ParagioPagurus: wordldwide review and new species 311 lemaitre r. — the genus Paragiopagurus lemaitre, 1996 (Crustacea, Decapoda, anomura, Paguroidea, Parapaguridae): a worldwide review and summary, with descriptions of five new species, in aHYONG S. t., CHaN t.-Y., COrbari l. & NG P. K. l. (eds), Tropical Deep-Sea Benthos 27. muséum national d’Histoire naturelle, Paris : 311-421 (mémoires du muséum national d’Histoire naturelle ; 204). iSbN : 978-2-85653-692-6. The genus Paragiopagurus Lemaitre, 1996 (Crustacea, Decapoda, Anomura, Paguroidea, Parapaguridae): A worldwide review and summary, with descriptions of five new species rafael Lemaitre Smithsonian institution, National museum of Natural History, Department of invertebrate Zoology, 4210 Silver Hill road, Suitland, mD 20746, USa lemaitrr@si.edu ABSTRACT A review of the deep-water hermit crab species of the genus Paragiopagurus Lemaitre, 1996 from the world oceans is presented. The core specimen base for this study has come primarily from the abundant collections of species of this genus obtained during French campaigns over the last four decades, and complemented with numerous specimens from many other deep-sea expeditions and depo- sited in various museum holdings around the world. Paragiopagurus is one of the most speciose genus among the Parapaguridae Smith, 1882, although it is considered a phylogenetically heterogeneous assemblage and does not appear to have an apomorphy of its own. Bathymetrically, the species range in depth from 36 to 2034 m, although they occur most frequently between 200 and 1000 m. The spe- cies utilize as housing, gastropod shells (or rarely scaphopod shells, siliceous sponges, or hollow pieces of wood) that may or may not be colonized by actinians or zoanthids. In this review, 24 species are recognized, of which five are new, P. laperousei n. sp., P. orthotenes n. sp., P. oxychelos n. sp., P. trilineatus n. sp., and P. umbonatus n. sp. The new species are fully described and illustrated. All previously known species of the genus are diagnosed or redescribed, and previously published illustrations of important taxonomic characters assembled and complemented, when useful, with new illustrations. The treatment of each species includes a full synonymy, materials examined (type and non-types), colouration, habitat or type of housing used, distribution, and remarks on taxonomy and morphologi- cal affinities. Colour photographs are included for 14 of the species. Parapagurus curvispina de Saint Laurent, 1974, a species tentatively moved after its description to Sympagurus Smith, 1883 and then to Paragiopagurus, is herein transferred with certainty to Oncopagurus Lemaitre, 1996. Parapagurus spinimanus Balss, 1911, a species that had been incorrectly placed in Paragiopagurus, is herein moved to Sympagurus. Parapagurus sculptochela Zarenkov, 1990, a taxon previously considered a junior synonym of Paragiopagurus boletifer rafael lemaitre312 (de Saint Laurent, 1972), is herein resurrected as a valid species of Paragiopagurus. The bathymetric and geographic distributions of Paragiopagurus species are summarized and briefly discussed, including a summary table, graph, and map with generalized distribu- tion patterns. RÉSUMÉ Le genre Paragiopagurus Lemaitre, 1996 (Crustacea, Decapoda, Anomura, Paguroidea, Parapaguridae) : une révison mondiale avec la description de cinq nouvelles espèces. Une révision mondiale des pagures profonds appartenant au genre Paragiopagurus Lemaitre, 1996 est présentée ici. Le cœur de cette étude repose essentiellement sur les abondantes collections d’espèces de ce genre collectées au cours des campagnes françaises durant les 40 dernières années et fut complété par de nombreux spécimens de plusieurs autres expéditions profondes, déposés dans différents muséums à travers le monde. Paragiopagurus est un des genres comprenant le plus d’espèces dans la famille des Parapaguridae Smith, 1882, bien qu’il soit considéré comme un ensemble phylogénétiquement hétérogène mais ne présentant pas d’apomorphies. Au niveau bathymétrique, les espèces sont distribuées de 36 à 2034 m de profondeur bien que la majorité d’entres elles soient présentes entre 200- 1000m. Les espèces utilisent des coquilles de Gastéropodes comme habitations (ou plus rarement des coquilles de scaphopodes, des éponges siliceuses ou des cavités dans des morceaux de bois) pouvant être ou pas colonisées par des actinaires ou zoanthaires. Dans cette révision, 24 espèces sont décrites dont 5 espèces nouvelles, P. laperousei n. sp., P. orthotenes n. sp., P. oxychelos n. sp., P. trilineatus n. sp., and P. umbonatus n. sp. Ces nouvelles espèces sont entièrement décrites et illustrées. Toutes les espèces précédemment décrites sont diagnosées ou re-décrites et les illustrations des caractères taxonomiques importants, déjà publiées, sont ici assemblées et com- plétées si nécessaire avec de nouvelles illustrations. La description de chaque espèce inclut une synonymie complète, la liste du matériel examiné (type et non-types), les couleurs, les habitats utilisés, la distribution et les remarques sur leurs affinités taxonomiques et mor- phologiques. Des photos couleurs sont présentes pour 14 de ces espèces. Parapagurus curvispina de Saint Laurent, 1974, une espèce déplacée après sa description dans les genres Sympagurus Smith, 1883 puis Paragiopagurus, est ici transférée avec certitude dans le genre Oncopagurus Lemaitre, 1996. Parapagurus spinimanus Balss, 1911, une espèce qui n’a pas été placée correctement dans le genre Paragiopagurus, est ici transferée dans le genre Sympagurus. Parapagurus sculptochela Zarenkov, 1990, un taxon précédemment consi- déré comme un synonyme Paragiopagurus boletifer (de Saint Laurent, 1972) est réactualisé comme une espèce valide appartenant au genre Paragiopagurus. Les distributions bathymétriques et géographiques des espèces de Paragiopagurus sont résumées et discutées brièvement, grâce à un tableau-résumé, graphe et carte présentant les patrons de distributions de ces espèces. INTRODUCTION When de Saint Laurent (1972) re-established the family Parapaguridae Smith, 1882, she included a total of 34 species in four genera. Of these, 31 species were placed in the genus Parapagurus Smith, 1879, of which three were subdivided into a total of 13 subspecies (seven for Parapagurus pilosimanus Smith, 1879, and three each for P. acutus de Saint Laurent, 1972 and P. bicristatus (A. Milne-Edwards, 1880)), and three species in the monotypic genera Tylaspis Henderson, 1885, Probeebei Boone, 1926, and Typhlopagurus de Saint Laurent, 1972. Although de Saint Laurent’s study was based on numerous samples obtained during earlier, classic deep-sea expeditions of the late 19th and first half of the 20th centuries, the specimens available to her for many of the taxa were few and often insufficient or difficult to use in ascertaining diagnostically useful specific characters, or for the understanding of morphological variations as well as geographic and bathymetric distributions. Since de Saint Laurent’s study, however, a wealth of new parapagurid specimens and data from numerous modern expeditions in the Atlantic, Pacific, and Indian Oceans has become available. This new material has been used in a number of revisionary or broadly focused taxonomic studies (e.g., Lemaitre 1986, 1989, 1990, 1994, 1996, 1997, 1998, 1999, 2004a, 2004b; Lemaitre & McLaughlin 1992), and in descriptions of new genera or species (Lemaitre 1993, 1998, 2004a, 2006a, 2006b; Lemaitre & Poupin 2003; Osawa 1995, 1996). The genus Paparapagurus, in particular, has undergone a number of revisions and taxonomic changes, with two of the species placed in that genus by de Saint Laurent (1972) moved to the monotypic genera Bivalvopagurus Lemaitre, 1993 and Tsunogaipagurus Osawa, 1995, and the remaining species either left in Parapagurus, or placed in the resurrected genus Sympagurus Smith, 1883, or moved ParagioPagurus: wordldwide review and new species 313 to one of the three genera, Strobopagurus Lemaitre, 1989, Oncopagurus Lemaitre, 1996, or Paragiopagurus Lemaitre, 1996. As result, the numbers of genera and species of Parapaguridae have increased by 250% and 220% respectively. The family currently includes 10 genera with a total of 75 valid species and at least eight recognized but undescribed species (McLaughlin et al. 2010), rather than the four genera and 34 species proposed for this family by de Saint Laurent (1972). Detailed, worldwide taxonomic reviews have been published so far for two of the most speciose genera, Parapagurus and Sympagurus, each containing 17 species (Lemaitre 1999, 2004a). The genus Strobopagurus has also been reviewed, although it contains only three species (Lemaitre 2004b). Among the non-monotypic genera of Parapaguridae, Paragiopagurus is the only one that has not been found to have an apomorphy of its own, although it contains a good number of species that tolerate shallower water (as shallow as 36 m) than most other species in other genera. This genus, as currently defined, is likely to represent a phylogenetically heterogeneous assemblage. In contrast, Oncopagurus species share the presence of an upwardly curved, slender epistomial spine (Lemaitre 1996), and Sympagurus species all posses a vestigial pleurobranch on the eight thoracic somite above each fifth pereopod (Lemaitre 2004a). The species of Strobopagurus are distinguished by their broad shield, stout ocular peduncles and corneas, straight dactyls of the ambulatory legs (second and third pereopods), and in males, the strongly twisted distal segments of the second gonopods. Aside from occurring nearly exclusively in deep water (> 1000 m), species of Parapagurus share a combination of characters such as the reduced ocular peduncles and corneas, deeply divided quadriserial gills, great variability in the proportions of segments of the right cheliped depending on size and sex, a broad (high) propodus on the fourth pereopod, and well developed male first and second gonopods (Lemaitre 1999). Since the genus Paragiopagurus was proposed by Lemaitre (1996), three new species have been described in this genus, including P. ventilatus Lemaitre 2004c, the first paguroid known to be associated with hydrothermal vents (Lemaitre 2004c), and two species have been transferred to other parapagurid genera. Furthermore, the large number of new specimens of many species, including information on colourations, that has been obtained and studied since 1996 has allowed a better evaluation of the morphological boundaries and variations of several poorly known species, clarification of several taxonomic problems, and has also provided a much clearer insight into their overall distributions. Additionally, five new undescribed species have been discovered during this study and found to be abundantly represented in these new collections. Including those five new species, a total of 24 valid species of Paragiopagurus are recognized in the review presented herein. Clearly, species of Paragiopagurus are among the most abundant and visible component of the macrocrustacean fauna on the outer continental shelf to upper slope zones of many ocean regions. Given all these developments and the accumulation of significant information since the genus was proposed in 1996, a worldwide taxonomic and distributional summary of all the species in this genus is now needed. All but eight of the 24 species discussed herein have been previously documented and illustrated in detail. Of those eight, five are new species that are fully described in this study, two (P. rugosus de Saint Laurent, 1972, and P. tuberculosus de Saint Laurent, 1972) were poorly known based on few badly preserved types and are redescribed; and one (Parapagurus sculptochela Zarenkov, 1990) is resurrected as a valid species from the synonymy of P. boletifer (de Saint Laurent, 1972). THE RIGHT CHELIPED AND THE MALE GONOPODS As in many parapagurids, the morphology of the right cheliped of Paragiopagurus species is frequently influenced by size and sex of specimens. Morphological variations of the right cheliped in species of this genus are similar to those documented for species of Parapagurus, Sympagurus, and Strobopagurus (see Lemaitre 1986, 1999, 2004a, 2004b). In contrast to species of other parapagurid genera, however, most species of Paragiopagurus can be differentiated by several characters of the right cheliped, and in particular of the chela. The shape of the chela is operculate in 13 of the 24 species of Paragiopagurus treated herein. The term “operculate” is used to describe a chela that serves as cover and functions as a kind of operculum for the opening of the housing of the hermit crab. An operculate chela is subcircular or ovate in shape, more or less dorsoventrally flattened (more so on the lateral half), with a sharply defined dorsolateral margin, and a dactyl set at a strongly oblique angle relative to the longitudinal axis of the palm. The dorsomesial margin of the dactyl is rafael lemaitre314 also sharply defined and often produced. The dorsal surfaces of the right chela can be smooth, armed with strong spines, or have setae that vary in density and type (e.g., simple, plumose, bristle-like). An intriguing feature of several species of Paragiopagurus is the armature exhibited on the ventral surfaces of the right chela. In some species, the armature consists of prominent, dense cylindrical, mushroom-like or mammilliform tubercles, in others of low, distinct tubercles, and yet in others, of spines. The intriguing right chela of one species, P. ruticheles (A. Milne-Edwards, 1891), is unique among parapagurids in having transverse furrows on the ventral surfaces of the palm and fingers of the right chela, and on the ventrolateral face of the carpus of the right cheliped. Although there is some degree of morphological variation related to size, the armature of the right chela serves to clearly distinguish several species. The presence in males of structurally modified first and second pleopods that function as gonopods, is one of the defining characters of the family Parapaguridae, although the reduction or loss of one or both pleopods seen in some species is considered an evolutionary tendency common in the Paguroidea in general (de Saint Laurent 1972). Herein, the first and second pleopods in males, when present, and whether paired or unpaired, are called gonopods regardless of degree of development. Species of several parapagurid genera invariably have well developed paired first and second gonopods, such as those of Parapagurus, Sympagurus, Strobopagurus, and the monotypic Tylaspis, Bivalvopagurus, and Tsunogaipagurus (see Lemaitre 1989, 1993, 1999, 2004a, 2004b; Osawa 1996). In Probeebei, the first gonopods are present and the second gonopods are absent, whereas in Typhlopagurus it is the reverse, the first are absent and the second are present. In contrast, the degree of development of the first and second gonopods, or their presence and whether they are paired or not, is variable within Oncopagurus and Paragiopagurus. Of the 24 species of Paragiopagurus 12 have well developed, paired first and second gonopods, whereas in the other 12 species the degree of development and presence of the first and/or second gonopods is variable. Of the latter 12 species, males of eight (P. acutus, P. bicarinatus, P. hirsutus, P. hobbiti, P. ruticheles, P. ventilatus, and two of the new species described herein), lack first gonopods, and there is only a uniramous, short and unpaired left second gonopod; in one species (P. insolitus), the first gonopods are absent, and the second pleopods are paired albeit moderately developed; and in one of the new species described herein, there are paired first gonopods, and only a uniramous, short and unpaired left second gonopod. MATERIALS AND METHODS For each species, the following information is included: full synonymy, material examined (including types), diagnosis, description or redescription, colouration (when known), habitat, distribution, and remarks. More complete sets of illustrations are presented for those species being described or redescribed, than for those where sufficiently detailed information exists in the recent literature. For the latter, only illustrations of the most important diagnostic features are included, assembled from previously published figures and supplemented with new figures when deemed useful. Unless otherwise stated, colour photographs are of specimens as fresh as possible. The parapagurid terminology used follows that defined or adopted by Lemaitre (1999, 2004a) in broad reviews of the genera Parapagurus and Sympagurus. The numbers or range in millimeters (mm) that follow the number and sex of specimens in the material examined sections, are measurements for shield length (sl), measured from the tip of the rostrum to the midpoint of the posterior margin of the shield. The list of material examined is arranged by regions from East to West, and within each ocean basin from North to South. Within each region, the expeditions and stations are listed chronologically. Unless otherwise stated, all specimens listed under type material have been examined. In the systematic part, the species are discussed in the same order as they appear in the key for their identification and do not suggest phylogenetic relationships. As usual, the key presented is meant as an aid for identifications, and the user should carefully compare with the descriptive text, figures and photographs, for confirmation. A large portion of the specimens used in this study have come from French campaigns and remain deposited in the Muséum national d’Histoire naturelle, Paris (MNHN), with some duplicates, where possible, deposited in the National Museum of Natural History, Smithsonian Institution, Washington D. C. (USNM). The rest of the material has come from the following museums or institutions: Bernice P. Bishop Museum, Honolulu, Hawaii (BPBM); Natural History Museum ParagioPagurus: wordldwide review and new species 315 and Institute, Chiba (CBM); Instituto de Ciencias del Mar-Decapoda, Barcelona (ICMD); Florida Museum of Natural History, University of Florida, Gainesville (UF); National Taiwan Ocean University, Keelung (NTOU); Queensland Museum, Brisbane (QM); Nationaal Natuurhistorisch Museum, Leiden (ex Rijksmuseum van Natuurlijke Historie, RMNH); Musée Océanographique, Monaco (MO); Rosenstiel School of Marine and Atmospheric Science, University of Miami (UMML); and Zoological Museum, Moscow State University (ZMMU). Other abbreviations used are: CC, otter trawl; CH, shrimp trawl; CP, beam trawl; CURASUB, Substation Curaçao submarine, Willemstaad; DW, Warèn dredge; N.O., Navire Océanographique; stn, station; spm, specimen; USFC, United States Fish Commission; CANCAP, Canarian-Cape Verdean Expeditions of the RMNH; TC, is used for the R.V. “Townsend Cromwell”, United States National Marine Fisheries Service, the vessel used to obtain most of the specimens from the Hawaiian Islands. The station data for the French campaigns and their acronyms, from which Paragiopagurus material was examined, can be found in the following publications, unpublished reports, or online in the MNHN website (http://www.mnhn.fr/ musorstom/). The original format for reporting latitude and longitude in these official sources or unpublished personal communications, has been retained. BATHUS 1-4: Richer de Forges & Chevillon (1996). BERYX 11: Lehodey et al. (1992), P. Maestrati (pers. comm.). BENTHAUS: B. Richer de Forges (pers. comm.). BENTHEDI: A. Crosnier (pers. comm.). BIOCAL: Richer de Forges (1990). BIOGEOCAL: Richer de Forges (1990). BOA 0, 1: B. Richer de Forges (pers. comm.). BORDAU 1, 2: A. Crosnier (pers. comm.). CHALCAL 1, 2: Richer de Forges (1990). CORAIL 2: Richer de Forges (1991). CORINDON: B. Richer de Forges (pers. comm.). EBISCO: P. Maestrati (pers. comm.). French Polynesia, “Marara”: Poupin et al. (1990), Poupin (1996a). HALICAL 1: P. Maestrati (pers. comm.). HALIPRO 1: Richer de Forges & Chevillon (1996). LITHIST: P. Maestrati (pers. comm.). Madagascar, N.O. “Vauban”: Crosnier (1978), P. Maestrati (pers. comm.). MUSORSTOM 1, 2: Forest (1981, 1985). MUSORSTOM 4-6: Richer de Forges (1990). MUSORSTOM 7: Richer de Forges & Menou (1993). MUSORSTOM 8: Richer de Forges et al. (1996). MUSORSTOM 8-10: B. Richer de Forges (pers. comm.). NORFOLK 1, 2: B. Richer de Forges (pers. comm.). SALOMON 1, 2: B. Richer de Forges (pers. comm.). SANTO: P. Bouchet (pers. comm.). SMIB 2-4: Richer de Forges (1990). SMIB 5: Richer de Forges (1993). SMIB 6: P. Maestrati (pers. comm.). SMIB 8: Richer de Forges & Chevillon (1996). SMIB 10: A. Crosnier (pers. comm.). TAIWAN 2000-2007: A. Crosnier, T.