A Revision of the Indo-West Pacific Species of Palicid Crabs
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Crustacea Decapoda: A revision of the Indo-west Pacific species of palicid crabs (Brachyura Palicidae) Peter Castro In: A. Crosnier (ed), Resultats des Campagnes MUSORSTOM, Volume 21. M&moires du Museum national d'Histoire naturelle, 184: 437-610. Paris ISBN: 2-85653-526-7 JESULTATS DES CAMP AGNES MUSORSTOM, VOLUME 21 — RESULT ATS DES CAMPAGNES MUSORSTOM, VOLUME 21 — RESULT 10 Crustacea Decapoda: A revision of the Indo-west Pacific species of palicid crabs (Brachyura Palicidae) Peter CASTRO Biological Sciences Department California State Polytechnic University Pomona, California 91768-4032, USA ABSTRACT The taxonomy of the crabs belonging to the family Palicidae Bouvier, 1898 from the Indo-west Pacific region is revised. On the basis of extensive material collected by French expeditions in the Coral Sea and other regions of the Pacific and Indian oceans, as well as material from numerous museums, including most of the types, the present study recognizes two subfamilies, 10 genera, and 43 species. Of these taxa, four are new genera: Exopalicus, Miropalicus, Paliculus, and Rectopalicus. Manella is synonymized with Crossotonotus A. Milne Edwards, 1873. Parapleurophricoides Nobili, 1906, sometimes believed to be a palicid, is a xanthoid and it is removed from the Palicidae. Nine nominal species described by previous authors are synonymized and an additional 17 species are described. RESUME Crustacea Decapoda : Revision des especes indo-ouest pacifiques de crabes palicides (Brachyura Palicidae). La taxonomie des crabes de la famille Palicidae Bouvier, 1898, provenant de l'lndo-ouest Pacifique est revisee. Basee sur l'abondant materiel recolte par les expeditions francaises en mer de Corail et dans d'autres regions des oceans Pacifique et Indien, ainsi que sur du materiel provenant de nombreux museums renfermant la plupart des types, la presente etude reconnait deux sous-families, dix genres et 43 especes. Parmi ces taxa, quatre sont des genres nouveaux : Exopalicus, Miropalicus, Paliculus, et Rectopalicus. Le genre Manella est mis en synonymie avec Crossotonotus A. Milne Edwards, 1873. Parapleurophricoides Nobili, 1906, parfois considere comme 6tant un palicide, est un xanthide et est retire de la famille Palicidae. Neuf especes decrites anterieurement sont mises en synonymie, tandis que 17 autres sont decrites comme nouvelles. CASTRO, P., 2000. — Crustacea Decapoda : A revision of the Indo-west Pacific species of palicid crabs (Brachyura Palicidae). In: A. CROSNIER (ed.), Resultats des campagnes MUSORSTOM, Volume 21. Memoires du Museum national d'Histoire naturelle, 184: 437-610. Paris ISBN: 2-85653-526-7. 438 P. CASTRO TABLE OF CONTENTS INTRODUCTION 439 PRESENTATION AND TERMINOLOGY 440 SYSTEMATIC ACCOUNT 443 Family PALICIDAE Bouvier, 1898 443 Key to the subfamilies of the family Palicidae 445 Subfamily PALICINAE Bouvier, 1898 445 Key to the Indo-west Pacific genera of the subfamily Palicinae 448 Genus PSEUDOPALICUS Moosa & Serene, 1981 448 Key to the species of Pseudopalicus 451 Pseudopalicus acanthodactylus sp. nov 452 Pseudopalicus amadaibai (Sakai, 1963) 455 Pseudopalicus declivis sp. nov 458 Pseudopalicus glaber sp. nov 462 Pseudopalicus oahuensis (Rathbun, 1906) 465 Pseudopalicus sexlobatus (Kensley, 1969) 467 Pseudopalicus investigatoris (Alcock, 1900) 469 Pseudopalicus serripes (Alcock & Anderson, 1895) 473 Pseudopalicus sp 474 Pseudopalicus pictus sp. nov 476 Pseudopalicus macrotneles sp. nov 480 Pseudopalicus undulatus sp. nov 483 Genus PARAPALICUS Moosa & Serene, 1981 486 Key to the species of Parapalicus 488 Parapalicus ambonensis Moosa & Serene, 1981 489 Parapalicus armatus sp. nov 492 Parapalicus clinodentatus sp. nov 495 Parapalicus denticulatus sp. nov 499 Parapalicus inermis sp. nov 502 Parapalicus elaniticus (Holthuis, 1977) 505 Parapalicus inanis sp. nov 506 Parapalicus microphthalmus sp. nov 509 Parapalicus nanshaensis Dai andXu, 1991 510 Parapalicus piruensis Moosa & Serene, 1981 512 Parapalicus trispiralis sp. nov 513 Parapalicus trituberculatus (Chen, 1981) 517 Parapalicus unidentatus (Zarenkov, 1968) 519 Genus MIROPALICUS new genus 520 Miropalicus vietnamensis (Zarenkov, 1968) 522 Genus PALICULUS new genus 525 Key to the species of Paliculus 527 Paliculus kyusyuensis (Yokoya, 1933) 527 Paliculus foliatus sp. nov 530 Paliculus sp 532 Genus RECTOPALICUS new genus 533 Key to the species of Rectopalicus 535 Rectopalicus woodmasoni (Alcock, 1900) 535 Rectopalicus amphiceros sp. nov 538 Rectopalicus ampullatus sp. nov 541 INDO-WEST PACIFIC SPECIES OF PALICID CRABS 439 Genus EXOPALICUS new genus 543 Exopalicus maculatus (Edmondson, 1930) 546 Genus NEOPALICUS Moosa & Serene, 1981 548 Key to the species of Neopalicus 550 Neopalicus contractus (Rathbun, 