig. 10B, C) (ventral plates slightly medially, almost completely covered by visible dorsally in Dromidia, Fig. 5B, C); vestigial abdomen, and with triangular tip (deeply hol­ pleopods absent (Fig. IOC) (P13-P15 present, in lowed and remaining always partly visible when Dromidia, Fig. 5C); male telson rounded at tip abdomen folded, and with truncate tip, in (Fig. 10B, C) (ending as sharp spine inDromidia, Exodromidia). The shape of ventral uropods, Fig. 5B, C); abdominal holding provided by partly or totally concealed, also distinguishes the tuberculate, sharp prominence on P2 coxa two genera: showing as rounded/ovate plates, not (Fig. 10B, D) (a strong spine on P2 coxa, directed really movable in Lamarckdromia n. gen. backwards and partly overhanging abdomen in (Fig. 10B, C), as narrow and elongated plates in Dromidia, Fig. 5A, B); and sternite 4 bluntly Exodromidia. triangular (Fig. 10A, D) (rounded at tip in 8) Elaledromia (type species: Dromia bicavernosa Dromidia, Fig. 5A, B). Zietz, 1887 by monotypy). InD. globosa-, aper­ 5) Epipedodromia André, 1932 (type species: tures of spermathecae ending between P2, Platydromia thomsoni Fulton & Grant, 1902). In together on central prominence (Fig. 10A), while

ZOOSYSTEMA • 2003 • 25 (1) 73 Guinot D. & Tavares M.

in Haledromia the apertures end as far forward as shaggy coat of setae, and the deflexed front por­ PI, on large tubercle. tion of carapace is concealed by a transverse 9) Platydromia (type species: Dromidia spongiosa fringe of longer setae, giving it a unique appear­ Stimpson, 1858), which has also ventral andance. Epipedodromia thomsoni (Fulton & Grant, concealed uropods. InD. globosa: sternite 3 with 1902) has a similar hairy ridge, forming a ledge a small part visible, particularly in males that limits the anterior margin of the flattened (Fig. 10A, D) (largely exposed in both sexes of carapace. Platydromia, Figs 15A; 16); sternite 4 triangular Lamarckdromia globosa n. comb, is only known (Fig. 10A, D) (wide and with convex lateral mar­ from Australia. It has direct development and gins in both sexes ofPlatydromia)-, male telson broods its young (Hale 1941: 281, figs 15, 16, as rounded at tip (Fig. 10B, C) (ending in acute tip Dromidiopsis excavata-, McLay et al. 2001: 742, as in Platydromia, Fig. 15B, C); and apertures of Dromidiopsis globosa). spermathecae ending together, on central promi­ nence between P2, Fig. 10A (ending together, on slight prominence between chelipeds in Genus Lewindromia n. gen. Platydromia, Fig. 16). (Fig. 11) 11) Speodromia (type species: Dynomene platy­ Dromia — Riippell 1830 pro parte-. 16. — H. Milne arthrodes Stebbing, 1905 by monotypy). In Edwards 1837 pro parte-. 170. — Alcock 1900pro D. globosa-, female uropods not visible dorsally (vis­ parte-. 139. (Non Dromia Weber, 1795). ible beneath setae inSpeodromia, after Stebbing Dromidia — Borradaile 1903a pro parte-. 299. — Ihle 1905: 60, pi. 17, asDynomene platyarthrodes,-, ac­ 1913 pro parte-. 31. — Edmondson 1922: 34. — cording to McLay 1993: 182, table 5, the uropods Barnard 1950pro parte-. 319, 323. — Garth 1973: 316. — Lewinsohn 1977: 9; 1979pro parte-. 2. — are not visible in both sexes); apertures of sper­ Tirmizi & Kazmi 1991 pro parte-. 27. (Non Dromidia mathecae between P2 (between PI inSpeodromia)-, Stimpson, 1858). and P4 and P5 reduced, with P4 short and stout, Cryptodromiopsis — McLay 1993 pro parte-. 187, 192; P5 much longer than P4 and, when extended for­ 2001a pro parte-. 84. — McLay et al. 2001 pro parte-. ward, reaching as far as anterolateral tooth (P4 and 740, table 3- — Chen & Haibao 2002 pro parte-. 102, P5 dissimilar in size and shape inSpeodromia-, P4 541, 542. (NonCryptodromiopsis Borradaile, 1903). short, thick and three-sided; P5 much more slen­ Type SPECIES. — Cryptodromiopsis unidentata Riippell, der, specially last three articles). 1830 by present designation. Gender: feminine. 12) Tunedrom ia (type species: Petalom era ETYMOLOGY. — The genus Lewindromia n. gen. is yamashitai Takeda & Miyake, 1970, by original dedicated to Chanan Lewinsohn (1927-1983) in recognition for his contribution to the knowledge of designation). Tunedromia is now know n from the Indian Ocean dromiids. females and males (Takeda 2001). In D. globosa-, uropods showing as ventral plates, relatively SPECIES INCLUDED. — Cryptodromiopsis unidentata Riippell, 1830. developed but completely concealed (absent in Dromidia unidentata hawaiiensis Edmondson, 1922 both sexes of Tunedromia)-, and P5 propodus and Cryptodromia unilobata Campbell & Stephenson, with one distal spine opposing the dactylus and 1970 were synonymised with Dromidia unidentata by one single long outer propodal spine (multipleMcLay (1993: 192, 194). propodal spines opposing the dactylus and several DISTRIBUTION. — Indo-West Pacific: Red Sea and outer propodal spines inTunedromia). Indian Ocean, including sea mounts of the western Indian Ocean (see Zarenkov 1994), with a large exten­ Lamarckdromia globosa n. comb, has its sub- sion in the Pacific (Australia, Hawaii, Kermadec hepatic region deeply hollowed, which provided Islands, Easter Island, see Garth 1973: 316). the specific name ofexcavata given by Stimpson (1858: 77, asDromidia excavata Stimpson, 1858, D e s c r ip t io n junior synonym ofDromia globosa Lam arck, Carapace longer than wide, evenly convex; dorsal 1818). The body and legs are covered by a dense, surface with regions poorly defined; branchial

74 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

cx1 mxp3

cx1

cx2

¡4/5 cx2

5/6

A, B

cx1 cx2 4/5 e5

5/6 cx2 cx3

6/7 cx3

cx4

cx5

Fig. 11. — Lewindromia unidentata (Rüppell, 1830) n. comb.;A, B, Obock, Lewinsohn det.Dromidia unidentata,S about 30 mm width (MNHN B-6930), thoracic sternum, two front views, without abdomen and with abdomen;C, Red Sea, Aden and Obock, Nobili det. D. unidentata,9 24 x 24 mm (MNHN-B 6940), thoracic sternum and spermathecae. Abbreviations: a6, abdominal seg­ ment 6;cx1-cx5, coxae of P1-P5;e4, e5, episternites 4, 5 (in fact, episternite 4 not so visible when abdomen folded);mxp3, exter­ nal maxilliped; o, female gonopore;pr, holding prominence: blunt prominence on P1, carina ons, P2;aperture of spermatheca; t, telson;u, uropod; 4-8,sternites 4-8;4/5-7/8, thoracic sternal sutures415-7/8. Dotted line indicates difference in level. Scale bars: 2.5 mm.

ZOOSYSTEMA • 2003 • 25 (1) 75 Guinot D. & Tavares M.

groove marked, deeply notching external border. thick, as wide as long, with one distal spine Anterolateral margin not joining exorbital angle,opposing dactylus which is not markedly curved long, convex, entire, only with a very blunt toothand ends in long horny spine; two smaller outer behind level of branchial groove; posterolateral propodal spines. margins not noticeably convergent posteriorly.Male P5 coxa with mobile penial tube. Front narrow, with rostrum very small and blunt, Male G2 much longer than Gl, thin and long not visible dorsally, and two pseudorostral teeth; flagellum overreaching sterno-abdominal depres­ supraorbital, suborbital and exorbital teeth pres­ sion. ent. Antenna: urinal article noticeably developed and rather straight; basal article thick, with exo­ Carrying behaviour pod strongly developed and internal corner Individuals carry a wide range of camouflage slightly produced, much shorter than exopod; material (sponges, soft coral, compound or soli­ article 4 wide. Mxp3: coxae closely approximated. tary ascidians, actinians), and the cap is often Thoracic sternite 3 not visible. Sternite 4 very very large, so that the crab is “deeply embedded” narrow, forming acute plate, ending in sharp tip (McLay 2001a: 84). Lewindromia unidentata in both sexes. In females, sternites 7 and 8 tilted, n. comb, looks like “a hairy ball that fits tightly almost perpendicular in relation to precedent into its piece of camouflage” (McLay 2001a: 84). ones. Female sternal sutures 7/8 posteriorly wide See also Chen & Haibao 2002: pi. 5, fig. 2. apart, sharply close to each other at level of P3 where they are marked by thick ridge; apertures R em a rks of spermathecae ending together on slight promi­Dromia unidentata was transferred to Dromidia nence between P2 (Fig. 11C). No sternal parts by Kossmann (1880: 67) until McLay (1993: (very small episternite 4 may be discernible) 192) placed it in Cryptodromiopsis. The study of remaining visible when male abdomen applied Dromidia sensu nobis andCryptodromiopsis sensu against ventral surface. nobis, which are restricted to their type species, Male abdomen very long, without pleural parts, led us to conclude that D. unidentata does not reaching mxp3, completely covering narrow and belong to neither of these genera and that a new deep sterno-abdominal depression, and with all genus should be erected for it. That new genus is segments free; telson very long, with large base named herein Lewindromia n. gen. and regularly tapering, pointed at tip. Telson of We found it useful to compareLewindromia females semi-ovate. Male segment 6 with external n. gen. with the genera below; special reference is borders oblique and modified on edges. No ves­ made to the features other than those of the cara­ tigial pleopods in males. Uropods as dorsal plates, pace. vertically oriented in males (Fig. 11B), well- Lewindromia n. gen. is distinguished from developed in females. Uropods involved in hold­ Austrodromidia sensu nobis (Figs 1; 2) by the fol­ ing of abdomen. A very efficient abdominal lowing characters: 1) uropods showing as dorsal holding provided by long and prominent serru­ plates, vertically oriented (markedly reduced or lated carina on P2 coxa; presence of large blunt even obsolete ventral plates inAustrodromidia ); prominence on P 1 coxa. 2) male segment 6 with external borders oblique Chelipeds without epipod. PI, P2 and P3 not (external borders anteriorly hollowed and posteri­ knobbed nor nodose; propodus of P2 and P3 orly expanded in Austrodromidia ); and 3) sper­ without distal spine; inner margin of dactylus mathecae ending together on central prominence armed with spines. P4 and P5 reduced, P5 being between P2 (ending wide apart between P2 in much longer than P4 and, when extended for­ Austrodromidia ). ward, reaching about mid-length of anterior mar­ The differences between Lewindromia n. gen. gin of carapace. P5 coxa very developed in both and Cryptodromia (type species: C. coronata sexes. Propodus of P4 and P5 very short and Stimpson, 1858: 226, by original designation)

76 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

include: 1) apertures of spermathecae closely 2) pseudorostral teeth prominent, so that entire approximated (situated wide apart in front is strongly produced forward (lateral frontal Cryptodromia ; see the diagnosis by Stimpson lobes extremely low so that entire front is pro­ 1858: 225: “Fœminæ sterni sulci remoti, ad seg­ duced forward only slightly in Dromidiopsis)-, mentum pedum secundi paris tantum producti, 3) propodus of P4 and P5 subequal in size (of terminis in tuberculis”); 2) male telson much different size, that of P5 much longer in longer than wide (wider than long in Dromidiopsis)-, 4) when extended forward, P5 Cryptodromia); 3) male thoracic sternite 3 not reaching about mid-length of anterior margin of visible (visible dorsally in Cryptodromia); 4) male carapace (P5 much long, reaching as far as outer thoracic sternite 4 ending in acute tip (sternite 4 orbital angle inDromidiopsis)-, 5) female sternal anteriorly truncated inCryptodromia). sutures 7/8 sharply close to each other at level of Differences betweenLewindromia n. gen. and P3 where they are lined by thick ridge (separated Cryptodromiopsis sensu nobis (Fig. 4) include: wide apart, getting progressively close to each 1) thoracic sternite 1-3 concealed in males other inDromidiopsis)-, 6) apertures of spermathe­ (exposed in Cryptodromiopsis); 2) coxae of mxp3 cae at level of P2 (at level of PI or just behind placed close together (separated by distinct gap in them in Dromidiopsis)-, and 7) thoracic sternite 4 Cryptodromiopsis); 3) thoracic sternite 4 ending in ending in acute tip (sternite 4 truncated distally acute tip in males (sternite 4 not pointed in in Dromidiopsis). Cryptodromiopsis); 4) male telson much longer Lewindromia n. gen. andHomalodromia (Fig. 9), than wide (wider than long inCryptodromiopsis); both with uropods vertically oriented, differ from and 5) apertures of spermathecae at level of P2 (at each other, as follows: 1) apertures of sperma­ level of PI in Cryptodromiopsis). thecae together on slight prominence between P2 Lewindromia n. gen. is distinguished from (Fig. 11C) (apertures at each extremity of a tubu­ Dromidia sensu nobis (Fig. 5) by the following lar prominence forming bridge between P1 coxae characters: 1) male abdominal segment 6 with in Homalodromia); 2) sternite 4 forming acute external borders oblique and sinuous (external plate, ending in sharp tip (Fig. 1 IA, C) (bluntly borders subparallel in Dromidia); 2) male telson truncate at tip inHomalodromia)-, and 3) no ster­ much longer than wide, regularly tapering, and nal parts remaining visible when male abdomen the tip becoming very narrow (telson wider than applied against ventral surface (Fig. 11B); small long, ending by sharp spine inD ro m id ia ); episternite 4 hardly dicernible at lower plane or 3) male uropods as dorsal plates, vertically oriented, not visible (sternite 4 and episternite 4 remaining exposed and completely visible dorsally (almost visible in Homalodromia). totally concealed ventral plates inDromidia)-, Lewindromia n. gen. andLamarckdromia n. gen. 4) uropods involved in abdominal holding which share the thoracic sternite 4 showing as triangular consists of uropods fitting in front of serrulatedacute plate, but narrower inLewindromia n. gen. carina on P2 coxae; an additional strong, rounded The following characters readily distinguish prominence on P1 coxa (uropods not involved atLewindromia n. gen. fromLamarckdromia n. gen. all in abdominal holding; a strong spine directed (Fig. 10): 1) male abdominal segment 6 with backwards, partly overhanging abdomen in external borders oblique and sinuous (external Dromidia)-, and 5) thoracic sternite 4 tapering borders subparallel in Lewindromia n. gen.); and ending in acute tip (sternite 4 broad, ending 2) male telson much longer than wide, regularly in rounded tip inDromidia). tapering, the tip becoming narrow (telson wider Lewindromia n. gen. andDromidiopsis sensu than long, rounded at tip, inLewindromia nobis (Fig. 6), both with uropods oriented verti­n. gen.); 3) male uropods plates exposed, vertically cally, differ from each other as follows: 1) all oriented, involved in abdominal holding (show­ abdominal segments free (abdominal segments 5 ing as totally concealed ventral plates, not and 6 fused together in Dromidiopsis)-, involved, in Lewindromia)-, and 4) thoracic

ZOOSYSTEMA • 2003 • 25 (1) 77 Guinot D. & Tavares M.

sternite 4 forming acute plate, ending in sharp tip D e s c r ip t io n (with anterior margin triangular inLewindromia Carapace distinctly longer than wide, convex. n. gen.). Dorsal surface with regions poorly defined; Lewindromia n. gen. and Lauridromia (McLay branchial groove defined and posteriorly marked 1993: 145, table 2; see Guinot & Bouchard by blunt tooth. Anterolateral margin of carapace 1998, fig. 2A-C) share several characters: trian­ not joining exorbital angle and armed with two or gular shape of sternite 4, although narrower in three teeth. Posterolateral margin slightly shorter Lewindromia n. gen.; male abdomen completely than anterolateral margin and with blunt tooth just covering sterno-abdominal depression; vertically behind branchial groove. Front (Fig. 13) wide, ob­ oriented uropods; holding prominences on P2scurely tridentate: rostrum very small, directed and PI coxae. Lewindromia n. gen. differs from downwards, and not visible dorsally; two pseudo­ Lauridromia by: 1) all abdominal segments free rostral teeth, more or less developed, sometimes (segments 5 and 6 fused, at least in some eave-like. Supraorbital and suborbital teeth not extent, in Lauridromia ); 2) apertures of sper­ well marked; exorbital angle not produced. Antenna: mathecae grouped together on slight promi­ urinal article broader than long, with anterior part nence between P2 (Fig. 11C) (wide apart, at of beak small and downcurved; basal article with level of episternites 4, each at summit of tuber­ well-developed exopod; internal corner about as cle, in Lauridromia ); 3) epipod on cheliped long as exopod; antennal article 4 long and fitted in absent (present inLauridromia ); and 4) termi­ between. Mxp3: coxae closely approximated. nal apparatus of P5 consisting of only one dis­Thoracic sternite 3 partly visible at lower plane tal propodal spine opposing the dactylus, and (sternites 1-2 not exposed); sternite 4 narrow, two outer propodal spines small (two propodal with subparallel margins, anterior margin trun­ spines and several others well-developed spines cated (Fig. 12). Female sternal sutures 7/8 long, in Lauridromia). apertures of spermathecae placed apart, each on a Lewindromia n. gen. differs from all the above tubercle, between coxae of P2 (Fig. 12B). When genera by having an extremely long G2, which male abdomen folded against céphalothorax, ster­ overreaches the tip of telson when abdomen is nite 3 (partly) and large part of sternite 4 remain­ completely folded (G2 shorter, fitted inside ing visible; a minute part of episternite 4 exposed; sternoabdominal depression in the other genera).episternite 5 not visible. Male abdomen not completely covering sterno- abdominal depression. All abdominal segments Genus Mclaydromia n. gen. free in males and females. Male abdominal seg­ (Figs 12; 13) ment 6 abruptly constricted, edge markedly thickened. Telson rather long, with enlarged base ? Dromidiopsis — Lewinsohn 1984pro parte. 102. and rounded anterior margin. No vestigial Dromidiopsis — McLay 1993 pro parte\ 135, 138; pleopods in males. Uropods showing as strongly 2001a pro parte. 79, 80. (NonDromidiopsis Borradaile, 1900 sensu nobis). salient dorsal plates in males, that are obliquely oriented. Uropods markedly involved in abdomi­ Type SPECIES. — Mclaydromia colini n. gen., n. sp., by present designation. nal holding. Abdominal holding provided by curved, serrate, strong flange on P2 coxa, which SPECIES INCLUDED. — Mclaydromia colini n. gen., n. sp.; Dromidiopsis Lewinsohn, 1984. is in close contact with uropod and fits into exca­ vation on lateral edge of abdominal segment 6; ETYMOLOGY. — We dedicate the new genus Mclay­ additionally, on coxa of PI, few tiny tubercles dromia n. gen. to our colleague Colin L. McLay (University of Canterbury, Christchurch, New placed closely together, or one more distinctly Zealand), for greatly improving our view of the marked tubercle, without contact with telson. Dromiidae. Gender: feminine. Chelipeds with an epipod. In both sexes cutting DISTRIBUTION. — New Caledonia and Madagascar. edges of fixed finger and dactylus of two che-

78 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

1-2 -1-2 mxp3 mxp3

cx1

cx1 e4

cx2 cx2

cx3 cx3

cx4

Fig. 12. — Mclaydromia colini n. gen., n. sp.; A, LAGON, stn 111, 25 m,S 10.4 x 9.1 mm, paratype (MNHN-B 26282), thoracic ster­ num with abdomen; B, LAGON, stn 569, 62 m, ovigerous 9 9.4 x 8.6 mm, paratype (MNHN-B 26289), thoracic sternum and sper­ mathecae. Abbreviations: a6, abdominal segment 6;cx1-cx4, coxae of P1-P4; e4, episternite 4 (very small); mxp3, external maxilliped (coxae separated for clarity); o, female gonopore;pr, holding prominence;s, aperture of spermatheca; t, telson; u, uro- pod; 1-2 sternites 1-2 (normally not exposed);3, sternite 3; 4-8, sternites 4-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 2.5 mm.

lipeds armed halfway with strong molariform R em a r k s tooth (may be bifid), which is directed backwards In assigning with doubt his new speciesD. dubia to and followed by the usual interlocking distal teeth. Dromidiopsis, Lewinsohn (1984: 102, 104, fig. 2b, P2 and P3 short and stout, not knobbed nor c) stressed its peculiar features, particularly the nodose; propodus of P2 and P3 without distal fingers and dactyli of chelipeds. The presence of a spine; inner margin of dactylus armed with molariform tooth on two cutting edges in spines. P4 and P5 reduced, P5 longer than P4; Mclaydromia n. gen. appears unique amongst the propodus of P4 and P5 very short, subequal in Dromiidae (which is sometimes present in the size. Subcheliform apparatus formed by one small Dynomenidae) and may indicate a specialized distal spine opposing short and curved dactylus; feeding habit (McLay 1993: 139). A proximal tooth an outer propodal spine may be present on P5. may be present along the cutting edge of dactylus in Male P5 coxa with mobile penial tube. a few drom iid genera (as inEpigodromia McLay, Male G2 with a needle-like flagellum, long but 1993), but the conditionoí Mclaydromia n. gen., completely included in sterno-abdominal depres­ with two molariform teeth halfway on prehensile sion. margin of fixed finger and dactylus, is exceptional. Mclaydromia n. gen. can be readily distinguished Carrying behaviour from Dromidiopsis sensu nobis (Fig. 6) as follows: Sponges (McLay 1993: 139, under Dromidiopsis 1) abdominal segments free (abdominal segments dubia). 5 and 6 fused together inDromidiopsis );

ZOOSYSTEMA • 2003 • 25 (1) 79 Guinot D. & Tavares M.