-Y. Chan (pers. comm.). VOLSMAR: Laboute et al. (1989). rafael lemaitre316 SPECIES LIST Paragiopagurus acutus (de Saint Laurent, 1972) P. bicarinatus (de Saint Laurent, 1972) P. boletifer (de Saint Laurent, 1972) P. bougainvillei (Lemaitre, 1994) P. diogenes (Whitelegge, 1900) P. fasciatus Lemaitre & Poupin, 2003 P. hirsutus (de Saint Laurent, 1972) P. hobbiti (Macpherson, 1983) P. insolitus Lemaitre, 1997 P. laperousei n. sp. P. macrocerus (Forest, 1955) P. orthotenes n. sp. P. oxychelos n. sp. P. pacificus (Edmondson, 1925) P. pilimanus (A. Milne-Edwards, 1880) P. rugosus (de Saint Laurent, 1972) P. ruticheles (A. Milne-Edwards, 1891) P. schnauzer Lemaitre, 2006 P. sculptochela (Zarenkov, 1990) P. trilineatus n. sp. P. tuberculosus (de Saint Laurent, 1972) P. umbonatus n. sp. P. ventilatus Lemaitre, 2004 P. wallisi (Lemaitre, 1994) SYSTEMATIC PART Family PARAPAGURIDAE Smith, 1882 Genus PARAGIOPAGURUS Lemaitre, 1996 Parapagurus – de Saint laurent, 1972 (in part). Sympagurus – lemaitre, 1989: 36 (in part). Paragiopagurus lemaitre, 1996: 207; 2000: 218 (type species by original designation: Sympagurus diogenes Whitelegge, 1900, gender masculine). — mclaughlin, 2003a: 118 (key). — Poore, 2004: 282. — Campos et al., 2005: 140. — mclaughlin et al., 2007: 317; 2010: 39. — De Grave et al., 2009: 26. DIAGNOSIS — Eleven pairs of biserial or quadriserial, phyllobranchiate gills. Shield about as broad as long; dorsal surface often with irregularly-shaped, weakly calcified areas medially. Corneas weakly to moderately dilated. Fourth segment of antennal peduncle armed with dorsodistal spine. Epistomial spine straight, or absent. Right chela usually with well deli- mited dorsomesial and dorsolateral margins; often operculate. Left cheliped well calcified, or sometimes weakly calcified on merus and carpus. Ambulatory legs (second and third pereopods) with dactyls broadly curved. Fourth pereopod with propodal rasp consisting of 1 or more rows of ovate corneous scales. Second pleonal somite with left pleuron terminating ParagioPagurus: wordldwide review and new species 317 in small subtriangular lobe. Males with weakly to moderately developed paired first and second pleopods modified as gonopods; or lacking first gonopods and with paired second pleopods modified as gonopods; or lacking first gonopods and with unpaired, rudimentary second left gonopod. Females with vestigial second right pleopod. HABITAT — Most species of this genus have been found using gastropod shells which frequently have a small actinian or zoanthid living on the shell, or occasionally with an actinian covering the entire shell. One species (P. schnauzer) has been found living in scaphopod shells; another species (P. hobbiti) in siliceous sponges; and at least a juvenile of one species (P. tuberculosus) has been found in a hollow piece of wood. DISTRIBUTION — Indian Ocean; western and central Pacific Ocean; and western and eastern Atlantic Ocean. Depth: 36 to 2034 m. SPECIES INCLUDED — See species list. REMARKS — Based on the original description, and without examining the type and only existing material of Parapagu- rus curvispina de Saint Laurent, 1974, Lemaitre (1989) first assigned this taxon to Sympagurus Smith, 1883, and subse- quently he (Lemaitre 1996) reassigned it to Paragiopagurus. Examination of the type of P. curvispina during the present study revealed that it has a curved epistomial spine, a character that is unique to species of Oncopagurus. Thus, de Saint Laurent’s (1974) taxon is herein correctly placed in Oncopagurus as O. curvispina n. comb. As discussed by Lemaitre (2004a: 93) and surmised by Zhadan in a manuscript that regrettably was never published (listed as “in press” in Lemaitre (2004a), M. Türkay, pers. comm.), Parapagurus spinimanus Balss, 1911 was incorrectly assigned by Lemaitre (1996: 207) to Paragiopagurus based exclusively on information from the literature. Examination of the holotype and only known specimen of Balss’ species, showed that it belongs correctly in Sympagurus. A recent phylogenetic study of anomurans (Tsang et al. 2011) used four genera and six species of parapagurids: Paragiopagurus (P. acutus, P. ventilatus, P. boletifer), Oncopagurus (O. orientalis de Saint Laurent, 1972), Strobopagurus (S. gracilipes (A. Milne-Edwards, 1891)), and Sympagurus (S. burkenroadi Thompson, 1943). Although the study did not focus on generic relationships, the consensus trees suggest that Paragiopagurus is more closely related to Oncopagurus than to the other genera included in the analyses. KEY TO SPECIES OF PARAGIOPAGURUS 1. Antennal acicles extending at most to mid-length of ocular peduncles, with mesial margins unarmed; telson with spines on lateral and posterior margins ................................................................................................. P. pacificus – Antennal acicles extending well beyond mid-length of ocular peduncles, with mesial margins armed with spines; telson with spines on posterior margin only ...................................................................................................... 2 2. Ocular acicles simple, terminating in spine ................................................................................................................. 6 – Ocular acicles multifid (at least on one side) ............................................................................................................... 3 3. Fourth pereopod with propodal rasp consisting of 1 row (at least distally) of ovate scales . ............................ 4 – Fourth pereopod with propodal rasp consisting of more than 1 row of ovate scales ........ P. trilineatus n. sp. 4. Palm of right chela with ventromesial margin well delimited by row of spines, mesial face expanded distally (Figure 6C) ......................................................................................................................................................... P. bicarinatus – Palm of right chela with ventromesial margin not delimited by row of spines, mesial face not expanded distally ........................................................................................................................................................................................ 5 5. Dorsal and often ventral faces of right palm armed with strong to moderately strong spines, and dense bristle-like setae (Figures 7B-D, 1C) ................................................................................................................. P. hirsutus rafael lemaitre318 – Dorsal and ventral faces of right palm smooth or at most with few weak spines or tubercles, and lacking dense setae (Figures 8B, C, 1D) ............................................................................................................................. P. acutus 6. Right cheliped with transverse furrows on ventral surface of chela and ventrolateral face of carpus (Figures 10B, D, E) .............................................................................................................................................................. P. ruticheles – Right cheliped without transverse furrows on chela or carpus ............................................................................... 7 7. Corneas reduced, subconical (Figure 11B, C); ocular peduncles diminishing in width distally, length (includ- ing corneas) less than half that of shield; uropods and telson nearly symmetrical; gill lamellae deeply divided (Figure 11A) ......................................................................................................................................................... P. schnauzer – Corneas not reduced or subconical; ocular peduncles not diminishing in width distally, length (including corneas) more than half that of shield; uropods and telson distinctly asymmetrical; gill lamellae undivided (e.g., Figures 2A, 30A) or at most weakly divided distally (e.g., Figures 2B, 13A, 16A) ......................................... 8 8. Ventral surface of right palm with numerous large tubercles, or with distinct low, well-spaced, blister-like tubercles, or densely covered with small spines .............................................................................................................. 9 – Ventral surface of right palm smooth or at most with scattered small spines or tubercles ............................. 12 9. Ventral surface of right palm densely covered with small spines often with minute corneous-tips (Figures 13E, 15E) ................................................................................................................................................................... P. rugosus – Ventral surface of right palm not covered with small spines, instead armed with numerous large tubercles or distinct low, well-spaced, blister-like tubercles ......................................................................................................... 10 10. Ventral surface of right palm with distinct low and well-spaced, blister-like tubercles (Figure 18B, D, E); dorsal surface of palm densely covered with mostly plumose setae on entire surface ........ P. umbonatus n. sp. – Ventral surface of right palm without low, blister-like tubercles; dorsal surface of palm with scattered or at most moderately dense setation ......................................................................................................................................... 11 11. Ventral surface of right palm armed with numerous large mushroom-like or mammilliform tubercles (Figure 21D, E, F) ................................................................................................................................................... P. boletifer – Ventral surface of right palm with numerous large tubercles not mushroom- or mammilliform-like (Figure 15C, D) ............................................................................................................................................................. P. sculptochela 12. Antennal acicles exceeding distal margins of corneas by 0.3 to 0.5 length of acicle (Figure 23A); shield distinctly broader than long .......................................................................................................................... P. macrocerus – Antennal acicles at most reaching distal margin of corneas; shield longer than broad or about as long as broad ......................................................................................................................................................................................... 13 13. Ocular acicles terminating in long slender spine extending to nearly mid-length or more of ocular pe- duncles ...................................................................................................................................................................................... 14 – Ocular acicles terminating in short spine reaching to at most 0.2 length of ocular peduncles ...................... 15 14. Right chela not operculate, palm with dorsolateral margin at most weakly delimited; dactyl of fourth pereo- pod with corneous claw masked by long plumose setae arising near base of claw (Figure 25D, E); epistome with median region strongly produced anteriorly, somewhat pyramid-shaped (Figure 24C) ........... P. insolitus – Right chela operculate, palm with dorsolateral margin well delimited by row of spines; dactyl of fourth pereopod without long setae masking corneous claw; epistome with median region not produced anteriorly P. orthotenes n. sp. 15. Dorsomesial margin of dactyl of right chela finely serrate, drawn out as thin, somewhat translucent ridge with transverse lines suggesting spine fusion (Figure 32D) .......................................................... P. oxychelos n. sp. – Dorsomesial margin of dactyl of right chela not as above ....................................................................................... 16 16. Right chela operculate, with dorsolateral margin sharply defined, and dactyl set at strongly oblique angle (distinctly > 45°) to longitudinal axis of palm (e.g., Figure 34C, D) ........................................................................... 17 – Right chela not operculate, with dorsolateral margin rounded, and dactyl set at weakly oblique angle ParagioPagurus: wordldwide review and new species 319 ( distinctly < 45°) to longitudinal axis of palm (e.g., Figure 46B, C) ............................................................................ 21 17. Left cheliped with distinct tubercles on dorsal surfaces of merus and palm, and with row of 8–10 (less in juveniles) strong spines on dorsal margin (Figure 34E); palm of right cheliped with numerous tubercles (Figure 34C); second and third pereopods each with 2 red stripes on lateral faces of meri, carpus, propodi and dactyls (Figure 1I) ............................................................................................................................................................... P. fasciatus – Left cheliped without tubercles on dorsal surfaces of merus and palm, at most with scattered small spines, and with row of weak spines on dorsal margin .............................................................................................................. 18 18. Dorsal surface of right chela smooth, or at most armed with minute spines or tubercles .............P. diogenes – Dorsal surface of right chela armed with many distinct spines or tubercles ...................................................... 19 19. Gill lamella undivived distally (Figure 37A) or with oblique distal margin; dorsal surfaces of fingers of right cheliped armed with weak, well spaced spines or tubercles; eyes pink, second and third pereopods white ex- cept for orangish dorsal margin of carpi, and orangish spots on dorsal margins of propodi and dactyls (Figure 35B–D) ...................................................................................................................................................... P. laperousei n. sp. – Gill lamella divived distally (Figures 42A, 43A ); dorsal surfaces of fingers of right cheliped armed with strong low, mammilliform tubercles or spines; colour not as above ........................................................................ 20 20. Posterior margin of telson having several large corneous spines with distal portions abruptly curved ven- trolaterally (Figure 43H); second and third pereopods with numerous small reddish spots on cream or yel- lowish background, and lateral faces of carpi of left cheliped and second and third pereopods each with large red spot (persisting for several years in preservative; Figure 35F) ..................................................... P. bougainvillei – Posterior margin of left lobe of telson without large corneous spines abruptly curved ventrolaterally (Figure 42I); second and third pereopods uniformly pinkish or orangish, and without large spot on lateral faces of carpi of left cheliepd and second and third pereopods (Figure 35E) ........................................................... P. wallisi 21. Propodal rasp of fourth pereopod with 1 row of ovate scales (at least distally) ............................................... 22 – Propodal rasp of fourth pereopod with 2 or more rows of ovate scales (at least distally) ............................... 23 22. Chelipeds and ambulatory legs covered with numerous tufts of setae; ocular acicles with transverse striae; bacteriophore setae present on outer faces of basis and ischium of third maxillipeds, and mesial and ventral faces of merus and ischium of chelipeds (except in juveniles) ................................................................ P. ventilatus – Chelipeds and ambulatory legs at most with scattered short setae; ocular acicles without transverse striae; bacteriophore setae absent from third maxillipeds and chelipeds .............................................................. P. hobbiti 23. Dorsal surface of right chela with strong, well spaced spines; telson strongly asymmetrical ........................... .................................................................................................................................................................................. P. pilimanus – Dorsal surface of right chela with scattered small spines or tubercles or with numerous blunt tubercles or spines; telson weakly asymmetrical ........................................................................................................... P. tuberculosus Paragiopagurus pacificus (Edmondson, 1925) Sympagurus pacificus edmondson, 1925: 28, fig. 5 e-g, pl. 2C (type locality: off laysan, Hawaiian islands,). — eldredge, 1967: unpaginated list. — lemaitre, 1989: 37; 1994: 412. Parapagurus pacificus – de Saint laurent, 1972: 108. Paragiopagurus pacificus – lemaitre 1996: 207. — mclaughlin et al., 2005: 246; 2010: 39. — Castro, 2011: 30. TYPE MATERIAL — Holotype, Hawaiian Islands, off Laysan, 365.8 m, male (“total length 13 mm, length of carapace 5 mm, width of front border of carapace 2 mm” – Edmondson 1925: 28) (BPBM C-1816); presumably lost. rafael lemaitre320 MATERIAL EXAMINED — None. DISTRIBUTION — Known only from the holotype collected off Laysan, one of the northwestern Hawaiian Islands, at a depth of 365.8 m. DESCRIPTION — See Edmondson (1925: 28). COLOUR — Unknown. HABITAT — Unknown. REMARKS — This species was tentatively placed in Paragiopagurus by Lemaitre (1996). The holotype and only specimen collected has not been found, and is presumed lost. Although Edmondson (1925) did not indicate the sex of the holotype, de Saint Laurent (1972) did examine the type specimen and mentioned that it was a male in poor condition. Unfortuna- tely, a sufficiently detailed diagnosis of this species is not possible because Edmondson’s (1925) description lacks details of most critical diagnostic features, his figures are sketchy and could apply to a number of species of Paragiopagurus or even Oncopagurus. Edmondson’s figures, if accurate, show several unique features that may set his species apart from other Paragiopagurus species, including the short antennal acicles that are shown to extend only to midlength of the ocular peduncles, and the armature of the telson, shown with spines throughout the terminal and lateral margins. Alternatively, Edmondson’s specimens could be juveniles of one of the other known species of Paragiopagurus. Paragiopagurus trilineatus n. sp. Figures 1a , 2-5 Paragiopagurus sp. 3 – mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, Tonga, BORDAU 2, stn CP 1593, 19°06’S, 174°18’W, 436-442 m, 14.06.2000, male 7.1 mm (MNHN-IU-2008-15886). Paratypes: Solomon Islands. SALOMON 2: stn CP 2597, 9°05.69’S, 158°15.98’E, 728-777 m, 11.08.2004, 6 males 4.2-7.1 mm, 2 ovigerous females 5.4, 6.1 mm, 1 female 3.4 mm (MNHN-IU-2012-995). Fiji. MU- SORSTOM 10: stn CP 1327, 17°13.26’S, 177°51.62’E, 370-389 m, 07.08.1998, 1 ovigerous female 4.6 mm, 2 females 4.3, 5.7 mm (MNHN-IU-2008-15884). – stn DW 1375, 18°18,43’S, 178°07.37’W, 409-480 m, 17.08.1998, 1 male 8.1 mm (MNHN-IU-2008-15885). – BORDAU 1: stn CP 1395, 16°45.13’S, 179°59.20’E, 423-500 m, 23.02.1999, 1 female 5.1 mm (MNHN-IU-2008-15878). – stn DW 1432, 17°19.90’S, 178°44.25’W, 477-493 m, 02.03.1999, 1 male 8.8 mm (MNHN- IU-2008-15879). – stn CP 1433, 17°19.93’S, 178°42.89’W, 488-500 m, 02.03.1999, 5 males 2.5-4.7 mm, 4 females 2.3- FIGURE 1 Coloration (in life or fresh except where noted). A, Paragiopagurus trilineatus n. sp., preserved male (7.2 mm) mNHN-iU-2008-15883, tonga , bOrDaU 2 stn CH 1596, 371-437 m. B, P. bicarinatus (de Saint laurent, 1972), taiwan, taiWaN 2000 stn CP 23, 880-1070 m (specimen not located , from mclaughlin et al. 2007: 325, unnumbered photo.). C, P. hirsutus (de Saint laurent, 1972), taiwan, taiWaN 2003 stn CP 212, 223-260 m (NtOU, from mclaughlin et al. 2007: 327, unnumbered photo.). D, P. acutus (de Saint laurent, 1972), tokyo bay, Japan (Cbm-ZC 9898, from mclaughlin et al. 2010: 102, fig. 19b). E, P. rugosus (de Saint laurent, 1972), female (9.2 mm) UF 12147, Hawaiian islands, French Frigate Shoals, baited traps, 185-251 m. F, P. boletifer (de Saint laurent, 1972), taiwan, taiWaN 2003 stn CP 212, 223-260 m (NtOU, from mclaughlin et al. 2007: 318, unnumbered photo.). G, P. boletifer, anterior portion, ventral view (right second and third pereopods shown partially), austral islands, French Polynesia (mNHN-Pg 5137, from lemaitre 1994: fig. 28c, as Sympagurus boletifer). H, P. oxychelos n. sp., paratype male (3.5 mm) NtOU a01347, taiwan, taiWaN 2003 stn CP 212, 223-260 m. I, P. fasciatus lemaitre & Poupin, 2003, austral islands, French Polynesia (mNHN-Pg 6916, from lemaitre & Poupin 2003: fig. 4). (Photographs: a, r. lemaitre; b, C, F, H, t.-Y. Chan; D, t. Komai; e, G. Paulay; G, i, J. Poupin). ParagioPagurus: wordldwide review and new species 321 A B E C D G F I H rafael lemaitre322 FIGURE 2 Paragiopagurus trilineatus n. sp. A, B, F, paratype female (8.0 mm) USNm 1016948, Fiji, bOrDaU 1 stn CP 1433, 480-500 m. C-E, G, holotype male (7.1 mm) mNHN-iU-2008-15886, tonga, bOrDaU 2 stn 1593, 436-442 m. A, B, lamella of posterior arthrobranch. C, shield and cephalic appendages, dorsal view. D, right antennal peduncle and anterior portion of branchiostegite, lateral view. E, telson, dorsal view. F, distal margin of telson, dorsal view. G, left second pleopod, lateral view. Scale bars 0.25 mm for a, b; 2 mm for C; 1 mm for D; 0.5 mm for e-G. A B FG E C D A, B C D E-G ParagioPagurus: wordldwide review and new species 323 8.0 mm (USNM 1016948, ex MNHN-Pg6993). – stn CP 1448, 16°45.04’S, 179°58.97’E, 410-500 m, 04.03.1999, 1 male 4.1 mm, 1 female 3.1 mm (MNHN-IU-2008-15881). – stn DW 1488, 19°01.33’S, 178°25.16’W, 500-516 m, 10.03.1999, 1 male 6.3 mm (MNHN-IU-2008-15882). Vanuatu. BOA 1: stn CP 2413, 15°39.33’S, 15°41.5’E, 268-445 m, 05.09.2005, 1 female 6.8 mm (MNHN-IU-2012-996). Tonga. BORDAU 2: stn CH 1596, 19°06’S, 174°18’W, 371-437 m, 14.06.2000, 10 male 2.5-7.2 mm, 1 ovigerous female 5.2 mm, 1 female 4.4 mm (MNHN-IU-2008-15883). New Caledonia. EBISCO: stn CP 2611, 19°36.446’ S, 158°40.838’ E, 492-506 m, 10.19.2005, 1 male 4.1 mm, 1 female 4.5 mm (MNHN-IU-2012-997). