The Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae) Lipke B. HOLTHUIS National Museum of Natural History, P.O. Box 9517, 2300 RA, Leiden (The Netherlands) Holthuis L. B. 2002. — The Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae). Zoosystema 24 (3) : 499-683. ABSTRACT A revision is provided of the Indo-Pacific species of the subfamily Scyllarinae. All of these species were formerly placed in the genus Scyllarus Fabricius, 1775, but a closer study revealed that several genera could be distinguished within the subfamily. The 13 new genera now recognized in the Indo-Pacific biogeographic region are as follows: Acantharctus n. gen., Antarctus n. gen., Antipodarctus n. gen., Bathyarctus n. gen., Biarctus n. gen., Chelarctus n. gen., Crenarctus n. gen., Eduarctus n. gen., Galearctus n. gen., Gibbularctus n. gen., Petrarctus n. gen., Remiarctus n. gen. and Scammarctus n. gen. Diagnoses and keys are provided for all the genera and their species. New and insufficiently known species have been described extensively, for the others additional mor- phological details are given. New species are: Bathyarctus chani n. gen., n. sp., B. steatopygus n. gen., n. sp., Petrarctus veliger n. gen., n. sp., Chelarctus crosnieri n. gen., n. sp., Eduarctus pyrrhonotus n. gen., n. sp., E. marginatus n. gen., n. sp., n. gen., n. sp. and n. gen., n. sp. KEY WORDS E. perspicillatus E. reticulatus Crustacea, Furthermore efforts were made to provide each species with a complete syn- Decapoda, onymy, a description of the colour, its biology, habitat and geographical dis- Scyllarinae, tribution. All the material examined is listed in detail. Where appropriate, Indo-Pacific, new genera, remarks are provided on nomenclature, published data on the larval develop- new species. ment and other topics. ZOOSYSTEMA • 2002 • 24 (3) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 499 Holthuis L. B. RÉSUMÉ Les cigales scyllarines indo-pacifiques (Crustacea, Decapoda, Scyllaridae). Une révision des espèces appartenant à la sous-famille Scyllarinae est propo- sée. Auparavant toutes ces espèces étaient classées dans le genre Scyllarus Fabricius, 1775 mais une étude plus précise a montré que plusieurs genres pouvaient être distingués dans la sous-famille. Les 13 nouveaux genres recon- nus maintenant dans la région indo-pacifique sont les suivants : Acantharctus n. gen., Antarctus n. gen., Antipodarctus n. gen., Bathyarctus n. gen., Biarctus n. gen., Chelarctus n. gen., Crenarctus n. gen., Eduarctus n. gen., Galearctus n. gen., Gibbularctus n. gen., Petrarctus n. gen., Remiarctus n. gen. et Scammarctus n. gen. Des diagnoses sont données pour tous les genres ainsi que des clés pour leurs espèces. Les espèces nouvelles ou mal connues ont été décrites en détails ; pour les autres, des détails morphologiques complémen- taires ont été fournis. Les nouvelles espèces sont : Bathyarctus chani n. gen., n. sp., B. steatopygus n. gen., n. sp., Petrarctus veliger n. gen., n. sp., Chelarctus crosnieri n. gen., n. sp., Eduarctus pyrrhonotus n. gen., n. sp., E. marginatus n. gen., n. sp., E. perspicillatus n. gen., n. sp. et E. reticulatus n. gen., n. sp. De MOTS CLÉS Crustacea, plus on s’est efforcé de donner la synonymie complète de chaque espèce, sa Decapoda, coloration, sa biologie, son habitat et sa distribution géographique. Tout le Scyllarinae, matériel examiné est mentionné avec les détails complets relatifs aux récoltes. Indo-Pacifique, nouveaux genres, Quand cela s’est avéré nécessaire des remarques ont été faites sur la nomencla- nouvelles espèces. ture, le développement larvaire ainsi que d’autres sujets. INTRODUCTION Crosnier, I can now present the results of this work. The long period over which this study was Forty years ago, in 1960, during a year’s stay at the made is the cause that several inconsistencies in U. S. National Museum in Washington, D. C., I the treatment do occur for which I apologize. For started a study intended to revise the family the same reason the illustrations are of very diverse Scyllaridae Latreille, 1825. I had not finished the origin: many different artists have provided the job when I returned to my position at the Leiden drawings of the complete animals, the photo- Museum. Again in Leiden, so many chores had graphs were made mostly at the Muséum national accumulated during my absence that I did not d’Histoire naturelle, Paris, and most of the simple have the opportunity to immediately continue my line drawings were made by the author. I hope work on the Scyllarid lobsters. In 1985 finally the that nonetheless the present paper will form a revision of the Ibacinae could be published. worthwhile contribution to the study of the Indo- Unfinished manuscripts of the other subfamilies