2) propodus of P4 and P5 subequal in size (dis­ phaerodromia (apertures forward, between P2, in similar, much longer on P5, inDromidiopsis); Mclaydromia n. gen.); and 3) fingers of chelipeds 3) when extended forward, P5 barely overreach­ normally toothed along prehensile margin in ing last lateral tooth of carapace (much long, Hemisphaerodromia (molariform teeth on fixed reaching about outer orbital angle inD rom i­ finger and dactylus in Mclaydromia n. gen.). diopsis)-, 4) female sutures 7/8 wide apart, The following characters readily distinguish oblique, getting progressively close to each other Mclaydromia n. gen. fromLewindromia n. gen. as they run forward over thoracic sternites (get­ (Fig. 11), with vertically oriented dorsal uropods: ting much closed to each other at level of P3 and 1) sternite 4 with anterior margin truncated subparallel in Dromidiopsis)-, 5) apertures of (acutely produced in Lewindromia n. gen.); spermathecae at level of P2 and placed wide apart 2) male abdominal segment 6 abruptly constrict­ from each other, on tubercles (at level of PI or ed (with external borders sinuous, oblique, in just behind and lying close together, on central Lewindromia n. gen.); 3) male telson not longer prominence, inDromidiopsis)-, and 6) male than wide and rounded at tip (very long and uropods showing as dorsal plates that are oblique­ pointed inLewindromia n. gen.); and 4) propo­ ly oriented (vertically oriented in Dromidiopsis). dus of P4 and P5 not much different in size, a The following characters readily distinguish distal spine opposing the curved dactylus, and an Mclaydromia n. gen. fromCryptodromiopsis sensu outer propodal spine may be present on P5 (P5 nobis (Fig. 4), which also has obliquely oriented much longer than P4 and, when extended for­ dorsal uropods: 1) male segment 6 with external ward, reaching about mid-length of anterior mar­ borders deeply hollowed and thickened (sub­ gin of carapace, a propodal distal spine opposing parallel on anterior half inCryptodromiopsis)-, the dactylus which is not strongly curved, and 2) apertures of spermathecae apart, each on a two outer propodal spines inLewindromia n. gen.). tubercle, at level of P2 (ending together on slight tubercle between chelipeds in Cryptodromiopsis)-, and 3) P2 and P3 not knobbed (knobbed in Mclaydromia colini n. sp. Cryptodromiopsis). (Figs 12; 13) Mclaydromia n. gen. is very close toH em is­ phaerodromia on account of similarities on theDromidiopsis dubia — McLay 1993: 138, fig. 15c. (NonDromidiopsis ¿/«Ait Lewinsohn, 1984). frontal and orbital regions and of subcheliform nature of the P4 and P5. Additionally, they share T ype MATERIAL. —New Caledonia. LAGON, stn 619, 22°3.2’S, 166°54.2’E, 27-42 m, 06.VIII.1986, holo- a sternite 4 that is anteriorly truncated and with type $ 16.2 X 13.2 mm (MNHN-B 22546); stn 111, subparallel lateral borders; male abdominal seg­ 22°24.30’S, 166°47.70’E, 25 m, 22.VIII.1984, ment 6 broad, and with external borders deeply p a ra ty p e $ 10.4 X 9.1 mm (MNHN-B 26282); hollowed and thickened to receive coxal promi­ stn 569, 22°48.80’S, 166°58.90’E, 62 m, 17.VII.1985, paratype ovigerous Î 9.4 X 8.6 mm (MNHN-B nence of P2; dorsal uropods salient, that are 26289); stn 215, 21°52.90’S, 165°49.90’E, 14 m, obliquely oriented and completely involved in 21.IX.1984, paratype ovigerous Î 12.0 X 10.3 mm abdominal holding (Guinot & Bouchard 1998: (MNHN-B 26284); stn 303, 22°38’S, 166°49.10’E, fig. 3C, D, H. monodus)-, apertures of spermathe­ 30-35 m, 27.XI.1984, paraty p e $ 11.3 X 9.7 mm (MNHN-B26280); stn 104, 22°26’S, 166°40.40’E, cae apart, at about the level of P2 (see Fig. 7B for 24 m, 22.VIII.1984, paratype ovigerous Î 9.6 X H. monodus, and Fig. 12B for M . colini n. gen., 8.5 mm (MNHN-B 26281). n. sp.). The following characters distinguish the Although we have examined all the material from New Caledonia referred by McLay (1993), only the specimens two genera: 1) thoracic sternite 3 not exposed in selected for the type series are given herein. For the loca­ Hemisphaerodromia (partly visible at lower plane lity details of remaining specimens see McLay ( 1993: 138). in Mclaydromia n. gen., sternites 1-2 not ETYMOLOGY. — The present species is dedicated to exposed, however); 2) apertures of spermathecae Colin L. McLay (University of Canterbury, located at level of episternites 5 inH em is­ Christchurch, New Zealand).

80 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

Type locality. — N ew C aledonia, 2 2 °3 .2 ’S, 166°54.2’E, 27-42 m.

Distribution. — Only known from New Caledonia.

D escription Carapace noticeably longer than wide, covered with a fine tomentum. Tomentum more devel­ oped on flanks of carapace, chelipeds, and legs. Cardiac region ill defined; branchial groove well recognizable, a blunt small tooth just behind; B regions of carapace not defined. Front wide. Median frontal tooth (rostrum) small but well recognizable. Lateral frontal teeth (pseudorostral) prominent, blunt, so that entire front is produced forward. Supraorbital tooth quitely distinct. Exorbital angle without tooth. Orbital fissure Fig. 13. — Mclaydromia colini n. gen., n. sp., LAGON, stn 111, very deep. Suborbital margin abruptly interrupt­ 25 m, S 10.4 x 9.1 mm, paratype (MNHN-B 26282), views of ed, forming suborbital lobe, and leaving deep and frontal border of carapace; A, dorsal view; B, anterodorsal view. broad hiatus between suborbital lobe and anten­ Scale bar: 2.5 mm. na. Two marked but blunt teeth on anterior half of anterolateral margin: first tooth lying aboutsingle spine propodal, stronger than on P4; a level of orbital fissure; second tooth much larger minute spine on distal outer propodal margin. and placed at higher level than first; a smaller, Abdomen composed of six free segments, plus rounded third anterolateral tooth visible. telson. Male telson about as long as broad, Subhepatic tubercle strong in adults, low but well rounded distally. Abdominal segment 2 with recognizable in young. wing-like lateral expansion covering base of Carpus of cheliped ornamented with several penial tube issued from P5 coxa. Abdominal seg­ tubercles on dorsal surface and with strong tuber­ ment 6 abruptly constricted, considerably nar­ cle on external distal margin; remaining cheliped rower than segment 5. Uropod plates salient, articles smooth. Cutting edge of dactylus with visible dorsally. Abdominal holding consisting of minute proximal teeth followed by molariform uropod plate fitting in front of curved serrate tooth backwards directed, opposed to similar flange on base of P2 coxa; additionally, on PI tooth on cutting edge of fixed finger; distally, the coxa a marked tubercle, may be a group of very usual interlocking teeth. Cheliped with well- small and closely approximated granules, without developed epipodite. contact with telson. Rather dense tomentum covering P2 to P5, their margins fringed with closely placed plumose R e m a r k s setae. P2 and P3 robust. Upper margin of merus Dromidiopsis dubia was described from a single with a rounded tubercle placed distally; upper male (the holotype) from Madagascar. The com­ surface of remaining articles smooth, no obviousparison of the holotype and two additional adult tubercles or conspicuous elevations. Dactylus males from Madagascar with several individuals slightly shorter than propodus, inner margin from New Caledonia, previously identified as armed with acute spines, claw strong; condyles of D. dubia by McLay (1993: 138), has shown that P2 and P3 very strong, rounded. P5 longer than the New Caledonian specimens should no longer P4. Dactylus of P4 curved, opposed by small be attributed to D. dubia. The material belongs propodal spine; no spine on outer propodal mar­to a new species, named hereinMclaydromia gin. Dactylus of P5 strongly curved, opposed by colini n. gen., n. sp.

ZOOSYSTEMA • 2003 • 25 (1) 81 Guinot D. & Tavares M.

In addition to male holotype (10.5X 9 mm) of tubercle is well recognizable although being very D. dubia Lewinsohn, 1984, the following materiallow, and the two teeth on carpus of cheliped are has been examined: absent. The development of carapace teeth varies Madagascar. Near Tany Kely, 13°27’S, 48°10’E, with age in both species. A colour photograph of 30 m, 13.VIII.1971, A. Crosnier coll., holotype a specimen from Maldive Islands is given by S 10.5 X 9 mm (MNHN-B 6894). — Near Debelius (1999: 249, as D. dubia). Nosy-Bé, Tany Kely, north-west coast, 23 m, Two individuals ofM. colini n. gen., n. sp. were 30.IX.1970, P. Laboute coll., S 12.5 X 11.5 mm carrying a sponge cap. (MNHN-B 22592). — 13°40.3’S, 47°48’E, 32 m, 05.XII.1972, A. Crosnier coll., S 15 X 13 mm (MNHN-B 22593). Genus Moreiradromia n. gen. Mclaydromia colini n. gen., n. sp. andM. dubia (Figs 14; 28H) n. comb, can be differentiated as follows: 1) in Dromidia — Stimpson 1858pro parte: 225. (Non M. colini n. gen., n. sp. the blunt lateral frontal teeth Dromidia Stimpson, 1858, type species:Dromia hir­ (pseudorostral teeth) are prominent, so that entire sutissima Lamarck, 1818 by original designation). front is produced forward, while inM . dubia Dromidia (restricted synonymy) — Henderson 1888 n. comb, the lateral frontal teeth are short and pro parte. 12. — Bouvier 1896pro parte. 20 (53). — rounded so that entire front is produced only Borradaile 1903a pro parte. 299, 301. — Moreira 1901: 34; 1905: 136. — Rathbun 1937: 32. — slightly forward; and 2) in M. colini n. gen., n. sp. Williams 1965: 143; 1984: 255. — Coelho & Ramos the dorsal face of carpus of cheliped is ornamented 1972: 177. — Coelho & Ramos-Porto 1989: 215. — with conspicuous tubercles, whereas inM. dubia Forest 1974pro parte. 89, footnote. — Powers 1977: n. comb, the dorsal surface of carpus is smooth. 19- — Manning & Chace 1990: 43. The above characters are constant throughout the Evius Moreira, 1912: 322 (type species Evius ruber material examined, regardless sex and age of the Moreira, 1912 by monotypy. Name pre-occupied by Evius Walker, 1855 [type species: Phalaena hippia individuals examined. Full-grown individuals of (Stoll, 1790), Lepidoptera]). — Rathbun 1937: 30, Mclaydromia colini n. gen., n. sp. andM. dubia footnote. — Franco 1998: 11. n. comb, can also be separated by the following Cryptodromiopsis — McLay 1993 proparte\ 187. — additional characters: 1) in Mclaydromia colini Melo 1996: 67. — Hendrickx 1997: 17. — Debelius n. gen., n. sp. suborbital margin abruptly inter­ 1999: 80 (photograph ofC. antillensis). — Melo & rupted, forming well individualized suborbital Campos 1999proparte\ 275, 279. — McLay etal. 2001 proparte\ 733, 743, tables 2, 3. (Non Cryptodromiopsis lobe and leaving deep and broad hiatus between Borradaile, 1903, type species: Cryptodromiopsis tridens suborbital lobe and antenna (suborbital marginBorradaile, 1903 by monotypy). gently interrupted, leaving only a narrow sinus Type SPECIES. — Dromidia antillensis Stimpson, 1858 between suborbital lobe and antenna M.in dubia by present designation. n. comb.); 2) inM. colini n. gen., n. sp. a small, SPECIES INCLUDED. — Dromidia antillensis Stimpson, rounded but well visible third anterolateral tooth 1858;D. sarraburei Rathbun, 1910 (notlarraburev, see (only a small elevation behind second antero­Boyko 1998: 234). lateral tooth of carapace inM. dubia n. comb.); ETYMOLOGY. — The present new genus is named after and 3) in M . colini n. gen., n. sp. subhepatic Carlos Moreira (1869-1946) in recognition for his sig­ tubercle strong (very low inM. dubia n. comb.). nificant contributions to Brazilian carcinology. The original description ofDromidiopsis dubia Gender: feminine. mentions the absence of subhepatic tubercle onDISTRIBUTION. — Western Atlantic and Eastern carapace and the smooth cheliped, except two Pacific regions. weakly developed tubercles on distal margin of carpus (Lewinsohn 1984: 102). We confirm D e s c r ip t io n these features in male holotype (10.5X 9 mm) Carapace longer than wide or slightly wider than but, in a larger male (15 X 13 mm), a subhepatic long, convex. Regions weakly defined, except for

82 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

B

1-2 mxp3 1

cx1

v5

cx2

v4

v3 cx3 I

cx4

cx5

1-2

jrixp3 jp-Jf

cx3

~ u

cx4 cx2

cx5 cx3

Fig. 14. — A, B, Moreiradromia antillensis (Stimpson, 1858) n. comb.;A, South America, Calypso, 9 1 4 x 1 5 mm (MNHN-B 12717), thoracic sternum, two front views, and spermathecae;B, Florida, S 1 6 x 1 5 mm (MNHN-B 12753), ventral surface of abdomen, with vestigial pleopods on segm ents 3-5;C, Moreiradromia sarraburei (Rathbun, 1910) n. comb., San José Island, Diguet coll., McLay det. Cryptodromiopsis larraburei, S 18.4 x 19 mm (MNHN-B 12761), thoracic sternum with abdomen (tilted for clarity). Abbreviations: a6, abdominal segment 6;cx1-cx5, coxae of P1-P5;g, granules on P1 coxa;mxp3, external maxilliped; o, female gonopore;pr, holding spiniform prominence;s, aperture of spermatheca; t, telson; u, uropod; v3-v5, vestigial pleopods 3-5; 1-2, sternites 1-2; 3-8,sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 1 mm.

ZOOSYSTEMA • 2003 • 25 (1) 83 Guinot D. & Tavares M.

branchial groove. Front dentate, rostrum directed conspicuous; epsiternite 5 with a few granules downward. Anterolateral margin toothed, teeth (M. sarraburei n. comb.). almost spiniform, a tooth just behind branchial Chelipeds rather short and stout, without epipod. groove. Subhepatic region with tooth, visible P2 and P3 rather slender, smooth, not nodose; dorsally. Supra- and suborbital teeth present, no distal spine on propodus; inner margin of exorbital absent; a deep fissure separating supra- dactylus spinulated. P4 and P5 reduced, P5 and suborbital margins. Ocular stalk short and much longer than P4, both with a subcheliform stout. Antenna: urinal article rather straight, so apparatus formed by two distal propodal spines urinal opening aligned with horizontal axis ofopposing dactylus; an outer propodal spine, and article; basal article with exopod noticeably devel­ one or two spines on margins of P5 dactylus. oped, reaching antennal article 4, and internalMale P5 coxa with mobile penial tube (Fig. 28H). angle weakly produced, much shorter than exo­ G2 with a styliform, needle like flagellum. pod. Mxp3: coxae separated by gap. Thoracic sternites 1-2 visible at lower plane. Carrying behaviour Thoracic sternite 3 inserted between mxp3 Sponges, zoanthoid polyps, compound ascidians, coxae, exposed but more or less covered by occasionally sea anemones. telson (M. antillensis n. comb., Fig. 14A; M. sarraburei n. comb., Fig. 14C). Sternite 4 R em a rks narrowing distally, ending in gently rounded/ As pointed out by Rathbun (1937), the descrip­ truncate and raised tip [M. antillensis n. comb.), tion as a new genus and species,Evius ruber slightly concave and not raised {M. sarraburei Moreira, 1912, was based on a megalopa stage. n. comb.), so coxae of PI close to each other.Recently Guinot & Tavares (2000) elucidated Female sutures 7/8 extremely long, getting another case of a dromiid genus based on a mega­ progressively close to each other as they run lopa, Conchoedromia Chopra, 1934. The descrip­ forward over thoracic sternites; apertures of tion of dromiid genera based on the megalopa spermathecae situated together and beyond stage stems from the fact that the dromiid larvae articular condyle of P1 on anterior part of ster­were poorly known at that time. The only dromi­ nite 4, raised (M. antillensis n. comb., Fig. 14A) id postlarvae known prior to 1934 were those of or not(M. sarraburei n. comb.). Austrodromidia octodentata (Haswell, 1882) and Male abdomen long, almost reaching mxp3, leav­ Stimdromia lateralis (Gray, 1831) (Guinot & ing no parts of sternite 4 visible; all abdominal Tavares 2000). Had the dromiid megalopa been segments free. Male abdominal segment 6 long, better documented at that time, someone with length as much as three quarters of width; telson Moreira’s or Chopra’s experience would certainly longer than wide, rounded and bluntly tipped. have recognized his material as a megalopa stage. Male uropods showing as well-developed dorsal According to Rathbun (1937: 31, pi. 8, figs 1, 2), plates, visible in dorsal view of abdomen, Evius ruber is the megalopa ofDromia erythropus obliquely oriented and very movable, not G. Edwards, 1771. A detailed study by Franco involved in holding of abdomen. Presence of ves­(1998) revealed, however, that A. ruber w í s actu­ tigial pleopods in males (Fig. 14B): P13-P15 ally described from a megalopa of D rom idia showing as vestigial buds, tiny and may be obso­ antillensis. lete in larger individuals. Abdominal holding Although the availability of the name Evius provided by main spine that projects from P2 Moreira, 1912 is not affected by the fact that it coxa and overhangs abdominal segment 6; base was applied to a “stage in the life cycle” (ICZN of P2 coxa may bear other small and not efficient 1999: article 17.3), Evius Moreira, 1912 is actu­ tubercles [M. sarraburei n. comb.); on coxae of ally pre-occupied by Evius Walker, 1855 (type PI (M. sarraburei n. comb.), P3 and even P4, species: Phalaena hippia (Stoll, 1790), Lepi­ presence of tubercles or granules, more or less doptera). Consequently, a new name,Moreira-

84 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

dromia n. gen., is herein established to replace coxae of mxp3 closely approximated in Evius Moreira, 1912. Dromidia)-, and 8) P4 and P5 with two distal Originally described in Dromidia, the position of propodal spines opposing the dactylus (a single D. antillensis and D. sarraburei remained undis­ distal spine opposing the very long dactylus in puted until McLay (1993: 187, 188) transferred Dromidia). The abdominal holdings are similar both species to Cryptodromiopsis. He provided no in both genera, being provided on the P2 coxae morphological information to justify such a deci­by spine which overhangs abdomen. In sion and limited himself to state that “the Moreiradromia n. gen., however, the coxal spine Atlantic species formerly known asDromidia is much smaller, other tubercles may be present antillensis Stimpson, 1858, as well as the closely on P2 coxa and episternite 5{M. sarraburei related D. larraburei [sic\ Rathbun, 1910, from n. comb.), and tubercles or granules exist on PI the Pacific, should now be referred to as coxae (M. sarraburei n. comb.), P3 and even P4 Cryptodromiopsis antillensis (Stimpson, 1858) and coxae (see Guinot & Bouchard 1998: 624, 628, C. larraburei [sic] (Rathbun, 1910)” (McLay fig. 4C). 1993: 188). Nevertheless, we agree with McLay The following characters distinguishM oreira­ that the American species do not belong in dromia n. gen. fromCryptodromiopsis sensu nobis Dromidia sensu nobis. (Fig. 4): 1) male vestigial P13-P15 present (absent A study of species assigned to D rom idia and in Cryptodromiopsis)-, 2) thoracic sternites 1-2 Cryptodromiopsis (now restricted to their type visible dorsally, thoracic sternite 3 exposed, species, Dromidia hirsutissima and C rypto­ remaining dorsally partly visible or almost comple­ dromiopsis tridens, respectively) showed that a tely covered by male abdomen, when folded, in new genus should be created for the American M. sarraburei n. comb. (Fig. 14C); sternite 4 species antillensis (Western Atlantic) andsarrabu­ always covered (sternites 1-3, anterior portion of rei (Eastern Pacific). That new genus is herein sternite 4 and episternites 4 and 5 visible in referred as Moreiradromia n. gen. Cryptodromiopsis)-, 3) holding of abdomen Several morphological features support the cre­ provided mostly by small spine on P2 coxa, ation ofMoreiradromia n. gen. and distinction overhanging abdominal segment 6, without the from D rom idia sensu nobis: 1) male uropods involvement of uropods (a serrated granulous showing as movable and well-developed dorsal prominence on P2 coxa inCryptodromiopsis)-, plates in Moreiradromia n. gen. (as ventral plates, 4) female sutures 7/8 getting progressively close almost completely concealed in dorsal view, and to each other as they run forward over thoracic almost immovabe inDromidia, Fig. 5); 2) male sternites and ending well beyond articular abdomen narrow (relatively wide inDromidia)-, condyle of P 1 (separated wide apart, getting close 3) male abdomen when folded nearly reaching to each other abruptly at level of P2 and ending coxae of mxp3, leaving no parts of sternite 4 visi­ at level of articular condyle of PI in ble (anterior portion of sternite 4 and episterniteCryptodromiopsis)-, 5) apertures of spermathecae 4 visible in Dromidia)-, 4) male abdominal seg­ and thoracic sternite 4 placed about at same ment 6 long, with length as much as three quar­ plane (apertures and sternite 4 placed at different ters of width (short, length half the width, in planes; sternite 4 directed downward, in Dromidia)-, 5) male telson longer than wide, Cryptodromiopsis)-, 6) female thoracic sternite 4 rounded and bluntly tipped (telson much wider narrowing progressively forward, ending in gen­ than long, ending in spine Dromidia)-,in 6) male tly rounded tip, so that PI coxae get close to each P13-P15 showing as vestigial buds (Fig. 14B), other (wider, noticeably truncated, so that PI always present but hard to locate due to their tiny coxae are separated from each other wide apart in size (P13-P15 longer, easy to locate inDromidia)-, Cryptodromiopsis)-, 7) urinal article of antenna 7) thoracic sternite 3 present, so a narrow gap rather straight, so that the urinal opening is between coxae of mxp3 (sternite 3 not visible, aligned with horizontal axis of article (upturned