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — Western Pacific: from Solomon Islands, Fiji, Vanuatu, Tonga, and New Caledonia. Depth: 370 to 516 m. DESCRIPTION — Eleven pairs of biserial or quadriserial gills (Figure 2A, B) with lamellae weakly divided distally. Shield (Figure 2C) approximately as broad as long; dorsal surface weakly calcified along midline, just posteriorly to rostrum, and anterior and anterolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections bluntly subtriangular, sometimes with small terminal spine. Ante- rolateral margins sloping. Ventrolateral margins with small spine on one or both sides. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, with longitudinal row of short setae on dorsal surfaces. Corneas moderately dilated. Ocular acicles subtriangular, about 0.25 times as long as ocular peduncles, each terminating in strong multifid spine; separated basally by about half width of 1 icicle. Antennular peduncles exceeding distal margin of corneas by 0.75 or more length of ultimate segment; ventral flagellum with about 7 articles. Ultimate segment about twice as long as penultimate segment, with scattered setae dorsally. Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 3 or 4 spines, and strong spine proximally. Antennal peduncles (Figure 2D) reaching to, or slightly exceeding distal margin of corneas. Fifth segment unarmed, with few, short bristle-like setae on dorsolateral distal angle, and mesial margin. Fourth segment with strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong spine reaching to about midpoint of acicle; mesial margin with strong, usually slender spine on dorsolateral distal angle. First segment with lateral surface armed with small spine; ventromesial angle produced, with row of 3-5 spines laterally. Antennal acicle nearly straight or weakly sinuous (dorsal view), reaching to or at most slightly exceeding distal margin of cornea (or falling just short in small specimens sl < 3. 0 mm), and terminating in strong spine; mesial margin with few setae, armed with row of 9 to 13 strong spines diminishing in strength distally. Flagellum exceeding distal end of right cheliped; with scattered short (< 1 article in length) setae. Mandible (Figure 3A) with 3-segmented palp. Maxillule (Figure 2B) with external lobe of endopod moderately developed, internal lobe with 3 long setae. Maxilla (Figure 3C) with endopod slightly exceeding distal margin of scaphognathite. First maxilliped (Figure 3D) with endopod exceeding distal margin of exopod. Second maxilliped (Figure 3E) without distinguishing characters. Third maxilliped (Figure 3F) with dactyl about 2.5 as long as wide, and capus and propodus about twice as long as wide; crista dentata with about 12 or 13 corneous-tipped teeth decreasing in size distally; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine short, sharp, or occasionally absent. Chelipeds markedly dissimilar. Right cheliped (Figure 4A-C) massive, chela, carpus and merus sparsely setose; chela somewhat dorsoventrally flattened, operculate. Fingers slightly curving ventromesially, each terminating in small, usually blunt corneous claw; cutting edges with row of calcareous teeth, that of dactyl also with short row of small corneous spines distally; dorsal faces with numerous small spines or tubercles, and few tufts of setae near cutting edges; ventral faces smooth or with low tubercles. Dactyl subequal in length to palm, set at weakly oblique angle to longitudinal axis of palm; mesial margin well defined by row of spines; ventromesial face concave. Fixed finger broad at base. Palm broader rafael lemaitre324 than long; lateral margin well delimited by row of spines; dorsomesial margin delimited by row of small spines; mesial face rounded, with small spines or tubercles; dorsal surface entirely covered with numerous small spines or tubercles; ventral surface usually smooth mesially, and bearing tubercles laterally. Carpus covered with many small spines or tubercles on dorsal surfaces; ventrolateral margin relatively well defined by row of spines; dorsodistal and ventromesial distal margins each with row of small spines. Merus with small spines or tubercles on dorsal surfaces, and row of small spines on dorsodistal margin; ventral surface nearly smooth; ventromesial distal margin with row of spines. Ischium unarmed except for setae on dorsal margin and ventral surface; coxa with row of spines on ventrodistal margin, and ventromesial row of long setae. Left cheliped (Figure 4D) usually well calcified. Fingers terminating in sharp corneous claws, slightly gaping when closed; dorsal and ventral surfaces unarmed except for tufts of setae; cutting edge of dactyl with row of small calcareous FIGURE 3 Paragiopagurus trilineatus n. sp., paratype male (8.1 mm) mNHN-iU-2008-15885, Fiji, mUSOrStOm 10 stn DW 1375, 409-480 m, left mouthparts, internal view. A, mandible. B, maxillule. C, maxilla. D, first maxilliped. E, second maxilliped. F, third maxilliped. Scale bars 0.5 mm for a-D; 1 mm for e, F. F E A-D E D F C B A ParagioPagurus: wordldwide review and new species 325 FIGURE 4 Paragiopagurus trilineatus n. sp., holotype male (7.1 mm) mNHN-iU-2008-15886, tonga, bOrDaU 2 stn 1593, 436-442 m. A, right cheliped, dorsal view. B, chela of same, mesial view. C, same chela, lateral view. D, left cheliped, dorsal view. Scale bar 2 mm. ADB C rafael lemaitre326 FIGURE 5 Paragiopagurus trilineatus n. sp., holotype male (7.1 mm) mNHN-iU-2008-15886, tonga, bOrDaU 2, stn 1593, 436-442 m. A, right second pereopod, lateral view. B, dactyl of same, mesial view. C, third right pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of left fourth pereopod, lateral view. F, propodus and dactyl of left fifth pereopod, lateral view. G, anterior lobe of sternite Xii (of third pereopods), ventral view. Scale bars 2 mm for a-D; 0.25 mm for e-G. E-G A-D F E D C B G A ParagioPagurus: wordldwide review and new species 327 teeth, cutting edge of fixed finger with row of closely-set small corneous spinules. Dactyl shorter than palm. Palm with dorsomesial and dorsomedian rows of small spines; mesial face setose. Carpus with 1 or 2 irregular rows of small spines on dorsal margin; dorsomesial and dorsolateral faces setose; ventral face setose. Merus unarmed except for dorsal and ventral bristle-like setae, and row of spines on ventrolateral distal margin. Ischium unarmed except for dorsal and ventral setae. Coxa with row of spines on ventrodistal margin, and ventromesial row of long setae. Ambulatory legs (Figure 4A-D) similar from right to left, usually exceeding right cheliped by about half length of dactyl of legs when fully extended. Dactyl approximately 1.7 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, ventromesial margin unarmed or with 1-3 minute corneous spinules; lateral and mesial faces with shallow, longitudinal sulcus on proximal half (deeper on mesial face). Propodus with setae or rows of short transverse setae on dorsal margin. Carpus with small dorsodistal spine, and short setae on dorsal margin; carpus of first leg with irregular row of small spines (stronger on right carpus) on dorsal margin. Merus with setae on dorsal margin; merus of second pereopod with ventrodistal row of small spines. Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods) setose, with or without short, blunt subterminal spine (Figure 5G). Fourth pereopod (Figure 5E) subchelate. Dactyl broadly curved, terminating in sharp corneous claw; with ventrolateral row of small corneous spinules. Propodus longer than wide, rasp consisting of 2 or 3 rows of rounded scales (smaller specimens sl < 4.0 mm sometimes with 1 row of scales distally). Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 5F) chelate. Propodal rasp extending posteriorly to or beyond mid-length of segment. Uropods and telson markedly asymmetrical. Telson (Figure 2E, F) lacking lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 rounded projections by wide, shallow cleft, left projection often considerably produced; rounded projections each armed with several rows of slender, corneous spines, those on left projection in females often numerous, with some strong and curved ventrally (Figure 2F). Male lacking first gonopods; with rudimentary, uniramous, bud-like, unpaired left second gonopod (Figure 2G). Female lacking rudimentary second right pleopod. COLOUR (preserved, Figure 1a) — Ocular peduncles with ventral surface dark orange; dorsal surface light orange. Shield cream with light orange areas near anterior and anterolateral margins. Right cheliped with chela cream to white dorsally except for light orange near lateral, mesial, distal and proximal margins, and short orange stripe medially near proximal margin; carpus with 3 distinct, broad, dark orange stripes dorsally. Left cheliped with chela cream; carpus with dorso- lateral orange stripe. Ambulatory legs with meri, carpi, and propodi each with orange stripe laterally, that on propodi narrowing and fading to cream at midpoint. All colours fading in time to straw white. HABITAT — Gastropod shells, often with actinians or zoanthinds growing on shells. REMARKS — This new species is one of eight in Paragiopagurus in which males lack first gonopods, and have rudimen- tary, unpaired left second gonopods (Figure 2G), the others being P. acutus, P. bicarinatus, P. hirsutus, P. hobbiti, P. ortho- tenes n. sp., P. ruticheles, and P. ventilatus. All these species, however, are otherwise very different morphologically and in colouration. The multifid condition of the ocular acicles of P. trilineatus n. sp. is shared only with three species: P. acutus, P. bicarinatus, and P. hirsutus. Among other differences, P. trilineatus n. sp. differes form those three species by having two or three rows of scales on the propodal rasp of the fourth pereopod instead of only one row. ETYMOLOGY — The specific name is derived from the Latin tri, three, and lineatus, linear, and refers to the three distinct orange stripes on the carpus of the right cheliped of this new species. rafael lemaitre328 Paragiopagurus bicarinatus (de Saint Laurent, 1972) Figures 1b, 6 Parapagurus acutus bicarinatus de Saint laurent, 1972: 113 (type locality: Philippines). Sympagurus acutus bicarinatus – lemaitre, 1989: 37; 1994: 412. Paragiopagurus bicarinatus – lemaitre, 1996: 214, fig. 27. — rahayu, 2000: 397. — Davie 2002: 89. — mclaughlin et al., 2007: 325, 2 unnumbered figs.; 2010: 39. — Wang, 2008: 759. TYPE MATERIAL — Holotype, Philippines, southern Luzon, 5 mi (9.2 km) NW Matocot Point, Albatross, stn 5289, 13°41’50”N, 120°58’03”E, 315 m, 22.07.1908, male 5.6 mm (USNM 168310). [See Lemaitre (1996: 166) for comments on original material used in the description of this species by de Saint Laurent (1972).] MATERIAL EXAMINED — the type material (see above). Solomon Islands. SalOmON 1: stn DW 1762, 8°39.9’S, 160°03.9’e, 396-411 m, 27.09.2000, 1 male 2.3 mm (mNHN-Pg 6766). – stn CP 1783, 8°32.8’S, 160°41.7’e, 399-700 m, 29.09.2000, 1 male 3.9 mm (mNHN-Pg 6767), 2 females 1.9, 3.2 mm (mNHN-Pg 6768). – stn CP 1794, 9°16.1’S, 160°07.7’e, 494-504 m, 30.09.2000, 1 male 2.8 mm (mNHN-Pg 6769). Vanuatu. SaNtO: stn at 18, 15°41.3’S, 167°02.6’e, 321-336 m, 21.09.2006, 1 male 2.6 mm (mNHN-iU-2012-998). Tonga. bOrDaU 2: stn CP 1526, 21°16’S, 174°59’W, 463-464 m, 02.06.2000, 1 ovigerous female 2.9 mm (mNHN-Pg 6765). – stn CP 1545, 21°17’S, 175°17’W, 444-447 m, 05.06.2000, 3 males 2.3-2.9 mm, 3 ovigerous females 2.7-4.1 mm (mNHN-iU-2008-15724). – stn CP 1593, 19°06’S, 174°18’W, 436-442 m, 14.06.2000, 4 males 2.1-5.6 mm, 5 ovigerous females 2.0-3.0 mm, 3 females 1.7-4.7 mm (mNHN-iU-2008-15735). – stn CP 1643, 21°05’S, 175°22’W, 487 m, 22.06.2000, 1 male 2.0 mm (mNHN- iU-2008-15727). Chesterfield Islands, and Lansdowne and Fairway Banks. COrail 2: stn De 16, 20°48’S, 160°56’e, 500 m, 21.07.1988, 3 males 2.2-4.2 mm, 2 ovigerous females 2.2, 3.2 mm (USNm), 1 male 2.9 mm (mNHN-Pg 6761). New Caledonia. mUSOrStOm 4: stn CP 171, 18°57.80’S, 163°14.00’e, 435 m, 17.09.1985, 1 female 1.8 mm, 2 ovigerous females 1.8, 2.1 mm (USNm). – stn CP 179, 18°56.60’S, 163°13.70’e, 480 m, 8.09.1985, 1 male 2.5 mm (USNm), 2 females 2.1, 2.5 mm (USNm). – stn CC 201, 18°55.80’S, 163°13.80’e, 500 m, 20.09.1985, 2 ovigerous female 2.7, 2.9 mm (USNm). – mUSOrStOm 5: stn DC 378, 19°53.74’S, 158°38.30’e, 355 m, 20.10.1986, 2 males 2.4, 2.5 mm (USNm). – ebiSCO: stn DW 2497, 24°44.54’S, 159°43.111’e, 485-500 m, 10.06.2005, 1 female 4.5 mm (mNHN-iU-2012-999). – stn DW 2565, 20°22.31’S, 158°40.51’e, 414-419 m, 10.13.2005, 1 male 1.9 mm, 1 ovigerous female 2.2 mm, 1 female, 2.8 mm (mNHN-iU-2012-1000). – stn DW 2585, 19°35.492’S, 158°43.422’e, 543-544 m, 10.15.2005, 2 males 3.8, 4.8 mm (mNHN-iU-2012-1001). – stn DW 2598, 19°43.016’S, 158°40.292’e, 496-505 m, 10.17.2005, 1 male 3.8 mm (mNHN-iU-2012-1002). – stn CP 2599, 19°42.500’S, 158°44.099’e, 586-600 m, 10.18.2005, 1 male 3.4 mm (mNHN-iU-2012-1003). – stn CP 2600, 19°39.203’S, 158°45.249’e, 603-630 m, 10.18.2005, 1 male 2.6 mm (mNHN-iU-2012-1004). – stn DW 2601, 19°36.400’S, 158°42.775’e, 500-547 m, 10.18.2005, 2 males 2.9, 3.7 mm (mNHN-iU-2012-1005). – stn DW 2603, 19°37.761’S, 158°43.898’e, 568-570 m, 10.18.2005, 1 male 2.3 mm (mNHN-iU-2012-1006). – stn DW 2607, 19°34.300’S, 158°40.489’e, 400-413 m, 10.18.2005, 1 female 2.8 mm (mNHN-iU-2012-1007). – stn CP 2614, 19°37.766’S, 158°45.246’e, 608-642 m, 10.19.2005, 1 female 3.3 mm (mNHN-iU-2012-1008). – stn DW 2617, 20°05.805’S, 160°23.043’e, 427-505 m, 10.20.2005, 1 male 2.0 mm, 1 ovigerous female 2.0 mm, 1 male 2.4 mm (mNHN-iU-2012-1009). – stn CP 2620, 20°05.864’S, 160°22.318’e, 522-623 m, 10.20.2005, 1 male 1.9 mm, 1 female 2.5 mm (mNHN-iU-2012-1010). – stn DW 2625, 20°04.872’S, 160°20.039’e, 627-741 m, 10.20.2005, 1 male 2.1 mm (mNHN-iU-2012-1011). – stn DW 2638, 20°47.933’S, 161°00.210’e, 418-421 m, 10.22.2005, 1 ovigerous female 1.9 mm (mNHN-iU-2012-1012). For additional material examined, inclu- ding specimen illustrated in Fig. 5, see lemaitre (1996). DISTRIBUTION — Western Pacific: Taiwan, South China Sea (off Hong Kong), Philippines, Solomon Islands, Vanuatu, Tonga, New Caledonia, Chesterfield Island and Lansdowne and Fairway Banks, and Australia. Depth: 274 to 1070 m. DIAGNOSIS — Gills with lamellae undivided distally. Shield (Figure 6A) as long as broad; dorsal surface weakly calcified medially; rostrum broadly rounded, with low dorsal ridge; anterior margins weakly concave; lateral projections subtrian- gular, terminating in small spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, each terminating in strong multifid spine; corneas slightly dilated. Maxillule with internal lobe of endopod bearing 4 long seta. Antennular peduncles exceeding distal margin of corneas by 0.75 length of ultimate segment. Antennal peduncles reaching distal margin of corneas; second segment with dorsolateral distal angle produced, terminating in strong simple spine reaching to midpoint of antennal acicle; acicle reaching distal margin of corneas, mesial margin of each with 10 to ParagioPagurus: wordldwide review and new species 329 FIGURE 6 Paragiopagurus bicarinatus (de Saint laurent, 1972), male (4.1 mm) Qm, off Queensland, australia. A, shield and cephalic appendages. B, right cheliped. C, chela of same. D, telson. all in dorsal view except C, mesial view. Scale bars 1 mm. (adapted from lemaitre 1996). A D B, C D A B C rafael lemaitre330 13 spines; flagellum with setae < 1 to 4 flagellar articles in length. Chelipeds markedly dissimilar, with moderately dense setae. Right cheliped (Figure 6B, C) with palm broader than long, fingers curved ventromesially, dactyl with concave ven- tromesial face; dorsal face of palm with numerous small spines or tubercles, ventral face smooth or with scattered small tubercles; palm with dorsolateral, dorsomesial, and ventromesial margins each well delimited by row of strong spines, and ventromesial face expanded distally; carpus with numerous small tubercles or spines on dorsal surface, and well delimited dorsolateral margin with row of spines. Left cheliped with lateral face of carpus usually weakly calcified; carpus with dorsal row of 6 to 15 small spines, and strong dorsodistal spine. Ambulatory legs with dactyls having ventromesial row of 3 or 4 small corneous spines, and dorsal and dorsomesial rows of long setae; carpi each with small dorsodistal spine; carpus of second pereopod with 2 small dorsal spines (spines sometimes obsolete). Anterior lobe of sternite XII (of third pereopods) setose, armed with 1 marginal spine. Fourth pereopod with propodal rasp consisting of 1 row of ovate scales. Uropods and telson markedly asymmetrical; telson lacking transverse suture separating anterior and posterior lobes; posterior lobes separated by shallow U-shaped median cleft, terminal margins armed with often long, curved cor- neous spines. Males lacking first gonopods, with unpaired, uniramous second left gonopod. Female with vestigial right second pleopod. COLOUR (Figure 1b) — Shield and cephalic appendages, including ocular peduncles, and chelipeds pinkish-white. Ambu- latory legs slightly darker light pink. HABITAT — Gastropod shells. REMARKS — See Paragiopagurus acutus. Paragiopagurus hirsutus (de Saint Laurent, 1972) Figures 1C, 7 Parapagurus acutus hirsutus de Saint laurent, 1972: 113, fig. 19 (type locality: tosa bay, Japan). — miyake, 1982: 196 (list). Sympagurus acutus hirsutus – lemaitre, 1989: 37; 1994: 412. Paragiopagurus hirsutus – lemaitre, 1996: 217, fig. 28; 2000: 221, fig. 71. — rahayu, 2000: 397. — Forest & mclay, 2001: 702, fig. 11. — Davie, 2002: 90. — asakura et al., 2006: 222, fig. 5. — mclaughlin et al., 2007: 327, 2 unnumbered figs; 2010: 39. — Wang, 2008: 759. — Webber et al., 2010: 226. TYPE MATERIAL — Holotype, Japan, Tosa Bay, 11.1963 (day unknown), male, coll. K. Sakai (MNHN-Pg 2225). [See Le- maitre (1996: 166) for comments on original material used in the description of this species by de Saint Laurent (1972)]. MATERIAL EXAMINED — the type material (see above). Taiwan. taiWaN 2000: stn CP 58, 24°35.1’N, 122°05.8’e, 221-254 m, 04.08.2000, 1 female 3.8 mm (mNHN-iU-2012-1013). – taiWaN 2003: stn CP 212, 24°34.60’N, 122°5.84’e, 223-260 m, 26.08.2003, 3 males 4.7-5.4 mm, 6 males 3.4-4.9 mm, 1 female 4.6 mm, 1 male 5.3 mm (mNHN-iU-2012-1014). – taiWaN 2004: stn CP 270, 24°32.7’N, 122°2.26’e, 340-407 m, 02.090.2004, 37 males 2.7- 6.6 mm, 33 females 2.8-5.8 mm, 4 males 4.3-5.5 mm, 3 females 4.9-5.4 mm (mNHN-iU-2012-1015). Philippines. mUSOrStOm 1: 1976, stn CP 42, 13°55’S, 120°29’e, 379-407 m, 24.03.1976, 1 male 5.5 mm (mNHN-iU-2008-160931). Solomon Islands. SalOmON 1: stn DW 1825, 9°50.5’S, 160°58.0’e, 340-391 m, 04.10. 