Pacific members of the subfamily Scyllarinae. remained in an advanced state of preparation. The general morphology of the Scyllarinae and Then Alain Crosnier of the Paris Museum offered their place among the other subfamilies of me for study the Scyllarid material collected by Scyllaridae have been extensively dealt with in my the French MUSORSTOM expeditions in the 1985 paper, which also carries a schematic figure Philippines, New Caledonia and other areas. This showing the terms used for ridges, tubercles and gave me the incentive to take up again the study other anatomical features of carapace and cephalon. of the subfamily Scyllarinae. Thanks to the conti- In the lists of material examined, the names of nuous encouragement and help by Alain the cruises are in capital letters and the names of 500 ZOOSYSTEMA • 2002 • 24 (3) Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae) the vessels in italic letters. The measurements, RMNH Rijksmuseum van Natuurlijke Historie unless otherwise specified, refer to the carapace (National Museum of Natural History), Leiden; length. SUF Shimonoseki University of Fisheries, Some of the figures, drawn a long time ago, have Shimonoseki; no scale bar. USNM National Museum of Natural History, Readers requiring details on the French cruises Washington D. C.; UZM Universitetets Zoologiske Museum, carried out in the south-west Pacific since 1976 Copenhagen; can refer to the web site: http://www.tropical VNIRO All-Russia Research Institute of Marine deepseabenthos.org. Fisheries and Oceanography, Moscow; There is a rather extensive literature on Scyllarine WAM Western Australian Museum, Perth; ZLK Zoological Laboratory of Kyushu larvae, but most of the material studied there was University, Fukuoka; obtained from the plankton, so that the specific ZMA Zoological Museum, Amsterdam; identity often could not or only tentatively be ZMB Zoologisches Museum der Humboldt Universität, Berlin; ascertained. This is quite understandable as the ZMH Zoologisches Museum, Hamburg; number of scyllarine species is quite extensive and ZMM Zoological Museum, Moscow; new species are still being discovered. Some ZRC Zoological Reference Collection, Raffles authors managed to describe the complete series Museum, University of Singapore; ZSM Zoologische Staatssammlung, Munich; of larvae of a certain species from plankton mate- cl. carapace length; rial (e.g., Eduarctus martensii (Pfeffer, 1881) n. juv. juvenile; comb. by Phillips & McWilliam 1986). Ritz ov. ovigerous; (1977) managed to obtain all larval stages of P.1-P.5 first to fifth pereiopods; tl. total length. Petrarctus demani (Holthuis, 1946) n. comb. by culturing the species in the laboratory. Where larval stages have been (correctly or incor- rectly) definitely assigned to a species, they usual- SYSTEMATICS ly are listed in the present paper. But “Scyllarus spec. A.”, “Scyllarus spec. B.”, etc., are usually not Order DECAPODA Latreille, 1802 included even when they were accompanied by a Suborder PLEOCYEMATA vague guess as to the identity of the species. Burkenroad, 1963 Family SCYLLARIDAE Latreille, 1825 ABBREVIATIONS AHF Allan Hancock Foundation, University of Subfamily SCYLLARINAE Latreille, 1825 Southern California, Los Angeles (at present housed in the Los Angeles County Scyllarides Latreille, 1825: 278. Museum); AM Australian Museum, Sydney; Scyllarinae – Holthuis 1985: 3, 10, 12. ANS Academy of Natural Sciences, Philadelphia; The family Scyllaridae is such a well-defined and BM The Natural History Museum (formerly British Museum (Natural History)), homogeneous entity that only as late as 1985 a London; proposal was published to subdivide it into subfa- BPBM Bernice Pauahi Bishop Museum, Honolulu; milies. The subfamily Scyllarinae was then consi- Mbogor Zoological Museum, Bogor; dered to consist of a single genus Scyllarus MCZ Museum of Comparative Zoology, Harvard University, Cambridge; Fabricius, 1775, and only the study of the pre- MG Muséum d’Histoire naturelle, Geneva; sent very extensive material showed that the old MNHN Muséum national d’Histoire naturelle, Paris; genus can be subdivided. Suggestions that groups MS Senckenberg Museum, Frankfurt am Main; of species can be recognized within the old genus MT Afrika Museum, Tervuren; NTOU National Taiwan Ocean University, Keelung; Scyllarus have already been made by previous PN collection P. Naiyanetr, Department of authors like Ortmann (1897: 270) and Bouvier Biology, Chulalongkorn University, Bangkok; (1917: 105, 106), but the outlines of these ZOOSYSTEMA • 2002 • 24 (3) 501 Holthuis L. B. groups were not very sharp and no names were the Crustacea, H. Milne Edwards (1837) reco- suggested for them. Bouvier’s remarks did not gnized three genera