ZOOSYSTEMA • 2003 • 25 (1) 85 Guinot D. & Tavares M.

urinal article, with opening placed above axis of n. status), and Hypoconcha (Hypoconchinae urinal article in Cryptodromiopsis); 8) exopod of n. subfam., see Figs 19; 20; 28K). Nevertheless, basal antennal article noticeably developed, this rather poor American dromiid fauna, with reaching article 4, much longer than internal cor­two endemic genera H ( ypoconcha and ner (exopod comparatively poorly developed, Moreiradromia n. gen.), is unique. As currently much shorter than internal corner which is pro­defined, Dromia is composite, and the generic duced, in Cryptodromiopsis); and 9) P4 and P5 status of a number of species currently assigned with two distal propodal spines opposing the to Dromia s.s., including the single Western dactylus (only one distal propodal spine in Atlantic species in the genus(D. erythropus Cryptodromiopsis). G. Edwards, 1771), deserves more attention. The following characters distinguishM oreira­ A colour photograph ofMoreiradromia antillensis dromia n. gen. fromDromidiopsis sensu nobis: n. comb, showing a specimen from Dominica is 1) uropods as dorsal plates obliquely oriented in given by Debelius (1999: 80 as Cryptodromiopsis Moreiradromia n. gen. (vertically oriented in antillensis). Dromidiopsis, Fig. 6); 2) male abdomen with all segments free (segments 5-6 fused in Dromidiopsis)-, 3) sternite 4 completely covered, Genus Platydromia Brocchi, 1877 when male abdomen folded and reaching coxae (Figs 15; 16) of mxp3 (extreme anterior part of sternite 4 and a Platydromia Brocchi, 1877: 53, 54. (NonPlatydromia small epsisternite 4 remaining uncovered in Fulton & Grant, 1902a: 57, replacement name: Dromidiopsis)-, 4) apertures of spermathecae on Epipedodromia André, 1932: 180 footnote, see McLay prominence placed well beyond articular condyle 1993: 124, 224, table 8, type species: Platydromia thomsoni Fulton & Grant, 1902 by monotypy). of PI (ending between PI or just behind them, together on central prominence, D in ro m i­ Dromien - Vélain 1878: 67, 73. diopsis)-, 5) P4 and P5 with two distal propodal Platydromia — André 1932: 178, 179. spines opposing the dactylus (one distal spine Dromidia — Stimpson 1858pro parte. 225; 1907: 170. opposing the dactylus in Dromidiopsis) ; 6) male — Henderson 1888: 12. — Bouvier 1896pro parte. vestigial P13-P15 present inMoreiradromia n. gen. 21 (54). — McLay 1993 pro parte. 121, 183, 224, 225. (Non Dromidia Stimpson, 1858 sensu nobis, (absent in Dromidiopsis). Other differences type species: Dromia hirsutissima Lamarck, 1818 by between Moreiradromia n. gen. andDromidiopsis original designation). are the abdominal holdings, provided in Cryptodromiopsis — Barnard 1947pro parte. 369; 1950: Dromidiopsis by strong dentate crest on P2 coxae 329- (Non Cryptodromiopsis Borradaile, 1903 sensu acting with uropods (Fig. 6B), but by a spine on nobis, type species: C. tridens Borradaile, 1903 by P2 coxa which overhangs abdominal segment 6, monotypy). without involvement of the uropods, in ^Parasphaerodromia Spiridonov, 1992: 69. Moreiradromia n. gen. (Fig. 14C). Pseudodromia — Macpherson 1988: 61. (Non Carrying behaviour has been reported for both Pseudodromia Stimpson, 1858). species currently included in Moreiradromia T ype SPECIES. — Dromidia spongiosa Stimpson, 1858 n. gen.,M. antillensis n. comb, andM. sarraburei which is a senior synonym ofPlatydromia depressa n. comb. Individuals of the two species cover Brocchi, 1877, type species ofPlatydromia Brocchi, themselves with sponges, zoanthoid polyps, 1877 by monotypy. Gender: feminine. Several specimens ofPlatydromia depressa Brocchi, compound ascidians, and occasionally sea 1877, from Saint-Paul Island (M. de 1’Isle coll., 600- anemones (Brusca 1980; Williams 1984; 1876) (MNHN-B 8739), may be considered syntypes. Hendrickx 1997). A lectotype, a male 9X10 mm, is selected herein Only three dromiid genera are known from the (MNHÑ-B 27934); the remaining specimens are paralectotypes. Other specimens from the same expe­ Atlantic and Eastern Pacific:Dromia (Dromiinae dition (see Vélain 1878) are labeled “Saint-Paul Island, n. status), Moreiradromia n. gen. (Dromiinae Vélain coll., 178-1875” (MNHN-B 12724).

86 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

a6 1-2 mxp3

cx1

cx2 e5 5/6

cx3

cx4 cx5

cx3

cx4

cx5

Fig. 15. — Platydromia spongiosa (Stimpson, 1858) n. comb.; A, B, Saint-Paul Island, Brocchi det.Platydromia depressa Brocchi, 1877, Vélain coll., 178-1875,S 11 x 14 mm (MNHN-B 12724); A, thoracic sternum, two front views, note sternite 4 medially hol­ lowed; B, dorsal view of abdomen; C, Amsterdam Island, Ams-A4, 11.11.1971, Beurois coll.,S 1 3 x 1 6 mm (MNHN-B 12728), ventral view of abdomen, note uropods not visible dorsally and oriented forward, and the vestigial pleopods showing only as papillae on segments 3-4. Abbreviations: a2-a6, abdominal segm ents 2-6; cx1-cx5,coxae of P1-P5; e4, e5, episternites 4, 5; mxp3, external maxilliped; pt, penial tube; t, telson;u, uropod; v3-v5, vestigial pleopods 3-5; 1-2, sternites 1-2; 3-8,sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 1 mm.

ZOOSYSTEMA • 2003 • 25 (1) 87 Guinot D. & Tavares M.

SPECIES INCLUDED. — Dromidia spongiosa Stimpson, telson of females without acute tip. Segment 6 1858. with external borders parallel. In males, vestigial Whether other species ofD rom idia or Crypto­ dromiopsis should be transferred to Platydromia pleopods present on segments 3-5 (P15 as small requires further investigation. For example, the status lobes and P14 and P13 as short papillae, difficult of C. lepidota Barnard, 1947, based on an immature to see, Fig. 15C). Uropods as ventral plates, not specimen and assigned toDromidia by McLay (1993: visible dorsally, oriented forward instead laterally; 183-185), is uncertain. uropods not involved in holding of abdomen. No DISTRIBUTION. — South Africa and south western efficient holding of abdomen: at the more least, a Indian Ocean. very low prominence on coxa of P2 just in con­ tact with external margin of abdominal segment D e s c r ip t io n 6 which is not modified. Female gonopore on P3 Carapace wider than long, convex; dorsal surface coxa relatively large. with not well-defined regions; branchial grooveChelipeds without epipod. PI, P2 and P3 very weakly marked. Anterolateral margin long, short and stout, not knobbed nor nodose; propo­ strongly convex, entire, only undulated, with dus of P2 and P3 without distal spine; inner mar­ only blunt knob behind level of branchial groove;gin of dactylus armed with spines. P4 and P5 posterolateral margins very short, markedly con­ reduced, P5 being slightly longer and more slen­ vergent posteriorly. Front narrow, with rostrumder. Propodus of P4 and P5 short and with only in lower plane and deflexed, and two low one distal spine opposing dactylus which is short pseudorostral lobes; only a slight supraorbital and ends in horny spine. tooth, no suborbital and exorbital teeth. Male P5 coxa with mobile penial tube (Fig. 15A). Antenna: urinal article noticeably developed, with anterior part of beak rounded, wider and Carrying behaviour shorter than posterior ones; basal article very Com pound ascidians. short, with exopod thick; internal angle acutely produced. Mxp3 with coxae not closely approxi­ R em a rks mated. Dromidia spongiosa differs substantially from Thoracic sternites 1 and 2 narrrow and hardly D. hirsutissima, type species of D rom idia (see visible; sternite 3 largely visible dorsally, at level above), so that it is necessary to separate it in a not much lower than sternite 4. Thoracic sternitedifferent genus. ForD. spongiosa we rehabilitate 4 broad, deeply medially hollowed in males, with the genus Platydromia Brocchi, 1877, its type lateral borders convex and anterior margin medi­ species P. depressa being a junior synonym of ally concave in both sexes (Figs 15A; 16). In Dromidia spongiosa (McLay 1993: 121, 183). males, thoracic sternites 7 and 8 perpendicular in The species described as Dromidia spongiosa by relation to preceding ones; in females, sternites 5-Stimpson has been known under five specific 8 perpendicular in relation to sternite 4. Female names and placed in at least six genera (McLay sternal sutures 7/8 very long, with apertures of 1993: 183). In the present study D. spongiosa is spermathecae ending together on slight promi­ excluded from Dromidia, its original (Stimpson nence between chelipeds (Fig. 16). When male 1858) and last genus (McLay 1993: 183,184), as abdomen is applied against ventral surface, stern­ well as from Cryptodromiopsis where it was placed ites 1-2 partly discernible, sternite 3 and anterior by Barnard (1947, 1950). It is nowPlatydromia part of sternite 4 remaining visible, and no epis- spongiosa (Stimpson, 1858) n. comb. ternites exposed. We agree with McLay (1993: 184) who placed Male abdomen very broad, with distinct pleural Pseudodromia inermis Macpherson, 1988 (p. 61, parts, not covering whole sterno-abdominalfigs 6, 7) in the synonymy of hisDromidia spon­ depression, and with all segments free; telson giosa. McLay also synonymised with D. spongiosa broadly triangular, ending in sharp tip (Fig. 15B, C); Parasphaerodromia subglobosa Spiridonov, 1992

88 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

(p. 70, fig. 2a-d), from the South Indian Ocean, indicated with reduced uropods (see below). Platydromia spongiosa n. comb, differs from Dromidia sensu nobis (Fig. 5) in particular by 1-2 having the frontal, orbital and anterolateral mxp3 margins without teeth, a thick, sponge-like tomentum, and no spine on P2 coxa. Other characters which distinguish Platydromia from cx1 Dromidia sensu nobis are: 1) front lobate (tri- dentate in Dromidia); 2) sternite 3 visible (not visible dorsally in Dromidia); 3) male sternite 4 relatively wide, with convex lateral margins and medially cleft anterior margin (with oblique cx2 lateral margins and rounded tip in Dromidia); 4) male abdomen that leaves uncovered ster­ nite 3 and part of sternite 4 (Figs 15A; 16) 5/6 (covering whole abdominal depression with cx3 exception of extreme anterior part of sternite 4 in Dromidia , Fig. 5A, B); 5) uropods showing as small, totally concealed, ventral plates (Fig. 15B, C) (oblique and intercalary ventral cx5 plates, almost entirely concealed inDromidia , Fig. 5B, C); and 6) no efficient abdominal holding (efficient holding by strong coxal spine cx1 on P2 in Dromidia , Fig. 5B). Platydromia is easily distinguishable from Cryptodromiopsis sensu nobis (Fig. 4) by a num ­ cx2 ber of characters, chiefly: 1) round shape of cara­ pace, longer than wide (distinctly wider than long in Cryptodromiopsis); 2) uropods show ing as totally concealed ventral plates (salient dorsal cx3 plates in Cryptodromiopsis); 3) male telson ending in acute tip (triangular tip in Cryptodromiopsis); 4) male segment 6 broad (narrower in Cryptodromiopsis); 5) male sternite 4 broad, deeply hollowed medially (narrower, not hol­ lowed medially in Cryptodromiopsis); and 6) an inconspicuous low prominence on P2 coxa (a ser­ cx5 rated prominence, and uropods not involved in Cryptodromiopsis). Platydromia and Lamarckdromia n. gen. (type species: Dromia globosa), both with ventral and Fig. 16. — Platydromia spongiosa (Stimpson, 1858) n. comb., Saint- Paul Island, Vélain coll., 178-1875, Brocchi det.Platydromia de­ concealed uropods, differ from each other by sev­ pressa Brocchi, 1877, 9 12 x 1 6 mm (MNHN-B 12724), thoracic eral features of the carapace and pereopods, and sternum, two front views, and spermathecae. Abbreviations:cx1- cx5, coxae of P1-P5;e4, e5,episternites 4,5;gy, episternal gynglyme specially by: 1) sternite 3 largely visible in receiving sternal articular condyle of the pereopod;mxp3, external Platydromia (only a small part visible in male maxilliped; o, female gonopore;s, aperture of spermatheca; 1 -2, sternites 1 -2; 3-8,sternites 3-8; 4/5-7/8,thoracic sternal sutures Lamarckdromia n. gen., Fig. 10A, D); 2) sternite 4/5-7/8. Dotted line indicates difference in level. Scale bar: 2 mm.

ZOOSYSTEMA • 2003 • 25 (1) 89 Guinot D. & Tavares M.

4 wide, with convex lateral margins in both sexes Dromia species by several morphological fea­ (triangular inLamarckdromia n. gen., Fig. 10A, tures (in particular the abdominal holding with D); 3) male telson ending in acute tip (rounded a coaptation by engagement between P2 coxae at tip in Lamarckdromia n. gen., Fig. 10B, C); and the modified edges of abdominal seg­ and 4) apertures of spermathecae ending together ment 6, and no involvement of uropods, see on slight prominence between chelipeds (endingBouchard 2000) and larval development (see together on central prominence between P2 inMcLay et al. 2001). Its worldwide geographical Lamarckdromia n. gen., Fig. 10A). distribution includes all the three major oceans; The carapace and legs of Platydromia spongiosa it is found as deep as 520 m. are completely covered with “a dense and firm envelope of pubescence, sponge-like in appear­ ance”, “distinctly marked with shallow pits or Genus Pseudodromia Stimpson, 1858 depressions” (Stimpson 1907: 171, asDromidia Pseudodromia Stimpson, 1858: 226. — Henderson spongiosa), or with “a very short close and thick 1888: 15- — Alcock 1900pro parte-. 149. — Stebbing pile” (Barnard 1950: 330, as Cryptodromiopsis 1900: 23. — Stimpson 1907: 177. — Barnard 1950 spongiosa). By comparison, the carapace and legs pro parte-. 315. — Gordon 1950pro parte-. 209. — Kensley 1977pro parte-. 183; 1978: 257; 1980pro of Dromidia hirsutissima are “covered with short parte-. 25. — McLay 1993: 125, 175, table 4. — stiff pile, and long dense fibrous and shaggy McLay et al. 2001: 741, table 3. — Guinot 1995: 186. brown or yellow hairs” (Barnard 1950: 320), — Guinot & Bouchard 1998: 626, table 3. — Stewart et al. 2001: 136. hence the name ofhirsutissima given by Lamarck and the common name of “shaggy Type SPECIES. — Pseudodromia latens Stimpson, 1858 sponge crab” (Barnard 1950). The same commonby original designation (Stimpson 1858: 226). Gender: feminine. name was given to Cryptodromiopsis plumosa Lewinsohn, 1984 by Hoover (1998: 266, fig. n. n.;SPECIES INCLUDED. — Pseudodromia latens Stimpson, 1858;Dromia rotunda MacLeay, 1838 (see Ng & see below under Stebbingdromia plumosa Ahyong 2001); Pseudodromia trepidus Kensley, 1978. n. comb.). InLamarckdromia globosa n. comb, Pseudodromia cacuminis Kensley, 1980, from the the whole body and legs are covered by a dense South Atlantic (Vema Seamount), does not belong to shaggy coat of hairs, and the deflexed front Pseudodromia (McLay 1993: 176). portion of carapace is concealed by transverse D istribution . — South Africa. fringe of longer setae, giving it a characteristic appearance. D e s c r ip t io n The soft and areolated tomentum ofPlatydromia Carapace more or less longer than wide, extreme­ spongiosa n. comb, somewhat resembles that of ly narrow anteriorly, convex; dorsal surface “Dromia” wilsoni (Fulton & Grant, 1902b: 61, smooth, without defined regions; cervical groove as Cryptodromia wilsoni). The two species may not present, branchial groove strongly marked. be confused but differ by: 1) shape of carapace, Anterolateral margin very long, usually without rounded and not toothed laterally inP. spon­ tooth; posterolateral margin short. Front very giosa n. comb, (distinctly wider and armed narrow, tridentate, with rostrum markedly laterally in D. wilsoni)-, 2) uropods, only ventral deflexed and hardly or not visible dorsally at all, in P. spongiosa n. comb, (very salient dorsal may be elongated (P. trepidus) and with two plates in D. wilsoni). We agree with McLay in pseudorostral teeth; supraorbital tooth marked or that the generic status of “D rom ia” wilsoni not; suborbital and exorbital teeth absent. needs a re-appreciation (see McLay 1991: 470, Antenna: urinal article with anterior part of beak figs 7, 8, as Petalomera wilsoni-, 2001a: 84, as much shorter than posterior one and with broad Dromia wilsoni-, N g et al. 2000: 160, fig. 2b, as tip; basal article with exopod enlarged and inter­ D. wilsoni-, McLay et al. 2001: 742, table 1, as nal angle not elongated. Mxp3 with coxae closely D. wilsoni). “Dromia” wilsoni differs from other approximated.