2000, 1 female 4.8 mm (mNHN-Pg 6883). – SalOmON 2: stn DW 2229, 7°33.16’S, 157°42.35’e, 223-275 m, 10.26.2004, 1 female 3.5  mm mNHN-iU-2012-1016. Fiji. bOrDaU 1: stn CP 1448, 16°45.04’S, 179°58.97’e, 410-500 m, 04.03.1999, 1 male 5.1 mm (mNHN- iU-2008-15717). – stn DW 1453, 16°45.03’S, 179°59.30’e, 414-510 m, 04.03.1999, 1 female 3.4 mm (mNHN-iU-2008-15718). Vanuatu. mUSOrStOm 8: stn CP 973, 19°21.30’S, 169°27.03’e, 460-480 m, 22.09.1994, 1 male 5.6 mm (mNHN-Pg 6859). – stn DW 977, 19°24.89’S, 169°28.61’ e, 410-505 m, 22.09.1994, 1 female 3.6 mm (mNHN-Pg 6860). – stn CP 1119, 15°08.32’S, 166°53.35’e, 254-300 m, 9.10.1994, 2 males 2.5, 2.7 mm, 1 ovigerous fe- male 2.5 mm (USNm 1184070). – SaNtO: stn at 101, 15°04.5’S, 166°52.4’e, 356-375 m, 15.10.2006, 2 males 5.9, 6.1 mm,1 female 2.6 mm mNHN- iU-2012-1017. Tonga. bOrDaU 2: stn DW 1593, 19°06’S, 174°18’W, 436-442 m, 14.06.2000, 1 ovigerous female 4.4 mm (mNHN-iU-2008-15734). New Caledonia. N.O. “Vauban”: stn Dr 15, 22°49’S 167°12’e, 390-395 m, 10.04.1978, 2 males 3.4, 3.5 mm mNHN-iU-2012-1018. – stn Dr 16, 22°46’S ParagioPagurus: wordldwide review and new species 331 167°1’e, 390-400 m, 10.04.1978, 1 male 2.9 mm mNHN-iU-2012-1019. – biOCal: stn DW 44, 22°47.30’S, 167°14.30’e, 440 m, 30.08.1985, 1 male 5.5 mm mNHN-iU-2012-1020, 1 male 4.3 mm (USNm). – stn CP 45, 22°47.34’S, 167°14.80’e, 430 m, 30.08.1985, 2 males 4.3, 6.6 mm, 1 ovigerous female 5.2 mm, 2 females 4.2, 4.5 mm mNHN-iU-2012-1021. – stn DW 77, 22°15.32’S, 167°15.40’e, 440 m, 04.09.1985, 1 female 2.0 mm mNHN- iU-2012-1022. – mUSOrStOm 4: stn DW 156, 18°54.0’S, 163°18.80’e, 530 m, 15.09.1985, 2 females 3.9, 5.8 mm (USNm). – stn CP 170, 18°57.00’S, 163°12.60’e, 485 m, 17.09.1985, 7 males 3.0-6.9 mm, 1 ovigerous female 4.9 mm, 2 females 3.1, 3.4 mm (mNHN-iU-2012-1023). – stn CC 177, FIGURE 7 Paragiopagurus hirsutus (de Saint laurent, 1972). A-C, male (2.7 mm) mNHN, Vanuatu, mUSOrStOm 8 stn CP 1119, 254-300 m. D-F, male (6.1 mm) Qm, off Queensland, australia. A, anterior portion of shield and cephalic appendages. B, right cheliped. C, right chela of same. D, right cheliped. E, chela of same. F, telson. all in dorsal view except C, ventral view, and e, mesial view. Scale bars 1 mm for a-C; 2 mm for D-F. (adapted in part from lemaitre 1996). A B, C D-F BAF E CD rafael lemaitre332 18°54.60’S, 163°10.00’e, 540-600 m, 18.09.1985, 1 female 4.5 mm (USNm). – stn CP 179, 18°56.60’S, 163°13.70’e, 480 m, 18.09.1985, 6 males 2.7- 6.3 mm, 4 ovigerous females 2.1-3.0 mm, 2 females 2.8, 5.2 mm mNHN-iU-2012-1024. – stn DW 196, 18°55.00’S, 163°23.70’e, 460 m, 20.09.1985, 1 male 3.3 mm (USNm), 2 males 3.0, 3.1 mm (USNm). – stn DW 197, 18°51.30’S, 163°21.00’e, 560 m, 20.09.1985, 1 male 3.6 mm (USNm). – stn CC 201, 18°55.80’S, 163°13.80’e, 500 m, 20.09.1985, 1 male 3.9 mm, 1 ovigerous female (damaged), 1 female 4.5 mm, 9 males 2.8-7.5 mm, 5 ovigerous females 4.8-6.3 mm, 2 females 3.3, 5.4 mm mNHN-iU-2012-1025, 1 male 6.0 mm (USNm). – stn DW 222, 22°57.60’S, 167°33.00’e, 440 m, 30.09.1985, 3 males 1.8-3.6 mm mNHN-iU-2012-1026. – stn DW 223, 22°57.00’S, 167°30.00’e, 545-560 m, 30.09.1985, 1 male 5.5 mm, 3 females 4.3-4.6 mm, 3 ovigerous females 5.3-6.7 mm, 1 ovigerous female 4.5 mm mNHN-iU-2012-1027. – stn DW 228, 22°47.00’S, 167°18.20’e, 420 m, 30.09.1985, 1 fe- male 3.9 mm mNHN-iU-2012-1028. – stn DW 229, 22°51.60’S, 167°13.50’e, 460 m, 30.09.1985, 1 male 4.6 mm, 7 males 3.3-5.2 mm, 1 female 3.7 mm mNHN-iU-2012-1029. – stn DW 230, 22°52.50’S, 167°11.80’e, 390-420 m, 30.09.1985, 1 male 3.9 mm, 2 ovigerous females 3.6, 4.3 mm (USNm), 2  males 2.2, 3.3 mm, 2 females 3.7, 3.8 mm, 1 female 3.6 mm mNHN-iU-2012-1030. – Smib 2: stn DW 1, 22°52.70’S, 167°12.60’e, 438-444 m, 17.09.1986, 1 male 4.5 mm, 1 female 4.2 mm mNHN-iU-2012-1031. – stn DW 3, 22°56.00’S, 167°14.80’e, 412-428 m, 17.09.1986, 1 male 3.1 mm, 1 female 3.6 mm mNHN-iU-2012-1032. – stn DW 19, 22°52.40’S, 167°11.90’e, 435 m, 20.09.1986, 1 ovigerous female 3.7 mm mNHN-iU-2012-1033. – mUSOrStOm 5: stn DW 388, 20°45.35’S, 160°53’e, 500-510 m, 22.10.1986, 1 female 2.9 mm mNHN-iU-2012-1034. – CHalCal 2: stn CC 1, 24°54.96’S, 168°21.9’e, 500-580 m, 28.10.1986, 6 males 3.7-5.2  mm, 1 ovigerous female 3.9 mm mNHN-iU-2012-1035. – stn CC 2, 24°55.48’S, 168°21.29’e, 500-610 m, 28.10.1986, 3 males 3.9-4.5 mm, 2 ovigerous females 3.8, 4.1 mm, 1 female 2.4 mm mNHN-iU-2012-1036. – stn DW 72, 24°54.50’S, 168°22.30’e, 527 m, 28.10.1986, 2 males 4.2, 5.1 mm, 1 ovigerous female 3.7 mm mNHN-iU-2012-1037. – Smib 3: stn DW 1, 24°55.70’S, 168°21.80’e, 520 m, 20.05.1987, 2 males 3.7, 3.8 mm, 1 ovigerous female 3.6 mm mNHN-iU-2012-1038. – stn DW 2, 24°53.40’S, 168°21.70’e, 530 m, 20.05.1987, 1 female 5.2 mm mNHN-iU-2012-1039. – stn DW 3, 24°55.00’S, 168°21.70’e, 513 m, 20.05.1987, 2 males 4.0, 4.1 mm, 1 ovigerous female 4.2 mm mNHN-iU-2012-1040. – stn DW 5, 24°54.90’S, 168°21.60’e, 502-512 m, 21.05.1987, 3 males 3.1-4.9 mm, 3 ovigerous females 3.3-3.4 mm mNHN-iU-2012-1041, 1 male 3.5 mm (USNm). – stn DW 6, 24°56.40’S, 168°21.20’e, 505 m, 21.05.1987, 1 ovigerous female 3.7 mm mNHN- iU-2012-1042. – stn DW 7, 24°54.60’S, 168°21.30’e, 505 m, 21.05.1987, 4 ovigerous females 3.0-4.9 mm mNHN-iU-2012-1043. – stn DW 21, 22°59.20’S, 167°19.00’e, 525 m, 24.05.1987, 1 female 4.3 mm (USNm). – stn DW 22, 23°03.00’S, 167°19.10’e, 503 m, 24.05.1987, 2 males 2.7, 5.8 mm, 1 female 3.0  mm mNHN-iU-2012-1044. – stn DW 25, 22°56.10’S, 167°16.20’e, 437 m, 24.05.1987, 3 males 3.5-4.0 mm (USNm). – stn DW 26, 22°55.10’S, 167°15.50’e, 450 m, 24.05.1987, right cheliped only mNHN-iU-2012-1045. – stn DW 28, 22°47.00’S, 167°11.80’e, 394 m, 25.05.1987, 2 males 3.9, 4.0 mm (USNm). – stn DW 29, 22°46.70’S, 167°11.70’e, 405 m, 25.05.1987, 5 males 3.7-4.5 mm, 1 ovigerous female 3.6 mm, 1 female 3.1 mm mNHN- iU-2012-1046. – mUSOrStOm 6: stn DW 428, 20°23.54’ S, 166°12.57’e, 420 m, 17.02.1989, 1 male 2.5 mm mNHN-iU-2012-1047. – stn CP 467, 21°05.13’S, 167°32.11’e, 575 m, 21.02.1989, 1 male 2.7 mm (USNm). – stn DW 487, 21°23.30’S, 167°46.40’e, 500 m, 23.02.1989, 1 female 3.0 mm mNHN-iU-2012-1048. – Smib 4: stn DW 34, 24°55.00’S, 168°22.00’e, 515 m, 07.03.1989, 1 male 4.2 mm, 1 female 4.1 mm mNHN-iU-2012-1049. – stn DW 36, 24°55.60’S, 168°21.70’e, 530 m, 07.03.1989, 1 male 4.6 mm mNHN-iU-2012-1050. – stn DW 38, 24°54.5’S, 168°22.0’e, 510 m, 07.03.1989, 1 male 3.9 mm (USNm). – stn DW 55, 23°21.40’S, 168°04 e, 260 m, 09.03.1989, 1 male 4.3 mm mNHN-iU-2012-1051. – stn DW 58, 22°59.60’S, 167°24.20’e, 560 m, 09.03.1989, 1 male 3.5 mm, 1 male 5.5 mm mNHN-iU-2012-1052. – stn DW 63, 22°58.70’S, 167°21.10’e, 520 m, 10.03.1989, 2 males 4.0, 4.5 mm mNHN-iU-2012-1053. – stn DW 65, 22°55.30’S, 167°14.5’e, 400-420 m, 10.03.1989, 1 male 5.1 mm mNHN-iU-2012-1054, 1 male 3.9 mm, 1 female 3.0 mm (USNm). – stn DW 67, 22°55.10’S, 167°15.60’e, 450-460 m, 10.03.1989, 1 male 3.7 mm (USNm). – stn DW 68, 22°55.00’S, 167°16.00’e, 440 m, 10.03.1989, 1 male 5.5 mm (USNm). – Smib 8: stn DW 147, 24°54.90’S, 168°21.85’e, 508-532 m, 27.01.1993, 3 males 3.1-4.5 mm, 1 ovigerous female 3.4, 1 female 3.2 mm (mNHN-Pg 6861). – stn DW 149, 24°54.94’S, 168°21.82’e, 508-510 m, 27.01.1993, 1 female 5.0 mm, 2 ovi- gerous females 3.2, 3.7 mm (mNHN-Pg 6862). – stn DW 150, 24°54.30’S, 168°22.23’e, 519-530 m, 27.01.1993, 3 males 2.2-4.0 mm (mNHN-Pg 6863), 2 males 3.7, 3.5 mm (mNHN-Pg 6864). – stn DW 152, 24°54.35’S, 168°22’e, 514-530 m, 27.01.1993, 1 male 2.9 mm, 2 females 3.5, 4.1 mm (mNHN-Pg 6865), 1 males 3.9 mm (mNHN-Pg 6866). – stn DW 189, 23°17.60’S, 168°05.51’e, 400-402 m, 31.01.1993, 1  female 2.1 mm (mNHN-Pg 6867). – stn DW 197, 22°51.27’S, 168°12.54’e, 414-436 m, 01.02.1993, 2 males 3.5, 4.2 mm, 1 female 2.9 mm, 8 spms unsexed in housing (mNHN-Pg 6869). – stn DW 198, 22°51.59’S, 167°12.44’e, 414-430 m, 01.02.1993, 4 males 3.5-4.2 mm, 1 female 3.0 mm, 11 spms in housing (mNHN-Pg 6870). – stn DW 199, 22°51.63’S, 168°12.22’e, 408-410 m, 01.02.1993, 14 spms in housing (mNHN-Pg 6871). – batHUS 2: stn DW 718, 22°46’S, 167°14’e, 430-436 m, 11.05.1993, 1 male 2.3 mm, 2 ovigerous females 3.1, 3.9 mm, 2 females 2.8, 3.2 mm (mNHN-Pg 6842). – stn DW  719, 22°47.57’S, 167°14.58’e, 444-455 m, 11.05.1993, 18 spms in housing (mNHN-Pg 6843). – stn DW 720, 22°51.62’S, 167°16.40’e, 530-541 m, 11.05.1993, 4 males 3.4-5.6 mm, 1 female 3.5 mm, 13 spms in housing (ZmmU, ex mNHN-Pg 6844). – stn DW 721, 22°53.82’S, 167°17.42’e, 525-547 m, 11.05.1993, 2 males 3.4, 4.7 mm, 1 female 4.0 mm (mNHN-Pg 6845). – stn DW 723, 22°50.21’S, 167°26.84’e, 430-433 m, 11.05.1993, 2 females 2.2, 3.7 mm (mNHN-Pg 6846). – stn DW 729, 22°52.42’S, 167°11.90’e, 349-400 m, 12.05.1993, 31 spms in housing (mNHN-Pg 6847). – stn CP 736, 23°03.38’S, 166°58.96’e, 356-464 m, 13.05.1993, 11 spms in housing (mNHN-Pg 6848). – stn CP 738, 23°02.09’S, 166°56.61’e, 358-647 m, 13.05.1993, 1 male 4.0 mm, 1 ovigerous female 4.4 mm, 1 female 3.7 mm (mNHN-Pg 6849). – batHUS 3: stn DW 817, 23°42,38’S, 168°15,51’e, 405-410 m, 28.11.1993, 1 male 4.0 mm, 1 ovigerous female 3.6 mm (mNHN-Pg 6850). – stn DW 819, 23°45.11’S, 168°16.27’e, 478-486 m, 28.11.1993, 1 female 2.2 mm (mNHN-Pg 6851). – stn DW 838, 23°00,81’S, 166°55,87’e, 400-402 m, 30.11.1993, 1 female 2.7 mm (mNHN-Pg 6852). – batHUS 4: stn DW 908, ParagioPagurus: wordldwide review and new species 333 18°58.38’S, 163°10.7’e, 502-527 m, 04.08.1994, 1 male 2.6 mm, 1 female 4.9 mm (parasitized) (mNHN-Pg 6853). – stn CP 909, 18°57.64’S, 163°10.30’e, 516-558 m, 04.08.1994, 2 males 4.9, 5.0 mm (mNHN-Pg 6854). – stn DW 914, 18°48.79’S, 163°15.23’e, 600-616 m, 05.08.1994, 1 male 5.6 mm, 2 ovi- gerous females 3.9, 4.5 mm (mNHN-Pg 6855). – stn DW 929, 18°51.55’S, 163°23.2’e, 502-516 m, 07.08.1994, 2 males 3.6, 5.2 mm, 4 females 2.5- 4.4 mm (mNHN-Pg 6856). – Smib 10: stn DW 203, 24°56’S, 168°22’e, 502-508 m, 10.01.1995, 2 males 2.4, 2.9 mm, 1 female 2.6 mm (mNHN-Pg 6872). – stn DW 204, 24°57’S, 168°21’e, 513-553 m, 10.01.1995, 1 male 3.2 mm (mNHN-Pg 6873). – stn DW 207, 508-553 m, 1 female 3.2 mm (mNHN-Pg 6874). – stn DW 215, 24°56’S, 168°21’e, 508-553 m, 11.01.1995, 1 female 4.4 mm (mNHN-Pg 6824). – litHiSt: stn DW 7, 24°55.36’S, 168°21.58’W, 504-530 m, 11.08.1999, 1 ovigerous female 4.4 mm (mNHN-iU-2008-15738). – stn CP 8, 24°54,24’S, 168°21.35’W, 540 m, 11.08.1999, 2 males 1.9, 3.4 mm (mNHN-iU-2008-15739). – berYX 11: stn CP 21, 24°45.03’S, 168°06.80’e, 430-450 m, 17.10.1992, 5 males 4.3-6.2 mm, 5 females 4.7-6.0 mm (mNHN-Pg 6857), 1 male 4.7 mm, 2 females 5.0, 5.5 mm (mNHN-Pg 6858). – NOrFOlK 1: stn DW 1680, 24°45’481’S, 168°10.060’e, 385-392 m, 22.06.2001, 1 male 2.7 mm (mNHN-iU-2008-15741). – stn DW 1698, 24°40.186’S, 168°39.545’e, 562-576 m, 23.06.2001, 1 female 3.7 mm (mNHN- iU-2008-15732). – stn DW 1732, 23°20.114’S, 168°16.024’e, 347-1063 m, 27.06.2001, 1 male 3.2 mm (mNHN-iU-2008-15744). – NOrFOlK 2: stn DW 2034, 23°40.64’S, 167°41.36’e, 485-505 m, 22.10.2003, 1 male 6.7 mm (mNHN-iU-2008-16095). – stn DW 2087, 24°56.2’S, 168°21.66’e, 518-586 m, 28.10.2003, 2 males 4.4, 4.4 mm, 1 female 2.9 mm (mNHN-iU-2008-16096). – stn DW 2091, 24°45.36’S, 168°06.24’e, 600-896 m, 29.10.2003, 1 male 5.0 mm (mNHN-iU-2008-16097). – stn DW 2137, 23°01.18’S, 168°22.70’e, 547-560 m, 03.11.2003, 1 female 4.7 mm (mNHN-iU-2008-160986). – stn CP 2146, 22°50.17’S, 167°17.35’e, 518 m, 04.11.2003, 2 males 5.4, 5.6 mm (mNHN-iU-2008-16099). – stn DW 2148, 22°44.20’S, 167°15.97’e, 386-391 m, 04.11.2003, 1 female 3.9 mm (mNHN-iU-2008-16100). – stn DW 2155, 22°52.40’S, 167°13.46’e, 453-455 m, 05.11.2003, 1 male 4.7 mm (mNHN- iU-2008-16101). – stn DW 2156, 22°54.19’S, 167°15.13’e, 468-500 m, 05.11.2003, 1 male 3.5 mm (mNHN-iU-2008-16102). – ebiSCO: stn DW 2535, 22°17’S, 159°28’e, 490 m, 10.10.2005, 1 ovigerous female 3.6 mm mNHN-iU-2012-1055. – stn DW 2578, 20°21’S, 158°40’e, 440-505 m, 14.10.2005, 1  female 2.6 mm mNHN-iU-2012-1056. – stn CP 2579, 20°21’S, 158°40’e, 440 m, 14.10.2005, 1 male (damaged), 1 female 3.3 mm mNHN- iU-2012-1057. – stn DW 2607, 19°33’S, 158°40’e, 400-413 m, 18.10.2005, 1 male 3.3 mm mNHN-iU-2012-1058. – stn CP 2614, 19°39’S, 158°47’e, 608-642 m, 19.10.2005, 1 female 3.0 mm mNHN-iU-2012-1059. – stn DW 2617, 20°06’S, 160°22’e, 427-505 m, 20.10.2005, 1 male 5.3 mm mNHN- iU-2012-1060. – stn DW 2619, 20°06’S, 160°2’e, 490-550 m, 20.10.2005, 2 males 2.7, 2.8 mm mNHN-iU-2012-1061. – stn DW 2625, 20°05’S, 160°20’e, 627-741 m, 20.10.2005, 1 male 2.1 mm, 1 female 2.4 mm mNHN-iU-2012-1062. For additional material examined, including specimen illustrated in Figure 6D-F, see lemaitre (1996). DISTRIBUTION — Indo-West Pacific: Taiwan, South China Sea (off Hong Kong), Philippines, Solomon Islands, Fiji,Vanuatu, Tonga, New Caledonia, Australia, New Zealand, and questionably from off Durban, South Africa (see Lemaitre 1996). Depth: 221 to 1063 m. DIAGNOSIS — Gill lamellae undivided distally. Shield (Figure 7A) about as long as broad; dorsal surface weakly calcified medially; rostrum broadly rounded, with low dorsal ridge; lateral projections subtriangular, terminating in small spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, each terminating in strong multifid spine (rarely simple on one side); corneas slightly dilated. Maxillule with internal lobe of endopod bearing 3 or 4 long setae. Epistomial spine straight, frequently bifid. Antennular peduncles exceeding distal margin of corneas by 0.75 length of ultimate segment. Antennal peduncles reaching distal margin of corneas; second segment with dorsolateral distal angle produced, terminating in strong simple spine reaching to midpoint of antennal acicle; acicles reaching distal margin of corneas, mesial margin of each with 8-13 spines; flagellum with long setae 3 or 4 flagellar articles in length. Chelipeds markedly dissimilar, surfaces with moderately dense setae interspersed with numerous long, stiff setae. Right cheliped (Figure 7B-E) with palm broader than long, fingers curved ventromesially, dactyl with longitudinal row of blunt spines on ventral face; dorsal face of palm with strong spines (at least proximomedially), ventral face with numerous tubercles; palm with dorsolateral and dorsomesial margins each well delimited by row of strong spines, and ventromesial face rounded; carpus about as long as chela or longer than chela (Figure 7D) in large males (sl > 6. 0mm), with numerous small tubercles or spines on dorsal surface. Left cheliped with lateral face of carpus usually weakly calcified; with dorsal row of 2-6 small, well-spaced spines, and strong dorsodistal spine on carpus. Ambulatory legs with dactyls having ventromesial row of 8-13 corneous spines, and dorsal and dorsomesial rows of long setae; carpi each with small dorsodistal spine; carpus of second pereopod with dorsal margin armed with 1 to 4 small spines. Anterior lobe of sternite XII (of third pereopods) setose, armed with 1 or 2 marginal spines. Fourth pereopod with propodal rasp consisting of 1 row of ovate scales. Uro- pods and telson (Figure 7F) markedly asymmetrical; telson lacking transverse suture separating anterior and posterior rafael lemaitre334 lobes, left side up to 1.5 times as long as right side; posterior lobes separated by U-shaped median cleft, terminal margins armed with often long, curved corneous spines. Males lacking first pleopods, with unpaired, uniramous second left pleo- pod. Females with vestigial right second pleopod. COLOUR (Figure 1C) — Shield whitish, tinged with light orange or reddish-orange; ocular peduncles and acicles light orange or reddish-orange, with darker orange tint adjacent to corneas; antennule with ultimate segment and dorsal ramus of flagellum yellowish; antennal flagellum yellowish. Chelipeds with chelae whitish-orange, spines darker; dactyl with darker orange tint basally; carpi whitish centrally, with dark orange reddish-orange band distally and proximally, spines also reddish-orange; meri with similar pattern to carpi. Ambulatory legs with dactyls light reddish-orange, darker basally, tips white; propodi, carpi and meri, each with distal and proximal band of light orange or reddish-orange, and median band white. HABITAT — Gastropod shells. REMARKS — See Paragiopagurus acutus. Paragiopagurus acutus (de Saint Laurent, 1972) Figures 1D, 8 Parapagurus acutus acutus de Saint laurent, 1972: 113, figs 7, 18 (type locality: Philippines). — miyake, 1982: 196 (list). Sympagurus acutus acutus – lemaitre, 1989: 37; 1994: 412. Paragiopagurus acutus – lemaitre, 1996: 211, figs 25, 26; 1997: 586. — Spiridonov & Zhadan, 1999: 628. — rahayu, 2000: 396. — Davie, 2002: 89. — asakura et al., 2006: 221. — mclaughlin et al., 2007: 329, unnumbered fig.; 2010: 39, fig. 19b. — Wang, 2008: 759. — tsang et al., 2011: 5, figs 2, 4 (phylogenetic trees). TYPE MATERIAL —Holotype, Philippines, between Marinduque and Luzon, 9.2 mi (17 km) NW of W San Andreas Island, Albatross, stn 5222, 13°38’30”N, 121°42’45”E, 357 m, 24.04.1908, male 5.6 mm (USNM 168309). [See Lemaitre (1996: 166) for comments on original material used in the description of this species by de Saint Laurent (1972)]. MATERIAL EXAMINED — the type material (see above). Taiwan. taiWaN 2001: stn CP 68, 24°49.6’N, 122°00.8’e, 370 m, 06.05.2001, 2 males 5.3- 5.5 mm, 1 female 4.1 mm (mNHN-Pg 6756). – stn CP 73, 24°52.9’N, 122°01.0’e, 220-330 m, 07.05.2001, 7 males 5.4-6.4 mm (mNHN-Pg 6762). – stn CP 74, 24°50.8’N, 121°59.3’e, 220 m, 07.05.2001, 16 males 3.3-5.4 mm, 9 females 3.2-4.3 mm, 12 ovigerous females 3.6-4.8 mm (mNHN-Pg 6763), 1  female 3.5 mm (mNHN-Pg 6757). – stn CP 80, 24°50.9’N, 121°59.4’e, 194-214 m, 08.05.2001, 1 female 4.0 mm (mNHN-Pg 6758). – stn CP 85, 24°00.5’N, 122°00.5’e, 255-390 m, 09.05.2001, 3 males 5.3-5.9 mm, 1 female 4.8 mm (mNHN-Pg 6755), 2 males 6.0, 6.4 mm (mNHN-Pg 6764). – stn  CP  89, 24°53.6’N, 122°01.4’e, 310-420 m, 09.05.2001, 2 males 5.3, 5.5 mm, 2 ovigerous females 3.5, 4.1 mm (mNHN-Pg 6816). – stn CP 95, 24°55.8’N, 122°05.7’e, 269-360 m, 18.05.2001, 1 male 5.0 mm (mNHN-Pg 6759), 2 males 5.3, 6.1, 1 ovigerous female 3.9 mm, 1 female 3.9 mm (mNHN-Pg 6817). – stn CP 109, 24°48.3’N, 122°84.0’e, 246-256 m, 20.05.2001, 3 males 5.5-6.3 mm, 1 ovigerous female 4.5 mm (mNHN-Pg 6818). – stn CP 115, 24°53.9’N, 122°02.0’e, 381-440 m, 21.05.2001, 1 male 4.8 mm, 1 ovigerous female 4.2 mm (mNHN-Pg 6819). – taiWaN 2002: stn CP 162, 22°09.64’N, 120°37.86’e, 190-200 m, 25.05.2002, 1 male 3.8 mm (mNHN-Pg 6820). – Dasi fishing port, Yilan County: commercial trawlers, no date, 1 male 5.9 mm (NtOU). Philippines. mUSOrStOm 1: stn CP 26, 14°01’S, 120°17’e, 189 m, 22.03.1976, 2 males 1.5, 3.1 mm, 1 female 2.5 mm (mNHN- Pg 6723). – stn CP 42, 13°55’S, 120°29’e, 379-407 m, 24.03.1976, 1 male 5.9 mm. – mUSOrStOm 2: stn CP15, 13°55’S, 120°28’e, 326-330 m, 21.11.1980, 1 male 5.8 mm (mNHN-Pg 6409). – stn CP 49, 13°38’S, 121°44’e, 416-425 m, 26.11.1980, 1 male 4.3 mm (mNHN-Pg 6410). Indonesia. COriNDON: stn CH 240, 00°38’S, 119°33’e, 675 m, 05.11.1980, 1 female 5.9 mm (mNHN-iU-2008-16082). Solomon Islands. SalOmON 1: stn CP 1783, 8°32.8’S, 160°41.7’e, 399-700 m, 29.09.2000, 1 female 3.4 mm (mNHN-Pg 6748). – stn CP 1800, 9°21.4’S, 160°23.9’e, 357-359 m, 01.10. 2000, 11 males 1.9- 4.9 mm, 2 ovigerous females 3.1, 3.5 mm, 6 females 1.9-3.1 mm (mNHN-Pg 6749). – stn CP 1801, 9°25.0’S, 160°25.9’e, 264-273 m, 01.10. 2000, 1 ovi- gerous female 4.4 mm (mNHN-Pg 6750). – stn CP 1804, 9°32.0’S, 160°37.4’e, 309-328 m, 02.10. 2000, 14 males 2.2-2.4 mm, 8 ovigerous females ParagioPagurus: wordldwide review and new species 335 2.5-3.3 mm, 6 females 2.2-3.8 mm (mNHN-Pg 6751). – stn CP 1805, 9°35.0’S,160°42.7’e, 367-500 m, 02.10. 2000, 1 male 4.5 mm (mNHN-Pg 6752). – stn CP 1859, 9°32.6’S, 160°37.3’e, 283-305 m, 07.10. 2000, 1 female 5.6 mm (mNHN-Pg 6753). – stn CP 1860, 9°22’S, 160°31’e, 620 m, 07.10. 2000, 12 males 2.2-4.9 mm, 1 ovigerous female 4.5 mm (mNHN-Pg 6754). – SalOmON 2: stn CP 2210, 7°33.5’S, 157°42.3’e, 240-305 m, 10.26.2004, 30 males 1.6-5.0 mm, 6 ovigerous females 2.4-3.8 mm, 5 females 2.5-4.4 mm mNHN-iU-2012-1063. – stn CP 2260, 8°04.45’S, 156°55.87’e, 399-427 m, 11.03.2004, 1 male 4.7 mm mNHN-iU-2012-1064. Fiji. mUSOrStOm 10: stn CP 1341, 16°52.51’S, 177°43.66’e, 500-614 m, 10.08.1998, 1 male 6.1 mm (mNHN-iU-2008-15712). – stn DW 1345, 17°14.92’S, 178°29.50’e, 660-663 m, 11.08.1998, 1 male 5.5 mm (mNHN-iU-2008-15733). – stn CP 1368, 18°10.92’S, 178°23.47’e, 380-469 m, 15.08.1998, 2 males 6.2, 6.4 mm (mNHN-iU-2008-15707). – stn DW 1376, 18°18.67’S, 178°09.07’e, 497-504 m, 17.08.98, 3 males 2.4-5.1 mm, 5 ovigerous females 2.7-2.8 mm mNHN-iU-2012-1065. – bOrDaU 1: stn CP 1396, 16°38.98’S, 179°57.16’W, 591-596 m, 24.02.1999, 1 male 4.1 mm, 3 females 4.9-5.1 mm (mNHN-iU-2008-15720). – stn CP 1409, 16°01.88’S, 179°29.83’W, 557-558 m, 26.02.1999, 1 female 4.7 mm (mNHN-iU-2008-15723). – stn CP 1420, 550-687 m 1 female 3.8 mm (mNHN-iU-2008-15719). – stn DW 1424, 17°16.61’S, 179°01.25’W, 385- 416 m, 01.03.1999, 1 female 4.0 mm (mNHN-iU-2008-15729). – stn CP 1427, 17°16.26’S, 179°01.06’W, 364-369 m, 01.03.1999, 1 female 2.2 mm (mNHN-iU-2008-15722). – stn DW 1453, 16°45.03’S, 179°59.30’e, 414-510 m, 04.03.1999, 1 male 2.6 mm (mNHN-Pg 6742). – stn DW 1499, 18°39.82’S, 178°26.78’W, 389-400 m, 12.03.1999, 1 male 3.9 mm, 2 females 2.1, 2.4 mm (mNHN-iU-2008-15715). – stn CP 1500, 18°41.74’S, 178°26.20’W, 366- 389 m, 12.03.1999, 1 female 1.6 mm (mNHN-iU-2008-15716). Vanuatu. mUSOrStOm 8: stn DW 1014, 17°54.53’S, 168°19.08’e, 495-498 m, 27.09.1994, 1 male 4.0 mm (USNm 1184061). – stn CP 1054, 16°27.95’S, 167°57.44’e, 522-527 m, 1.10.1994, 1 male 5.6 mm (mNHN-Pg 6724). – stn DW 1060, 16°13.82’S, 167°20.80’e, 375-397 m, 2.10.1994, 2 males 1.8, 1.9 mm, 1 ovigerous female 3.8 mm, 1 female 1.9 mm (mNHN-Pg 6725). – stn DW 1065, 16°16.12’S, 167°21.43’e, 360-419 m, 02.10.1994, 3 males 2.3-2.7 mm, 4 ovigerous females 2.2-2.5 mm, 2 females 2.2, 2.6 mm (USNm 1184062). – stn CP 1087, 15°10.18’S, 167°14,07’e, 394-421m, 6.10.1994, 1 male 3.8 mm (mNHN-Pg 6726). – stn CP 1088, 15°09.23’S, 167°15.13’e, 425-455 m, 06.10.1994, 1 male 2.7 mm (USNm 1184063). – stn CP 1091, 15°10.24’S, 167°13.01’e, 344-350 m, 6.10.1994, 3 males 2.0, 2.9 mm, 1 ovigerous female 3.1 mm, 2 females 2.3, 2.8 mm (mNHN-Pg 6727). – stn CP 1123, 15°07.19’S, 166°55.20’e, 262-352 m, 9.10.1994, 4 males 2.7-4.3 mm (mNHN-Pg 6728). – stn CP 1124, 15°01.72’S, 166°56.51’e, 532-599 m, 9.10.1994, 1 male 7.2 mm (mNHN-iU-2008-16084). – stn CP 1136, 15°40.62’S, 167°01.60’e, 398-400 m, 11.10.1994, 2 males 2.0, 6.2 mm, 3 ovigerous females 3.1-3.5 mm, 1 damaged female (mNHN-Pg 6729), 4 males 2.9-3.9 mm, 1 ovigerous female 2.6 mm (mNHN-Pg 6730). – stn CP 1137, 15°41.52’S, 167°02.67’e, 360-371 m, 11.10.1994, 4 males 2.7-4.4 mm (mNHN-Pg 6731). – bOa 0: stn CP 2314, 15°03’S, 166°54’e, 430-455 m, 16.11.2004, 1 male 5.0 mm mNHN-iU-2012-1066. – stn CP 2328, 15°43’S, 167°02’e, 314-547 m, 18.11.2004, 2 males 4.4, 5.6 mm mNHN-iU-2012-1067. – stn CP 2329, 15°43’e, 167°04’e, 514-609 m, 18.11.2004, 2 males 6.6, 6.8 mm mNHN-iU-2012-1068. – stn CP 2330, 15°44’S, 167°02’e, 295-890 m, 18.11.2004, 1 female 2.6 mm mNHN-iU-2012-1069. – bOa 1: no station data, 2 males 2.9, 5.1 mm mNHN-iU-2012-1070. – stn CP 2413, 15°39.337’ S, 167°01.475’ e, 268-445 m, 05.09.2005, 3 males 1.7-2.5 mm, 3 ovigerous females 3.4, 3.5 mm mNHN-iU-2012-1071. – stn CP 2414, 15°41.28’S, 167°02.897’e, 309-402 m, 05.09.2005, 2 males 2.4, 4.1 mm, 1 ovigerous female 3.8 mm mNHN-iU-2012-1072. – stn CP 2416, 15°04.35’S, 166°51.78’e, 350-400 m, 06.09.2005, 1 male 4.7 mm, 1 female 5.5 mm, 1 ovigerous female 3.8 mm mNHN-iU-2012-1073. – stn CP 2428, 15°05.73’S, 166°52.65’e, 323-397 m, 08.09.2005, 1 female 4.3 mm mNHN-iU-2012-1074. – stn CP 2448, 15°08.031’S, 166°50.853’e, 297-387 m, 10.09.2005, 5 males 1.7-6.0 mm, 1 ovigerous female, 4.3 mm mNHN-iU-2012-1075. – stn CP 2461, 