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Decapoda: Anomura: Paguroidea) and Descriptions of New Taxa

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Decapoda: Anomura: Paguroidea) and Descriptions of New Taxa THE RAFFLES BULLETIN OF ZOOLOGY 2009 THE RAFFLES BULLETIN OF ZOOLOGY 2009 Supplement No. 20: 159–231 Date of Publication: 31 Jul.2009 © National University of Singapore A NEW CLASSIFICATION FOR THE PYLOCHELIDAE (DECAPODA: ANOMURA: PAGUROIDEA) AND DESCRIPTIONS OF NEW TAXA Patsy A. McLaughlin Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, Washington, 98221-4042, USA Email: hermit@fi dalgo.net Rafael Lemaitre Smithsonian Institution, National Museum of Natural History, Department of Invertebrate Zoology, P.O. Box 37012, Washington, D.C. 20013-7012, USA Email: [email protected] ABSTRACT. – A new classifi cation is presented based on the results of the recently completed cladistic analysis of the Pylochelidae. The subfamilies Pylochelinae and Pomatochelinae are retained, the latter with the genera Pylocheles and Cheiroplatea; however, the subgenera Xylocheles and Bathycheles are elevated to generic rank together with the nominal subgenus Pylocheles. In addition, one new species, B. phenax, is described in Bathycheles and B. profundus is shown to be conspecifi c with B. integer. The subfamilies Parapylochelinae, Cancellochelinae, Trizochelinae, and Mixtopagurinae are reduced to ranks of tribes and included in the subfamily Trizochelinae. A new genus Forestocheles is proposed in the tribe Trizochelini. Within the genus Trizocheles, subspecifi c rank for T. spinosus bathamae is deemed unjustifi ed and this taxon is placed in synonymy with the nominal subspecies T. spinosus spinosus. The correct identity of Trizocheles balssi is established and the species mistakenly thought to represent that taxon is described as T. hoensonae, new species. Trizocheles gracilis is found to be conspecifi c with T. boasi and an additional new species, T. mendanai, is added to the genus. The superfamilial ranks of Cheiroplateoidea, Pomatocheloidea, Pylocheloidea, and Cancellocheloidea proposed by Watabe (2007) are rejected, as is Birgusoidea. KEY WORDS. – Decapoda, Anomura, Paguroidea, Pylochelidae, new classification, Pylochelinae, Pomatochelinae Trizochelinae, tribes, new genus, new species. INTRODUCTION exploratory cruises undertaken by the Muséum national d’Histoire naturelle (MNHN) and the Offi ce de la Recherche Although the first pylochelid genera and species were Scientifi que et Technique Outre-Mer (ORSTOM), now the described in the late nineteenth century, because of the Institut de Recherche pour le Developpement (IRD) has cryptic habitats of these unusual paguroids they were rarely been examined. This increased abundance of study material, collected and consequently known from very few specimens. together with recent advances in cladistic methodology and Prior to the monographic review of Forest (1987a) only 19 computer generated phylogenetic analyses, made it possible species in fi ve genera had been described. In contrast to the for Lemaitre et al. (2009) to evaluate the interspecifi c and total of 60 specimens reported in all the previous literature intergeneric relationships within the Pylochelidae. From those accounts, Forest was able to examine more than 400 results and from detailed morphological investigations, we individuals collected from approximately 200 sites around are able to propose a new classifi cation, amplify some of the the world. Despite his revisionary efforts, which included existing descriptions and describe new taxa. the establishment of six subfamilies, Forest recognized the heterogeneity that still existed within the Pylochelidae Bate, Additionally, fossilized carapaces thought to be paguroids 1888, and suggested that future study might show that each recently have been recovered from the reefal and yellow algal subfamily should be afforded familial rank. limestones, respectively, of the Felsenkalke Formation, which date back to the Jurassic Period (van Bakel et al., 2008). Of It has only been recently that the substantial amount these, two are believed to represent new pylochelid genera, of supplemental material gathered during subsequent one each assigned to the tribes Trizochelini Forest, 1987a 159 McLaughlin & Lemaitre: New classifi cation of Pylochelidae and Mixtopagurini Bouvier, 1895 (as subfamilies). This is of the cornea measured across the dorsal surface. A number the fi rst report of fossil pylochelids and the earliest evidence of morphological characters used by Lemaitre et al. (2009) of paguroids in the fossil record, which lends support to in their cladistic matrix, but not considered by Forest the proposition that symmetry rather than the hypothesized (1987a) in his monograph, have been added to the generic asymmetry is the ancestral state of the Paguroidea. diagnoses. In addition to