90 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

Thoracic sternite 3 not visible dorsally. Thoracic not hollowed inAustrodromidia)-, 3) no epistern­ sternite 4 very narrow, deeply hollowed medially ites visible when male abdomen flexed (epistern­ in males, with lateral borders convex and anterior ites 4 and 5 visible inAustrodromidia)-, 4) male margin pointed and deflexed. In females, thoracic telson ending in acute tip (bluntly triangular or sternites 7 and 8 very tilted in relation to preced­ rounded in Austrodromidia)-, 5) male segment 6 ing ones. Female sternal sutures 7/8 long, withwith external borders may be slightly concave apertures of spermathecae ending together on (deeply hollowed laterally in Austrodromidia)-, prominence between P2. When male abdomen 6) female sternal sutures 7/8 ending together at applied against ventral surface, anterior part of level of P2 (apart in Austrodromidia)-, and sternite 4 remaining visible and no episternites 7) absence of an abdominal holding (abdominal exposed. Sterno-coxal depressions absent. holding by prominence on P2 coxa in Male abdomen completely covering sterno- Austrodromidia). abdominal depression and with all segments free; In Pseudodromia (as well in Ascidiophilus) the male telson relatively long, ending in sharp tip; absence of an abdominal holding system seems to telson of females without acute tip. Segment 6 be related to the complete protection by a host, becoming more or less narrow posteriorly. Nothe body being almost entirely enclosed by an vestigial pleopods present on segments 3-5 in ascidian (Guinot & Bouchard 1998). males. Uropods as relatively small but distinct Using DNA sequence data, Stewart et al. (2001: ventral plates, not visible dorsally (P. latens, 136) questioned the monophyly ofPseudodromia P. trepidus ) or hardly visible (P. rotunda). but confirmed the specific status of two close Absence of any efficient holding of abdomen. species, P. rotunda and P. latens. Chelipeds without epipod. PI, P2 and P3 not knobbed nor nodose; propodus of P2 and P3 without distal spine; inner margin of dactylus Genus Stebbingdromia n. gen. with spines, longer distally. P4 reduced, P5 dis­ (Figs 17; 18) tinctly longer than P4 and being about of same Dromidiopsis — Lewinsohn 1984pro parte. 104, fig. 3. size than P2. Propodus of P4 and P5 without (NonDromidiopsis Borradaile, 1900; type species: spines opposing dactylus; outer propodal spines Dromia australiensis Haswell, 1882). absent; presence of lateral propodal spines, one Cryptodromiopsis — McLay 1993 pro parte. 188, 190; on P4 and two on P5; P5 dactylus straight. 2001a: 85, 86. — Hoover 1998: 266. — ?Chen & Male P5 coxa with mobile and very long penial Haibao 2002 pro parte-. 102, 541, 542. (Non Cryptodromiopsis Borradaile, 1903, type species: C. tri­ tube. dens Borradaile, 1903). T y p e AND ONLY SPECIES. — Dromidiopsis plumosa Carrying behaviour Lewinsohn, 1984. Compound ascidians. ETYMOLOGY. — The new genus is established in hon- nor ofThomas Roscoe Rede Stebbing (1835-1926) for R em a r k s his contributions to carcinology. Gender: feminine. Pseudodromia is distinguished by having uropods DISTRIBUTION. — Stebbingdromia plumosa n. comb., that show as ventral plates, sometimes partly visi­ the only species in the genus, is found from the ble dorsally. This is the major difference from Seychelles Islands (type locality) to Chesterfield Ascidiophilus, since there are no vestigial pleopods Islands, Guam and Hawaii. in the males of both genera (Guinot 1995: 186). Pseudodromia differs from Austrodromidia sensu D e s c r ip t io n nobis (Figs 1; 2) as follows: 1) sternite 3 not visi­ Carapace distinctly wider than long; dorsal sur­ ble dorsally (visible dorsally, clearly or slightly in face smooth, with regions not defined. Branchial Austrodromidia)-, 2) sternite 4 very narrow and groove marked and forming laterally very deep hollowed medially in males (truncate at tip and notch, but without tooth. Anterolateral margins

ZOOSYSTEMA • 2003 • 25 (1) 91 Guinot D. & Tavares M.

subparallel except anteriorly, armed with two dorsal plates, always visible dorsally, deeply teeth behind exorbital tooth; posterolateral mar­ inserted medially between segment 6 and telson, gin very short. Front projecting beyond orbits; horizontally oriented, relatively movable and rostrum acute, markedly deflexed but visible dor­ slightly projecting beyond outline of abdomen. sally; two acute pseudorostral teeth. Orbital mar­ Abdominal holding by distinctly tuberculate crest gin eave-like; supraorbital margin notched; on P2 coxa; uropods weakly involved in holding; supraorbital tooth absent or faintly marked; in addition, presence of rounded granular promi­ infraorbital tooth triangular, separated from exor­ nence on P 1 coxa. bital angle and from inner infraorbital tooth byChelipeds well-developed, with an epipod; sexual deep notches; exorbital tooth dentiform. dimorphism marked: in males, fingers strongly Proepistome raised, in front of wide epistome. downcurved, markedly gaping; dactylus with Orbits directed horizontally, deep. Ocular stalk external surface concave and cutting edge armed very long and narrow, curved. Antenna: urinal with small, molariform proximal tooth; cutting article wider than long, with posterior part of edge of fixed finger armed with several pointed beak broad; basal article with exopodal scale well- teeth (details not visible on the sketch by developed, enlarged; internal angle similarly Lewinsohn 1984: fig. 3c). P2 and P3 neither developed and markedly produced, touching lobed nor nodose; propodus long and bearing a front; following article completely included small distal propodal spine; inner margin of between these two lateral extensions of basal dactylus with several spines. P4 and P5 reduced, article. Mxp3 with coxae not completely approxi­ very short, practically similar in size, P5 only mated. slightly longer; dactylus strongly curved; subche- Sternite 3 not visible anteriorly but discernible on liform apparatus formed by multiple spines: three each side of sternite 4; sternite 4 with triangular (P4) or two (P5) distal propodal spines opposing anterior part in contact with coxae of mxp3. dactyli, and two (P4) or three (P5) spines on When male abdomen applied against ventral sur­ outer propodal margins; a large spine on outer face, sternite 4 completely covered except lateral­ dactylus margin on P5; namely a total of six ly, i.e. episternite 4; episternite 5 remaining spines on P5; P4 and P5 without spines on inner exposed. In females, thoracic sternum regularly margin of dactylus. sloping backwards, posterior part more bent; in Male P5 coxa with mobile penial tube. males, posterior sternites, i.e. posterior part of Male G 1 without apical plate, only with a setose sternite 6, and sternites 7 and 8, abruptly tilted, blunt knob. G2 without exopod, and unusual in so they are nearly perpendicular to preceding the Dromiidae (verified in two males, from ones. Female sutures 7/8 short, with apertures of Seychelles, MNHN-B 8572 and Chesterfield spermathecae apart on very slight prominence Islands, MNHN-B 22562): very short, (shorter between P3 (even in mature females, see McLay than G l), stout, regularly tapering to pointed tip, 2001a: 85, 86), i.e. not far from gonopores onwithout styliform, needle-like flagellum; third P3; these apertures (hidden beneath sperm plug distal part corneous. in the two examined females) completely exposed and perhaps relatively large. Carrying behaviour Abdomen long, reaching mxp3, and relatively Despite the long covering of plumose setae, wide (probably with pleural parts), triangular in pieces of sponge are used for camouflage (Hoover shape, with all segments free; telson rather devel­ 1998: 266). oped, bluntly triangular. Segment 6 much wider than long, with external margins slightly concave. R em a rks In males, pleopods present on somites 3-5, show­Because Lewinsohn (1984: 106) was fully aware ing as uniramous vestiges, rather long and dissim­ that the dromiid genera needed to be reviewed, ilar. In males, uropods showing as elongated he was uncertain of whether his new species

92 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

mxp3

cx1 mxp3

cx1 cx2

4/5

cx2

cx3

cx2

cx2 4/5 cx3 5/6

5/6 cx3

cx4 cx4

cx5 cx5

Fig. 17. — Stebbingdromia plumosa (Lewinsohn, 1984) n. comb.; A, B, Chesterfield Islands, CORAIL 2, stn DW 84, McLay det. Cryptodromiopsis plumosa, 11.3 S x 13.3 mm (MNHN-B 22562); A, thoracic sternum, two front views; B, abdomen, dorsal view; C, Hawaii, Oahu, McLay det. C. plumosa, ovigerous 9 9.5 x 10.5 mm (QM W21890), thoracic sternum and spermathecae; note large apertures of spermathecae at level of P3, concealed by sperm plug. Abbreviations: a6, abdominal segment 6;cx1-cx5, coxae of P1- P5; e4, e5, episternites 4, 5; g, granules on rounded prominence on P1 coxa;mxp3, external maxilliped; o, female gonopore; pr, holding prominence;pt, penial tube; sp, sperm plug; t, telson;u, uropod; 1-3, sternites 1-3; 4-8, sternites 4-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bar: 2 mm.

Dromidiopsis plumosa (“different from all other 85, 86) toCryptodromiopsis. As a result, known species”) should have been placed in D. plum osa has already been placed in three Dromidiopsis. Lewinsohni species was eventually different genera. transferred by McLay (1991: 470) first to Stebbingdromia plumosa n. comb, is the only Dromidia and later (McLay 1993: 138; 2001a: dromiid (actually Dromiacea) with a relatively

ZOOSYSTEMA • 2003 • 25 (1) 93 Guinot D. & Tavares M.

short male G 2 (slightly shorter than G l) that is occurs only inMoreiradromia n. gen. (Fig. 14B, stout and lacks the styliform distal part C, may be obsolete in larger individuals) and in (Fig. 18B). In all members of Dromiacea the G2 Dromia pro parte. We observed a long P15 and is typically as long as or longer than G 1 and ends obscure P14 and P13 in D. personata (Linnaeus, in neddle-like flagellum (Figs 20B; 23B). 1758) (Fig. 27A), more or less short P13-P15 in In the description for the first time of a female D. bollorei Forest, 1974 (Fig. 27B) andD. mar­ of Stebbingdromia plumosa n. comb., an ovi- morea Forest, 1974, and P13-P15 showing as short gerous one, McLay (2001a: 86) remarked that papillae in “Dromia” wilsoni (Fulton & Grant, female sternal sutures 7/8 “end more poste­ 1902). An analogous condition as Stebbing­in riorly than in other species of the genus dromia n. gen. is found in the subfamily Sphaero­ [Cryptodromiopsis]. This means that the ends of dromiinae n. subfam. (see Fig. 22: but biramous the sutures [apertures of spermathecae] are just vestigial P13-P15). below the female gonopores, only about 1 mmAs presently redefined, neither Dromidiopsis away”. This may be regarded as an ancestral sensu nobis,Cryptodromiopsis sensu nobis, nor condition, such as the arrangement of spines onDromidia sensu nobis can receiveD. plumosa. pereopods (distal propodal spine on P2 and P3; Stebbingdromia n. gen can be readily distin­ numerous spines on the subcheliform apparatus guished from Dromidiopsis sensu nobis (Fig. 6), of P4 and P5). as follows: 1) abdominal segments free (abdo­ The apertures of spermathecae are not readily minal segments 5 and 6 fused together in visible in the two available females of Dromidiopsis)-, 2) when extended forward, P5 Stebbingdromia plumosa n. comb, (ovigerous reaching about mid-length of lateral margin of female, 9.5 X 10.5 mm, Hawaii, Oahu, QM W carapace (P5 extremely long, reaching about 21890; female 6.8 X 7.2 mm, Guam, Apra outer orbital angle inDromidiopsis)-, 3) male G2 Harbour, ZRC 2000.2112), as both individuals short and stout, without styliform flagellum still carry sperm plugs (Fig. 17C). We suspect (long and needle-like inDromidiopsis)-, 4) P13-P15 that the spermathecal apertures are relatively as vestiges in males (P13-P15 absent in large in Stebbingdromia plumosa n. comb., and, if Dromidiopsis)-, 5) merus and carpus of P2 and P3 confirmed, this will be another distinctive feature stout, noticeably high; propodus and dactylus of the genus. The apertures of spermathecae show long and thin (merus, carpus, propodus, and as very minute pores in the Dromiacea, with few dactylus looking similar inDromidiopsis)-, and exceptions: for example Sternodromia spinirostris 6) female sternal sutures 7/8 ending between P3 (Miers, 1881) shows narrow oblique slits (see (between PI or just behind them, together on Forest 1974: fig. 6C, pi. 4, fig. 3). The shapes of central prominence inDromidiopsis). In addition, G2 (thick) and spermathecal apertures (if large) the fronto-orbital region is dissimilar in the two might be regarded as an indication that both Gl genera. and G2 are involved in the insemination process. Differences betweenStebbingdromia n. gen. and Two characters ofD. plumosa, female sternal Cryptodromiopsis sensu nobis (Fig. 4) include: sutures 7/8 ending between P3 (versus ending 1) thoracic sternite 3 medially concealed in more anteriorly, as far between or in front of PI,Stebbingdromia n. gen. (exposed in Crypto­ in the other Dromiinae n. status) and male ves­ dromiopsis)-, 2) coxae of mxp3 almost approxi­ tigial pleopods combined with intercalary dorsal mated (a distinct gap between them in uropods, make this species so different that none Cryptodromiopsis)-, 3) male thoracic sternite 4 of the existing dromiid genera could accommo­ ending in acute tip (truncate inC rypto­ date it. dromiopsis)-, 4) uropods oriented horizontally and Male vestigial pleopods combined with dorsal weakly involved in abdominal holding (more salient uropods are relatively rare in the subfamily and oriented obliquely, far from prominence on Dromiinae n. status (Table 1). This conditionP2 coxa, in Cryptodromiopsis)-, 5) P13-P15 as

94 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

Pt

cx5 J a5

a4

a3

Fig. 18. — Stebbingdromia plumosa (Lewinsohn, 1984) n. comb., Chesterfield Islands, CORAIL 2, stn DW 84, McLay det. Cryptodromiopsis plumosa,S 11.3 x 13.3 mm (MNHN-B 22562); A, coxa of P5, with penis as mobile tube;B, G2, without styliform flagellum (corneous part stippled); C, abdomen, ventral view. Abbreviations:a3-a6, abdominal segments 3-6; cx5, coxa of P5; pt, penial tube; t, telson;u, uropod; v3-v5, vestigial pleopods 3-5. Scale bars: 1 mm.

vestiges (P13-P15 absent in Cryptodromiopsis ); slightly beyond outline of abdomen (strongly 6) female apertures of spermathecae between P3 salient and movable inDromia , and with a char­ (together on slight tubercle between chelipeds in acteristic small beak overhanging telson, this beak Cryptodromiopsis ); and 7) male G2 shorter than being probably used as stop system for their Gl and without styliform flagellum (long and movement, see Bouchard 2000); 4) apertures of needle-like in Cryptodromiopsis). spermathecae apart, between P3 (placed more Differences, other than the carapace and legs, anteriorly inDromia ); and 5) G2 short and stout, between Stebbingdromia n. gen. andDromia s.s. not needle-like (with styliform flagellum in include: 1) when folded, male abdomen covering Dromia). whole sternite 4 (anterior part of sternite 4 Differences, other than the carapace and legs, exposed in Dromia); 2) telson longer than wide, between Stebbingdromia n. gen. andM oreira­ triangular (wider than long, rounded/truncate at dromia n. gen. (Fig. 14) include: 1) thoracic tip in Dromia); 3) dorsal uropods projecting only sternite 3 weakly visible dorsally (exposed, may

ZOOSYSTEMA • 2003 • 25 (1) 93 Guinot D. & Tavares M.

be partly covered by male abdomen, when folded, n. subfam.), the apertures of spermathecae are in Moreiradromia n. gen.); 2) sternite 4 anteriorly minute, a condition that is always connected with triangular (truncate inMoreiradromia n. gen.); a styliform and needle-like G2. Stebbingdromia 3) male abdominal segment 6 much wider than n. gen. is so far the only exception, and it is here­ long (very long, length as much as three quarters in tentatively referred to the Dromiinae n. status. of width inMoreiradromia n. gen.); 4) abdominal Stebbingdromia n. gen. shares (plesiomorphically?) holding by tuberculate crest on P2 coxa; presence the following features with the Sphaerodromiinae of round prominence on P1 coxa (provided with n. subfam.: 1) short female sternal sutures 7/8, spine overhanging coxa of P2, which may bear ending between P3, so that the apertures of sper­ other smaller tubercles, and presence of tubercles mathecae lie beside female gonopores (also found or granules on PI, P3 and even P4 coxae in in other basal Podotremata,viz. the H o m o ­ Moreiradromia n. gen.); 5) apertures of sperma­ lodromiidae and Dynomenidae). InStebbing­ thecae apart, between P3 (together on promi­ dromia n. gen., the very visible female sutures 7/8 nence placed well beyond articular condyle of P 1 and the completely exposed apertures of sper­ in Moreiradromia n. gen.); and 6) G2 short, not mathecae (Fig. 17C) actually differ from those needle-like (long and with a styliform flagellum found inSphaerodromia (Fig. 21C) and Eodromia in Moreiradromia n. gen.). (Fig. 24C), in which sutures 7/8 and the sper­ Differences betweenStebbingdromia n. gen. and mathecae are usually concealed under a lateral Hemisphaerodromia (Fig. 7B) include: 1) thoracic heightening of sternite 8; 2) uropods (Figs 17B; sternite 4 ending in acute tip in males and, when 18B) showing as dorsal plates, deeply inserted abdomen flexed, completely covered, except between abdominal segment 6 and telson, more small episternite 4 (rounded, and anterior and or less included in outline of abdomen, and not episternal parts exposed in Hemisphaerodromia)-, markedly salient. In Sphaerodromia (Figs 21 A; 2) telson long, in contact with coxae of mxp322) and Eodromia (Fig. 24A, B), however, the (shorter, remote from mxp3 in H em is­ uropods are immobile and inserted between tel­ phaerodromia)-, 3) male segment 6 with external son and segment 6, while inStebbingdromia margins slightly concave (deeply hollowed in n. gen. they remain independent and relatively Hemisphaerodromia)-, 4) male pleopods P13-P15 movable. The uropods do not play role in the present as uniramous vestiges, rather long, holding of abdomen inSphaerodromia or (absent inHemisphaerodromia)-, 5) abdom inal Eodromia whereas in Stebbingdromia n. gen. the holding by tuberculate crest on P2 coxa; uropods role of uropods in the abdominal holding is weakly involved; presence of an additional weak; 3) similar spinulation of walking legs, in prominence on PI coxa (only a serrated promi­particular the long propodi of P2 and P3, which nence on P2 coxa tigthly encircled in the space are armed with one distal propodal spine (consid­ just behind uropods, markedly involved in ered primitive by McLay 1993: 192 in Stebbing­ abdominal holding, inHemisphaerodromia)-, dromia plumosa n. comb, andSphaerodromia 6) female sternal sutures 7/8 ending apart onspp.); dactyli of P2 and P3 with numerous spines slight prominence between P3 (ending together regularly arranged on inner margin; 4) subcheli- on slight tubercle behind P2 in H em is­ form apparatus of P4 and P5, which consists of a phaerodromia)-, and 7) male G2 shorter than Gl large number of propodal spines opposing the and without styliform flagellum (long and nee- dactylus. In Stebbingdromia n. gen., however, the dle-like in Hemisphaerodromia). absence of spines on inner margin of P4 and P5 It is clear that Stebbingdromia n. gen. is quite dactyli differs from Sphaerodromia and Eodromia, unique. For instance, in all the families of characterized by the presence of spines; and 5) Dromiacea, i.e. Homolodromiidae, Dynomenidae complete male pleopodal formula (i.e., vestigial and Dromiidae (Dromiinae n. status, Hypo­ P13-P15 combined with dorsal uropods, conchinae n. subfam., and SphaerodromiinaeFig. 18C). The only other known Dromiidae

96 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

with vestigial male pleopods combined with GENUS INCLUDED. — Hypoconcha Guérin-Méneville, dorsal uropods are the dromiine genera 1854 (type species: Cancer parasiticus Linnaeus, 1763, senior synonym ofCancer sabulosus Herbst, 1799, see Moreiradromia n. gen. (Fig. 14B) andDromia Holthuis 1962). pro parte (see Patterns of uropods and vestigial male pleopods 3-5 Fig. 27; Table 1). D e s c r ip t io n Nevertheless, several features do not permit the Carapace generally rounded, hourglass-shaped; assignation ofStebbingdromia n. gen. to the dorsal surface flattened, very thin and partly Sphaerodromiinae n. subfam. They are as fol­ membranous; on each posterior side, large soft lows: 1) thoracic sternum, in particular the shape area with special texture; branchiostegal region of sternite 4; 2) male abdomen, in particular seg­ usually soft and of different texture, perhaps ment 6; 3) G2 shorter than Gl and not needle­ constituting ecdysis area. Cervical groove well like in Stebbingdromia n. gen.; 4) coxal structures marked, on two median gastric pits; cardiac of the pereopods for abdominal holding, with a region completely delimitated by well-defined tuberculate crest on P2 coxa and a round promi­ cardiac groove. Branchial groove deep and sepa­ nence on PI coxa, relatively inefficient in rating hard part of carapace from soft posterior Stebbingdromia n. gen. (Fig. 17A, B) (versus a part. Margin of anterior half of carapace usually prominence on P2 coxae involving telson in ante­hairy, appearing as “ciliated”; posterolateral bor­ rior part, and an inefficient prominence on P3der often markedly concave. Front and lateral coxae covered by abdomen in Sphaerodromia, margins greatly expanded, covering all parts of Fig. 21 A, B); 5) male coxa of P5 with long, inde­ head, except antennal flagella, and displacing eyes pendent penial tube inStebbingdromia n. gen. in ventral location. Front semicircular or slightly (Figs 17A; 18A) (male P5 coxa modified into truncate in outline, markedly deflected in large hard process which encloses penis in triangular ventral plate connected with proepis­ Sphaerodromia, Figs 23A; 28E-G); and 6) mobile tome. Eyes, antennules, antennae and mouth uropods (deeply inserted and immobile in parts deeply settled in depressions. Sphaerodromia ). Other differences refer to orbits Basal article of antennule well-developed. First and eyes, antennae, front, proepistome, and che- article of antenna beak-like; basal article notice­ lipeds (in particular fingers). ably developed, exopodal scale relatively small The chelipeds of Stebbingdromia n. gen. are pecu­ and internal angle markedly produced; following liar for the Dromiinae n. status. They are article deeply inserted inside basal article; remain­ markedly sexually dimorphic, with fingers ing articles very small; flagellum long, setose. strongly downcurved, very widely gaping, the Orbits small, concealed beneath body. Mxp3 cutting edge of dactylus concave and armed with operculiform; coxa developed and closely approx­ several long teeth in males. They do not conformimated; merus subtriangular or trapezoidal; exo­ with the chelipeds found in the Hypoconchinae pod noticeably wide, specially in proximal part; n. subfam. or the Sphaerodromiinae n. subfam. crista dentata (on ischium) with small number of (see below). corneous teeth. The identity of the Cryptodromidiopsis plumosa Ventral surface and legs solid, sometimes hairy recorded by Chen & Haibao (2002: 102, fig. 41) (not inHypoconcha parasitica (Linnaeus, 1763)). could not be verified. Gynglymes of sternites 1-3 largely spaced and stepped at lower plane. Sternite 3 only hardly vis­ ible (only represented by a small median hollow) Subfamily H ypoconchinae n. subfam. or not visible. Sternite 4 showing as well calcified, (Figs 19; 20; 28K) narrow and elongated plate, in close contact (except medially) with coxae of mxp3, sometimes

Type GENUS. — Hypoconcha Guérin-Méneville, 1854 appearing fixed to them. Episternites 4 and 5 by present designation. Gender: feminine. broad and well calcified. In both sexes, posterior

ZOOSYSTEMA • 2003 • 25 (1) 97 Guinot D. & Tavares M.

o er

mxp3

4/5 cx2

details

cx3

6/7/ details

cx4

cx5

Fig. 19. — A, B, Hypoconcha californiensis Bouvier, 1898, Gulf of California, San José Island (MNHN-B 22066); A,17x18 9 mm, syntype, thoracic sternum, two front views; note sternites 7 and 8 strongly tilted and perpendicular to preceding ones;S 20.3 B, x 19.4 mm, syntype, ventral view of abdomen, with asymmetrical, uniramous vestigial pleopods and uropods showing as ventral plates; C, H. panamensis Smith, 1869, Lower California, Bouvier det.H. digueti Bouvier, 1898, Diguet coll., 9 about 32 mm width (MNHN-B 22071), abdomen, ventral view; note uropods showing as setiferous ventral lobes. Abbreviations:a3-a6, abdominal segments 3-6; cx1-cx5,coxae of P1-P5;e4, e5, episternites 4, 5; mxp3,external maxilliped; o, female gonopore;pi, female pleopod (setae incompletely figured); s, aperture of spermatheca; t, telson; u, uropod; v3-v5, vestigial pleopods 3-5; 3-8, sternites 3-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: A, C, 2.5 mm; B, 1mm.