16°35.11’S, 167°59.68’e, 582-614 m, 13.09.2005, 1  male 5.3 mm mNHN-iU-2012-1076. – stn CP 2471, 16°25.05’S, 167°52.44’e, 591-627 m, 14.09.2005, 1 male 6.9 mm mNHN-iU-2012-1077. – stn CP 2479, 16°43.25’S, 167°50.86’e, 350-358 m, 15.09.2005, 1 male 3.9 mm mNHN-iU-2012-1078. – SaNtO: stn at 01, 15°32.4’S, 167°16.4’e, 167-367 m, 14.09.2006, 1 male 4.1 mm mNHN-iU-2012-1079. – stn at 08, 15°40.5’S, 167°01.5’e, 366-389 m, 17.09.2006, 1 male 1.9 mm, 1 ovigerous female 2.5 mm, 6 males 2.7, 2.9 mm, 1 female 2.2 mm mNHN-iU-2012-1080. – stn at 09, 15°41.5’S, 167°01.3’e, 481 m, 17.09.2006, 1 male 2.8 mm mNHN- iU-2012-1081. – stn at 58, 15°33.0’S, 167°19.3’e, 364-390 m, 03.10.2006, 1 male 5.3 mm, 1 ovigerous female 5.1 mm mNHN-iU-2012-1082. – stn at 73, 15°40.8’S, 167°00.5’e, 514-636 m, 07.10.2006, 3 males 2.4-7.9 mm mNHN-iU-2012-1083. – stn at 105, 15°04.2’S, 166°57.0’e, 408-444 m, 15.10.2006, 1 male 5.6, 1 female 4.5 mm mNHN-iU-2012-1084. – stn at 121, 15°38.7’S, 167°01.2’e, 275-290 m,19.10.2006, 3 males 2.8-2.7 mm mNHN- iU-2012-1085. tonga. bOrDaU 2: stn DW 1508, 21°02’S, 175°19’W, 555-581 m, 31.05.2000, 1 male 3.7 mm (mNHN-Pg 6745), 1 male 3.7 mm (mNHN- Pg 6746). – stn CP 1510, 21°05’S, 175°23’W, 461-497 m, 31.05.2000, 4 males 2.1-2.8 mm, 2 ovigerous females 2.4, 2.8 mm (mNHN-Pg 6743). – stn DW 1548, 20°38’S, 175°03’W, 476-478 m, 06.06.2000, 1 female 3.1 mm (parasitized) mNHN-iU-2012-1086. – stn CP 1556, 20°11’S, 174°45’W, 589-591 m, 07.06.2000, 2 males 3.4, 4.1 mm (mNHN-iU-2008-15711). – stn CH 1563, 19°52’S, 174°39’W 362-388 m, 08.06.2000, 1 male 5.6 mm (mNHN-iU-2008-15725). – stn CH 1564, 19°52’S, 174°39’W, 371-387 m, 08.06.2000, 2 males 5.1, 5.5 mm (mNHN-iU-2008-15708), 4 males 5.9-6.3 mm (mNHN-iU-2008-15710). – stn CP 1592, 19°08’S, 174°17’W, 391-426 m, 14.06.2000, 1 female 2.7 mm (mNHN-Pg 6747). New Caledonia. Smib 2: stn DW 23, 22°31.10’S, 167°36.50’e, 410-420 m, 20.09.1986, 1 male 4.6 mm,1 female 4.3 mm (USNm). – mUSOrStOm 6: stn DW 487, 21°23.30’S, 167°46.40’e, 500 m, 23.02.1989, 1 male 2.5 mm (USNm). – batHUS 1: stn CP 670, 20°54.05’S, 165°53.38’e, 394 m, 14.03.1993, 2 ovigerous females 3.0, 3.4 mm (mNHN-Pg 6709). – stn CP 683, 20°34.93’S, 165°07.24’e, 380-400 m, 16.01.1993, 1 male 4.0 mm (mNHN-Pg 6710). – stn CP 695, 20°34.59’S, 164°57.88’W, 410-430 m, 17.03.1993, 3 males 2.2-3.5 mm, 1 ovigerous female 3.7 mm, 2 females 3.1, 3.3 mm (mNHN-Pg 6711). – batHUS 4: stn CP 910, 18°59.32’S, 163°08.47’e, 560-608 m, 05.08.1994, 1 male 5.5 mm (mNHN-Pg 6713). – stn CP 911, 18°57.80’S, 163°08.47’e, 558-566 m, 05.08.1994, 1 male 5.6 mm (mNHN-Pg 6714). – stn CP 946, 20°33.81’S, 164°58.35’e, 386-430 m, 10.08.1994, 2 males 2.7, 3.0 mm (mNHN-Pg 6716), 2 ovigerous rafael lemaitre336 females 3.1, 5.0 mm (mNHN-Pg 6715). – stn CP 949, 20°32.01’S, 164°56.85’e, 616-690 m, 10.08.1994, 2 males 2.3, 2.9 mm, 1 ovigerous female 3.0 mm (mNHN-Pg 6712). – HaliPrO 1: stn CP 854, 21°40.90’S, 166°38.34’e, 650-780 m, 19.03.1994, 1 male 5.2 mm (mNHN-Pg 6717). – stn CP 866, 21°26.91’S, 166°17.23’e, 550-600 m, 22.03.1994, 1 male 2.9 mm (mNHN-Pg 6718), 1 male 6.2 mm (mNHN-Pg 6719), 1 male 6.0 mm (mNHN-Pg 6720). – stn CP 869, 21°14.84’S, 165°55.49’e, 450-490 m, 23.03.1994, 1 male 3.0 mm (mNHN-Pg 6721). – stn CH 870, 21°15.06’S, 165°55.43’e, 450-500 m, 23.03.1994, 1 male 5.6 mm (mNHN-Pg 6722). – NOrFOlK 2: stn CP 2139, 23°00.68’S, 168°22.60’e, 372-393 m, 03.11.2003, 1 male 2.8 mm (mNHN- iU-2008-16085). – ebiSCO: stn DW 2602, 19°37.662’S, 158°43.625’e, 547 m, 10.18.2005, 1 female 4.6 mm mNHN-iU-2012-1087. – stn DW 2607, 19°34.300’S, 158°40.489’e, 400-413 m, 10.18.2005, 1 female 2.7 mm mNHN-iU-2012-1088. – stn DW 2610, 19°34’S, 158°41’e, 486-494 m, 19.10.2005, 3 males 3.1-4.9 mm, 1 female 4.6 mm mNHN-iU-2012-1089. For additional material examined see lemaitre (1996). DISTRIBUTION – Western Pacific: Japan, Taiwan, China Sea (off Hong Kong), Philippines, Indonesia, Solomon Islands, Fiji, Vanuatu, Tonga, New Caledonia, and Western Australia. Depth: 161 to 780 m. FIGURE 8 Paragiopagurus acutus (de Saint laurent, 1972). A, B, male (5.1 mm) mNHN, Fiji, mUSOrStOm 10 stn DW1376, 497-504 m. C-E, holotype male (5.6 mm) USNm 168309, Philippines, albatross stn 5222, 357 m. A, anterior portion of shield and cephalic appendages. B, right cheliped. C, right cheliped. D, chela of same. E, telson. all in dorsal view except D, mesial view. Scale bars 1 mm for a, e; 2 mm for b-D. (adapted in part from lemaitre 1996). E C, D B A E A BC D ParagioPagurus: wordldwide review and new species 337 DIAGNOSIS – Gills with lamellae undivided distally. Shield (Figure 8A) as long as broad; dorsal surface weakly calcified medially; rostrum broadly rounded, with low dorsal ridge; lateral projections subtriangular, terminating in small spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, terminating in strong multifid spine; cor- neas slightly dilated. Maxillule with internal lobe of endopod bearing 4 long seta. Epistomial spine straight. Antennular peduncles exceeding distal margin of corneas by 0.75 length of ultimate segment. Antennal peduncles reaching distal margin of cornea; second segment with dorsolateral distal angle produced, terminating in strong simple spine reaching to midpoint of antennal acicle; acicles reaching distal margin of corneas, mesial margin with 9 to 14 spines; flagellum with short and long setae < 1 to 4 flagellar articles in length respectively. Chelipeds markedly dissimilar, with moderately dense setae. Right cheliped (Figure 8B-D) with chela proportions varying from broader than long to longer than broad (up to 1.8 times as long a broad) in larger specimens (sl ≥ 6.0 mm); fingers curved ventromesially, dactyl with strongly concave and smooth ventromesial face; dorsal face of palm with scattered small spines or tubercles, ventral face smooth or with small tubercles; palm with dorsolateral and dorsomesial margins each well delimited by row of spines, ventromesial mar- gin rounded, occasionally with row of 2 or 3 blunt spines proximally; carpus with numerous small spines or tubercles on dorsal surface, and well delimited dorsolateral margin with row of spines. Left cheliped usually weakly calcified on lateral face of carpus; carpus with dorsal row of 4 to 6 small, well-spaced spines, and strong dorsodistal spine. Ambulatory legs with dactyls having ventromesial row of 7-15 small, often minute corneous spines, and dorsal and dorsomesial rows of long setae; carpi each with dorsodistal spine; carpus of second pereopod with dorsal row of 7 small spines. Anterior lobe of sternite XII (of third pereopods) setose, armed with 1 spine. Fourth pereopod with propodal rasp consisting of 1 row of ovate scales. Uropods and telson (Figure 8E) markedly asymmetrical, with asymmetry more pronounced in larger specimens and sexually dimorphic; telson lacking transverse suture separating anterior and posterior lobes; posterior lobes separated by shallow U-shaped median cleft, terminal margins armed with often long, curved corneous spines; large males (sl ≥ 8.0 mm) with left side of telson as much as twice as long as right side; in large females (sl ≥ 6.0 mm) pos- terior lobes armed with several rows of corneous spines on distal margin, rows of spines often extending to dorsodistal surface. Males lacking first gonopods, with unpaired, uniramous second left gonopod. Females with vestigial right second pleopod. COLOUR (Figure 1D) — Shield cream or light orange with darker regions near frontal margin. Cephalic appendages cream to light orange; ocular acicles and second segments of antennal peduncles sometimes pinkish. Chelipeds generally white or cream; carpus of right and left chelipeds with orangish region on dorsal surfaces proximally often extending distally on carpus of right along all of dorsal margin; carpus of left cheliped often light pink. Ambulatory legs generally white or pinkish; dorsal margins of meri and carpi, and distolateral regions of meri orangish; proppodi and dactyls white or pin- kish, with proximal surfaces light orange. HABITAT — Gastropod shells. REMARKS — As pointed out by Lemaitre (1996), Paragiopagurus acutus is most similar to P. bicarinatus and P. hirsutus, all three of which have multifid ocular acicles. Paragiopagurus acutus can be separated from P. bicarinatus by the shape and armature of the mesial face of the right palm. The mesial face of P. acutus is not expanded distally, and the ventrome- sial margin is rounded; the mesial face of P. bicarinatus is expanded distally, and the ventromesial margin is well delimited by a row of spines. Paragiopagurus acutus differs from P. hirsutus by the armature and setation of the right chela. Addi- tionally, P. acutus reaches a much larger size than P. bicarinatus or P. hirsutus, and exhibits a stronger degree of sexual dimorphism in the telson. As mentioned under P. trilineatus n. sp., that species also has multifid ocular acicles. However, P. trilineatus n. sp. is distinguished from P. acutus and others with multifid ocular acicles, primarily by the number of rows of scales on the rasp of the fourth pereopod; there is one row in P. acutus, P. bicarinatus and P. hirsutus, and two or three rows in P. trilineatus n. sp. rafael lemaitre338 Paragiopagurus ruticheles (A. Milne-Edwards, 1891) Figures 9, 10 Eupagurus ruticheles a. milne-edwards, 1891: 133 (type locality: near Graciosa, azores). Parapagurus ruticheles – de Saint laurent, 1972: 112. Sympagurus ruticheles – lemaitre, 1989: 37; 1990, 235, figs 11, 12; 1994: 412. — ingle, 1993: 36. — Parin et al., 1997: 163. Paragiopagurus ruticheles – lemaitre, 1996: 207. — Zhadan, 1997: 69. — d’Udekem d’acoz, 1999: 176. — Davie, 2002: 89. — mclaughlin et al., 2005: 246; 2010: 39. — Castro, 2011: 30. TYPE MATERIAL — Syntypes, Azores, near Graciosa, Atlantic Ocean, “L'Hirondelle ”, stn 234, 39°01’40”N, 30°15’40”W, 454 m, 19.08.1888, 2 males 3.6, 5.9 mm, 1 ovigerous female 3.2 mm (MO). MATERIAL EXAMINED — the type material (see above). Madagascar. N.O. “Vauban”: stn P1-Ch. 2, 12°53.3’S, 48°09.4’e, 480-520 m, 04.03.1971, 7  males 3.5-4.2 mm (mNHN-Pg 5788). Taiwan. taiWaN 2004: stn CP 270, 24°32.7’N, 122°2.26’e, 340-407 m, 02.09.2004, 13 males 2.4-4.7 mm, 7  females 3.3’4.0 mm (mNHN-iU-2008-16087). Solomon Islands. SalOmON 2: stn DW 2229, 6°35.83’S, 156°20.05’e, 315-349 m, 10.29.2004, 1 male 2.2 mm mNHN-iU-2012-1090. Wallis and Futuna Islands. mUSOrStOm 7: stn DW 569, 12°30.0’S, 176°51.2’W, 300-305 m, 21.05.1992, 2 females 1.6, 2.9 mm (USNm). Vanuatu. VOlSmar: stn DW 38, 22°21.60’S, 168°43.10’e, 420 m, 08.06.1989, 1 male 1.8 mm (USNm). – stn DW 51, 20°58’5S, 170°03’4 e, 450 m, 04.07.1989, 4 ovigerous females 3.0-3.9 mm (USNm). New Caledonia. mUSOrStOm 5: stn DW 301, 22°06.90’S, 159°24.60’e, 487- 610 m, 12.10.1986, 2 males 2.2, 2.8 mm, 4 females 1.7-3.0 mm (USNm). – stn DW 305, 22°09.27’S, 159°24.42’e, 430-440 m, 12.10.1986, 1 male 1.8 mm (USNm). – stn DW 306, 22°07.66’S, 159°21.40’e, 375-415 m, 12.10.1986, 2 males 2.5, 3.9 mm, 1 female 2.4 mm, 1 spm sex indet. 1.2 mm (USNm). – stn DW 338, 19°51.60’S, 158°40.40’e, 540-590 m, 15.10.1986, 1 male 3.6 mm (USNm). – Smib 3: stn DW 5, 24°55.00’S, 168°21.70’e, 502-512 m, 20.05.1987, 1 male 3.8 mm, 1 female 3.1 mm (USNm). – mUSOrStOm 6: stn DW 428, 20°23.54’S, 166°12.57’e, 420 m, 17.02.1989, 1 male 2.2 mm (USNm). – stn DW 460, 21°01.72’S, 167°31.45’e, 420 m, 20.02.1989, 1 male 3.3 mm (USNm). – Smib 8: stn DW 150, 24°54.30’S, 168°22.23’e, 519-530 m, 27.01.1993, 1 female 3.4 (mNHN-Pg 6891). – batHUS 2: stn DW 719, 22°47.57’S, 167°14.58’e, 444-455 m, 11.05.93, 2 females 2.8, 2.9 mm (mNHN-Pg 6886). – batHUS 3: stn DW 838, 23°00.81’S, 166°55.87’e, 400-402 m, 30.11.1993, 2 males 2.2, 2.7 mm 1 female 2.4 mm (mNHN-Pg 6887). – batHUS 4: stn DW 923, 18°51.51’S, 163°24.17’e, 470-502 m, 06.08.1994, 1 male 3.5 mm (mNHN-Pg 6888). – NOrFOlK 2: stn DW 2032, 23°39’S, 167°43’e, 420-450 m, 22.10.2003, 1 ovigerous female 5.0 mm (mNHN-iU-2008-16089). – stn CP 2050, 23°42’S, 168°16’e, 377 m, 24.10.2003, 1 male 3.3 mm (mNHN-iU-2008-16088). – stn  CP  2114, 23°45’S, 168°17’e, 390-398 m, 31.10.2003, 1 male 3.8 mm (mNHN-iU-2008-16090). – stn  2115, 23°45’S, 168°17’e, 377-401 m, 31.10.2003, 1 male 8.7 mm (mNHN-iU-2008-7559). – stn DW 2136, 23°01’S, 168°23’e, 402-410 m, 03.11.2003, 1 male 5.1 mm (mNHN-iU-2008-16092). – stn DW 2147, 22°49.80’S, 167°16.09’e, 496 m, 04.11.2003, 1 female 4.1 mm mNHN-iU-2012-1091. – ebiSCO: stn DW 2534, 22°18.004’S, 159°28.136’e, 340-390 m, 10.10.2005, 1 male 1.8, 1 ovigerous female 2.8 mm mNHN-iU-2012-1092. – stn DW 2577, 20°20.317’S, 158°41.62’e, 280-399 m, 10.14.2005, 1 male 2.0 mm mNHN-iU-2012-1093. – stn DW 2578, 20°19.785’S, 158°39.714’e, 180-440 m, 10.14.2005, 1 male 3.0 mm mNHN-iU-2012-1094. – stn CP 2579, 20°19.744’S, 158°39.721’e, 180-440 m, 10.14.2005, 1 male 2.2 mm mNHN-iU-2012-1095. – stn DW 2580, 20°21.024’S, 158°39.800’e, 180-448 m, 10.14.2005, 1 female 2.3 mm mNHN-iU-2012-1096. – stn DW 2606, 19°36.805’S, 158°42.062’e, 320- 442 m, 10.18.2005, 3 males 2.7-2.8 mm, 1 female 2.7 mm mNHN-iU-2012-1097. – stn DW 2607, 19°34.300’S, 158°40.489’e, 340-400 m, 10.18.2005, 1 male 2.5 mm mNHN-iU-2012-1098. – stn DW 2610, 19°33.147’S, 158°40.618’e, 160-486 m, 10.19.2005, 3 males 3.1-4.0 mm, 1  female 3.7 mm mNHN-iU-2012-1099. – stn DW 2617, 20°05.805’S, 160°23.043’e, 270-427 m, 10.20.2005, 1 male 2.6 mm mNHN-iU-2012-1100. – stn DW 2631, 21°03.78’S, 160°44.408’e, 345-372 m, 10.21.2005, 1 female 3.2 mm mNHN-iU-2012-1101. – berYX 11: stn CP 21, 24°45.03’S, 168°06.80’e, 430-450 m, 17.10.1992, 1 female 3.3 mm (mNHN-Pg 6889), 1 female 5.4 mm (mNHN-Pg 6890). Tonga. bOrDaU 2: stn CP 1510, 21°05’S, 175°23’W, 461-497 m, 31.05.2000, 1 male 3.4 mm, 2 ovigerous females 2.9 mm, 3.8 mm (mNHN-iU-2008-15713). – stn CP 1643, 21°05’S, 175°22’W, 487 m, 22.06.2000, 1 ovi- gerous female 3.6 mm (mNHN-iU-2008-15726). Hawaiian Islands. bUrCH: stn 79036,21°16.6’N, 157°54.2’W, 265 m, 28.05.1979, 1 female 3.2 mm (bPbm S10989), 1 male 1.3 mm (bPbm). – West lanai: stn mac 3, 20°48’N, 157°01’W, 256-274 m, 11.10.1967, 6 males 3.0-3.9 mm, 8 females 2.1- 3.7 mm (bPbm). – NOaa/NmFS “townsend Cromwell”: stn tC 33-15, 21°03.2’N, 156°43.5’W, 241-254 m, 31.10.1967, 1 male 4.0 mm (bPbm S11004). – stn tC 33-23, 21°05.0’N, 156°32.6’W, 314-380 m, 03.11.1967, 1 male 3.6 mm (bPbm S10955). – stn tC 33-34, 20°02.7’N, 155°53.9’W, 344-356 m, 08.11.1967,1 male 5.7 mm (bPbm). – stn tC 33-38, 20°41.3’N, 156°41.3’W, 289-296 m, 09.11.1967, 3 males 2.7-5.1 mm, 1 ovigerous female 3.3 mm, 1 female 3.6 mm (bPbm), 6 males 3.1-4.8 mm, 2 ovigerous females 3.4, 3.7 mm, 1 female 3.7 mm (bPbm), 1 ovigerous female 3.5 mm, 1 female 3.6 mm (bPbm). – stn tC 35-15, 21°05.0’N, 156°32.0’W, 360 m, 01.04.1968, 4 males 3.1-6.4 mm (bPbm), 1 female 3.2 mm (bPbm). – stn tC  40- 91, 21°03.8’N, 156°32.8’W, 274 m, 25.11.1968, 2 ovigerous females 3.9, 5.4 mm (bPbm). French Polynesia. beNtHaUS: stn DW 1897, 27°34.27’S, 144°26.68’W, 480-700 m, 08.11.2002, 2 males 4.6, 4.8 mm, 1 ovigerous female 4.4 mm, 5 females 3.6-4.8 mm (mNHN-Pg 6895). – stn DW 1940, ParagioPagurus: wordldwide review and new species 339 23°49.28’S, 147°41.45’W, 100-460 m, 15.11.2002, 1 male 3.8 mm, 1 ovigerous female 3.8 mm, 1 female 2.4 mm (mNHN-Pg 6896). – stn DW 1951, 23°49.08’S, 147°53.38’W, 206-450 m, 17.11.2002, 1 female 1.6 mm (mNHN-Pg 6897). – stn DW 1999, 22°25.12’S, 151°22.15’W, 270-500 m, 23.11.2002, 1 female 2.7 mm (mNHN-Pg 6898). – stn DW 2000, 22°25.11’S, 151°21.88’W, 270-480 m, 23.11.2002, 1 male 3.4 mm (mNHN-Pg 6899). – stn DW 2009, 22°31.98’S, 151°19.85’W, 320-450 m, 24.11.2002, 1 male 4.1 mm (mNHN-Pg 6900). For additional material examined, including specimen illustrated in Figures 8, 9, see lemaitre (1990 as Sympagurus ruticheles, 1996). DISTRIBUTION — Western Indian Ocean: Madagascar. Western and central Pacific: Taiwan, Solomon Islands, Wallis and Futuna Islands, Vanuatu, Tonga, New Caledonia, Southern Australia, Hawaiian Islands, French Polynesia, and Nazca and Sala-y-Gómez Ridges. Eastern Atlantic: from southern coast of Portugal to Senegal, including the Azores, Madeira and the Canary Islands. Depth: 100 to 1440 m. FIGURE 9 Paragiopagurus ruticheles (a. milne-edwards, 1891), male (3.4 mm) rmNH, northeastern atlantic, CaNCaP iV stn 4.029. A, shield and cephalic appendages. B, right antennal peduncle, lateral view. C, right third pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of right fourth pereopod, lateral view. F, telson, dorsal view. Scale bar 3 mm for a, C, D; 2 mm for b; 1 mm for e, F. (adapted from lemaitre 1990). F E C, D B A E F D A CB rafael lemaitre340 DIAGNOSIS — Gills with lamellae undivided distally. Shield (Figure 9A) about as broad as long, dorsal surface usually weakly calcified medially and anteriorly; rostrum broadly rounded, with short dorsal ridge; lateral projections subtrian- gular, with small terminal spine. Ocular peduncles more than half length of shield; acicles terminating in strong spine, mesial margins nearly parallel; corneas slightly dilated. Antennular peduncles exceeding distal margins of corneas by slightly less than length of ultimate segment. Antennal peduncles (Figure 9B) not exceeding distal margins of corneas; acicle not exceeding distal margin of cornea, mesial margin of each armed with 7-14 spines. Epistomial spine straight, oc- casionally bifid. Right cheliped (Figure 10B-E) elongate, with transverse furrows on ventral surfaces of chela and ventrola- teral face of carpus; palm with well delimited dorsomesial, ventromesial and dorsolateral margins each armed with spines (usually having corneous tips), and concave mesial face often expanded distally; carpus with well delimited dorsolateral margin armed with corneous-tipped spines. Left cheliped (Figure 10A) weakly calcified on merus and carpus; unarmed except for scattered small spines on dorsal surface of palm and dorsal margin of carpus. Ambulatory legs (Figure 9C, D) with dactyls each having dorsal and distal dorsomesial row of setae, and ventromesial row of about 13 spinules; merus of right third pereopod usually with dorsal row of small spines. Anterior lobe of sternite XII (of third pereopods) unarmed. Fourth pereopod (Figure 9E) with strongly curved dactyl; propodal rasp consisting of 1 row of ovate scales. Uropods and telson (Figure 9F) strongly asymmetrical; telson with transverse suture separating anterior and posterior lobes; poste- rior lobes separated by shallow median cleft, terminal margins armed with strong often curved corneous spines. Males lacking first gonopods; with unpaired, uniramous second left gonopod. Females lacking vestigial right second pleopod. FIGURE 10 Paragiopagurus ruticheles (a. milne-edwards, 1891), male (3.4 mm) rmNH, northeastern atlantic, CaNCaP iV stn 4.029. A, left cheliped, dorsal view. B-E, right cheliped in lateral (b), dorsal (C), ventral (D), and mesial (e) views. Scale bars 2 mm. (adapted from lemaitre 1990). EDCBA ParagioPagurus: wordldwide review and new species 341 COLOUR — Overall, evenly cream or light yellow. HABITAT — Gastropod shells. REMARKS — Paragiopagurus ruticheles is the most broadly distributed species of Paragiopagurus, occurring in the wes- tern Indian Ocean, western, central and South Pacific Ocean, and the eastern Atlantic. Although broadly distributed, so far this species has not been found in the eastern Pacific or western Atlantic. Morphologically, P. ruticheles is the most distinct among species of Paragiopagurus, and even all parapagurids. The transverse furrows on the ventral surfaces of the chela and ventrolateral face of carpus of the right cheliped, immediately distinguishes P. ruticheles from all others in the genus and family. Despite being widely distributed, minimal morphological variations have been observed in all the material examined. The color pattern is also indistinct in fresh specimens. As commonly encountered in many parapagu- rid species and even paguroids in general, the right chela is broader in females than in males. Paragiopagurus schnauzer Lemaitre, 2006 Figures 11, 12 Paragiopagurus schnauzer lemaitre, 2006b: 524, figs 5-8 (type locality : New Caledonia). — mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, New Caledonia, BIOGEOCAL, stn CP 214, 22°43.09’S, 166°27.19’E, 1590-1665 m, 09.04.1987, male 3.3 mm (MNHN-IU-2008-16047). Paratypes, Loyalty Islands. HALIPRO 2: stn BT 105, 25°45’S, 162°50’E, 1200- 1218 m, 27.11.1996, 1 male 3.2 mm (MNHN-IU-2008-15792). