Crustacea, Decapoda)
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European and Mediterranean Thalassinidea (Crustacea, Decapoda) Nguyen NGOC-HO Muséum national d’Histoire naturelle, Département Milieux et Peuplements aquatiques, 61 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Ngoc-Ho N. 2003. — European and Mediterranean Thalassinidea (Crustacea, Decapoda). Zoosystema 25 (3) : 439-555. ABSTRACT All genera and species from Europe and the Mediterranean are diagnosed or redescribed, differentiating characters are illustrated and an identification key is provided. A review of literature and biological information is included, and a classification, with two new genera, is proposed. The superfamily Axioidea comprises the Axiidae, with Axius stirhynchus, Calocarides coronatus, Levantocaris hornungae; and the Calocarididae with Calastacus laevis and Calocaris macandreae. The Callianassoidea include the Callianassidae, Ctenochelidae, Laomediidae and Upogebiidae. The Callianassidae contain three subfamilies. The Callianassinae have three species placed in Callianassa: C. acanthura, C. subterranea and C. truncata; three other species are assigned to a new genus, Pestarella n. gen.: P. candida n. comb., P. tyrrhena n. comb. and P. whitei n. comb. Pestarella n. gen. species have an operculiform Mxp3, a telson rounded in posterior half, and no Plp1-2 in male. The little known species Calliapagurops charcoti is placed in the Callichirinae. The Eucalliacinae contain two species: Calliax lobata and Calliaxina punica n. comb., type species of a new genus, Calliaxina n. gen. Species of Calliaxina n. KEY WORDS gen. possess a rostrum with pointed tip, an operculiform Mxp3 with exopod, Crustacea, Decapoda, the P1 equal and similar, and Plp1-2 with appendix interna in both male and Thalassinidea, female. The Ctenochelidae have one subfamily, Gourretinae with Gourrretia Europe, denticulata. The Laomediidae are represented by one species, Jaxea nocturna, Mediterranean, new genera, and the Upogebiidae by seven species: Gebiacantha talismani, Upogebia revision. deltaura, U. mediterranea, U. nitida, U. pusilla, U. stellata and U. tipica. ZOOSYSTEMA • 2003 • 25 (3) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 439 Ngoc-Ho N. RÉSUMÉ Thalassinidea (Crustacea, Decapoda) d’Europe et de Méditerranée. Une diagnose (ou redescription) de tous les genres et espèces d’Europe et de la Méditerranée est présentée et les caractères différentiels sont figurés. Une revue des données bibliographiques et biologiques est incluse ; une classifica- tion, comportant deux genres nouveaux, et une clé sont proposées. La super- famille Axioidea comprend les Axiidae, avec Axius stirhynchus, Calocarides coronatus, Levantocaris hornungae, et les Calocarididae, avec Calastacus laevis et Calocaris macandreae. Les Callianassoidea se composent des Callianassidae, Ctenochelidae, Laomediidae et Upogebiidae. Les Callianassidae contiennent trois sous-familles. Les Callianassinae ont trois espèces placées dans Callianassa: C. acanthura, C. subterranea et C. truncata ; trois autres sont attri- buées à un nouveau genre, Pestarella n. gen. : P. candida n. comb., P. tyrrhena n. comb. et P. whitei n. comb. Les espèces de Pestarella n. gen. ont un Mxp3 operculiforme, un telson arrondi dans sa partie postérieure et pas de Plp1-2 chez les mâles. L’espèce peu connue, Calliapagurops charcoti, est placée dans les Callichirinae. Les Eucalliacinae contiennent deux espèces : Calliax lobata et Calliaxina punica n. comb., espèce type d’un genre nouveau, Calliaxina n. gen. Les espèces de Calliaxina n. gen. possèdent un rostre à extrémité pointue, MOTS CLÉS un Mxp3 operculiforme muni d’exopode, les P1 égaux et les Plp1-2 avec Crustacea, Decapoda, appendix interna dans les deux sexes. Les Ctenochelidae ont une sous-famille, Thalassinidea, Gourretinae, avec Gourrretia denticulata. Les Laomediidae sont représentées Europe, par une espèce, Jaxea nocturna, et les Upogebiidae par sept espèces : Gebiacantha Méditerranée, nouveaux genres, talismani, Upogebia deltaura, U. mediterranea, U. nitida, U. pusilla, U. stellata révision. et U. tipica. INTRODUCTION The first objective of this paper is to define the species. All species and genera from Europe and The European and Mediterranean Thalassinidea the Mediterranean are diagnosed, with redescrip- are a small group of decapod crustaceans currently tions where necessary, and illustrated; extensive comprising 23 species, excluding a new axiid that synonymies are given from taxonomic and biolo- would have been described by M. de Saint gical literature. Characteristics of higher taxa can Laurent. They have never been subject to an be found in the Discussion and in the Key. overall revision; uncertainty and confusion still A classification of the