98 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

thoracic sternites 7 and 8 tilted; in females, ante­ ingly to its curvature at right angles, abdomen rior sternites 4 and 5 forming horizontal plate,flexed halfway and normally applied to thoracic sternites 6-8 bent at right angles, so they are per­ sternum. Female abdomen becoming expanded pendicular to preceding ones; sternite 6 small, in ovigerous specimens, with formation of brood rejected laterally, its raised anterior part often sur­ chamber. rounding apertures of spermathecae. Sternal Chelipeds stout, epipod present, podobranch suture 4/5 horizontal, well-marked laterally, its lacking. Pereopods capable of being folded trace medially discernible; suture 5/6 oblique, compactly against body and partly concealed by clearly visible; sutures 6/7 and 7/8 very oblique. carapace, with a perfect complementarity of Female sutures 7/8 relatively short, only present diverse parts. Fixed finger and dactylus of che­ on bent surface of posterior sternites, always end­lipeds armed on prehensile margin with comple­ ing apart. Apertures of spermathecae located only mentary teeth and close along whole length. P2 slightly beyond level of condyle of P3, not very and P3 not lobed nor nodose; propodus short, far from female gonopores on P3 (Fig. 19A). never armed with a distal spine; dactylus not Sterno-abdominal depression not noticeably deepstrongly curved, without spines or with only very and located posteriorly. A large portion of tho­small spines on inner margin. P4 and P5 both racic sternite 4 not concealed by male abdomen, reduced (but not coxae) and oriented in a differ­ when flexed; episternite 4 greatly exposed. ent way than P2 and P3 i.e. directed subdorsally Abdomen with all segments free, usually short or dorsally, such as in the Dromiinae n. status, (never attaining coxae of mxp3), broad, notice­ but each markedly dissimilar from the other. P4 ably widened at level of segments 5 and 6, trian­ subdorsal, noticeably robust, much shorter than gular shaped, and with first segments remaining preceding legs and P5; merus very stout. P5 dorsal, even in males; pleurae may be distinct. In completely dorsal, much longer than P4. In both both sexes abdomen bent at right angles about in P4 and P5, carpus relatively long; propodus short the middle and disposed into two different and stout. Dactylus of P4-P5 with a peculiar planes, so that the posterior part of abdomen lies dactylus, which is upturned, crescent and lunate, flat on ventral surface of céphalothorax. This extremely mobile, placed in a deep notch of abdominal curvature, less pronounced in males propodus, and ending in corneous hook. than in females, probably connected with the Male P5 coxa with mobile penial tube (Figs 20A; inclination of two last thoracic sternites, 7 and 8,28K). with regard to preceding ones. Telson triangular, G l not completely closed, without apical plate. broader than long. Uropods showing as minuteG2 very long, styliform, without exopod (Fig. 20B). ventral plates in both sexes, never visible dorsally, very narrow, rather immovable in males, sexually Carrying behaviour dimorphic (showing as small and more setiferous See under Remarks and under Discussion, Shell- lobes in females). Presence of uniramous vestigial carrying behaviour. male pleopods, varying along the species: P13-P15 showing as elongated and asymmetrical lobes R em a r k s (H . californiensis Bouvier, 1898; Fig. 19B), or This unusual crab has been known for a long only as short papillae (H . panamensis Sm ith, time under the name “Faux Bernard l’Hermite”, 1869), or indistinct, at least in small specimensgiven to H. parasitica (Linnaeus, 1763) by (H . parasitica, H. arcuata Stimpson, 1858). Nicolson (1776: 338, pi. 6, fig. 3) (see Rodriguez Abdomen holding may be provided by structure 1993: 44). Lamarck (1818: 264) considered this on PI coxa, which bears a series of spinous tuber­ species new but did not describe it. The genus cles, the strongest of which overhangs telsonHypoconcha was established by Guérin-Méneville (H . californiensis)-, more often, theses structure (1854: 333-343, pi. 5, as H. sabulosa), when he are absent (H . arcuata, H. panamensis)-, accord­ gave a new key for the “Dromiens”, already well-

ZOOSYSTEMA • 2003 • 25 (1) 99 Guinot D. & Tavares M.

defined by H. Milne Edwards (1837). Guérin- mathecae are located beyond level of condyle of Méneville (1854) added Hypoconcha to the two P3, not very far from female gonopores on P3, other known dromiacean genera,Dromia and however (Fig. 19A). Dynomene. Hypoconcha was defined by the flat­ In the Hypoconchinae n. subfam. the uropods tened carapace, with dorsal surface partly mem­ show as ventral plates (not visible dorsally in both branous and soft, and by posterior legs ending inmales and females) and occur along with vestigial crescent-shaped dactylus (carapace more or less P13-P15 in males. The male pleopodal formula is inflated and calcified, and P4 and P5 reduced complete, such as in the Sphaerodromiinae and subcheliform inDromia, and only P5 modi­ n. subfam. (intercalary dorsal plates) and in the fied in Dynomene). The precise observations and Dromiinae n. statuspro parte (see Patterns of figures of Hypoconcha by Guérin-Méneville uropods and vestigial male pleopods 3-5; (1854: pi. 5, fig. 4) referred to the abdomen Table 1). A combination of characters similar to longer than the carapace and halfway flexed, the that found in the Hypoconchinae n. subfam., i.e. uropods as ventral plates, and the crescent and vestigial pleopods combined with ventral “retractile” dactylus on P4 and P5 to firmly hold uropods, occurs (with certainty) inDromidia, the shell. Exodromidia and Platydromia. Members of the subfamily Hypoconchinae In the Hypoconchinae n. subfam. neither the n. subfam. are easily recognized by a number of uropod nor any other apparent structure holds features: 1) carapace generally rounded, hour- the abdomen folded beneath céphalothorax. In glass-shaped; dorsal surface flattened, very thin H. californiensis, however, the PI coxa bears a and partly membranous; ecdysis area probably series of spiniform tubercules, the strongest one represented by whole branchiostegal region, at overhanging telson margin. As a result of being least; 2) front and lateral margins greatly expand­ disposed onto two planes, the abdomen appar­ ed, covering all parts of head, except antennal fla­ ently has its posterior part remaining pressed gella, and displacing eyes in ventral location; against thoracic sternum in normal flexion. How front semicircular or slightly truncate in outline; the curious keels on P1-P2 coxae ofH. cali­ 3) male abdomen widely triangular and flexed at forniensis (Fig. 19A) play role in holding of right angles in mid-length; 4) male pleopodal for­ abdomen is unknown. mula complete: P13-P15 generally as uniramous As described in H. arcuata by Kircher (1970: vestiges in males; 5) uropods showing always as figs 2c, 12e) and in H. parasitica, by Fang & ventral plates; 6) P4 and P5 very dissimilar, with Young (1980: 860, fig. 8A, D, H. as sabulosa peculiarly contorted and extremely mobile (Herbst, 1799)), the megalopa of the Hypo­ dactyli, which are fitted in deeply notched conchinae n. subfam. is the only one in the extremity of propodi (Guinot & Tavares 2000: family Dromiidae with a single, long, terminal 306, fig. 5); 7) condyle of propodi of P4 and P5 setum on dactylus of P5 (versus several feelers modified into prop-up plate, noticeably large, in the Dromiinae n. status, even inConchoe­ not concealed; gynglyme of propodi deeply cetes, see Sankolly & Shenoy 1968; Franco notched in order to receive propodal condyle; the 1998; Guinot & Tavares 2000). The single set prop-up plate/condyle blocking and prevent­ feeler on dactylus of P5 may well prove to be ing carpus from completely folding against another diagnostic character of the Hypo­ merus; 8) female thoracic sternites 7 and 8 tilted conchinae n. subfam. drastically, perpendicular in relation to preceding The larval and postlarval features (plesiomor- thoracic sternites; and 9) obligate carrier of a phies?) shared by Hypoconcha and Conchoecetes lamellibranch shell. (see McLay et al. 2001: 739, 744, table 2) are not In the Hypoconchinae n. subfam. the female exclusive to these genera and could not be regard­ sternal sutures are not as extended forward as in ed here as an indication of close relationship the Dromiinae n. status; the apertures of sper­ between Hypoconcha and Conchoecetes. Therefore,

100 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

the two genera are kept in different subfamilies, Hypoconchinae n. subfam. and Dromiinae n. status, respectively. Spears et al. (1992: 457) obtained interesting results from sequence-divergence estimates and phylogenies inferred by maximum parsymony analyses of 18S rRNA aligned sequences. As far as Hypoconcha [H. arcuata ] is concerned, their results suggested that “it seems unlikely that D rom idia [in fact Moreiradromia antillensis n. comb., Dromiinae n. status] andHypoconcha [Hypoconchinae n. subfam.], the [only] two dromiid genera used in this study, are as closely related as previously thought”. The uniqueness ofHypoconcha led McLay (1993: 229) to question whether it belongs or not to the Dromiidae. McLay (2001b: 8) expressed the opinion thatHypoconcha and Conchoecetes shared cx5 a common ancestor and belonged to the same particular grouping, whereas Desmodromia was to be kept amongst the other dromiids.Hypoconcha and Conchoecetes share a shell-carrying behaviour, but each with a different method of grapsing the shell (see under Shell-carrying behaviour) and (perhaps) some larval features (McLay et al. 2001). They also share several characters that is: thin tegument of dorsal carapace (partly membra­ nous inHypoconcha :), related to shell-carrying; strongly calcified and anteriorly truncated stern­ ite 4. A closer examination of the morphology of Hypoconcha and Conchoecetes reveals, however, that overall similarities are probably superficial Fig. 20. — Hypoconcha panamensis Smith, 1869;A, Panama, S 20 x 21 mm (MNHN-B 21597), coxa of P5, with mobile penial (Guinot & Tavares 2000) and that there is no tube; B, México, S 27 x 28 mm (EMU 2941), G2, without exo- reason to includeConchoecetes in the subfamily pod. Abbreviations:cx5, coxa of P5;pt, penial tube. Scale bars: 1 mm. Hypoconchinae n. subfam. The shape of male abdomen, the male uropods (salient dorsal uropods in Conchoecetes, Fig. 3B, C, narrow ventral pleopodal formula is found inHypoconcha (Fig. plates in Hypoconcha, Fig. 19B) and the condi­ 19B), but does not seem present inConchoecetes tion of sternites 7 and 8 (bent at right angles and(Fig. 3C). The similarities of thoracic sternum in bordered laterally by sutures 7/8 in female Hypoconcha and Conchoecetes are difficult to be Hypoconcha, Fig. 19A), the sternite 3 visible dor­ appraised: they are probably in close relationship sally in Conchoecetes (Fig. 3A, B) (hardly or not with the shell-carrying behaviour. In male visible dorsally in Hypoconcha, Fig. 19A), the Hypoconcha the well calcified, very flat sternite 4 location of spermathecal apertures (not very far (Fig. 19A) and the short abdomen which only from gonopores on P3 Hypoconcha in , Fig. 19A, occupies a posterior location (Rathbun 1937: between P2 in Conchoecetes, Fig. 3A) clearly dis­ pi. 11, fig. 2) are different from the condition of tinguish the two genera. A complete male Conchoecetes (Fig. 3A, B).

ZOOSYSTEMA • 2003 • 25 (1) 101 Guinot D. & Tavares M.

The taxonomic position ofDesmodromia (not GENERA INCLUDED. — Eodromia McLay, 1993 (type examined) is puzzling. As in Hypoconcha the cara­ species: Eodromia denticulata McLay, 1993 by mono­ typy); Sphaerodromia Alcock, 1899. pace, although not membranous on posterior half, is poorly calcified and flattened; the eaves over­ hang the eyes; and P4-P5 end in upturned dacty­ D e s c r ip t io n lus which supposedly holds a shell. Hypoconcha Carapace longer than wide or as long as wide, and Desmodromia differ from one another as fol­subglobose. Lateral margins subparallel; antero­ lows: 1) epipod present on PI (absent inDesmo­ lateral margin long, joining buccal cavern instead dromia)-, 2) narrow ventral uropods (dorsal and of exorbital angle, and separated from short pos­ well-developed in immature female ofD. tranterae terolateral margin by deep notch. Dorsal surface McLay, 2001); 3) female sternal sutures 7/8 ending with regions not defined or almost indistinct; rather posteriorly inHypoconcha (Fig. 19A) subhepatic area more or less inflated. Branchial (between P2 in Desmodromia)-, and 4) abdominal groove not marked. Front projecting well beyond holding never involving uropods and provided by orbits. Rostrum noticeably deflexed. Presence of structure on PI coxa, or, more usually, without two pseudorostral lobes extending uninterrupted­ coxal differentiation (provided by a differentiation ly around supraorbital margin. Orbits oblique, of P2 coxa involving uropods inDesmodromia). It deeply hollowed on lateral sides of carapace; is n o t kn o w n if inDesmodromia (such as in supra- and infraorbital margins entire, forming a Hypoconcha) the female thoracic sternites 7 and sort of eave, orbital border almost continuous. 8 are tilted, and if the male pleopodal formula is Proepistome widely triangular, in front of well- complete and the P5 coxa differentiated into pe­ defined epistome. Ocular stalk short and thick. nial mobile tube since males are unknown. TheAntennules with basal article strongly developed. lack of information on the morphology of maleAntenna: first article beak-like; basal article with abdomen and of details on the P4 and P5 grasping exopodal scale markedly developed, as long or apparatus make it difficult to compare it with the longer than following article; internal angle very specialized Hypoconcha. weakly or not produced. Mxp3 operculiform; Presently, the subfamily Hypoconchinae n. sub­ coxae approximated. fam. includes six species, all belonging to Thoracic sternum narrow. Gynglymes of thoracic Hypoconcha Guérin-Méneville, 1854:H. arcuata sternites 1-3 largely spaced from each other, Stimpson, 1858;H. californiensis Bouvier, 1898; stepped at lower plane. Sternites 1-3 not visible; H. lowei Rathbun, 1933; H. panamensis Smith, sternite 4 forming plate overhanging or just 1869; Cancer parasiticus Linnaeus, 1763; and touching bases of mxp3 (Fig. 21A, B). H. spinosissima Rathbun, 1933. Episternites 4 and 5 more or less elongated and The larval development is only known for twowide, their gynglymes in almost terminal loca­ species, H. parasitica (see Lang & Young 1980, as tion. When male abdomen flexed against ventral H. sabulosa) and H. arcuata (see Kircher 1970). It surface, anterior portion of sternite 4 and lateral is abbreviated to only three zoeal stages and a part (i.e. episternite 4) exposed, while episternite megalopa. We examined an ovigerous female 5 is completely covered by uropod and hardly or (6 mm length) ofH. parasitica (M N H N -B not visible at all (Fig. 21A). Sutures 4/5 and 5/6 28279) with less than 40 rather large eggs. very short, only lateral and not clearly visible; sutures 6/7 and 7/8 oblique. Female sternal sutures 7/8 short; apertures of spermathecae very Subfamily SPHAERODROMIINAE n. subfam. minute, behind level of P3 gonopore, located (Figs 21-24; 28E-G) laterally, and either completely exposed (Sphaerodromia pro parte, for example S. lamella­ Type GENUS. — Sphaerodromia Alcock, 1899 by pres­ ent designation (type species:Dromidia kendalli Alcock ta Crosnier, 1994), or concealed under the lateral & Anderson, 1894 by monotypy. Gender: feminine). heightening and fold of sternite 8Sphaerodromia (

102 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

cx1 cx1 e4

% -4/5, cx2 cx2

cx3

a6 cx3

cx4

Fig. 21. — Sphaerodromia ducoussoi McLay, 1991; A, B, Tuamotu, S 43.2 x 43 mm, holotype (MNHN-B 22172), thoracic sternum without and with abdomen;C, Tuamotu, Mururoa, 9 41.4 x 40.9 mm, paratype (MNHN-B 22173), spermathecae, behind level of P3. Abbreviations: a6, abdominal segment 6;cx1-cx4, coxae of P1-P4; e4, e5, episternites 4, 5; i, prominence on P3 coxa, covered by abdomen and inefficient for abdominal holding;o, female gonopore;pr, holding prominence;s, aperture of spermatheca; t, telson; u, uropod; 4-8,sternites 4-8;4/5-778, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 2.5 mm.

pro parte, for example S. ducoussoi McLay, 1991, 3-5 present, either biramous (Sphaerodromia , Fig. 21C; Eodromia , Fig. 24C). Fig. 22) or uniramous (Eodromia , Fig. 24A). Abdomen long but not reaching mxp3, once Male uropods as elongated dorsal plates, exposed folded; pleural parts well recognizable, all seg­ but deeply inserted between abdominal segment ments free; segment 6 noticeably expanded later­ 6 and telson (the base of which covering uropod), ally ( Sphaerodromia , Figs 21 A; 22; Eodromia , included in the outline of abdomen,viz. not real­ Fig. 24A, B); telson long. Vestigial male pleopods ly salient nor movable, occupying a relatively

ZOOSYSTEMA • 2003 • 25 (1) 103 Guinot D. & Tavares M.

B

a6

details

Fig. 22. — Sphaerodromia Alcock, 1899, biramous vestigial maie pleopods on segments 3-5 with details on each side; A,ducoussoi S. McLay, 1991, Tuamotu, Tuanake, 3 43.2 x 43 mm, holotype (MNHN-B 22172); B, S. nux Alcock, 1899, Madagascar, 3 60 x 67 mm (MNHN-B 6872). Abbreviations:a3, a6,abdominal segments 3, 6;t, telson;u, uropod; v3-v5, biramous vestigial pleopods 3-5. Scale bars: A, 2.5 mm; B, 2 mm. large portion of lateral margin of abdomen, play­fixed finger with marked proximal teeth and sev­ ing no role in holding of abdomenSphaero­ ( eral smaller ones. P2 and P3, very long, neither dromia, Figs 21 A; 22; Eodromia, Fig. 24A, B). lobed nor nodose; propodus very long, bearing Dimorphism of uropods marked. Female distal propodal spine. P4 and P5 reduced, shorter uropods deeply inserted, rather developed, occu­ than preceding ones, similar in size, oriented in a pying large part ( Sphaerodromia :) or whole part different way than P2 and P3, only P5 dorsal; {Eodromia) of abdominal external margin posteri­ subcheliform apparatus formed by multiple distal orly to telson, and well visible dorsally{Sphaero­ propodal spines opposing dactyli, three to five; dromia and Eodromia). no spines on outer propodal margin; presence of Holding of male abdomen not really efficient spines on inner margin of P4 and P5 dactylus. P5 when provided by granulous prominence on P2 without spines on outer dactylus margin. coxae involving telson in its anterior partMale coxa of P5 strongly modified, extended, {Sphaerodromia, where the abdomen is loosely without movable penial tube, in the two genera retained). Always, a prominence on P3 coxae, {Sphaerodromia, Figs 23A; 28E, F; Eodromia, covered by abdomen and inefficient (Fig. 2 IB). Fig. 28G). Epipod present on chelipeds; podobranch either Gl with well-developed apical plate. G2 long, present {Sphaerodromia) or absent{Eodromia). P2 with styliform flagellum, exopod present but of and P3 with epipods, with or without variable length {Sphaerodromia, Fig. 23B; pres­ podobranchs {Sphaerodromia) , or without ence to be verified inEodromia). epipods {Eodromia). Chelipeds stout, with fingers close along most ofCarrying behaviour length; dactylus with a large proximal tooth and Large pieces of sponges forSphaerodromia rest of prehensile margin very thin and smooth; (McLay & Crosnier 1991; McLay 1991, 1993;

104 ZOOSYSTEMA • 2003 • 25 (1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

Crosnier 1994); camouflage data not known in Eodromia.