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — New Caledonia region. Depth: 1200 to 1665 m. DIAGNOSIS — Gill lamellae deeply divided (Figure 11A). Shield (Figure 11B) distinctly longer than broad, weakly calcified medially and anteriorly. Rostrum broadly subtriangular, reaching well in advance of lateral projections, with short mid- dorsal ridge. Ocular peduncles stout, diminishing in width distally, somewhat inflated ventroproximally; with longitudinal row of long setae dorsally. Corneas reduced, subconical, terminating bluntly or sharply (Figure 11C). Ocular acicles sub- triangular, each terminating in strong spine. Antennular peduncles exceeding distal margins of corneas by 0.7 to nearly full length of penultimate segment. Antennal peduncles (Figure 11D) exceeding distal margins of corneas by 0.3 length of fifth segment; fourth segment lacking spine on dorsodistal margin; second segment with dorsodistal angle produced, ter- minating in strong spine; first segment with small spine on lateral face; antennal acicle exceeding distal margin of cornea by about half length of acicle, terminating in strong spine, armed mesially with 4-8 spines. Flagellum with numerous short and long setae < 1 to 4 flagellar articles in length. Maxillule with internal lobe with long, terminal seta. Third maxilliped with crista dentata with about 14 corneous-tipped teeth. Epistomial spine straight, simple. Chelipeds markedly dissimilar. Right cheliped (Figure 11E, F) operculate, with at most moderately dense setae (mostly plumose) on dorsal surfaces of carpus and chela; fingers slightly curved ventromesially; dactyl set at strongly oblique angle to longitudinal axis of palm, mesial margin well defined by row of spines or tubercles, dorsomesial surface with small spines, ventromesial face at most weakly concave; fixed finger broad basally, with scattered small tubercles on dorsal surface, unarmed on ventral surface except for a few tufts of setae; palm broadening distally, slightly broader than long, dorsolateral margin well defined by small spines at least on distal half, dorsomesial margin with irregular row of small spines, mesial face rounded; carpus with dorsal surface with few, widely separated small tubercles or spines. Left cheliped (Figure 11G) well calcified; palm dorsal surface usually with small median spine or tubercle proximally; carpus with strong dorsodistal spine. Ambulatory rafael lemaitre342 legs (Figure 12A-D) with dactyl having dorsal and distomesial rows of long setae, and ventromesial row of 4-6 small, cor- neous spinules; carpus with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) subsemicircular, sparsely setose, lacking spine. Fourth pereopod (Figure 12E) with propodal rasp consisting of 1 row (at least distally) of FIGURE 11 Paragiopagurus schnauzer lemaitre, 2006, holotype male (3.3 mm) mNHN-iU-2008-16047, New Caledonia, biOGeOCal stn CP 214, 1590-1665 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages, dorsal view. C, right ocular peduncle, lateral view. D, right antennal peduncle, lateral view. E, right cheliped, dorsal view. F, chela of same, lateral view. G, left cheliped, lateral view. Scale bars 0.25 mm for a, C; 1 mm for b, e-G; 0.5 mm for D. D B, E-G A-C EGF C A DB ParagioPagurus: wordldwide review and new species 343 ovate scales. Uropods and telson (Figure 12F) symmetrical or nearly so. Telson with weak or obsolete transverse suture; dorsal surface with low, blister-like tubercles; posterior margin separated by shallow, narrow U-shaped cleft, into rounded projections each armed with about 8-10 corneous spines, some often ventrally curved. FIGURE 12 Paragiopagurus schnauzer lemaitre, 2006, holotype male (3.3 mm) mNHN-iU-2008-16047, New Caledonia, biOGeOCal stn CP 214, 1590-665 m. A, right second pereopod, lateral view. B, dactyl of same, mesial view. C, right third pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of left fourth pereopod, lateral view. F, uropods and telson, dorsal view. G, left first gonopod, mesial view. H, left second gonopod, anterior view. Scale bars 1 mm for a-e; 0.5 mm for F; 1 mm for G, H. F E, G, H A-D HGF E D C B A rafael lemaitre344 Male with paired first and second gonopods well developed; first gonopod (Figure 12G) with concave distal lobe; second gonopod (Figure 12H) with or without rudimentary exopod, distal segment with row of short bristles on lateral margin medially, and long setae on distomesial face. Female unknown. COLOUR — Unknown. HABITAT — Scaphopod shells. REMARKS — This species is known only from the type material used by Lemaitre (2006b). It is unique among species of Paragiopagurus in having reduced, subconical corneas (Figure 11B, C), the length of ocular peduncles relative to the shield being less than half the shield length, the distal extension of the antennal acicles exceeding the distal margins of corneas by about half the length of the acicle, and in having nearly symmetrical uropods and telson. Paragiopagurus rugosus (de Saint Laurent, 1972) Figures 1e, 13, 14, 15e-G Parapagurus rugosus de Saint laurent, 1972: 112, figs 6, 21 (type locality: Hawaiian islands). Sympagurus rugosus – lemaitre, 1989: 37; 1994: 412. Paragiopagurus rugosus – lemaitre, 1996: 207. — mclaughlin et al., 2005: 246; 2010: 39. — Castro, 2011: 30. TYPE MATERIAL — Holotype, Hawaiian Islands, Kalohi Channel, between Molokai and Lanai, Albatross, stn 3859, 21°02’40”N, 156°44’ 20”W, 252-256 m, 09.04.1902, male 6.4 mm (USNM 168307). Paratypes, Hawaiian Islands. Albatross: stn 3859, 21°02’40”N, 156°44’ 20”W, 252-256 m, 09.04.1902, 1 male 7.3 mm, 1 female 7.8 mm (USNM 168308), cheliped and ambulatory legs of presumed paratype (no body) (MNHN-Pg 6885, see REMARKS). MATERIAL EXAMINED — the type material (see above). Hawaiian Islands. albatross: stn 3937, 25°52’05”N, 171°46’47”W, 238-271 m, 16.05.1902, 1 female 4.3 mm (USNm 1154275). – French Frigate Shoals: 23°79.13’N, 166° 39.33’W, baited traps, 185-251 m, 6.10. 2006: 1 male 11.6 mm (UF 12154), 1 female 9.2 mm (UF 12147). – bUrCH: stn 79036, 21°16.6’N, 157°54.2’W, 265 m, 28.05.1979: 1 female 4.5 mm (bPbm S10972). – NOaa/ NmFS “townsend Cromwell”: stn tC 35-15, 21°05’N, 156°32’W, 360.3 m, 01.03.1968, 1 ovigerous female 11.6 mm (USNm 1154277). – stn tC 33-26, 20°43.1’N, 156°51.3’W, 221-232 m, 04.11.1967, 1 male 10.1 mm (bPbm). Marquesas Islands. mUSOrStOm 9: stn DW 1165, 8°58.2’S, 140°05.0’W, 350- 360 m, 24.08.1997, 1 ovigerous female 4.4 mm (mNHN-Pg 6884). DISTRIBUTION — Central Pacific: Hawaiian and Marquesas Islands. Depth: 221 to 360 m. REDESCRIPTION — Eleven pairs of quadriserial gills (Figure 13A), with lamellae weakly divided distally. Shield (Figure 13B) approximately as broad as long; dorsal surface weakly calcified along midline posteriorly to rostrum, and anterior and anterolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections subtriangular, with small subterminal spine. Anterolateral margins sloping. Ventrolateral margins without spine. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, with longitudinal row of few, short setae on dorsal surface. Cornea weakly dilated. Ocular acicles subtriangular, each terminating in strong simple spine; separated basally by about half width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by about 0.75 length of ultimate segment; ventral flagellum with about 10-14 articles. Ultimate segment about twice as long as penultimate segment, with scattered setae dorsally. ParagioPagurus: wordldwide review and new species 345 FIGURE 13 Paragiopagurus rugosus (de Saint laurent 1972). A-C, E-G, ovigerous female (4.4 mm) mNHN-Pg 6884, marquesas islands, mUSOrStOm 9 stn DW 1165, 350- 360 m. D, holotype male (6.4 mm) USNm 168307, Hawaiian islands, albatross stn 3859, 252-56 m. H, I, male (10.1 mm) bPbm, Hawaiian islands, NOaa/NmFS “townsend Cromwell” stn tC 33-26, 221-232 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages. C, right antennal peduncle, lateral view. D, merus and chela of right cheliped, dorsal view (from de Saint laurent, 1972). E, right chela, ventral view. F, area of ventral surface of right palm at about midsection. G, left cheliped, dorsal view. H, left first gonopod, mesial view. I, left second gonopod, anterior view. Scale bars 0.5 mm for a; 2 mm for b, e, G; 1 mm for C, H, i; 0.5 mm for F. I H D C, F B, E, G A I HE A B F DCG rafael lemaitre346 Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 1 or 2 spines, and strong spine proximally. Antennal peduncles (Figure 13C) at most reaching to distal margins of corneas. Fifth segment unarmed, with few, short bristle-like setae on dorsolateral distal angle, and mesial margin. Fourth segment with 1 strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong simple or multifid spine extending for about 0.3 length of acicle and sometimes with 1 or more additional spines proximally on dorsal margin; mesial margin with small spine on dorsolateral distal angle. First segment with lateral surface armed with small spine; ventromesial angle produced, with row of 3 often strong spines laterally. Antennal acicle nearly straight or broadly curved outward (dorsal view), at most reaching distal margin of corneas, and terminating in strong spine; mesial margins setose, with row of 8-10 strong spines diminishing in strength distally. Flagellum exceeding distal end of right cheliped; with long setae 1-4 flagellar articles in length every 10-12 articles. Mandible with 3-segmented palp. Maxillule with external lobe of endopod slender, moderately developed, internal lobe with 4 long setae. Maxilla with endopod slightly exceeding distal margin of scaphognathite. First maxilliped with endopod exceeding distal margin of exopod. Second maxilliped without distinguishing characters. Third maxilliped with dactyl, carpus and propodus about 2.5 as long as wide; crista dentata with about 12-14 small teeth; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine short, straight. Chelipeds markedly dissimilar. Right cheliped (Figures 13D-F, 15E, F) massive, chela somewhat dorsoventrally flattened, operculate. Fingers slightly curving ventromesially, each terminating in small, usually blunt corneous claw; cutting edges with row of calcareous teeth, that of dactyl also with short row of small corneous spines distally. Dactyl longer than length to palm, set at strongly oblique angle relative to longitudinal axis of palm; dorsal surface with numerous small to strong corneous-tipped spines; ventral face with corneous-tipped spines irregularly arranged and often with longitudinal row of spines; ventromesial face concave proximally. Fixed finger broad at base, armed similarly to dactyl. Palm broader than long (broader in females than in males); dorsolateral and dorsomesial margins well delimited by row of spines often corneous- tipped; mesial face rounded, with spines or tubercles; dorsal surface sparsely setose, armed with dense, mostly strong and often corneous-tipped spines (spines stronger distolaterally); ventral surface (Figure 13E, F) sparsely setose, densely covered with small spines (sometimes fused) and most corneous-tipped. Carpus sparsely setose, subtriangular in cross- section; dorsal surface with numerous small spines on dorsal surface; ventrolateral margin relatively well defined distally by short row of spines; dorsodistal margin with row of small spines; ventromesial distal margin with row of spines. Merus unarmed dorsally except for 1 or 2 dorsodistal spines and small granules; ventromesial margin with row of blunt spines. Ischium unarmed except for setae on dorsal margin and ventral surface; coxa with row of small tubercles or spines on ventrodistal margin, and ventromesial row of long setae. Left cheliped (Figures 13G, 15G) usually well calcified; armature variable depending on size, with spines stronger in specimens sl > 10.0 mm. Fingers terminating in sharp corneous claws; with few small to strong spines on lateral (dactyl) and mesial (fixed finger) margins, dorsal and ventral surfaces with or without strong spines, with tufts of setae near cutting edges; cutting edge of dactyl with row of closely-set or fused small corneous spinules; cutting edge of fixed finger nearly straight. Dactyl about as long as palm. Palm sparsely setose; with dorsal, dorsolateral and mesial surfaces varying in armature from few small tubercles or spines to numerous strong, often corneous-tipped spines. Carpus densely setose dorsally; with dorsodistal spine, and sometimes small dorsolateral distal spine, or with row of strong spines on dorsal margin; dorsal and dorsomesial surfaces with few to many small, low spines or tubercles. Merus sparsely setose, unarmed or with small, low tuberlces on dorsal surface and row of small to strong spines on ventrolateral distal margin. Ischium unarmed except for dorsal and ventral setae. Coxa with ventrodistal margin unarmed or with small spine mesially, and ventromesial row of long setae. Ambulatory legs (Figure 14A-E) similar from right to left, usually exceeding right cheliped by about 0.3 length of dactyl of legs when fully extended. Dactyl approximately 1.4 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, ventromesial margin armed with irregular row of about 14-24 corneous spinules; mesial faces with shallow, longitudinal sulcus on proximal half. Propodus with long setae and short transverse ParagioPagurus: wordldwide review and new species 347 rows of short bristle-like setae on dorsal surface; ventrodistal margin with scattered setae distally. Carpus with row of small dorsodistal spine, and setae or tufts of setae on dorsal margin; dorsal margin with row of small spines (stronger on larger specimens and on second pereopod than on third, spines not always clearly visible in lateral view). Merus unarmed (second pereopod) or with row of small blunt spines (third pereopod) and setae on dorsal margin; ventral margin unarmed or with row of small, blunt tubercles. Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods) unarmed except for setae. FIGURE 14 Paragiopagurus rugosus (de Saint laurent, 1972). A-D, F, G, ovigerous female (4.4 mm) mNHN-Pg 6884, marquesas islands, mUSOrStOm 9 stn DW 1165, 350-360 m. E, male (10.1 mm) bPbm, Hawaiian islands, NOaa/NmFS “townsend Cromwell” stn tC 33-26, 221-232 m. A, second right pereopod, lateral view. B, dactyl of same, mesial view. C, third right pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and carpus of second right pereopod, mesial view. F, propodus and dactyl of fourth left pereopod, lateral view. G, propodus and dactyl of fifth left pereopod, lateral view. H, telson, dorsal view. Scale bars 2 mm for a-D; 3 mm for e; 1 mm for F-H. F-H E A-D F GH C E DB A rafael lemaitre348 Fourth pereopod (Figure 14F) subchelate. Dactyl broadly curved, terminating in sharp often slender, long corneous claw; with ventrolateral row of small corneous spinules. Propodus longer than high, rasp consisting of 1 row (at least distally) of ovate scales. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 14G) chelate. Propodal rasp extending posteriorly to or beyond mid-length of segment. Uropods and telson (Figure 14H) markedly asymmetrical. Telson with weak lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 broadly rounded projections by shallow U-shaped cleft; rounded projections each armed with row of often strong and curved, corneous spines. Male with paired first and second gonopods well developed. First gonopod (Figure 13H) with distal lobe subtriangular, and concave mesial face. Second gonopod (Figure 13I) with distal segment setose on distomesial face, and with bristle- like setae on mid-lateral margin. Female with vestigial right second pleopod. COLOUR (Figure 1e) — Shield overall whitish to light pink. Ocular peduncles whitish, with lateroventral faces dark orange. Antennular and antennal peduncles whitish. Chelipeds whitish or light pink , with many small, light orange spots; some spines on dorsal surfaces of chela and carpus orange. Ambulatory legs and fourth pereopods white with small, light orange spots. HABITAT — Gastropod shells. REMARKS — De Saint Laurent (1972) selected and listed the station data for the holotype. Although she mentioned that only three specimens, all from the Hawaiian Islands, were known, she did not indicate the station data or deposition for the two paratypes. The latter are deposited in the USNM and are from the same ALBATROSS station (stn 3859) as the FIGURE 15 A-D, F, right cheliped. E, right chela. G, left chela. A, B, Paragiopagurus diogenes (Whitelegge, 1900), male (12.4 mm) am P.9312, off New South Wales, australia. C, D, Paragiopagurus sculptochela (Zarenkov, 1990), paratype male (6.0 mm) ZmUm 4835, Nazca and Sala-y-Gómez ridges, r/V “Professor Shtockman” stn 1997, 330-350 m. E, F, G, Paragiopagurus rugosus (de Saint laurent, 1972), male (10.1 mm) bPbm, Hawaiian islands, NOaa/NmFS “townsend Cromwell” stn tC 33-26, 221-232 m. A, mesial view. B, F, dorsal view. C, E, ventromesial view. D, ventral view. Scale bars 5 mm. A FG EDC B ParagioPagurus: wordldwide review and new species 349 holotype (see Type Material). One lot, presumably also of paratype material from the same station and identified by M. de Saint Laurent as this species, was found at the MNHN (MNHN-Pg 6885), although with only dismembered appendages (chelipeds, and second and third pereopods) and without a body. Additional specimens are herein reported from the Hawaiian Islands, and also the Marquesas Islands, some 1000 km to the southeast. De Saint Laurent’s (1972) original description of P. rugosus, based on a few specimens from the Hawaiian Islands, was brief and lacked a number of important details that have now been included in the above redescription of this species. The additional specimens discovered during this study from the Hawaiian and Marquesas Islands show that the dense armature of the ventral surface of the right chela can vary from small spines or tubercles in small specimens (sl < 4.5 mm) to strong spines in large specimens (sl > 10.0 mm; Figures 13E, 15E, F). Also variable is the armature of the left cheliped, where the spines on the dorsal, lateral and mesial surfaces of chela, and dorsal margin of the carpus, can be few and weak, or numerous, strong and often bearing minute corneous tips (Figure 15G). The dorsal margin of the merus of the second and right pereopods may lack spines or have inconspicuous spines that are more clearly visible in mesial view, or in larger specimens (sl > 10.0mm) the dorsal margin has a row of distinct spines (Figure 14E). Paragiopagurus umbonatus n. sp. Figures 16-19 Paragiopagurus sp. 4 – mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, New Caledonia, SMIB 4, stn DW 36, 24°55.60’S, 168°21.70’E, 530 m, 07.03.1989, male 5.2 mm (MNHN-IU-2012-747). Paratypes: New Caledonia. MUSORSTOM 5: stn DW 272, 24°40.91’S, 159°43.00’E, 500-540 m, 09.10.1986, 1 juvenile male 2.9 mm (MNHN-IU-2012-742). – CHALCAL 2: stn CC 1, 24°54.96’S, 168°21.91’E, 500-580 m, 28.10.1986, 1 male 5.7 mm, 3 females 3.1-4.5 mm (USNM 1184056). – stn CC 2, 24°55.48’S, 168°21.29’E, 500-610 m, 28.10.1986, 2 males 4.3, 4.6 mm, 3 females 3.6-4.6 mm (MNHN-IU-2012-737). – stn DW 72, 24°54.50’S, 168°22.30’E, 527 m, 28.10.1986, 1 male 4.3 mm (USNM 1184057), 1 female 4.8 mm (MNHN-IU-2012-738). – stn DW 73, 24°39.90’S, 168°38.10’E, 573 m, 29.10.1986, 1 female 4.2 mm (MNHN-IU-2012-739). – stn DW 74, 24°40.36’S, 168°38.38’E, 650 m, 29.10.1986, 1 female 5.4 mm (MNHN-IU-2012-740). – stn DW 75, 24°39.31’S, 168°39.67’E, 600 m, 29.10.1986, 2 males 6.9, 7.6 mm, 2 females 4.3, 4.9 mm (USNM 1184058), 2 females 5.7, 6.0 mm (MNHN-IU-2012-741). – SMIB 3: stn DW 5, 24°54.90’S, 168°21.60’E, 502 m, 21.05.1987, 1 male 4.3 mm (MNHN-IU-2012-744). – stn DW 6, 24°56.40’S, 168°21.20’E, 505 m, 21.05.1987, 1 male 3.9, 1 female 3.7 mm (USNM 1184059). – stn DW 7, 24°54.60’S, 168°21.30’E, 505 m, 21.05.1987, 2 males 4.5, 4.8 mm, 1 ovigerous female 4.5 mm, 2 females 3.4, 3.5 mm (MNHN-IU-2012-745). – stn DW 29, 22°46.70’S, 167°11.70’E, 405 m, 25.05.1987, 1 male 3.9 mm (MNHN-IU-2012-743). – SMIB 4: stn DW 36, 24°55.60’S, 168°21.70’E, 530 m, 07.03.1989, 1 male 5.1 mm, 2 females 3.7, 4.5 mm (MNHN-IU-2012-746). – stn DW 39, 24°56.20’S, 168°21.50’E, 525-560 m, 07.03.1989, 1 male 5.5 mm, 1 female 4.2 mm (USNM 1184060). – BERYX 11: stn CP 07, 24°56.20’S, 168°21.60’E, 510-550 m, 15.10.1992, 1 female 3.4 mm (MNHN-IU-2008-15809). – stn DW 10, 24°53.15’S, 168°21.60’E, 560-600 m, 15.10.1992, 2 males 1.9, 2.1 mm (MNHN-IU-2008-15800). – SMIB 8: stn DW 147, 24°54.90’S, 168°21.85’E, 508-532 m, 27.01.1993, 1 male 4.5 mm, 2 females 4.7, 5.2 mm (MNHN-IU-2008-15810). – stn DW 149, 24°54.94’S, 168°21.82’E, 508-510 m, 27.01.1993, 2 females 3.1, 3.2 mm (MNHN-IU-2008-15811). – stn DW 150, 24°54.30’S, 168°22.23’E, 519- 530 m, 27.01.1993, 2 males 3.4, 3.5 mm (MNHN-IU-2008-15812), 6 males 3.5-5.4 mm, 5 females 2.5-3.6 mm (MNHN- IU-2008-15813). – stn DW 152, 24°54.35’S, 168°22.23’E, 514-530 m, 27.01.1993, 1 male 2.5 mm, 1 female 3.1 mm (MNHN- IU-2008-15815), 2 juveniles (MNHN-IU-2008-15814). – stn DW 153, 24°53.55’S, 168°21.33’E, 547-560 m, 27.01.1993, 1 female 3.9 mm (MNHN-IU-2008-15816). – BATHUS 3: stn DW 807, 23°40.20’S, 167°59.39’E, 420-438 m, 27.12.1993, 1 male 4.9 mm (MNHN-IU-2008-15817). – stn DW 819, 23°45,11’S, 168°16,27’E, 478 m, 28.11.1993, 1 male 1.7 mm, 2 females 2.9, 4.2 mm (MNHN-IU-2008-15808). – LITHIST: stn CP 08, 24°54.24’S, 168°21.35’E, 540 m, 11.08.99, 2 males 2.8, 3.5 mm (MNHN-IU-2008-15818). – NORFOLK 1: stn DW 1700, 24°39.459’S, 168°39.588’E, 572-605 m, 24.06.2001, rafael lemaitre350 FIGURE 16 Paragiopagurus umbonatus n. sp. A, paratype male (5.1 mm) mNHN-iU-2012-746, New Caledonia, Smib 4 stn DW 36, 530 m. B-G, holotype male (5.2 mm) mNHN-iU-2012-747, New Caledonia, Smib 4 stn DW 36, 530 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages. C, right antennal peduncle and anterior portion of branchiostegite, lateral view. D, anterior lobe of sternite Xii (of third pereopods), ventral view. E, telson, dorsal view. F, left first gonopod, mesial view. G, left second gonopod, anterior view. Scales bar 0.25 mm for a; 1 mm for b, F, G; 0.5 mm for C-e. D, E FE D C A B G A B F, G C ParagioPagurus: wordldwide review and new species 351 FIGURE 17 Paragiopagurus umbonatus n. sp., paratype male (5.1 mm) mNHN-iU-2012-746, New Caledonia, Smib 4 stn DW 36, 530 m, left mouthparts, internal view. A, mandible. B, maxillule. C, distal end of endopod of same. D, maxilla. E, first maxilliped. F, second maxilliped. G, third maxilliped. Scale bars 0. 