the new classifi cation and keys, abbreviated redescriptions are presented for previously well known species and more detailed redescriptions for those MATERIALS AND METHODS taxa where additional information is now available. When possible the species accounts are accompanied by photographs Specimens utilized in this reappraisal have come principally of living specimens. Complete descriptions and detailed from the exceptionally large collections of the Muséum illustrations are presented for the new species. Terminology national d’Histoire naturelle, Paris, France, but these have for the descriptions follows that of McLaughlin (2003) and been supplemented by specimens from the National Museum McLaughlin et al. (2007a). The arrangement in the text of of Natural History, Smithsonian Institution, Washington, subfamilies, tribes, genera and species follows the keys and D.C., USA (USNM), the National Taiwan Ocean University, is not meant to imply phylogenetic relationships. Keelung, Taiwan, Republic of China (NTOU), the Raffl es Museum of Biodiversity Research, National University of Singapore, Republic of Singapore (ZRC), and the National CLASSIFICATION AND JUSTIFICATION Institute of Water and Atmospheric Research, Ltd. (NIWA) [formerly New Zealand Oceanographic Institute (NZOI)], At the time of Forest’s (1987a) review of the Pylochelidae, Wellington, New Zealand. Type specimens housed in the the family was still considered part of the superfamily Instituto de Oceanología de la Academia de Ciencias de Coenobitoidea, a subdivision of Forest’s Section Paguridea. Cuba, Havana, Cuba (IOACC), the Kitakyushu Museum Martin & Davis (2001) rejected Forest’s classification, of Natural History and Human History, Kitakyushu, Japan accepting instead an earlier proposition (McLaughlin, 1983b) (ZLKU), the Museum of Comparative Zoology, Harvard that paguroids formed a monophyletic taxon, thus they University, Cambridge, Massachusetts, USA (MCZ), the abandoned Coenobitoidea and combined all existing families National Institute of Water and Atmospheric Research; in the superfamily Paguroidea. McLaughlin et al. (2007b) the Natural History Museum, London, United Kingdom confi rmed the monophyly of the Paguroidea but found, as (NHM), the South African Museum, Cape Town, South had Richter & Scholtz (1994), that the Pylochelidae was Africa (SAM), and the Zoological Museum University paraphyletic according to the defi nition of paraphyly given Copenhagen, Denmark (ZMUC), have been examined as by Hennig (1966), i.e., defi ned only by plesiomorphies. As well as those in the Muséum national d’Histoire naturelle and may be seen from Lemaitre et al.’s (2009) strict consensus National Museum of Natural History. Additional identifying cladogram of relationships among the genera of the institutional abbreviations used in the text are IM, Indian Pylochelidae, weighted against homoplasy, (Fig. 1), three Museum, Calcutta, India; NMCR, National Museum of the distinct evolutionary branches are clearly distinguished, one Philippines, Manila, Philippine Islands; NMNZ, National representing the subfamily Pylochelinae Bate, 1888 (branch Museum of New Zealand (now Museum of New Zealand Te A), the second the subfamily Pomatochelinae Stebbing, Papa Tongarewa), Wellington, New Zealand; ZSI, Zoological 1914 (branch B), and the third including the remaining four Survey of India. MUSORSTOM is the acronym for the joint other subfamilies (branch C), three of which are represented expeditions of the MNHN and ORSTOM; EBISCO is the by single species. In Lemaitre et al.’s unweighted analysis acronym for Exploration de la Biodiversité et ISolement (cladogram not shown), the Pylochelinae and Pomatochelinae en Mer du Corail; Panglao is the Philippine island around received strong Bremer support, eight and 16, respectively, which certain expeditions surveyed. Specific collection whereas the interfamilial relationships of the other subfamilies gear used precedes the station number; gear abbreviations were only weakly supported, if at all. In the weighted are: CC, otter trawl (shrimp); CP, beam trawl; DC, Charcot analysis, fi ve synapomorphies attest to the monophyly of dredge; DW, Warén dredge; BS, benthic sample. Latitudes the Pylochelinae: 1) reduction or loss of the rostrum; 2) and longitudes are given only for the start of each gear the loss of the epipod from the second maxilliped; 3, 4) the deployment. Additional abbreviations used in the text are Stn. cheliform terminations of the second and third maxillipeds; for station, R.V. for research vessel, coll. for collector, and and 5) the prominent elevation of the dorsodistal facet of the ovig. for ovigerous. Data for the Royal Indian Marine Survey carpus of each cheliped. Continuity of the linea transversalis vessel INVESTIGATOR has been taken from Anonymous is the only synapomorphy uniting the
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