R e m a r k s Differences between Sphaerodromia and other dromiids were already discussed (McLay & Crosnier 1991; McLay 1991, 1993). They concern the shape of carapace, the presence of cx5 podobranchs on pereopods, the presence of dis­ tal propodal spine on P2 and P3, and the arrangement of spines to form the subchelate mechanism of P4 and P5. The condition is similar in Eodromia , except that podobranchs and epipods on P2 and P3 are lost, an absence that must be regarded as a more advanced char­ acter state (McLay 1993: 131). A unique char­ acter of the Sphaerodromiinae n. subfam. is the long P2 and P3 propodus (with distal spine) and the not strongly curved dactylus. The propodus is shorter and without distal spine, and dactylus curved in the Dromiinae n. status. In the Sphaerodromiinae n. subfam. and Dromiinae n. status, the inner margin of dacty­ lus on P2 and P3 bears several spines (whereas it is smooth, or nearly smooth, in the Hypo­ conchinae n. subfam.). The study in this review of morphological struc­ tures often neglected before revealed that the sphaerodromiine genera share a combination of characters found nowhere else within the Dromiidae. In Sphaerodromia and Eodromia , the coxa of P5 is extended to form conical expansion in which the penis is completely enclosed. The coxa and penis thus form a single structure (Figs 23A; 25B, D; 28E-G). In contrast, in the Dromiinae n. status (Fig. 28H-J) and Hypo­ conchinae n. subfam. (Figs 20A; 28K) the male Fig. 23. — Sphaerodromia nux Alcock, 1899, Madagascar,6 60 x coxa of P5 is not modified to enclose penis and67 mm (MNHN-B 6872);A, coxa of P5, modified and elongat­ ed into process enclosing penis and not forming penial tube; there are two independent structures: the B, G2, with long exopod. Abbreviations:cx5, coxae of P5; unmodified P5 coxa and the long, sclerotized e, elongation of P5 coxa enclosing penis. Scale bars: 1 mm. penis emanating from male gonopore. The dromiine and hypoconchine movable structures, in the vicinity of female gonopores on P3, that is, here named “penial tube”, end in soft tip (see behind level of P3. In Sphaerodromia lamellata , Patterns of P5 coxa and penis; Fig. 28). the suture 7/8 is shorter than inS. ducoussoi The Sphaerodromiinae n. subfam. (Figs 21C; (Fig. 21C) or S. kendalli (Alcock & Anderson, 24C) have short female sternal sutures 7/8, so 1894), and the spermathecal aperture lies posteri­ the apertures of spermathecae lie, always laterally, orly and is not concealed under raised part of

ZOOSYSTEMA • 2003 • 25 (1) 103 Guinot D. & Tavares M.

U

cx2

cx3

cx4

cx5

Fig. 24. — Eodromia denticulata McLay, 1993, Indonesia, Kai Islands, Tanimbar Island, KARUBAR (MNHN-B 26327); A, B, S 4.9 x 4.5 mm, abdomen, ventral and dorsal views; note elongated uniramous vestigial pleopods on segments 3-5; C, ovigerous 9 5 x 4.5 mm, thoracic sternum and spermathecae, with aperture at level of P3. Abbreviations:a3-a6, abdominal segments 3-6; cx2- cx5, coxae of P2-P5; o, female gonopore;s, aperture of spermatheca; t, telson;u, uropod; v3-v5, vestigial pleopods 3-5; 6-8, stern­ ites 6-8; 6/7, 7/8,thoracic sternal sutures 6/7, 7/8. Scale bars: 0.5 mm.

106 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

sternite 8. Conversely, all the Dromiinae n. status number of characters unique among the show extremely long female sutures 7/8, so that Dromiidae. The Sphaerodromia species are rela­ the spermathecae open far beyond level of P3, tively large (width more than 65 mm inS. nux sometimes at level of PI or beyond. Therefore, in Alcock, 1899), whileEodromia denticulata the dromiine females the sternites 7 and 8 occupy McLay, 1993 is very small, with ovigerous much of the ventral surface of céphalothorax, and females measuring only 4.5 mm width. In the thoracic sternum appears dramatically dis­ Sphaerodromia and Eodromia the basal antennal torted. The only exception is Stebbingdromia article bears a long exopodal scale, extending plumosa n. comb., where the spermathecae open “beyond joint of segments three or four” between P3 (Fig. 17C). In the dromiine genera (Me Lay 1993: 127), and its internal corner is the spermathecae open apart or together, each not or only weakly produced. spermathecal aperture being often positioned onIn Sphaerodromia (Fig. 21 A, B) and Eodromia, more or less prominent tubercle; in a few generathe long male abdomen covers most part of tho­ the spermathecae open on a single tubercle. It is racic sternum, except anterior part of sternite 4 worth noting that the position of the apertures ofwhich is in contact with mxp3. InSphaerodromia spermathecae in relation to female gonopores on(Figs 21A; 22) and Eodromia (Fig. 24A, B), the P3 coxae and the kind of penial structure seem to male abdominal segment 6 is expanded laterally. be related. Thus, in considering the sphaero- In Sphaerodromia and Eodromia, no specialized dromiine condition, the location of the aperturesstructures are found for an efficient abdominal about level of coxae of P3 seems to be connected holding, so the male abdomen is rather loosely with a short and non-articulated penis. retained beneath céphalothorax (the granulous Conversely, a spermathecal aperture positioned prominences found on P2 and P3 coxae are inef­ far beyond coxa of P3 appears to be connected ficient). with a long, sclerotized and movable penis Sphaerodromia and Eodromia share the subcheli- (penial tube). This condition is found in the form system formed by multiple distal propodal Dromiinae n. status (again, the only exception is spines on the dactyli. The minute apertures of the atypical Stebbingdromia plumosa n. comb., spermathecae, at level of P3, are more or less con­ which has short female sutures 7/8 and penial cealed under fold of sternite 8. tube; Figs 17A, C; 18A). Vestigial pleopods are always present in male The Hypoconchinae n. subfam. show a different Sphaerodromiinae n. subfam.: they are biramous combination: sternal sutures 7/8 relatively shortin Sphaerodromia (Fig. 22) and uniramous in (Fig. 19A) and the presence of a penial tube Eodromia (Fig. 24A). In male Sphaerodromiinae (Figs 20A; 28K). In the Hypoconchinae n. sub­ n. subfam. P13 to P15 occur with immovable fam. the female sternal sutures 7/8 are slightly uropods showing as intercalary dorsal plates. This extended forward, more than in the combination of characters (P13-P15 combined Sphaerodromiinae n. subfam. (where they are with dorsal uropods) is known from only a few behind level of P3) but not so far forward as in genera and species of Dromiinae n. status: the Dromiinae n. status. In this respect D rom ia pro parte (Fig. 27), Moreiradromia Dromiinae n. status and Hypoconchinae n. sub­n. gen. (Fig. 14B) and Stebbingdromia n. gen. fam. differ less from one another than from(Fig. 18C; Table 1). Sphaerodromiinae n. subfam. The G l of Sphaerodromia is provided by apical The Sphaerodromiinae n. subfam. also shares the plate, and the G2 (Fig. 23B) with exopod. absence of spines on outer margin of propodus ofMcLay (1993: 132) indicated that the first pairs P4 and on outer margin of dactylus of P5. of pleopods of the male paratype ofEodromia W ithin the Sphaerodromiinae n. subfam., denticulata (8.2 X 7.8 mm; MNHN-B 22545) Sphaerodromia and Eodromia seem to be very were “not properly developed”. This specimen closely related to one another as they share a has several pairs of pleopods and a P5 coxa not

ZOOSYSTEMA • 2003 • 25 (1) 107 Guinot D. & Tavares M.

extended: it seems to be an abnormal individual. 94, fig. 8), which is in accordance to the ple- In a male specimen (4.5 mm width; MNHN-B siomorphic condition of the Sphaerodromiinae 26327), the P5 coxa is elongated and both Gl n. subfam. and G2 are developed; an apical plate is clearly About the status ofSphaerodromia lethrinusae present on Gl as inSphaerodromia, whereas an Takeda & Kurata, 1976, see under Dromidiopsis exopod is not discernible on the G2, which is sensu nobis. typically prolonged by styliform flagellum. W hether Parasphaerodromia Spiridonov, 1992 Presently, the subfamily Sphaerodromiinae belongs to the Sphaerodromiinae n. subfam. is n. subfam. consist of six species including among not certain but seems unlikely. Based on the orig­ two genera: 1) Sphaerodromia (Sphaerodromia inal description ofParasphaerodromia subglobosa brizops McLay & Crosnier, 1991;Sphaerodromia Spiridonov, 1992, type species and the only rep­ ducoussoi ; Dromidia kendalli-, Sphaerodromia resentative of the genus, McLay (1993: 122, 183, lamellata-, Sphaerodromia nux)-, 2) Eodromia 184) synonymized the species with Dromidia (Eodromia denticulata). spongiosa, nowPlatydromia spongiosa n. comb, The close relationship betweenSphaerodromia (see above, Fig. 15), but furnished no arguments and Eodromia, both with many ancestral charac­ for doing so. As a result,Parasphaerodromia is teristics, was already noted by McLay (1993: 130, merged into Platydromia Brocchi, 1877. The 228). McLay et al. (2001: 741, table 3) recently synonymy between Parasphaerodromia subglobosa predicted that the larvae of these two genera Spiridonov, 1992 andPlatydromia spongiosa should also have primitive features. McLay n. comb, shall very likely to be confirmed in the (1993: 131) noted that the absence of epipods on future (see above, under Platydromia). O ur view P2-P3 and of podobranch on chelipeds must be is that the establishment of dromiid taxa necessi­ regarded as the more advanced character states of tates the adequate description and illustration of Eodromia. essential characters such as the thoracic sternum Our investigation suggests that the Sphaero­ and uropods; details of the carapace and legs dromiinae n. subfam. is a basal group within the alone are usually insufficient. Dromiidae. Several sphaerodromiine plesiomor- phic characters, that is, a male P5 coxa modified and extended in a process (in contrast to a mobile DISCUSSION penial tube in the Dromiinae n. status and Hypoconchinae n. subfam.), the male vestigial The status of the genus Frodrom ia M c L a y , pleopods on abdominal segments 3-5, the short 1993 (F ig s 25; 26) female sutures 7/8, and the apertures of sper­ T he genus Frodromia, which consists of two mathecae positioned near female gonopores onspecies (Petalomera atypica T. Sakai, 1936, type P3 coxae, are also found in the two other families species by original designation; andPetalomera of Dromiacea, Homolodromiidae Alcock, 1900 reticulata T. Sakai, 1974), is not more related to and Dynomenidae Ortmann, 1892. An exopod the Sphaerodromiinae n. subfam. than to present on G2, sometimes well-developed Dromiinae n. status. The sphaerodromiine char­ (,Sphaerodromia nux, Fig. 23B) or shorter acters o f Frodromia are as follows: elongated (S. brizops, S. lamellata), is shared with the Dyno­ shape of carapace, with subparallel lateral borders menidae, but not with the Homolodromiidae. (see McLay 1993: figs 6a, 17d); long male telson It is worth noting that the spermatozoal ultra­(Fig. 25A, C); vestigial male pleopods 3-5 present structure of Sphaerodromia (5. lamellata) allies the (Fig. 25C); male uropods showing as deeply genus more closely to the dynomenidM eta­ inserted and obliquely oriented dorsal plates; dynomene tanensis (Yokoya, 1933) than to the short female sternal sutures 7/8; apertures of sper­ advanced dromiid (dromiine) Stim d ro m ia mathecae lying not very far from P3 coxae, i.e. lateralis (Gray, 1831) (Guinot et al. 1998: 91, 93, close to female gonopores on P3 (Fig. 26); male

108 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

If cx3

Fig. 25. — Frodromia atypica (Sakai, 1936), New Caledonia, MUSORSTOM 4, CP 171, S 10.6 x 9.5 mm (MNHN-B 22559);A, B, tho­ racic sternum, with and without abdomen;C, abdomen, ventral view; note uropods showing as intercalary dorsal plates and elon­ gated uniramous vestigial pleopods on segments 3-5;D, P5 coxa, modified and elongated into process enclosing penis and not forming penial tube. Abbreviations:a3-a6, abdominal segments 3-6; cx1-cx5,coxae of P1-P5;e, elongation of P5 coxa enclosing penis; e4, e5, episternites 4, 5; mxp3, external maxilliped; pr, holding prominence on P1 and P2;t, telson;u, uropod; v3-v5, vestig­ ial pleopods 3-5; 1-3, sternites 1-3; 4-8, sternites 4-8; 4/5-7/8, thoracic sternal sutures 4/5-7/8. Dotted line indicates difference in level. Scale bars: 1 mm.

P5 coxa strongly modified, without “articulated”, (see McLay 1993: fig. 6d); presence of a promi­ movable penial tube (Fig. 25D); P2 and P3 with nence on P3 coxae, covered by abdomen, but long propodus and not strongly curved dactylus inefficient for abdominal holding.

ZOOSYSTEMA • 2003 • 25 (1) 109 Guinot D. & Tavares M.

Nevertheless, Frodromia is distinguished from the and whether or not this genus belongs to the Sphaerodromiinae n. subfam. by several charac­ Sphaerodromiinae n. subfam. deserves further ters: two pseudorostral teeth, one at each side of investigation.Frodromia seems apart within the rostrum (two pseudorostral lobes extending unin­Sphaerodromiinae n. subfam., but several of its terruptedly around supraorbital margin in features (such as loss of apical plate on G l and Sphaerodromia and Eodromia)-, sutures 7/8 almost exopod on G2, reduction of spines of P4 and P5 longitudinal (oblique inSphaerodromia and subcheliform apparatus, efficiency of abdominal Eodromia)-, exopod of antenna with internal angleholding) may be regarded as more advanced char­ produced (weakly or not produced in acter state, such as features of Eodromia com ­ Sphaerodromia and Eodromia)-, mxp3 with coxae pared with those of Sphaerodromia. It should be not approximated (approximated in Sphaero­ noted that Frodromia is the only case in the drom ia and Eodromia)-, sternites 1-3 visible Dromiidae where the female gonopore on P3 (Fig. 25A) (not exposed in Sphaerodromia and coxa opens very close to anterior border of coxa Eodromia)-, sternite 4 covered by abdomen when and at summit of tubular process, which is proxi- flexed (exposed in Sphaerodromia and Eodromia)-, mally armed with strong spur (Frodromia atypica, apertures of spermathecae raised, each on a tuber­ Fig. 26; F. reticulata, not examined, condition to be cule, and visible (Fig. 26) (often concealed in verified). In Frodromia atypica the P3 coxa is dimor­ Sphaerodromia)-, abdominal segment 6 not phic since the male does not possess such a process. expanded laterally (Fig. 25A, C) (noticeably expanded in Sphaerodromia and Eodromia)-, P a t t e r n s o f u r o p o d s a n d v e s t ig ia l m a l e abdominal holding efficient, by means of marked p l e o p o d s 3-5 granulo us prominence on PI coxae acting at baseUnlike other Decapoda, the uropod is never bi- of telson and of another prominence on P2 coxaeramous in the Brachyura (Podotremata, Hetero­ acting with (probably mobile) uropods (Fig. 25A, C) tremata and Thoracotremata). The brachyuran (abdominal holding not really efficient, noturopod shows a wealth of morphological varia­ involving uropods inSphaerodromia and tion, even if its primary function is related to the Eodromia)-, podobranchs absent on pereopods abdominal holding against thoracic sternum. (present in Sphaerodromia but absent in In the majority of the families of the Hetero­ Eodromia)-, fingers of chelipeds with prehensile tremata and Thoracotremata (Eubrachyura) the margins toothed (fingers of chelipeds, each with uropod shows as a socket situated on ventral sur­ one marked proximal tooth and close along mostface of abdominal segment 6 and is homologous of length of prehensile margins, and no otherto the ancient uropod. The socket is the hollow marked teeth on dactyl margin in Sphaerodromia part of the press-button mechanism (Guinot & and Eodromia)-, P2 and P3 without distal propo­ Bouchard 1998: 682, table 3). Conversely, the dal spine (with a distal propodal spine in uropod is diversely modified in the Podotremata. Sphaerodromia and Eodromia)-, subchelate mecha­ Always uniramous, it shows either as a ventral nism of P4 and P5 with dactyli opposed by one lobe or plate (Homolodromiidae, Dromiidaepro (on P4) or two (on P5) propodal spines (multipleparte), or as a calcified dorsal plate (Dyno­ distal propodal spines opposing dactyli, three to menidae, Dromiidae pro parte), or as a socket five, in Sphaerodromia and Eodromia)-, Gl sharply (Homoloidea, Lyreidinae), all homologous. The tipped (a well-developed apical plate in term “vestigial uropod” is mistaking as this struc­ Sphaerodromia)-, G2 without exopod (with exo­ ture actually represents a wealth of character pod, at least in Sphaerodromia). The camouflage states (Guinot & Tavares 2001: fig. 15). The of Frodromia is provided by ascidians (Sakai uropod is completely lost in the podotreme fami­ 1936; McLay 1993). lies Cymonomidae Bouvier, 1897, Phyllo­ The combination of characters ofFrodromia does tymolinidae Tavares, 1998, Cyclodorippidae not secure its position in any dromiid subfamily, Ortmann, 1892, and in the Raninoidea de Haan,

110 ZOOSYSTEMA • 2003 • 25(1) New sub familial arrangement for the Dromiidae (Crustacea, Decapoda)

cx1

cx2

m

cx3 '7/8

cx4

cx5

cx3 cx3 sr

Fig. 26. — Frodromia atypica (Sakai, 1936), New Caledonia, MUSORSTOM 4, CP 171, 9 8.5x 7.3 mm (MNHN-B 22560), thoracic sternum and sperm athecae with two detailed views (lower) of coxa of P3; note apertures of spermatheca behind level of female gonopores located near anterior border of coxa, at summit of tubular process. Abbreviations:cx1-cx5, coxae of P1-P5;o, female gonopore;s, aperture of spermatheca; sr, spur of P3 coxa; tb, tubular process of P3 coxa, with gonopore at its extremity;4-8, ster­ nites 4-8;4/5-7/8, thoracic sternal sutures 4/5-7/8. Scale bars: 1 mm.

1839 (Lyreidinae Guinot, 1993 excepted). The 1977 sensu Guinot & Tavares 2001 (Cymo­ total loss of the uropod may be interpreted as a nomidae, Phyllotymolinidae, Cyclodorippidae, synapomorphy of the Archaeobrachyura Guinot, Raninoideapro parte).

ZOOSYSTEMA • 2003 • 25 (1) 111 Guinot D. & Tavares M.