5 mm for a, b, D, e; 0.25 mm for C; 1 mm for F, G. F, G C A, B, D, E E FG B DC A rafael lemaitre352 2 males 3.7, 4.8 mm (MNHN-IU-2008-15819). – stn DW 1703, 23°44.05’S, 168°16.19’E, 423-437 m, 25.06.2001, 2 males 4.0, 6.2 mm (MNHN-IU-2008-15820). – NORFOLK 2: stn CP 2050, 23°42.17’S, 168°15.72’E, 377 m, 24.10.2003, 1 da- maged spm sex indet. (MNHN-IU-2008-16076). – stn DW 2067, 25°16.46’S, 168°55.76’E, 614-690 m, 26.10.2003, 1 female 7.9 mm (MNHN-IU-2008-16077). – stn DW 2081, 25°54.40’S, 168°21.64’E, 500-505 m, 28.10.2003, 1 female 4.5 mm (MNHN-IU-2008-16078). – stn DW 2091, 24°45.36’S, 168°06.24’E, 600-896 m, 29.10.2003, 1 female 3.9 mm (MNHN-IU-2008-16079). – stn DW 2098, 24°42.27’S, 168°06.10’ E, 550-668 m, 29.10.2003, 1 female 4.5 mm (MNHN- IU-2008-16080). Johnston Atoll. Hawaii Undersea Research Lab: dive 208, 16°41.5’N, 169°34.3’W, 381 m, 23.10.1983, 1 ovigerous female 7.5 mm (BPBM S10547). French Polynesia. BENTHAUS: stn DW 1897, 27°34.27’S, 144°26.68’W, 480-700 m, 08.11.2002, 2 males 6.7, 8.3 mm, 2 females 5.0, 5.4 mm (MNHN-IU-2008-15821). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — Western, central, and South Pacific, from New Caledonia to French Polynesia, including Johnston Atoll. Depth: 377 to 668 m. DESCRIPTION — Eleven pairs of quadriserial gills (Figure 16A) weakly divided distally. Shield (Figure 16B) approximately as broad as long; dorsal surface weakly calcified along midline, just posteriorly to rostrum, and anterior and anterolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections bluntly subtriangular, or with small terminal spine. Anterolateral margins sloping. Ventrola- teral margins without spine or occasionally with minute spine, on one or both sides. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, each with longitudinal row of short setae on dorsal surface. Corneas weakly dilated. Ocular acicles subtriangular, about 0.25 as long as ocular peduncles, each terminating in strong simple spine; separated basally by about half width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by 0.75 or more length of ultimate segment; ventral flagellum with about 10 articles. Ultimate segment about twice as long as penultimate, with scattered setae dorsally. Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 1 or 2 spines, and strong spine proximally. Antennal peduncles (Figure 16C) slightly exceeding distal margins of corneas. Fifth segment unarmed, with few, short bristle-like setae on dorsolateral distal angle, and mesial margin. Fourth segment with 1 (rarely with 2) strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong simple or multifid spine extending for about 0.3 length of acicle; mesial margin with small spine on dorsolateral distal angle. First segment with lateral surface armed with small spine; ventromesial angle produced, with row of about 5 spines laterally. Antennal acicle nearly straight or slightly curved outward (dorsal view), at most slightly exceeding distal margin of cornea, and terminating in strong spine; mesial margin setose, with row of 10 to 14 spines diminishing in strength distally. Flagellum exceeding distal end of right cheliped; with setae 1-4 articles in length. Mandible (Figure 17A) with 3-segmented palp. Maxillule (Figure 17B, C) with external lobe of endopod slender, moderately developed, internal lobe with 2 long setae. Maxilla (Figure 17D) with endopod slightly exceeding distal margin of scaphognathite. First maxilliped (Figure 17E) with endopod slightly exceeding distal margin of exopod. Second maxilliped (Figure 17F) without distinguishing characters. Third maxilliped (Figure 17G) with dactyl, carpus and propodus about 2.5 times as long as wide; crista dentata with about 17 small teeth; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine sharp, occasionally absent. Chelipeds markedly dissimilar. Right cheliped (Figure 18A-F) massive, chela somewhat dorsoventrally flattened, operculate. Fingers slightly curving ventromesially, each terminating in small, usually blunt corneous claw; cutting edges with row of calcareous teeth, that of dactyl also with short row of small corneous spines distally; dorsal faces with dense blunt to sharp spines usually with short plumose setae at bases; ventral faces naked. Dactyl subequal in length to palm, set ParagioPagurus: wordldwide review and new species 353 at strongly oblique angle relative to longitudinal axis of palm; mesial margin well defined by row of spines often corneous- tipped; ventromesial face concave; ventral face with or without longitudinal ridge of distinct low tubercles. Fixed finger broad at base. Palm broader than long; lateral margin well delimited by row of small spines often corneous-tipped; dorsomesial margin delimited by row of spines; mesial face rounded, with spines or tubercles; dorsal surface covered with short, dense plumose setae hiding scattered small spines or tubercles beneath; ventral surface nearly naked, with few to numerous usually well-spaced, low but distinct tubercles (usually larger and more numerous on medial and mesial portions). Carpus setose dorsally, with many small spines on dorsomesial face; ventrolateral margin relatively well defined by row of spines; dorsodistal margin unarmed or with row of small blunt spines; ventromesial distal margin with row of spines. Merus unarmed dorsally except for few setae and small spines or granules; ventromesial margin with row of spines, and in large male specimens (sl > 5.0 mm) with fringe of long, bristle-like setae. Ischium unarmed except for setae on dorsal margin and ventral surface; coxa with row of spines on ventrodistal margin, and ventromesial row of long setae. Left cheliped (Figure 18G) usually well calcified. Fingers terminating in sharp corneous claws; dorsal and ventral surfaces unarmed except for tufts of setae; cutting edge of dactyl with row of closely-set small corneous spinules, cutting edge of fixed finger with row of small calcareous teeth. Dactyl slightly longer or about as long as palm. Palm unarmed except for tufts of setae dorsally. Carpus with densely setose dorsal margin, dorsodistal spine, and sometimes small dorsolateral distal spine. Merus unarmed except for dorsal and ventral bristle-like setae, and row of spines on ventrolateral distal margin. Ischium unarmed except for dorsal and ventral setae. Coxa with ventrodistal margin unarmed or with small spine mesially, and ventromesial row of long setae. Ambulatory legs (Figure 19A-D) similar from right to left, usually exceeding right cheliped by about 0.3 length of dactyl of legs when fully extended. Dactyl approximately 1.7 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, ventromesial margin armed with irregular row of 14-23 corneous spinules; lateral and mesial faces with shallow, longitudinal sulcus on proximal half (deeper on mesial face). Propodus with setae or tufts of setae on dorsal margin. Carpus with small dorsodistal spine, and short setae or tufts of setae on dorsal margin; lateral face with median row of short tufts of setae. Merus with setae on dorsal margin; merus of second pereopod unarmed ventrodistally except for row of setae or bristles-like setae. Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods) setose, usually lacking spine or at most with minute subterminal spine (Figure 16D). Fourth pereopod (Figure 19E) subchelate. Dactyl broadly curved, terminating in sharp corneous claw; with ventrolateral row of small corneous spinules. Propodus longer than wide, rasp consisting of 1 row (at least distally) of rounded scales. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 19F) chelate. Propodal rasp extending posteriorly to or beyond mid-length of segment. Uropods and telson (Figure 16E) markedly asymmetrical. Telson lacking lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 broadly rounded projections by shallow U-shaped cleft; rounded projections each armed with row of often curved, corneous spines. Male with paired first and second gonopods well developed. First gonopod (Figure 16F) with distal lobe subtriangular, and concave mesial face. Second gonopod (Figure 16G) often with rudimentary exopod. Female with rudimentary second right pleopod. COLOUR — Unknown. HABITAT — Gastropod shells with anthozoan polyp. REMARKS — This new species resembles P. boletifer and P. sculptochela, particularly in the armature of the right palm. These three species have distinct tubercles on the ventral and mesial faces of the right palm. In P. umbonatus n. sp., the tubercles are low, rounded, and usually well-spaced and concentrated on the ventromedial and mesial faces (Figure 18B, D, E); in P. boletifer the tubercles are high and numerous, usually mammilliform in shape, and cover the entire ventral and mesial faces (Figure 21A, B, D); and in P. sculptochela, the tubercles are high and cylindrical in shape (Figure 15C, D). rafael lemaitre354 FIGURE 18 Paragiopagurus umbonatus n. sp. A-C, G, holotype male (5.2 mm) mNHN-iU-2012-747, New Caledonia, Smib 4 stn DW 36, 530 m. D, paratype female(3.7 mm) mNHN-iU-2012-746, New Caledonia, Smib 4 stn DW 36, 530 m. E, F, paratype female (6.0 mm) mNHN-iU-2012-741, New Caledonia, CHalCal 2 stn DW 75, 600 m. A, left cheliped, dorsal view. B, right cheliped, dorsal view. C, chela of same, ventral view. D, same chela, mesial view. E, F, right chela, ventral view. G, lateral view of chela shown in F. Scale bar 2 mm for a-D, F, G; 1 mm for e. D B AG EFC ParagioPagurus: wordldwide review and new species 355 FIGURE 19 Paragiopagurus umbonatus n. sp., holotype male (5.2 mm) mNHN-iU-2012-747, New Caledonia, Smib 4 stn DW 36, 530 m. A, second left pereopod, lateral view. B, dactyl of same, mesial view. C, third left pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of left fourth pereopod, lateral view. F, propodus and dactyl of left fifth pereopod, lateral view. Scale bars 2 mm for a-D; 0.5 mm for e, F. D F C E AB E, F A-D rafael lemaitre356 In P. umbonatus n. sp., the lateral and dorsomesial margins of the right palm are delimited by small spines (Figure 18A), whereas in P. boletifer and P. sculptochela, the spines on those margins are large (Figures 21C, 15C, D). The dorsal face of the palm is more densely covered with short, plumose setae in P. umbonatus n. sp. than in P. boletifer and P. sculptochela. At least in P. umbonatus n. sp. and P. boletifer, for which more abundant material exists, there is wide variation in density and strength of the tubercles and spines related to size, with smaller specimens generally having less, weaker tubercles or spines than larger specimens. However, the degree of variation is larger in P. boletifer than in P. umbonatus n. sp. In small specimens (sl < 3.0 mm) of P. boletifer the tubercles or spines are less numerous or weaker than in large specimens (Figure 21A, B, D). ETYMOLOGY — The specific name is derived from the Latin noun umbo, meaning boss, rounded protuberance, or knob, and refers to the characteristic tubercles found on the ventral surface of the right chela. Paragiopagurus boletifer (de Saint Laurent, 1972) Figures 1F, G, 20 , 21 Parapagurus boletifer de Saint laurent, 1972: 110, figs. 5, 20 (type locality: Japan, tosa bay). — miyake, 1978: 72; 1982: 120, pl. 40-fig. 4. — baba et al., 1986: 196, fig. 144. Sympagurus boletifer – lemaitre, 1989: 37; 1994: 382, figs 5, 6, 27a, b, 28b, c. — Poupin, 1996a: 20, pl. 9a; 1996b: 96. — Parin et al., 1997: 163. Paragiopagurus boletifer – lemaitre, 1996: 218. — Zhadan, 1997: 69. — Spiridonov & Zhadan, 1999: 627. — Davie, 2002: 90. — asakura et al., 2006: 212. — mclaughlin et al., 2007: 318, 2 unnumbered figs; 2010: 39. — Wang, 2008: 759. — Castro, 2011: 29. — tsang et al., 2011: 5, figs 2, 4 (phylogenetic trees). TYPE MATERIAL — Holotype, Japan, Tosa Bay, 250-300 m, 1963, male 8.0 mm (MNHN-Pg 2230). Paratype, Japan. Alba- tross: stn 3755, NW of Suno Saki, off Honshu Island, 52-77 fms (95-141 m), 19.05.1900, 1 ovigerous female 6.9 mm (USNM 168321). MATERIAL EXAMINED — the type material (see above). Comoros. beNtHeDi: stn F49, 12°55’ S, 44°57’ e, 300-450 m, 28.03.1977, 1 male 7.0 mm, 2 females 6.0, 6.3 mm (USNm). Madagascar. NW coast: 12°01.2’S, 49°25.8’e, 450 m, 27.03.1973, 1 male 6.0 mm (mNHN-iU-2012-748). Japan. tosa bay: 170 m, 12.12.1984, 1 male 6.4 mm (USNm). – 120 m, 05.02.1985, 2 ovigerous females 6.4, 7.2 mm (USNm 276068). Taiwan. taiWaN 2003: stn  CP  212, 24°34.60’N, 122°5.84’e, 223-260 m, 26.08.2003, 3 males 3.0-4.6mm, 3 females 4.2-4.8 mm (NtOU). Solomon Islands. SalOmON 1: stn CP 1831, 10°12.1’S, 161°19.2’e, 135-325 m, 05.10.2000, 1 female 5.9 mm (mNHN-Pg 6825). Vanuatu. mUSOrStOm 8: stn DW 965, 20°20.40’S, 169°51.36’e, 361-377 m, 21.09.1994, 1 male 3.0 mm (mNHN-Pg 6804). – stn DW 978, 19°22.62’S, 169°27.11’e, 413-408 m, 22.09.1994, 1 male 5.0 mm (mNHN-Pg 6805). – stn CP 980, 19°21.02’S, 169°25.22’e, 450 m, 22.09.1994, 1 ovigerous female 7.5 mm (mNHN-Pg 6806). – stn CP 982, 19°21.80’S, 169°26.47’e, 408 m, 23.09.1994, 1 female 3.6 mm (mNHN-Pg 6807). Tonga. bOrDaU 2: stn CP 1545, 21°17’S, 175°17’W, 444-447 m, 05.06.2000, 1 fe- male 3.2 mm (mNHN-iU-2008-15728). – stn DW 1586, 18°34’S, 173°55’W, 440-487 m, 13.06.2000, 2 males 7.5, 8.0 mm (mNHN-Pg 6411). – stn DW 1595, 19°03’S, 174°19’W, 523-806 m, 14.06.2000, 1 male 3.4 mm (mNHN-iU-2008-15721). – stn DW 1615, 23°03’S, 175°53’W, 482-504 m, 17.06.2000, 1 male 3.4 mm mNHN-iU-2012-1102, 1 ovigerous female 6.8 mm (mNHN-iU-2008-15709). New Caledonia. biOCal: stn DW 44, 22°47.30’S, 167°14.30’e, 440-450 m, 30.08.1985, 1 spm sex indet. 1.7 mm (USNm 1184064). – stn DW 82, 20°30.65’S, 166°50.30’e, 440 m, 06.09.1985, 1 male 5.7 mm (mNHN-iU-2012-749). – mUSOrStOm 4: stn DW 196, 18°55.00’S, 163°23.70’e, 460 m, 20.09.1985, 1 male 4.2 mm (mNHN-iU-2012-750). – stn DW 222, 22°57.60’S, 167°33.00’e, 440 m, 30.09.1985, 1 male 2.5 mm, 1 ovigerous female 3.9 mm (mNHN-iU-2012-751). – stn DW 223, 22°57.00’S, 167°30.00’e, 545-560 m, 30.09.1985, 1 male 5.6 mm (mNHN-iU-2012-752). – stn DW 230, 22°52.50’S, 167°11.80’e, 390-420 m, 30.09.1985, 1 juvenile 3.0 mm (mNHN-iU-2012-753). – mUSOrStOm 5: stn DW 300, 22°48.27’S, 159°23.94’e, 450 m, 11.10.1986, 1 ovigerous female 6.7 mm, 1 female 7.0 mm (mNHN-iU-2012-755). – stn DW 306, 22°07.66’S, 159°21.40’e, 375-415 m, 12.10.1986, 1 male 6.6 mm (USNm). – stn DC 379, 19°53.20’S, 158°39.50’e, 370-400 m, 20.10.1986, 1 female 2.6 mm (USNm 1184065). – Smib 2: stn DW 14, 22°52.60’S, 167°13.30’e, 405-444 m, 18.09.1986, 1 male 6.9 mm (mNHN-iU-2012-759). – CHalCal 2: stn DW 84, 23°23.80’S, 168°07.10’e, 170 m, 31.10.1986, 1 male 2.1 mm (USNm 1184066). – mUSOrS- tOm 6: stn DW 391, 20°47.35’S, 167°05.70’e, 390 m, 13.02.1989, 5 males 2.7, 6.2 mm, 5 females 3.4, 5.1 mm (ZmmU), 1 male 3.7 mm, 1 female 3.7 mm ParagioPagurus: wordldwide review and new species 357 (USNm 1184067). – stn DW 428, 20°23.54’S, 166°12.57’e, 420 m, 17.02.1989, 1 female 2.9 mm (USNm 1184068). – stn DW 460, 21°01.72’S, 167°31.45’e, 420 m, 20.02.1989, 1 male 6.3 mm (mNHN-iU-2012-754). – stn DW 471, 21°08.00’S, 167°54.10’e, 460 m, 22.02.1989, 1 female 7.0 mm (mNHN-Pg 6803). – stn DW 478, 21°08.96’S, 167°54.28’e, 400 m, 22.02.1989, 2 females 2.8, 6.3 mm (mNHN-iU-2012-756). – stn DW 485, 21°23.48’S, 167°59.33’e, 350 m, 23.02.1989, 1 female 4.6 mm (mNHN-iU-2012-757). – stn DW 487, 21°23.30’S, 167°46.40’e, 500 m, 23.02.1989, 1 male 3.5 mm (USNm 1184069). – Smib 4: stn DW 65, 22°55.30’S, 167°14.50’e, 400-420 m, 10.03.1989, 1 male 3.3 mm (mNHN-iU-2012-760). – Smib 8: stn DW 149, 24°54.94’S, 168°21.82’e, 508 m, 27.01.1993, 1 male 4.6 mm (mNHN-Pg 6808). – stn DW 152, 24°54.35’S, 168°22.23’e, 514-530 m, 27.01.1993, 1 male 4.4 mm, 1 female 3.0 mm (mNHN-Pg 6809). – stn DW 167, 23°38.13’S, 168°43.16’e, 430-452 m, 29.01.1993, 1 female 5.2 mm (mNHN-Pg 6810). – stn DW 179, 23°45.87’S, 168°16.95’e, 400-405 m, 30.01.1993, 1 male 2.3 mm (mNHN-Pg 6811), 2 females 4.5, 5.2 mm (mNHN-Pg 6812). – stn DW 187, 23°17.71’S, 168°05.61’e, 390-540 m, 31.01.1993, 1 males 2.6 mm, 1 female 6.4 mm (mNHN-Pg 6813). – stn DW 189, 23°17.60’S, 168°05.51’e, 400-402 m, 31.01.1993, 2 males 5.4, 5.9 mm, 1 female 4.2 mm (mNHN-Pg 6821). – stn DW 197, 22°51.27’S, 168°12.54’e, 414-436 m, 01.02.1993, 1 male 4.0 mm (mNHN-Pg 6814), 2 males 2.5, 3.1 mm (mNHN-Pg 6815). – stn DW 199, 22°51.63’S, 168°12.22’e, 408-410 m, 01.02.1993, 1 female 5.0 mm (mNHN-Pg 6822). – batHUS 2: stn DW 719, 22°47.57’S, 167°14.58’e, 444-455 m, 11.05.1993, 1 male 4.5 mm (mNHN-Pg 6783). – stn DW 723, 22°50.21’S, 167°26.84’e, 430-433 m, 11.05.1993, 1 male 3.7 mm (mNHN-Pg 6784). – stn DW 729, 22°52.42’S, 167°11.90’e, 349-400 m, 12.05.1993, 1 ovigerous female 5.0 mm, 1 female 3.0 mm (mNHN-Pg 6785). – stn DW 730, 23°02.56’S, 166°58.30’e, 350-400 m, 12.05.1993, 5 males 2.2-2.6 mm, 4 females 2.6-3.0 mm (mNHN-Pg 6786). – stn CP 736, 23°03.38’S, 166°58.96’e, 356-464 m, 13.05.1993, 1 ovigerous female 5.5 mm (mNHN-Pg 6787). – batHUS 3: stn CP 811, 23°41.42’S, 168°15.50’e, 383-408 m, 28.11.1993, 1 male 8.7 mm (mNHN-Pg 6788), 3 males 3.2-7.7 mm, 1 ovigerous female 4.7 mm, 5  emales 2.7-5.0 mm, 7 spms in housing (mNHN-Pg 6789). – stn CP 812, 23°43.38’S, 168°15.98’e, 391-440 m, 28.11.1993, 2 males 3.1, 8.4 mm, 1 fe- male 4.9 mm (mNHN-Pg 6790). – stn CP 813, 23°45.23’S, 168°16.57’e, 410-415 m, 28.11.1993, 1 male 4.4 mm (mNHN-Pg 6791). – stn CP 815, 23°47.00’S, 168°16.74’e, 460-470 m, 28.11.1993, 1 male 3.9 mm (mNHN-Pg 6792). – stn DW 817, 23°42.38’S, 168°15.51’e, 405-410 m, 28.11.1993, 5 females 2.4-5.0 mm (mNHN-Pg 6793). – stn DW 818, 23°43.89’S, 168°16.32’e, 394 m, 28.11.1993, 3 males 3.7-4.4 mm, 1 female 4.2 mm (mNHN-Pg 6794). – stn DW 829, 23°21.37’S, 168°01.84’e, 386-390 m, 29.11.1993, 1 male 6.2 mm, 1 ovigerous female 6.0 mm (mNHN-Pg 6795). – stn DW 830, 23°19.75’S, 168°01.45’e, 361-365 m, 29.11.1993, 1 male 6.2 mm (mNHN-Pg 6796). – stn DW 838, 23°00.81’S, 166°55.87’e, 400-402 m, 30.11.1993, 2 males 3.4, 4.5 mm, 2 females 2.7, 3.6 mm, 3 spms in housing (mNHN-Pg 6797). – stn CP 847, 23°02.53’S, 166°58.18’e, 405-411 m, 01.12.1993, 1 fe- male 5.0 mm (mNHN-Pg 6798). – HaliPrO 1: stn CP 877, 23°03.51’S, 166°59.20’e, 464-480 m, 31.03.1994, 1 male 3.2 mm (mNHN-Pg 6801). – batHUS 4: stn DW 931, 18°55.38’S, 163°24.36’e, 360-377 m, 07.08.1994, 1 ovigerous female 7.2 mm (mNHN-Pg 6799). – HaliCal 1: stn DW 02, 18°54’S, 163°24’e, 352-397 m, 23.11.1994, 1 male 3.0 mm, 1 female 4.2 mm (mNHN-Pg 6800). – litHiSt: stn DW 07, 24°55.4’S, 168°21.6’W, 504-530 m, 11.08.1999, 3 males 2.6-6.1 mm (mNHN-iU-2008-15740). – stn DW 13, 23°45.0’S, 168°16.7’W, 400 m, 12.08.1999, 2 males 4.0, 4.2 mm (mNHN- iU-2008-15743). – stn CP 14, 23°42.2’S, 168°15.8’W, 378-402 m, 12.08.1999, 1 ovigerous female 6.2 mm (mNHN-iU-2008-15739). – stn CP 15, 23°40.4’S, 168°15.0’W, 389-404 m, 12.08.1999, 1 male 4.2 mm, 5 ovigerous females 3.9-7.1 mm (mNHN-iU-2008-15742). – stn CP 16, 23°43.2’S, 168°16.2’W, 379-391 m, 12.08.1999, 2 males 6.3, 6.9 mm (mNHN-iU-2008-15730), 1 male 4.3 mm, 1 female 3.3 mm (mNHN-iU-2008-15731). – NOrFOlK 2: stn DW 2032, 23°39.14’S, 167°43.39’e, 420-450 m, 22.10.2003, 1 ovigerous female 5.0 mm mNHN-iU-2012-1103. – stn CP 2050, 23°42.17’S, 168°15.72’e, 377 m, 24.10.2003, 1 male 3.3 mm mNHN-iU-2012-1104. – stn CP 2114, 23°44.84’S, 168°16.88’e, 390-398 m, 31.10.2003, 1 male 3.8 mm mNHN- iU-2012-1105. – stn CH 2115, 23°44.90’S, 168°17.20’e, 377-401 m, 31.10.2003, 1 male 8.7 mm mNHN-iU-2012-1106. – stn DW 2136, 23°00.73’S, 168°22.68’e, 402-410 m, 03.11.2003, 1 male 5.1 mm mNHN-iU-2012-1107. – ebiSCO: stn DW 2496, 24°44.572’S, 159°42.846’e, 400-418 m, 10.06.2005, 1 male 3.2 mm, 1 ovigerous female 5.8 mm, 1 female 4.6 mm mNHN-iU-2012-1108. – stn DW 2504, 24°48.108’S, 159°46.151’e, 390-600 m, 10.07.2005, 2 males 1.7, 3.3 mm mNHN-iU-2012-1109. – stn CP 2505, 24°46.63’S, 159°41.607’e, 328-463 m, 10.07.2005, 1 female 5.1 mm mNHN-iU-2012-1110. – stn DW 2525, 22°46.650’S, 159°23.106’e, 408-410 m, 10.09.2005, 2 males 6.1, 8.1 mm, 1 ovigerous female 7.9 mm mNHN-iU-2012-1111. – stn  DW  2526, 22°47.492’S, 159°22.890’e, 330-340 m, 10.09.2005, 1 male 6.9 mm, 5 females 3.4, 6.8 mm mNHN-iU-2012-1112. – stn DW 2527, 22°45.380’S, 159°22.370’e, 330-340 m, 10.09.2005, 1 male 5.5 mm, 1 ovigerous female 7.0 mm, 1 female 5.6 mm mNHN-iU-2012-1113. – stn DW 2530, 22°46.911’S, 159°22.810’e, 338-343 m, 10.09.2005, 5 males 4.2-8.6 mm, 2 ovigerous females 6.3, 7.1 mm mNHN-iU-2012-1114. – stn DW 2534, 22°18.004’S, 159°28.136’e, 390-430 m, 10.10.2005, 4 males 3.7-8.0 mm, 4 ovigerous females 5.7-7.7 mm, 3 females 3.7-5.9 mm mNHN-iU-2012-1115. – stn DW 2535, 22°17.121’S, 159°27.919’e, 490 m, 10.10.2005, 1male 1.9 mm, 1 ovigerous female 5.7 mm mNHN-iU-2012-1116. – stn DW 2576, 20°19.774’S, 158°43.910’e, 390-394 m, 10.14.2005, 1 male 1.9 mm mNHN-iU-2012-1117. – stn DW 2606, 19°36.805’S, 158°42.062’e, 442-443 m, 10.18.2005, 2 males 2.7, 3.4 mm, 1 female 4.4 mm mNHN-iU-2012-1118. – stn DW 2608, 19°32.05’S, 158°39.736’e, 393-396 m, 10.19.2005, 1 female 5.1 mm mNHN-iU-2012-1119. – stn DW 2609, 19°33.040’S, 158°40.376’e, 431-436 m, 10.19.2005, 1 female 2.6 mm mNHN-iU-2012-1120. – stn DW 2612, 19°34.000’S, 158°40.316’e, 392 m, 10.19.2005, 2 females 6.0, 6.2 mm mNHN-iU-2012-1121. Loyalty Islands. No specific data, 3 males 4.8-5.5 mm, 2   emales 5.8, 5.9 mm, 1 juvenile 1.8 mm (mNHN-Pg 