European and exist for a number of species, especially in the Mediterranean Thalassinidea, which includes Upogebiidae. On the other hand, several impor- two new genera, is presented. Different views tant works on the taxonomy and phylogeny of the concerning taxa are discussed. Thalassinidea have been published in recent years: de Saint Laurent (1973), de Saint Laurent & Le Loeuff (1979), Sakai & de Saint Laurent (1989), MATERIALS AND METHODS Manning & Felder (1991), Poore (1994), Sakai (1999a), Tudge et al. (2000). Various classifica- All materials of European and Mediterranean tion systems have been proposed, sometimes with Thalassinidea currently present in the Muséum large differences, which meant that choices had to national d’Histoire naturelle, Paris (MNHN), be made concerning the taxonomy of the The Natural History Museum, London European and Mediterranean fauna. (NHML) and the Rijksmuseum of Natural 440 ZOOSYSTEMA • 2003 • 25 (3) European and Mediterranean Thalassinidea (Crustacea) History, Leiden (RMNH) were examined during with larvae, those preceded by the symbol ° refer the course of this study. Additional specimens to a mention of the species in a list, without came from the Museum of Comparative detail. Zoology, Harvard (MCZ), the Museum of Copenhagen (ZMUC), the Museum of Victoria ABBREVIATIONS (NMV), the Natural History Museum, Vienna A1 antennule; A2 antenna; (NHMW), the Senckenberg Museum, Frankfurt Md mandible; (SMF), the Swedish Museum of Natural History, Mx1 maxillule; Stockholm (SMNH), the Smithsonian Mx2 maxilla; Institution, Washington DC (USNM), as well as Mxp1-3 maxillipeds 1-3; P1-5 pereopods 1-5; from the personal collections of Cédric Plp1-5 pleopods 1-5. d’Udekem d’Acoz (d’Udekem d’Acoz, now depo- sited in the MNHN), and of Prof. Athanasios The following terminology used may need clarifi- Koukouras (A.U.TH, deposited in the Museum cation: of the Zoology Department, Aristoteleio Uni- For axiids, A2 acicle = antennal scale. versity of Thessaloniki). For callianassids (adapted from Manning & It has not been possible to locate the types for Felder 1991), dorsal oval (e.g., Figs 8A; 13A) is about a third of the species studied, nor was it defined by a groove or furrow that delimits an possible to ascertain that they were lost. An effort oval anterior portion of the carapace; posteriorly, has been made to clarify problems concerning the it is delimited by the cervical groove. identity of these species, but no neotypes have Mxp3: pediform (Fig. 21E) means ischium-merus been selected. length more than three times merus width; sub- The measurements given in the description are: pediform (Fig. 9D) means ischium-merus length carapace length (cl.) measured from the tip of the about two to three times merus width; operculi- rostrum (in the Callianassoidea) or the base of form (e.g., Figs 12G; 13G) means ischium-merus the rostrum, at the orbital margin (in the length less than twice merus width. Axioidea) to the posterior border of the carapace; Dorsal plate on uropodal exopod (e.g., Figs 8G; total length (tl.) measured from the tip of the ros- 9F) is an antero-dorsal thickening terminating trum (in the Callianassoidea) or the base of the posteriorly in a dorsal setal row near the posterior rostrum (in the Axioidea) to the posterior border border. of the telson. For upogebiids, lateral ridges (e.g., Figs 24A; Most specimens belonging to the MNHN and a 26A) are the upper longitudinal toothed crests of few from other institutions were measured. The the gastric region, on either side of the rostrum. carapace length is given for all whereas the total length is only given for types and figured speci- mens. DISCUSSION Figured specimens and appendages were stained with a weak solution of chlorazol black, some- Examination of the European and Mediterranean times with a drop of lactic acid added. Thalassinidea supports Poore’s (1994) view If not otherwise stated, the rostrum, the anterior about assigning the Axiidae and Callianassidae part of the carapace, the telson and the uropods into separate superfamilies, the Axioidea and are figured in dorsal view and appendages in lat- Callianassoidea. Members of the two families eral view. share a chelate P2 and the presence of an appen- In the list of synonymies, the original names dix interna on Plp3-5 but differ in many features. given to a species are presented first, in chronolog- A number of presumably plesiomorphic characte- ical order while other names are in alphabetical ristics are found in the Axioidea: linea thalassinica order. References preceded by an asterisk (*) deal absent; rostrum large and armed with teeth or ZOOSYSTEMA • 2003 • 25 (3) 441 Ngoc-Ho N. spines; antennal acicle large; Mxp3 slender with a The Ctenochelidae differ by having no dorsal strong tooth