The dromiacean uropod is generally simply tively. In Table 1 we consider the uropods of referred as “vestigial”. Such an oversimplification Dromidia, Lamarckdromia n. gen. andHypo­ does not reflect the diversity of uropod patterns concha as ventral. found within the group. The dynomenid uropod In dromiids the uropods are only rarely com­ always shows as a well-developed dorsal plate, pletely lost (as in Ascidiophilus). There are no sexually dimorphic, not markedly prominent, known cases of an uropod modified into socket. even in males, and not efficient in maintainingThe uropodal plate, whether dorsal, intercalary the broad abdomen. or ventral, is the transformation of the biramous The dromiid uropod is more diverse. It often uropod of the megalopa. The role of uropod in shows as a calcified and exposed dorsal plate, with abdominal holding deserves further investigation, a marked sexual dimorphism in terms of size and the lobe or ventral plate being too ventral to be shape. This plate is placed between segment 6 truly efficient. Conversely, uropods as dorsal and telson. It does not have the same place or plates have generally proved to be very efficient, orientation than a vestigial male pleopod thanacting together with coxal structures on thora- may remain on preceding segments. Among copods. The evolution of the biramous uropod dromiids, the dorsal uropods are more or less into dorsal plate is likely to represent a specialized salient and movable in males (and immature structure functionally adapted to act in the hold­ females) and very often play major role in the ing of abdomen, even though not used by all abdominal holding. The dromiid uropod can also species. show as a ventral plate, more or less developed Information on the kind of uropods is still lack­ and calcified, never visible dorsally in either sex, ing for a number of dromiid genera, and the pres­ sometimes very reduced and resembling a soft ence of uropods still needs confirmation in a few lobe (Austrodromidia australis, Fig. 1C), or some­ genera. In Tunedromia McLay, 1993 the uropods times indistinct (A. octodentata, Fig. 2B). are indicated as absent in both sexes (Takeda The dromiid uropod can also show as a calcified 2001: 220). The uropods are absent in females of platelet, obliquely oriented, and so deeply insert­ Epipedodromia André, 1932, but the male condi­ ed ventrally that it is no longer markedly salient, tio n is unknow n. InAustrodromidia M cLay, but still exposed dorsally ( Stebbingdromia n. gen., 1993, the uropods have been reported as Fig. 17B; Sphaerodromia, Figs 21A; 22; “reduced and concealed or absent” (McLay 1993: Eodromia, Fig. 24A, B; Frodromia, Fig. 25A, C). 185). Actually, the uropods ofA. australis, type This condition is herein named “intercalary species of Austrodromidia, and ofA. octodentata platelet” and has been indicated as a dorsal uro­ have not been properly described. InA. australis pod in Table 1. Even though visible dorsally, the uropods are very small ventral plates (Fig. IB, C), intercalary platelets are not involved (or only whereas in A. octodentata they show as indistinct, weakly so) in abdominal holding. nearly obsolete ventral plates (Fig. 2B), perhaps In some cases the differentiation between inter­an indication thatAustrodromidia is heteroge­ calary platelets and ventral plates is not easy. In neous. Dromidia hirsutissima (Fig. 5B, C) for instance, It is actually important that the uropod condition the uropod is oblique and deeply inserted ven­ and the male pleopodal formula be determined trally, actually quite similar to the intercalary jointly in the Dromiacea. It is still not known if platelet condition, but hardly visible dorsally. In the male P13-P15 remain as vestiges in many basal this particular, the uropod of D. hirsutissima Podotremata (plesiomorphy). The vestiges are: could be regarded as intermediate between dorsal 1) always biramous in Dynomenidae (uniramous and ventral plate. The ventral uropods of in Dynomene praedator A. Milne Edwards, 1879) Lamarckdromia globosa n. comb. (Fig. 10B, C) and in Dromiidaepro parte (Sphaerodromia)-, and and Hypoconcha spp. (Fig. 19B, C) are also very 2) uniramous in Homolodromiidae and different, rounded and narrow/oblique respec­ Dromiidae pro parte (at various stages of vestigial

112 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

T able 1 . — Pleopodal formula In the dromiid genera, Including the condition of the uropod, dorsal, ventral or absent. Genera In bold have a complete pleopodal formula. The condition Conchoecetes In needs to be verified. *PI3-PI5 are present InExodromidia spinosa, while only PI5 seems to be present InE. spinosissima and £ bicornis. The enigmatic genusFrodromia Is not listed.

Uropod as dorsal plate Uropod as ventral plate Uropod absent Male vestigial pleopods

D r o m iin a e n . s t a t u s Alainodromia absent Ascidiophilus absent Austrodromidia absent 7Barnadromia ? Conchoecetes absent Cryptodromia absent Cryp todromiopsis absent Desmodromia ? Drom ia absent or present Dromidia present Dromidiopsis absent Epigodromia absent 7 Epipedodromia ? Eudromidia ? Exodromidia present* Fultodromia absent Haledromia ? Hemisphaerodromia ? Homalodromia absent Lamarckdromia n. gen. absent Lauridromia absent Lewindromia n. gen. absent Mclaydromia n. gen. absent Moreiradromia n. gen. present Paradromia ? Petalomera absent Platydromia present Pseudodromia absent Speodromia ? Sternodromia absent Takedromia absent 7Tunedromia ? Stebbingdromia n. gen. present Hypoconchinaen. subfam. Hypoconcha present Sphaerodromiinaen. subfam. Eodromia present Sphaerodromia present

condition). The male pleopods 3-5 have been lost example in Stebbingdromia plumosa n. comb. in a number of Dromiidae and in all remaining(Fig. 18C). Podotremata (Homolidea and Archaeobra- Because not enough attention has been paid to chyura sensu Guinot & Tavares 2001). the male pleopodal formulae, the character The present study revealed the presence of vestig­ remains unknown in a number of dromiid gen­ ial male P13-P15 in several dromiid species. era: 1) Barnardromia (uropods reduced, not visi­ Because P13-P15 are frequently m uch reduced, ble); 2) Eudromidia (uropods very reduced, not sometimes to short papillae, they are easily over­ visible dorsally); 3) Haledromia (uropods very looked and remain undescribed. Rudimentary reduced, concealed); 4) Speodromia (uropods as P13-P15 are often dissimilar in length, as for ventral plates); 5) Epipedodromia (u ropods

ZOOSYSTEMA • 2003 • 25 (1) 113 Guinot D. & Tavares M.

details

a6

¿ o v3

Fig. 27. — Dromia Weber, 1795, uropods and vestigial male pleopods on segments 3-5 with details on each side;Dromia A, personata (Linnaeus, 1758), Cape Fréhel,S 47 x 59 mm (MNFIN-B 22049); B, D. bollorei Forest, 1974, Ivory Coast,S 71.5 x 85 mm, paratype (MNHN-B 21932). Abbreviations: a3-a6,abdominal segments 3-6; t, telson; u, uropod; v3-v5, vestigial pleopods 3-5. Scale bars: A, B, 5 mm; details, 1 mm.

indicated as absent); and 6) Tunedromia (uropods combined with dorsal plates are as follows: absent). In these genera (not examined) the Dromia pro parte, i.e., in D. personata (Linnaeus, presence of vestigial pleopods has not been inves­1758) (Fig. 27A),D. bollorei Forest, 1974 tigated. (Fig. 27B), D. marmorea Forest, 1974, and also When information on the type of uropod andin “Drom ia” wilsoni (Fulton & Grant, 1902), male pleopodal formula are combined, four pat­ Moreiradromia n. gen. (M. antillensis and terns emerge in the Dromiidae (Table 1): 1) ves­ M. Sarraburei, Fig. 14B, C), and Stebbingdromia tigial pleopods P13-P15 com bined w ith dorsal n. gen. {S. plumosa n. comb., Fig. 18C). plates; 2) P13-P15 combined with ventral plates; Vestigial pleopods combined with ventral 3) presence of uropods and no P13-P15; and 4) all uropods (i.e., a complete male pleopodal formu­ abdominal appendages completely lost. We la) occur in the Hypoconchinae n. subfam. found no case so far of vestigial pleopods occur­CHypoconcha, Fig. 19B) and (with certainty) in ring in the absence of uropods. only a very limited number of genera and species The condition P13-P15 combined with dorsal of the large subfamily Dromiinae n. status, as fol­ plates has been found in all Sphaerodromiinae n. lows: 1) in the monospecificDromidia (.D. hir­ subfam. but only in a few Dromiinae n. status. In sutissima, Fig. 5B, C); and 2) in Exodromidia the male Sphaerodromiinae n. subfam., P13-P15 spinosa (Studer, 1883), type species of occur along with intercalary plates, which are Exodromidia Stebbing, 1905 (p. 64). The other exposed dorsally. Pl3-Pi5 are biramous in two species ofExodromidia, E. spinosissima Sphaerodromia (Fig. 22), uniramous in Eodromia (Kensley, 1977) andE. bicornis (Studer, 1883) (Fig. 24A); they are also uniramous in Frodromia only have a long Pl5, whereas P13-P14 seem to be (Fig. 25C). The genera and species of Dromiinae absent (the inclusion of these two last species in n. status with (uniramous) vestigial pleopods Exodromidia has been already questioned; see

114 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

Guinot 1995: 187; Guinot & Bouchard 1998:special condition in the Leucosiidae Samouelle, 624; Bouchard 2000). 1819, see G uinot 1979a: fig. 45). In contrast, The present study of Pseudodromia and the Podotremata exhibit several markedly different Ascidiophilus has confirmed the absence of vestig­ patterns of coxal morphology, related to sperm ial male pleopods as well as the presence of ven­ transfer. tral uropods inPseudodromia, and the complete The Homolodromiidae and Dynomenidae share loss of both P13-P15 and uropods inAscidiophilus. a totally modified P5 coxa, which is modified in a The uropod condition remains vague in the continuous hard extension, simply enclosing dromiid genera whose uropods are described in penis which ends in small extrusion formed by the literature as “concealed” (see above and Table 1). the soft extremity of vas deferens (Guinot 1978: W ithin family Dromiidae a complete male 231; 1979a: 191, 194, figs 43C, G, 63A, B; pleopodal formula occurs in the Sphaero­ 1995: 174, fig. 4; McLay 1999: 460, fig. I4f). dromiinae n. subfam. (intercalary platelets, This first type of modification involves an elon­ exposed dorsally), in the Hypoconchinae n. sub­ gation at different degrees of the P5 coxa. As fam. (ventral narrow uropods), and in a very lim­ shown in Fig. 28, the modification involves ited number of Dromiinae n. status (Dromiinae either the whole coxa, which is more or less regu­ pro parte, see Table 1). larly tapered (Dynomenidae, Fig. 28C, D), or only anterior part, which is abruptly constricted P a t t e r n s o f P5 c o x a a n d pe n is just anteriorly to P5 articular condyle on thoracic Several morphological structures remain over­ sternite 8 (Homolodromiidae, Fig. 28A, B). The looked or very poorly investigated in the extension of the coxa takes different shapes Brachyura. Among these is the coxa of P5 in the which, in the case of the Homolodromiidae and male with its associated penis (Guinot 1978, Dynomenidae, is family specific. In the 1979a, b). In the Brachyura the male gonopore Homolodromiidae there is a reduced P5, opens on coxa of last thoracic pereopodcompletely rejected dorsally (P4 is only subdor­ (Podotremata and Hetrotremata) or on thoracicsal), whereas in the Dynomenidae there is an sternum (Thoracotremata). The vas deferens obliquely directed and very small last pereopod. always prolongs itself into penis, whose role is toIn Homolodromia A. Milne Edwards, 1880 penetrate inside the base of male first pleopod, or (Homolodromiidae) the tubular process is spe­ gonopod. Conversely to the podotreme condi­cially long and hard. The details of the P5 coxa tion, with a single basal opening, or foramen, for(length, direction) seem useful to separate the the introduction of both penis and second gono­dynomenid genera. pod, in the Eubrachyura (Hetrotremata and Two substantially different patterns were found Thoracotremata) the completely closed first during this study in the family Dromiidae, gonopod is provided with two distinct basal opposing the Sphaerodromiinae n. subfam. to foramina: one for the insertion of penis (“fora­the Dromiinae n. status and Hypoconchinae men proximal externe”) and the other for the n. subfam. Our view is that both patterns insertion of male second pleopod (“foramenevolved independently and thereby are not proximal interne”) (Guinot 1979a: 239, figs 60- homologous. 68). That supports the monophyly of the non-The two sphaerodromiine generaSphaerodromia podotreme brachyurans. (Figs 23A; 28E, F) and Eodromia (Fig. 28G), and The Eubrachyura show a rather conservative also Frodromia (Fig. 25B, D) share with the condition for the male P5 coxa, while diverse Homolodromiidae and the Dynomenidae a mod­ modalities for the protection of penis exist (see ified and elongated P5 coxa that encloses the Guinot 1969a-c, 1979a, b; Hartnoll 1975;penis. The Sphaerodromia pattern is very close Tavares 1992, 1997). The coxa itself is only to that found in the Homolodromiidae slightly modified or not modified at all (for the (Fig. 28A, B): a marked constriction of the coxal

ZOOSYSTEMA • 2003 • 25 (1) 115 Guinot D. & Tavares M.

Fig. 28. — Patterns of male coxa of P5 in the Dromiacea de Haan, 1833;A-G, pattern with coxa modified and elongated into process enclosing penis and not forming penial tube;A, B, Homolodromiidae Alcock, 1900;A, Homolodromia paradoxa A. Milne Edwards, 1880, Bahamas, Gerda, stn 190, 6 24 x 15 mm (RMNH); B, Dicranodromia mahieuxii A. Milne Edwards, 1883, Bay of Biscay, Le Travailleur, stn 9, S 9 x 6.5 mm, holotype (MNHN-B 21681);C, D, Dynomenidae Ortmann, 1892;C, Dynomene hispida Guérin- Méneville, 1832, New Caledonia, S 8.8 x 11.5 mm (MNHN-B 22091); D, Paradynomene tuberculata Sakai, 1963, New Caledonia, 11 mm width (MNHN-B 24780); E-G, Sphaerodromiinae n. subfam. (Dromiidae);E, Sphaerodromia ducoussoi McLay, 1991, Tuamotu, S 43.2 x 43 mm, holotype (MNHN-B 22172); F, Sphaerodromia lamellata Crosnier, 1994, New Caledonia,S 47.8 x 42.6 mm, holotype (MNHN-B 24724); G, Eodromia denticulata McLay, 1993, Kai Islands, KARUBAR, S 4.9 x 4.5 (MNHN-B 26327); H-K, pattern with coxa not modified but prolonging into mobile penial tube;H-J, Dromiinae de Haan, 1833 n. status (Dromiidae); H, Moreira­ dromia antillensis (Stimpson, 1858) n. comb., French Guiana, stn S 33, 1 8 x 1 8 mm (MNHN-B 22030); I, “Dromia” wilsoni (Fulton & Grant, 1902), Guézé coll., S 22 x 37 mm (MNHN-B 26254); J, Lauridromia dehaani (Rathbun, 1923), Gulf of Suez, Monod det. Dromia dormia,S 42 x 45 mm (MNHN-B 21998); K, Hypoconchinae n. subfam. (Dromiidae),Hypoconcha californiensis Bouvier, 1898, Gulf of California,S 20.3 x 19.4 mm, syntype (MNHN- B 22066). Corneous regions stippled. Scale bars: A, D-F, H-K, 1 mm; B, C, G, 0.5 mm. article just anterior to articular condyle of P5 Since the extended coxa is a sexual modification coxa, forming a narrow process, well demarcated of the male, the female does not have a modified from the rest of the coxa. coxa. An elongated P5 coxa is therefore a unique

116 ZOOSYSTEMA • 2003 • 25 (1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

characteristic of the male in the Dynomenidae, n. status there are long female sternal sutures 7/8 Homolodromiidae and Sphaerodromiinae and the apertures of spermathecae are located far n. subfam. anteriorly than the female genital openings on P3 The remaining Dromiidae, that is, the Dromiinae coxae. As in these dromiines the spermathecal n. status (Fig. 28H-J) and Hypoconchinae apertures end on a more or less prominent tuber­ n. subfam. (Figs 20A; 28K), fall into the second cule, it is possible that mating requires a parti­ pattern sharing an ordinary unmodified P5 coxa. cular arrangement of the male copulatory system, The vas deferens protrudes from gonopore to involving both gonopods and the penis. The only form long, sclerotized tube, which is completely exception is the atypical dromiine Stebbingdromia independent from coxa and movable; from tube plum osa n. comb., with its long penial tube emerges the soft tip of penis. InDromia person­ (Fig. 18A) and short female sutures 7/8 ending ata, for example, the soft male vas deferens passes near gonopores on P3 (Fig. 17C). The male G2 through the foramen in the coxa, becoming pro­of Stebbingdromia plumosa n. comb., stout and tected by a thick cuticle (Hartnoll 1975: 665, without needle-like flagellum (Fig. 18B), repre­ figs IA, 4A). It is not a simple extrusion or évagi­ sents a unique pattern that differs from that of all nation of the vas deferens, since it becomes a dif­ other Dromiidae and all Dromiacea as well. ferentiated and movable independent structure, Unlike the Dromiinae n. status, the Hypo­ supported by long calcified sclerite. We propose conchinae n. subfam. combines a penial tube to name it “penial tube”. The external sclerotized (Figs 20A; 28K) and short female sutures 7/8 penial tube is unique to members of the subfami­ that end near gonopores on P3 (Fig. 19A). lies Dromiinae n. status and Hypoconchinae In the Dromiinae n. status and Hypoconchinae n. subfam., and constitutes one of longest penialn. subfam. the penial tubes are always symmetri­ structure ever found in the Podotremata in par­ cal, and in the numerous species that were exam­ ticular and in the Brachyura in general. The only ined no significant differences were observed in other brachyuran crabs with a long penial tube is the calcification of the penial tube. the family Dorippidae MacLeay, 1838, where the It is evident that either the modification of the P5 vas deferens arises from coxal gonopore to form acoxa or the formation of an independent penial tube which may remain exposed and sclerotized, tube in the Podotremata is tied with the marked as in the subfamily Dorippinae (see Guinot change in the alignment of the arthrodial cavities 1979a: 195, fig. 46F; 1979b: figs 2, 3). Amongst of posterior pereopods. The last thoracomere(s) the Podotremata, the Raninoidea combine anare tilted, often almost perpendiculary to preced­ elongated P5 coxa and a more or less long anding ones. All the families of Podotremata have P4 soft penis (see Guinot 1979a: 194, fig. 43F). + P5 or P5 alone modified (Guinot 1991, 1995; No intermediate stages between the two patterns Guinot & Richer de Forges 1995; McLay 1993, (P5 coxa extended/P5 coxa not modified and 1999; Tavares 1993, 1996, 1998; Guinot & prolonging into sclerotized mobile penial tube)Tavares 2000). Either the whole P5 is mobile and were found amongst the Dromiacea. Although ofcompletely raised on the back of the crab and different shapes from group to group, the elon­ therefore a carrying behaviour is well-developed gated male P5 coxae are typically found in the (as in Homolodromiidae, Dromiidae, Homo­ most basal Podotremata (Homolodromiidae, lidae, Latreilliidae, Cyclodorippidae) (Guinot et Dynomenidae and Sphaerodromiinae n. subfam.). al. 1995), or it is only the P5 coxa (not the whole The Homolodromiidae, Dynomenidae and leg) which is dorsal, that is, a dorsal location of Sphaerodromiinae n. subfam. have all short the P5 arthrodial cavity. Even in the Dyno­ female sternal sutures 7/8. The long and mobile menidae, where P5 are carried alongside the body penial tube is present only in two dromiid and not capable of carrying a camouflaging subfamilies, the Dromiinae n. status and object, the P5 coxa (with its arthrodial cavitiy) is Hypoconchinae n. subfam. In the Dromiinaemoved dorsally.

ZOOSYSTEMA • 2003 • 25 (1) 117 Guinot D. & Tavares M.

In the two groups of basal Podotremata of an elongation of the male P5 coxa, on one or (Dromiacea, except Dynomenidae, and Homo- both sides. This occurrs in only a few genera: lidea) the thoracic pereopods may perform several Coenobita, tubes not present in all species functions: 1) P2 for walking and holding of(Coenobitidae Dana, 1851); somePorcellano­ abdomen; 2) P3 for walking, abdominal holding, pagurus Filhol, 1885, andSolitariopagurus and reproduction; 3) P4 for walking and, in some Tiirkay, 1986 (Paguridae Latreille, 1802; cases, for camouflage; and 4) P5 for reproduction McLaughlin pers. comm.; Lemaitre pers. and camouflage. This is perhaps why the walking comm.). is so slow in dromiacean crabs. The second type of coxal modification refers to It is not surprising to find a kind of penial tube inthe development of a “sexual tube”, known for the Dorippinae (Eubrachyura) where the two more than 150 years (Henderson 1888, 1893). posterior legs (P4 and P5) are dorsal and prehen­ The term “tube sexuel” seems to have been intro­ sile to hold a sessile animal or shell for camou­ duced by A. Milne Edwards & Bouvier (1892: flaging, as in Dromiinae n. status and 188). Currently, more than half of the genera in Hypoconchinae n. subfam. the family Paguridae and one of the two genera The shape of the P5 coxa provides an invaluable of the Coenobitidae show the development of tool for recognition of families among the fossilsome type of tube emanating from one or both Podotremata. Because the fragile mobile penial gonopores on the P5 coxae. The tubes are often tube of the Dromiinae n. status and Hypo­ asymmetrical, sometimes filamentous or coiled, conchinae n. subfam. is connected to the coxamembranous or slightly calcified (Saint Laurent- through a delicate membrane, it cannot be fos­ Dechancé 1966; McLaughlin & Hogarth 1998; silized and does not remain attached. In this case, McLaughlin 2000; McLaughlin & Lemaitre male fossil specimens will show only unmodified 2000, 2001; Lemaitre & Tudge 2001; coxa. Conversely, the hard elongated coxa of the McLaughlin pers. comm.). But the hermit crabs Homolodromiidae, Dynomenidae and Sphaero­ tubes are quite different from the brachyuran dromiinae n. subfam. is far more easily preserved tubes, because their function is not associated in the fossil record. As a result, a homolodromiid, with male pleopods (Lemaitre & Tudge pers. dynomenid, or sphaerodromiine fossil crab could comm.). It is not known how insemination takes be easily recognized by a greatly expanded P5 place in hermit crabs. It probably occurs exter­ coxa, whereas a dromiine or hypoconchine would nally because spermatophores have been seen present an unmodified coxa and no penial tube. attached to various parts of specimens in species The gonopores are very difficult to see in fossils, having sexual tubes. In hermit crabs without sex­ thus the shape of the male P5 coxa may be a good ual tubes (e.g. the diogenid Clibanarius Dana, character not only to separate the podotreme 1852 and the coenobitidBirgus Leach, 1815) the families but also to recognize the sex of individuals. sperm mass definitely is attached to the sternum. There is little or no information on the mecha­Both, the details on the copulatory behaviour nism for the discharge of the sperm into the known to date and the absence of male structures penial tube and, therefore, into gonopods, as wellto be used to introduce the sperm into the as on the mobility and ultrastructure of the P5 females, suggest that fertilization occurs externally coxa and penis (see Hartnoll 1975). (Lemaitre & Tudge pers. comm.).