1902) 550 Neopalicus jukesii (White, 1847) 554 Genus PALICOIDES Moosa & Serene, 1981 558 Key to the species of Palicoides 561 Palicoides longimanus (Miyake, 1936) 561 Palicoides whitei (Miers, 1884) 565 Subfamily CROSSOTONOTINAE Moosa & Serene, 1981 568 Key to the genera of the subfamily Crossotonotinae 569 Genus CROSSOTONOTUS A. Milne Edwards, 1873 569 Key to the species of Crossotonotus 571 Crossotonotus compressipes A. Milne Edwards, 1873 571 Crossotonotus spinipes (De Man, 1888) 574 Crossotonotus ceramensis (Moosa& Serene, 1981) 578 Crossotonotus lophocheir sp. nov 579 Genus PLEUROPHRICVS A. Milne Edwards, 1873 582 Key to the species of Pleurophricus 583 Pleurophricus cristatipes A. Milne Edwards, 1873 583 Pleurophricus longirostris (Moosa & Serene, 1981) 585 ECOLOGY AND OTHER ASPECTS OF THE BIOLOGY OF THE INDO-WEST PACIFIC PALICIDS 585 Habitat 585 Biogeography 586 Diet 592 Sexual dimorphism and mating behavior 592 Role of the fifth pair of pereopods 592 ACKNOWLEDGMENTS 593 REFERENCES 594 INDEX 606 INTRODUCTION Brachyuran crabs belonging the family Palicidae Bouvier, 1898 are a group rather neglected by taxonomists and ecologists alike, notwithstanding their sometimes frequent occurrence in several marine environments. The feeding habits, locomotion, reproduction, and larval development of palicids remain practically unknown. The evolution and biological significance of the reduced last pair of pereopods of most species remain vexing questions. The taxonomy of the family and its position within the Brachyura have also remained much neglected subjects. The family Palicidae consisted of four ill-defined genera until the revision of the Indo-west Pacific species by MOOSA & SERENE (1981): Palicus Philippi, 1838, Crossotonotus A. Milne Edwards, 1873, Pleurophricus A. Milne Edwards, 1873, and Manella Rathbun, 1906. A fifth genus, Parapleurophrycoides Nobili, 1906 was questionably included in the family by SERENE (1968, as Pleurophrycoides [sic]). Most species were grouped into one genus, Palicus, with RATHBUN (1918) recognizing 21 western Atlantic and eastern Pacific species in six "groups". Three more species from this area (all in Palicus) were subsequently described. An additional species, P. caronii (Roux, 1830), is Mediterranean and eastern Atlantic in distribution. In the Indo-west Pacific, 24 species (16 in Palicus) had been described until 1981 (SERENE, 1968; ZARENKOV, 1968). While describing two new western Pacific species, ZARENKOV (1968: 765) briefly commented on RATHBUN's groups of Palicus species without revising her classification. 440 P. CASTRO The first and until now the only major revision of Indo-west Pacific palicids was that of MOOSA & SERENE (1981). They separated the family into two subfamilies, Palicinae s. str. and Crossotonotinae Moosa & Serene, 1981. For the Palicinae, MOOSA & SERENE (1981) restricted Palicus for the Atlantic, Mediterranean, and eastern Pacific species, and established four new genera (Neopalicus, Palicoides, Parapalicus, and Pseudopalicus) for the 26 species they recognized from the Indo-west Pacific. Six new species were described from these genera. Within the subfamily Crossotonotinae, MOOSA & SERENE (1981) initially recognized four genera, Crossotonotus, Parapleurophrycoides, Pleurophricus, and Manella, with a total of six species. Parapleurophrycoides and its only species, however, were "discarded" since the description of the species was based on a juvenile "which cannot be used for the definition of the genus" (MOOSA & SERENE, 1981: 52). Unfortunately, the revision of MOOSA & SERENE (1981) was based on limited material. They examined specimens belonging to only nine of the 24 Indo-west Pacific species known at the time and the type material of only one of the nine species. In addition, the identities of several species (such as Parapleurophrycoides roseus Nobili, 1906) were left unresolved. Three Indo-west Pacific species of palicids were described after MOOSA & SERENE'S revision: Palicus trituberculatus Chen, 1981 (its description was apparently published before the revision), Palicus bidentatus Sakai, 1983, and Parapalicus nanshaensis Dai & Xu, 1991, bringing the total number of nominal species known from the region to 33. About 58% of all the known species of palicids (34 out of 58) are Indo-west Pacific in distribution. The study of a large collection of palicids from numerous French expeditions (mostly from the Coral Sea but also from several other areas of the Indo-west Pacific region) plus type material and other specimens from museums, institutions, and private collections from around the world provided the opportunity to undertake a more comprehensive revision of the Indo-west Pacific Palicidae. PRESENTATION AND TERMINOLOGY A list of complete references is given for each of the species treated, while references