The P5 coxa in the Anomura S h e l l - c a r r y in g b e h a v io u r H. Milne Edwards (1837: 239) described in the Shell-carrying is relatively rare in the dromiid Anomura the tubular prolongations on the coxaegenera. The posterior legs, P4 and P5, are gener­ of Coenobita Latreille, 1829. Actually, two types ally reduced and modified (oriented subdorsally of coxal modifications related to sperm transfer and/or dorsally, ending in special prehensile are shown by the hermit crabs. The first consists apparatus) for carrying an object. In the

118 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

commonest condition, P4 and P5 are roughlylar in size and shape, markedly modified but in similar, P5 being either slightly longer and more different ways (Guinot & Tavares 2000). slender, or markedly longer than P4 (in On a photograph ofAustrodromidia octodentata Pseudodromia). The grasping mechanism is repre­ published by Hale (1925: pi. 40A), the crab is sented by a subchelate ending, usually formed by shown sheltered beneath the valve with its one or several distal propodal spines opposing the cephalic parts situated near the umbo of shell. dactylus, the number and location being often Hypoconcha has an opposite posture inside the different on P4 and P5. shell (see Rathbun 1937: pi. 8, fig. 6, pi. 9, The camouflage among dromiids typically figs 4, 5 “in its natural habitat”). Its body is very includes soft material such as sponges, ascidians, small and thin in relation to the large shell which and weeds. Occasionally, hard material such as a is kept very close to carapace. The crab, which is large shell is encountered. Shell-carrying has been well concealed, probably walks slowly while car­ reported, for example, in the dromiineAustro­ rying the shell, perhaps trailing it, without stand­ dromidia octodentata (Hale 1925: 406, pi. 40A; ing much off the ground. The carapace can be 1927: 108, 109). Curiously, shell-carrying (using pressed against the inner concave surface shell, a Pecten shell) has never been observed again in with the setae aiding “in a tight seal”, and only this species. In Austrodromidia, P4 and P5 are not P2 and P3 are used in locomotion, altough the really different from the P4 and P5 of other crab may rest on the tips of chelipeds as well dromiids. They are therefore similarly shaped, (Wicksten 1986b: 19). The posture ofConchoe­ both reduced, and end in subchelate apparatus cetes under the shell is known by photographs of formed by one (on P5) or two (on P4) distallive C. artificiosus (Fabricius, 1798). The indi­ propodal spines opposing the dactylus; a long vidual is hiding under the shell but remains par­ spine on the outer margin of P5 dactylus may tially exposed (Ng et al. 2000: fig. lb); in another also be involved. photograph (Nishimura 1987: pi. M) the crab In other dromiids the two last pereopods show was completely concealed by the shell. a different pattern. Desmodromia (2001b: 1-8, Conchoecetes exhibits a particular combination of figs Id, 2d, 3), recently described for two P4-P5, dissimilar in position, size, and shape. Australian species collected without camouflage The P4, not dorsal, are markedly heavy, each with material, supposedly may carry bivalve shells a thick propodus bearing a mobile process which (McLay 2001b: 7, 8). On both P4 and P5, more or less sinks in a socket, and with a long, which do not look very dissimilar in shape, the curved dactylus. The P5 are very thin and end in dactyli are very small but stout, curved simply simple, upturned dactylus. This arrangement in and without opposing propodal spines. Thereby, Conchoecetes allows the bivalve shell to be inserted they do not form a subchelate mechanism such between the mobile process and the curved dacty­ as in most Dromiidae which carry soft material, lus. Detailed information, however, is lacking. even inAustrodromidia, which occasionally may This highly-specialized mechanism is the only of carry a shell. its kind in the Brachyura. All the species of Hypoconcha (Hypoconchinae The disposition inHypoconcha, also with P4-P5 n. subfam.) and Conchoecetes (Dromiinae n. sta­ different in position and size, is distinct. The P4 tus) are by far the best known examples of perma­ are not dorsal, and are robust but shorter than the nent shell-carrying behaviour in the family. P5. In both P4 and P5, the dactylus is crescent­ These species are adapted to hold and fit inside shaped, upturned and mobile in a notch at the the valve of a lamellibranch mollusk (Nicolson end of the stout propodus. The shell of the lamel­ 1776; Lamarck 1818; Crane 1937; Rathbunlibranch is held by the posterior legs and also by 1937; Schmitt 1965; Brusca 1980; Williams the angular abdomen inserted under the hinge 1984; Hendrickx 1997; Nget al. 2000: 157- (Rathbun 1937: 44, pi. 9, figs 4, 5; Wicksten 159). Their two last legs (P4 and P5) are dissimi­ 1986b: 19; Hendrickx 1997).H. arcuata clings

ZOOSYSTEMA • 2003 • 25 (1) 119 Guinot D. & Tavares M.

so tightly to its shell that the removal of the valve find crabs carrying a shell markedly larger than without crushing the crab is almost impossible carapace (Hendrickx pers. comm.; see Schmitt (Williams 1984: 257). It is not uncommon to 1965: fig. 62; Brusca 1980).

Key t o fa m il ie s o f D r o m ia c e a a n d subfamilies o f D r o m iid a e

The suprafamilial level, Homolodromioidea (with the single family Homolodromiidae) and Dromioidea (with families Dromiidae and Dynomenidae), is not specified in this key.

1. Both P4 and P5 modified, reduced, subdorsal or dorsal, with terminal prehensile apparatus. Pleopodal formula variable in males, either complete (vestigial pleopods P13-P15 combined with dorsal or ventral plates) or incomplete (no vestigial pleopods P13-P15). Uropods showing as dorsal plates or ventral lobes, absent exceptionally. Female sternal sutures 7/8 short or long, spermathecal apertures diversely located. Exopod present or absent on G2. Male P5 coxa modified and elongated, or unmodi­ fied and with independent penial tube ...... 2 — Only P5 modified, very reduced, obliquely directed, somewhat rigid, without real prehensile apparatus. Male pleopodal formula complete in males. P13-P15 as bira- mous vestiges in males (uniramous inDynomene praedator). Uropods always present, showing as large dorsal plates in both sexes, not efficiently involved in abdominal holding. Female sternal sutures 7/8 short, spermathecal apertures lying close to gonopores on P3. Exopod always present on G2. Male P5 coxa modified, elongated, prolonging into hard process enclosing penis ...... Dynomenidae

2. Mxp3 pediform. Male pleopodal formula always complete. P13-P15 as uniramous vestiges in males. Uropods showing as ventral lobes, never involved in abdominal holding. Male abdomen with distinct pleurae; telson very long, reaching mxp3. Endophragmal skeleton regularly layered; phragmae interfingered to each other. Female sternal sutures 7/8 short, spermathecal apertures lying close to gonopores on P3. Exopod absent on G2. Male P5 coxa modified, elongated, prolonging into hard process enclosing penis...... Homolodromiidae — Mxp3 operculiform. Male pleopodal formula variable. P13-P15 sometimes as vestiges in males. Uropods showing as salient dorsal plates (often efficiently involved in hol­ ding of abdomen), or as ventral plates, absent exceptionally. Male abdomen with pleurae sometimes distinct; telson variously shaped. Endophragmal skeleton not regularly layered; phragmae fused to each other. Female sternal sutures 7/8 short or long, spermathecal apertures close to or far from female gonopores on P3. Exopod present or absent on G2. Male P5 coxa modified or unmodified Dromiidae (3)

3. Male P5 coxa modified, elongated, prolonging into hard process enclosing penis. P2 and P3 propodus long and provided with distal spine; inner margin of dactylus of P2 and P3 with numerous spines. Male pleopodal formula complete. P13-P15 as bira- mous (or uniramous) vestiges. Uropods showing as dorsal plates, exposed but deeply inserted medially, intercalary. Female sternal sutures 7/8 short, spermathecal aper­ tures not reaching level of gonopores on P3. Exopod may be present on G2, length variable ...... Sphaerodromiinae n. subfam.

120 ZOOSYSTEMA • 2003 • 25(1) New subfamilial arrangement for the Dromiidae (Crustacea, Decapoda)

— Male P5 coxa unmodified, bearing long movable, “articulated”, sclerotized tube, from which emerges penis (“penial tube”). P2 and P3 propodus of variable length, usually without distal spine; inner margin of dactylus of P2 and P3 spinous or smoo­ th. Male pleopodal formula complete or incomplete. Female sternal sutures 7/8 short or long, spermathecal apertures beyond gonopores on P3, sometimes far beyond (except Stebbingdromia plumosa n. comb.). Exopod absent on G2...... 4

4. Front and lateral margins greatly expanded, covering all parts of head, except anten- nal flagella; eyes displaced in ventral location. Carapace hourglass-shaped; dorsal sur­ face flattened, very thin, membranous, specially in posterior half. Male abdomen widely triangular, generally flexed at right angles in the middle. Male pleopodal for­ mula complete: P13-P15 as uniramous vestiges in males; uropods always as ventral plates. Female sternal sutures 7/8 relatively short, entirely located on bent posterior part of sternum, spermathecal apertures showing as minute pores, always apart, slightly beyond female gonopores on P3. P2 and P3 propodus short, never armed with distal spine; inner margin of dactylus smooth or nearly smooth. P4 and P5 dissimilar, with peculiar, contorted dactylus, movable in a hollow at the end of propodus, and ending in corneous hook. Always carrying a valve of lamellibranch mollusk ...... Hypoconchinae n. subfam. — Front not expanded; eyes visible dorsally. Carapace dorsal surface convex, more or less calcified. Male abdomen relatively narrow and rather long, variously shaped, but never triangular. Male pleopodal formula variable, complete or incomplete. Uropods either as salient dorsal plates or as ventral plates, exceptionally absent. Female sternal sutures 7/8 long (exceptStebbingdromia plumosa n. comb.), spermathecal apertures generally showing as minute pores, far beyond gonopores on P3 (exceptS. plumosa n. comb.), and ending either apart or together. P2 and P3 propodus short, not armed with distal spine (except S. plumosa n. comb.); inner margin of dactylus with spines. P4 and P5 roughly similar in size (P5 often slightly longer than P4) and shape (except Conchoecetes, with P5 much smaller than P4 and both ending in different terminal apparatus) ...... Dromiinae n. status (5)

5. G2 long, with a needle-like flagellum. Female sutures 7/8 long, spermathecal aper­ tures generally far beyond gonopores on P3, showing as very minute pores (perhaps except Stebbingdromia plumosa n. comb.), exceptionally as slits Sternodromia ( ). Male pleopodal formula variable Dromiinae n. status (list of genera included, p. 49) — G2 short (slightly shorter than G l) and stout, without needle-like flagellum. Female sutures 7/8 short, spermathecal apertures beside gonopores on P3, shape of spermathecal apertures unknown. Male pleopodal formula complete Stebbingdromia n. gen.

Illustration Acknowledgements The numerous drawings, all original, were care­ We thank Penny Berents (AM), Paul F. Clark fully executed over a period of years by Michèle (BM), Peter Davie (QM), Michel E. Hendrickx Bertoncini (MNHN/CRNS, Paris). We express (EMU), Peter K. L. Ng (National University of our profond thanks for her invaluable contribu­Singapore) for kindly arranging the loan of crab tion, which has greatly added to the clarity of the material from their institutions. Thanks also to morphological interpretations. Patsy A. McLaughlin (Western Washington

ZOOSYSTEMA • 2003 • 25 (1) 121 Guinot D. & Tavares M.

University, USA), Rafael Lemaitre (Smithsonian Proceedings and Report o f the Royal Society o f South Institution, Washington DC) and Chris Tudge Australia 31: 173-191. B a ls s H. 1935- — Brachyura of the Hamburg (American University, Washington DC) for pro­ Museum Expedition to South-Western Australia, viding information on the sexual tubes in the 1905.Journal and Proceedings o f the Royal Society o f Anomura, and Hervé de Toulgoet (MNHN) for Western Australia 21: 113-151. providing information on the type species ofBARNARD K. H. 1947. — Description of new species of South African Decapod Crustacea, with notes on Evius Walker, 1855. We are grateful to Colin L. synonymy and new records. Annals and Magazine McLay (University of Canterbury, New Zealand) o f Natural History (11) 13 (102), 1946 (1947): for sending a female ofStebbingdromia plumosa 361-392. n. comb, and for making the proofs of a paperBARNARD K. H. 1950. — Descriptive catalogue of South African Decapod Crustacea. Annals o f the (McLay et al. 2001) available to us, and to Jean- South African Museum 38: 1-837. Marie Bouchard (Institut national de Recherche BARNARD K. H. 1954. — Notes sur une collection de pédagogique) for permission to publish one origi­ Crustacés Décapodes de la région malgache. Mémoires de l ’Institut scientifique de Madagascar (A) nal drawing from his doctoral dissertation. We 9: 95-104. are grateful to Peter Castro (California State BOAS J. E. V. 1880. — Studier over Decapodernes Polytechnic University, Pomona, California) Slægtskabsforhold. [With a French abstract: for correcting the English text and interesting Recherches sur les affinités des Crustacés décapodes: 163-207]. Koninklijke danske Videnskabelige Skrifter discussion. The manuscript was greatly improved (6) 1 (3): 25-210. by the comments of Rafael Lemaitre and Peter BORRADAILE L. A. I9OO. — On some Crustaceans K. L. Ng. Didier Geffard, Laurent Albenga and from the South Pacific. Part IV. The Crabs. Régis Cleva (MNHN) kindly provided technical Proceedings o f the Zoological Society o f London (4): 568-596. assistance on several occasions. This paper BORRADAILE L. A. 1903a. — Marine Crustaceans. IX. was done while M. T. held an appointment as The Sponge-crabs (Dromiacea),in GARDINER J. S. Invited Professor at the MNHN. Support from (ed.), The Tauna and Geography o f the Maldive and Universidade Santa Ursula and CNPq (National Laccadive Archipelagoes 2 (1). Cambridge University Press, Cambridge, UK: 574-578. Council for the Development of Science andBORRADAILE L. A. 1903b. — O n the genera of the Technology, Brasilia) is greatly appreciated. Dromiidae. Annals and Magazine o f Natural History (7) 11: 297-303. B o u c l l a r d J.-M. 2000. — Morphologie fonctionelle des systèmes de rétention de l ’abdomen chez les Brachyoures REFERENCES (Crustacea Decapoda). Microstructures ; implications phylogénétiques et systématiques. Thèse de Doctorat, A lcock A. W. 1899-— An Account o f the Deep-Sea Muséum national d’Histoire naturelle, Paris, Brachyura collected by the Royal Indian Marine France, 694 p. Survey Ship Investigator. Trustees of the Indian BOUVIER E.-L. 1896. — Sur l’origine homarienne des Museum, Calcutta, 85p. Crabes : étude comparative des Dromiacés vivants ALCOCK A . W. 1900. — Materials for a carcinological et fossiles. Bulletin de la Sociétéphilomatique (8) fauna of India. No. 5. Brachyura Primigenia or 8 (2) 1895 (1896): 34-110 (1-77). Dromiacea. Journal o f the Asiatic Society o f Benrnl BOUVIER E.-L. 1898. —Sur quelques Crustacés 68 (2): 123-169. anomoures et brachyures recueillis par M. D iguet A L C O C K A . W. 1901. — Catalogue of the Indian en Basse-Californie.Bulletin du Muséum d ’Histoire Decapod Crustacea in the Collection o f the Indian naturelle A (8): 371-384. Museum. Part I. Brachyura. Fasc. 1. Introduction BOUVIER E.-L. 1940. — Décapodes Marcheurs. Taune and Dromides or Dromiacea (Brachyura Primigenia). de Trance, 37. Lechevalier et Fils, Paris, 404 p. Trustees of the Indian Museum, Calcutta, viii + B o y k o C. B. 1998. — The correct original spelling of 80 p. Cryptodromidiopsis sarraburrei (Rathbun, 1910) not ANDRÉ M. 1932. — Crustacés recueillis par M. E. C. larraburei (Schmitt, 1921) (Decapoda, Aubert de la Riie aux îles Kerguelen, Saint-Paul et Brachyura, Dromiidae). Crustaceana 71 (2): de la Nouvelle-Amsterdam. Bulletin du Muséum 234-235. national d ’Histoire naturelle (2) 4 (2): 174-181. BROCCHI P. 1877. — Sur un Dromien nouveau, genre BAKER W. H. 1907. — Notes on South Australian Platydromia. Bulletin de la Société philomatique (6) Decapod Crustacea. Part 5. Transactions and 12, 1875 (1877): 53-54.

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H EN D ERSO N J. R. 1888. — Report on the Anomura 5. Crustacea, Stomatopoda, Isopoda and Decapoda. collected by H . M. S. Challenger during the years South African Journal o f Zoology 19 (3): 189-193. 1873-1876.Report on the Scientific Results ofithe K lR CH ER A. B. 1970. — The zoeal stages and glau- Exploring Voyage ofiH. M. S. Challenger 27 ( 1 ), xi + cothoë ofHypoconcha arcuata Stimpson (Decapoda: 221 p. Dromiidae) reared in the laboratory. Bulletin of H e n d e r s o n J. R. 1893. — A contribution to Indian Marine Science 20 (3): 769-792. carcinology. Transactions ofithe Linnean Society ofi KOSSMANN R. 1880. — Zoologische Ergebnisse einer London (2, Zoology) 5 (1): 325-458. Reise in die Küstengebiete des Rothen Meeres, HENDRICKX M. E. 1995. — Checklist of brachyuran Volume 2, Part 1, Section III, Malacostraca.Zoologische crabs (Crustacea: Decapoda) from the eastern tropi­ Ergebnisse im Aufträge der königlichen Academie der cal Pacific. Bulletin de l ’Institut royal de Belgique, Wissenschaften zu Berlin 1880: 67-140. Biologie 65: 15-150. LAMARCK J. B. P. A. DE 1818. —Histoire naturelle des HENDRICKX M. E. 1997. —Los Cangrejos Braquiuros anim aux sans vertèbres, présentant les caractères (Crustacea: Brachyura: Dromiidae, hasta Leucosiidae) généraux et particuliers de ces animaux, leur distribu­ del Pacifico Mexicano. Comisión Nacional para el tion, leurs classes, leurs familles, leurs genres, et la cita­ Conocimiento y uso de la Biodiversidad e Instituto tion des principales espèces qui s’y rapportent ; précédée de Ciencias del Mar y Limnología, UNAM, d ’une introduction offrant la détermination des carac­ México, 178 p. tères essentiels de l ’Animal, sa distinction du végétal et HERBST J. F. W. 1783. — Versuch einer des autres corps naturels, enfin, l ’Exposition des Naturgeschichte der Krabben und Krebse nebst einer principes fondamentaux de la zoologie, 5. Déterville systematischen Beschreibung ihrer verschiedenen Arten. et Verdière, Paris, 612 p. Lange, Berlin; Stralsund 1 (2-5): 87-182. LAMARCK J. B. P. A. DE 1838. — Histoire naturelle des H o l t h u i s L. B. 1962. — Dromia Niebet, 1795 anim aux sans vertèbres, présentant les caractères (Crustacea, Decapoda): proposed designation of a généraux et particuliers de ces animaux, leur distribu­ type-species under the plenary powers. Z. N. (S.) tion, leurs classes, leurs familles, leurs genres et la cita­ 1488. Bulletin ofi Zoological Nomenclature 19 (1): tion des principales espèces qui s’y rapportent ; précédé 51-57. d ’une introduction offrant la détermination des carac­ H O O V E R J. P. 1998. —Hawaii’s Sea Creatures: a tères essentiels de l ’Animal, sa distinction du végétal et Guide to Hawaii’s Marine Invertebrates. M utual des autres corps naturels, enfin l’Exposition des principes Publishing, Honolulu, Hawaii, 366 p. fondamentaux de la zoologie, 5 [Arachnides, Crustacés, ICZN 1999- — International Code ofi Zoological Annélides, Cirripèdes]. Baillière, Paris, 2e éd., 699 p. Nomenclature. 4*^ ed. International Trust for LANG W. H. & Y o u n g A. M. 1980. — Larval devel­ Zoological Nomenclature, London, 306 p. o p m e n t o f Hypoconcha sabulosa (D eca p o d a : I h l e J. E. W. 1913. — Die Decapoda Brachyura der Dromiidae). Fishery Bulletin 77 (4): 851-864. Siboga-Expedition. I. DromiaceaSiboga Expeditie LATREILLE P. A. 1802. — Histoire naturelle, générale et Monogr. 39b, livr. 71: 1-96. particulière, des Crustacés et des Insectes. Ouvrage Ja m ie s o n B. G. M., T u d g e C. C. & S c h e l t i n g a D. faisant suite aux Œuvres de Leclerc de Buffion, et par­ M. 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Submitted on 12 February 2002; accepted on 19 June 2002.

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