6780). Johnston Atoll. Hawaii Undersea research lab: dive 208, 16°41.5’N, 169°34.3’W, 381 m, 23.10.1983, 3 males 5.1-6.3 mm, 1 female 6.0 mm (bPbm S10549). For additional material examined, including specimens illus- trated in Figure 18, see lemaitre (1994 as Sympagurus boletifer, 1996). rafael lemaitre358 FIGURE 20 Paragiopagurus boletifer (de Saint laurent, 1972). A, G, ovigerous female (5.5 mm) mNHN-Pg 5137, French Polynesia, austral islands, stn 344. B-F, H, male (5.7 mm) mNHN-Pg 5137, French Polynesia, austral islands, stn 344. A, shield and cephalic appendages, dorsal view. B, right antennal peduncle, lateral view. C, left cheliped, dorsal view. D, left third pereopod, lateral view. E, dactyl of same, mesial view. F, G, propodus and dactyl of left fourth pereopod, lateral view. H, telson, dorsal view. Scale bars 3 mm for a, C-e; 1 mm for b, F-H. (adapted from lemaitre 1994). F-H D, E B A, C H F G E D BCA ParagioPagurus: wordldwide review and new species 359 FIGURE 21 Paragiopagurus boletifer (de Saint Laurent, 1972), right chela. A, sex indet. (1.7 mm) USNM 1184064, New Caledonia, BIOCAL stn DW 44, 440-450 m. B, juvenile male (2.1 mm) USNM 1184066, New Caledonia, CHALCAL 2 stn DW 84, 170 m. C-E, juvenile female (2.6 mm) USNM 1184065, New Caledonia, MUSORSTOM 5 stn DW 379, 370-400 m. F, ovigerous female (cl 7.2 mm) USNM 276068, Tosa Bay, Japan, 120 m. A, B, D, ventral view. C, dorsal view. E, lateral view. F, ventromesial view of area near base of fixed finger and ventromesial margin of chela. Scale bars 0.25 mm for A; 0.5 mm for B-E; 1 mm for F. F B-E A B D C FA E rafael lemaitre360 DISTRIBUTION — Western Indian Ocean: Comoros, Mozambique Channel, and Madagascar. Western, central and South Pacific: Japan, Taiwan, South China Sea, Solomon Islands, Vanuatu, New Caledonia, Australia, Johnston Atoll, French Polynesia, and Nazca and Sala-y-Gómez Ridges. Depth: 85 to 806 m. DIAGNOSIS — Gill lamellae undivided or weakly divided distally. Shield (Figure 20A) as long as broad; dorsal surface usually weakly calcified on half or more of surface; rostrum broadly rounded, with low dorsal ridge; anterior margins straight; lateral projections broadly subtriangular, terminating in spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, terminating in strong spine; corneas slightly dilated. Epistomial spine short, straight. Anten- nular peduncles exceeding distal margins of corneas by length of penultimate segment. Antennal peduncles (Figure 20B) at most reaching distal margins of cornea; third segment with strong ventromesial distal spine; second segment with dorsolateral distal angle produced, terminating in multifid spine (occasionally with additional small spine dorsally); acicle reaching distal margin of cornea, mesial margin of each armed with 11-14 spines; flagellum with setae arranged in series of long (4-8 articles in length) and short (about 1 article in length) setae about every 15-20 articles. Chelipeds strongly dissimilar. Right cheliped (Figure 21A-F) massive, operculate; chela about as broad as long, dorsal surface cove- red with numerous spines and dense plumose setae (especially on distal half and fingers); fingers curved ventromesially; ventral face of palm and fingers covered with numerous mushroom-like tubercles, or with less numerous mammilliform tubercles in small specimens (sl < 3.0 mm; see REMARKS); palm with dorsolateral margin well delimited by row of spines; carpus with numerous small tubercles or spines on dorsal surface. Left cheliped (Figure 20C) usually well calci- fied; palm with dorsomesial row of small tubercles; carpus with dorsodistal spine. Ambulatory (Figure 20D, E) legs with dactyl having ventromesial row of about 12 corneous spines, and dorsal and dorsomesial rows of long bristle-like setae; carpus with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) armed with marginal spine. Fourth pereopod (Figure 20F, G) with dactyl terminating in corneous claw (more slender and longer in females than in males); propodal rasp consisting of 1 row of ovate scales. Uropods and telson (Figure 20H) strongly asymmetrical, with weak transverse suture separating anterior and posterior lobes; posterior lobes separated by V-shaped median cleft, terminal margins armed with corneous spines (often strongly curved on left lobe). Male with paired first and second gonopods; first gonopod with concave distal lobe; second gonopod with distal segment nearly flat. Female with vestigial right second pleopod. COLOUR (Figure 1F, G) — Shield cream yellow tinged with orange. Ocular peduncles pale yellow or light pinkish-orange, darker on distal halves. Antennular peduncles pale yellow with flagella pale purple to pink or distal two segments and flagella blue. Antennal peduncles pale yellow or pinkish-orange; antennal flagella pale purple. Chela and carpus of right cheliped with dorsal and ventral surfaces orange, with white tubercles or spines; merus whitish with tinge of pale orange distally. Left cheliped and second to fifth pereopods uniformly pale purple or sometimes reddish. HABITAT — Gastropod shells. REMARKS (see also P. umbonatus n. sp. and P. sculptochela) — Although de Saint Laurent (1972) did not indicate an ety- mology for the species name boletifer, it is likely that the name was derived from boletus (Latin) or bolites (Greek), both meaning mushroom, and given in reference to the prominent mushroom-shaped tubercles found on the ventral side of the right chela that characterize this species. Lemaitre (1994) mistakenly included Hawaii in his overall distribution of Sympagurus boletifer (= Paragiopagurus boletifer); however, his report actually refers to specimens from Johnston Atoll, located more than 1000 km southeast of the Hawaiian Islands. Thus, this species has not been found yet in those islands. Examination of numerous specimens of Paragiopagurus boletifer during this study has shown that the shape and density of the armature of the ventral surface of the chela varies with size. In juvenile specimens (sl < 2.0 mm) the tubercles are rounded and low, well spaced, and few in number (Figure 21A); with increasing size (sl 2.0-3.0 mm) the tubercles increase ParagioPagurus: wordldwide review and new species 361 in number and prominence, although they are rounded or mammilliform (Figure 21B, D, E). In large specimens the ventral face of the chela has numerous, closely set mushroom-like tubercles (Figure 1G, 21F). Paragiopagurus sculptochela (Zarenkov, 1990) Figures 22, 15C, D Parapagurus sculptochela Zarenkov, 1990: 237, fig. 13 (type locality: Nazca and Sala-y-Gómez ridges). Paragiopagurus sculptochela – Zhadan, 1997: 69. TYPE MATERIAL — Holotype, Nazca and Sala-y-Gómez Ridges, R.V. “Professor Shtockman”, stn 1997, 27°07.2’S, 99°44.2’W, 330-350 m, 05.05.1987, male 7.5 mm, (ZMMU 4834, not examined). Paratypes, Nazca and Sala-y-Gómez Ridges. R.V. “Professor Shtockman”: stn 1986, 25°5.1’S, 97°28.5’W, 260-265 m, 04.05.1987, 1 female (ZMMU 4837, not examined). – stn 1997, 27°07.2’S, 99°44.2’W, 330-350 m, 05.05.1987,1 male 6.0 mm, 1 female 5.2 mm (ZMMU 4835). – stn 2028, 25°4.7’S, 97°30.2’W, 284-275 m, 09.05.1987, 1 male 6.2 mm (ZMMU 4836, not examined). For list of additional paratypes (not examined) see Zarenkov (1990). MATERIAL EXAMINED — None. DISTRIBUTION — So far known only from the Nazca and Sala-y-Gómez Ridges. Depth: 218 to 400 m. DIAGNOSIS — Gill lamellae undivided or weakly divided distally. Shield (Figure 20A) about as long as broad, weakly cal- cified medially and anteriorly. Rostrum broadly subtriangular, with short mid-dorsal ridge. Ocular peduncles more than half length of shield, corneas slightly dilated. Ocular acicles subtriangular, each terminating in strong spine. Antennular peduncles exceeding distal margin of corneas by slightly less than entire length of ultimate segment. Antennal peduncles (Figure 22B) at most reaching distal margin of corneas; fourth segment with strong dorsodistal spine; second segment with dorsodistal angle produced, terminating in strong multifid spine; first segment with small spine on lateral face; acicle not exceeding distal margin of cornea, terminating in strong spine, mesial margin armed with 6-8 strong spines; flagellum with long setae 3-5 flagellar articles in length, every 6-8 articles. Epistomial spine straight, simple. Chelipeds markedly dissimilar. Right cheliped (Figure 15C, D) massive, operculate, with moderately dense setae on dorsal surfaces of carpus and chela; fingers curved ventromesially; dactyl set at strongly oblique angle to longitudinal axis of palm; chela with dorsal surface covered with numerous spines; ventral face of palm and fingers covered with numerous strong tubercles, or thick, blunt spines; palm broader than long, with dorsolateral margin well delimited by row of blunt or sharp spines; carpus with numerous small tubercles or spines on dorsal surface. Left cheliped usually well calcified; palm at most with a few low, small tubercles on dorsomesial margin; carpus with dorsodistal spine. Ambulatory legs (Figure 22C-E) with dactyls each having ventromesial row of about 10-14 corneous spines, and dorsal and dorsomesial rows of long bristle- like setae; carpus with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) setose, armed with mar- ginal spine. Fourth pereopod (Figure 22F) with dactyl ending in sharp, corneous claw (longer in females than in males); propodal rasp consisting of 1 row of ovate scales. Telson and uropods strongly asymmetrical; telson (Figure 22G) without or at most with weak, transverse suture separating anterior and posterior lobes, posterior lobes separated by shallow, U-shaped median cleft, terminal margins armed with corneous spines (often strongly curved on left lobe). Male with paired first and second gonopods well developed; first gonopod (Figure 22H) with concave distal lobe; second gonopod (Figure 22I) with distal segment nearly flat and with row of short bristles on lateral margin medially, and long setae on distomesial face. Female with vestigial right second pleopod. COLOUR — Unknown. rafael lemaitre362 FIGURE 22 Paragiopagurus sculptochela (Zarenkov, 1990). A-G, paratype female (5.2 mm) ZmmU 4835, Nazca and Sala-y-Gómez ridges, r/V “Professor Shtockman” stn 1997, 300-350 m. H, I, male (6.0 mm) ZmmU 4835, Nazca and Sala-y-Gómez ridges, r/V “Professor Shtockman” stn 1997, 300-350 m. A, shield and cephalic appendages. B, right antennal peduncle, lateral view. C, second left pereopod, lateral view. D, third left pereopod, lateral view. E, dactyl of same, mesial view. F, propodus and dactyl of left fourth pereopod, lateral view. G, telson, dorsal view. H, male left first gonopod, mesial view. I, male left second gonopod, anterior view. Scale bars 1 mm for a; 2 mm for b, F-i; 0.5 mm for C-e. C-E A, B, F-I E H I G D F A CB ParagioPagurus: wordldwide review and new species 363 HABITAT — Not recorded by Zarenkov (1990), probably gastropod shells. REMARKS (see also P. umbonatus n. sp. and P. boletifer) — Based on a comparison of the types and only known speci- mens of Parapagurus scultptochela Zarenkov, 1990 with Lemaitre’s (1996) diagnosis of Paragiopagurus boletifer, Zhadan (1997) considered Zarenkov’s taxon a junior synonym of P. boletifer. At the time of Zhadan’s study, relatively few speci- mens of P. boletifer were available, and that synonymy appeared accurate given the overall similarity of these two taxa. Examination of a large number of specimens of P. boletifer available during this study from recent French expeditions has revealed the range of morphological variations in that species, particularly in the armature of the right cheliped, and show that Zarenkov’s taxon should be considered a valid species. Both P. sculptochela and P. boletifer are characterized by the dense, prominent armature on the ventral surfaces of the right chela. However, in P. sculptochela the ventral armature of the chela consists of strong, high semi-cylindrical tubercles or thick, strong spines (Figure 15C, D), whereas P. boletifer is armed with mushroom-like tubercles (adults) or low tubercles (specimens sl < 3.0 mm) (Figure 21E, F). In other albeit subtle features, the two species also differ, as in the spination of the antennal acicles (mesial margin with less spines, 6 to 8 in P. scultptochela, 11 to14 spines in P. boletifer), and the slenderness of the second and third pereopods (more slender in P. sculptochela than in P. boletifer). Paragiopagurus macrocerus (Forest, 1955) Figure 23 Parapagurus macrocerus Forest, 1955: 101, fig. 22, pl. 3-figs 1-7 (type locality: off banana, angola). — de Saint laurent, 1972: 116. — macpherson, 1983b: 12. Sympagurus macrocerus – lemaitre, 1989: 37; 1990: 233, figs 9, 10; 1994: 412. Paragiopagurus macrocerus – lemaitre, 1996: 207. — mclaughlin et al., 2010: 39. TYPE MATERIAL — Syntypes, West Africa, Angola, off Banana, stn AS 15, 05°50’S, 11°32’E, 215-220 m, 23.08.1948, 10 males, 8 females (MNHN, not examined). – stn AS 18, 05°46’S, 11°38’E, 145 m, 24.08.1948, 9 males, 1 female (MNHN, not examined), 1 male 5.3 mm (MNHN-IU-2008-15064). [see REMARKS]. MATERIAL EXAMINED — Gulf of Guinea. r.V. “John elliott Pillsbury”: stn 50, 04°58’N, 05°00’W, 128-192 m, 31.05.1964, 4 males 4.2-5.3 mm, 2 ovigerous females 4.0, 4.2 mm (USNm 1095968). Congo. stn aS 172, 05°15’S, 11°29’e, 225-240 m, 2.04.1948, 1 male 5.2 mm (mNHN-iU-2012-770). Angola. stn aS 18, 05°46’S, 11°38’e, 145 m, 24.08.1948, 1 male syntype 5.3 mm (mNHN-iU-2008-15064). – 11°50’S, 13°29.5’e, 190-200 m, 17.04.1968, 2 males 5.2-5.5 mm (mNHN-Pg 2688). Namibia. benguela i: stn P-5, 22°18’S, 13°07’e, 240 m, 10.11.1979, 1 male 7.7 mm (USNm). DISTRIBUTION — Eastern Atlantic, from Ivory coast in the Gulf of Guinea to Namibia. Depth: 128 to 280 m. DIAGNOSIS — Gill lamellae undivided distally. Shield (Figure 23A) broader than long; dorsal surface evenly calcified, often with low blister-like tubercles; rostrum broadly rounded, with low dorsal ridge often extending posteriorly to about mid-length of shield; lateral projections broadly rounded, unarmed. Ocular peduncles more than half length of shield; ocular acicles subtriangular, terminating in strong spine (rarely bifid on one side); corneas dilated. Maxillule with inter- nal lobe of endopod bearing long seta. Epistomial spine small, blunt. Antennular peduncles exceeding distal margin of corneas by slightly more than length of ultimate segment. Antennal peduncles (Figure 23B) exceeding distal margin of cornea by about 0.3 length of last (fifth) segment; fourth segment with small dorsolateral distal spine; second segment with dorsolateral distal angle produced, terminating in strong simple spine; acicle exceeding distal margin of cornea by 0.3-0.5 length of acicle, mesial margin with 7-10 spines on proximal 0.7; flagellum nearly naked, at most with few short se- tae less than half of 1 flagellar article in length. Chelipeds markedly dissimilar, surfaces moderately setose. Right cheliped rafael lemaitre364 FIGURE 23 Paragiopagurus macrocerus (Forest, 1955). A, B, D-K, male (5.3 mm) USNm 1095968 , Gulf of Guinea, r/V “John elliott Pillsbury” stn 50, 128-192 m. C, female (4.2 mm) USNm 1095968, Gulf of Guinea, r/V “John elliott Pillsbury” stn 50, 128-192 m. A, shield and cephalic appendages. B, right antennal peduncle, lateral view. C, D, right cheliped. E, left cheliped. F, second left pereopod, lateral view. G, dactyl of same, mesial view. H, propodus and dactyl of fourth left pereopod, lateral view. I, telson, dorsal view. J, male left first gonopod, mesial view. K, male left second gonopod, anterior view. Scale bars 4 mm for a, C-G; 2 mm for b; 1 mm for F, H, i, K; 0.5 mm for J. (adapted from lemaitre 1990). K J I H F, G D B A, C, E J KA D CE I H BFG ParagioPagurus: wordldwide review and new species 365 ( Figure 23C, D) elongate, proportions and armature affected by size and sex: carpus and chela usually broader in females than in males; fingers weakly curved ventrally; dactyl set at weakly oblique angle to longitudinal axis of palm; palm with well delimited dorsolateral margin, dorsal surface smooth except for medial, irregular rows of small spines at least proxi- mally; carpus with strong spine or tubercles dorsally. Left cheliped (Figure 23E) usually weakly calcified on merus and carpus dorsally; carpus with dorsal row of well-spaced spines, and dorsodistal spine. Ambulatory legs (Figure 23F, G) similar from right to left; dactyls unarmed on ventral and ventromesial margins, and with dorsal and dorsomesial rows of long setae; carpi each with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) setose, unarmed. Fourth pereopod (Figure 23H) with dactyl shorter than propodus; propodal rasp consisting of 1 row of ovate scales. Uro- pods and telson (Figure 23I) markedly asymmetrical; telson lacking transverse suture separating anterior and posterior lobes; posterior lobes separated by shallow U-shaped median cleft, terminal margins armed with strong, curved corne- ous spines (often more numerous in females than in males). Males with moderately developed first and second gonopods (Figure 23J, K); second gonopod with distal segment nearly flat, and long bristle-like setae distally. Females with vestigial right second pleopod. COLOUR — Unknown. HABITAT — Frequently found in gastropod shells covered with an unidentified actinian. REMARKS — In his description of this species, Forest (1955: 101) designated 30 syntypes from West Africa, Angola, off Banana: 10 males and nine females from stn AS 15, and 10 males and one female from stn AS 18. However, only one male syntype (5.3 mm) from stn AS 18 has been located (NMNH-IU-2008-15064). One additional non-type male (5.2 mm) from off Pointe-Noire, Congo, stn AS 172 was also located (NMNH-IU-2012-770). Paragiopagurus macrocerus is distinguishable from other congenerics primarily by the broad shield, dilated corneas, and the long antennal acicles that well exceed the distal margin of the corneas, and the shape of the dactyl and propodus of the fourth pereopod (dactyl is shorter than propodus and broad basally, and propodus is as high as long). Lemaitre (1990) noted that this species superficially resembles species of the parapagurid genus Strobopagurus, in having relatively short ocular peduncles and dilated corneas, a broad shield, long antennal acicles, and in the shape of the propodus and dactyl of the fourth pereopods. However, P. macrocerus differs from Strobopagurus species in important generic characters such as the shape of the dactyls of the second and third pereopods (curved in Paragiopagurus versus straight in Strobopagurus), and gonopod shape and development (second gonopods not twisted in Paragiopagurus versus twisted in Strobopagurus). Paragiopagurus insolitus Lemaitre, 1997 Figures 24, 25 Paragiopagurus insolitus lemaitre, 1997: 586, figs 7-10 (type locality : Kai islands, indonesia). — mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, Indonoesia, Kai islands, KARUBAR, stn DW 28, 05°31’S, 132°54’E, 448-467 m, 26.10.1991, male 2.2 mm (MNHN-IU-2008-15429). Paratypes, Indonoesia. KARUBAR: stn DW 28, 05°31’S, 132°54’E, 448-467 m, 26.10.1991, 3 male 1.3-1.8 mm, 1 ovigerous female1.7 mm (MNHN-IU-2008-15430), 1 male 2.0 mm, 1 ovigerous female 1.6 mm (USNM 276037). MATERIAL EXAMINED — the type material (see above). Wallis and Futuna Islands. mUSOrStOm 7: stn DW 494, 14°18.9’S, 178°03.0’W, 100-110 m, 10.05.1992, 1 male 1.6 mm (USNm). – stn DW 588, 12°17.3’S, 174°44.6’W, 490-500 m, 23.05.1992, 1 male 2.6 mm (USNm). – stn DW 598, 12°30.5’S, 174°18.4’W, 702-708 m, 24.05.1992, 1 male 2.2 mm (USNm). – stn DW 626, 11°53.6’S, 179°32.0’W, 597-600 m, 29.05.1992, 1 male 2.1 mm (USNm). rafael lemaitre366 FIGURE 24 Paragiopagurus insolitus lemaitre, 1997. A, B, E-H, holotype male (2.2 mm) mNHN-iU-2008-15429, indonesia, KarUbar stn DW 28, 448-467 m. C, D, paratype male (2.0 mm) USNm 276037, indonesia, KarUbar stn DW 28, 448-467 m. A, shield and cephalic appendages. B, right antennal peduncle, lateral view. C, epistome, ocular peduncles and acicles, anterior view. D, distal half of mandible, internal view. E, right cheliped, dorsal view. F, chela of same, mesial view. G, left cheliped, dorsal view. H, telson, dorsal view. Scale bars 0.5 mm for a, e-G; 0.25 mm for b-D; 0.5 mm for H. (adapted from lemaitre 1996). H D C B A, E-G G C H E DA FB ParagioPagurus: wordldwide review and new species 367 DISTRIBUTION — Western Pacific: Indonesia (Kai islands), and Wallis and Futuna. Depth: 100 to 708 m. DIAGNOSIS — Gill lamellae undivided distally. Shield (Figure 24A) slightly longer than broad; dorsal surface weakly cal- cified medially and elsewhere on scattered small areas; rostrum broadly rounded, with short, mid-dorsal ridge; late- ral projections subtriangular, terminating in small spine. Ocular peduncles more than half length of shield; corneas weakly dilated; ocular acicles subtriangular, terminating in long slender spine directed forwardly and reaching to nearly mid-length of ocular peduncles. Antennular peduncles exceeding distal margin of cornea by about 0.2 of penultimate segment. Antennal penduncles (Figure 24B) exceeding distal margin of corneas by about 0.2 length of fifth segment; fourth segment with strong dorsodistal spine; second segment with dorsolateral distal angle produced, terminating in strong simple spine; acicle terminating in strong spine, mesial margin armed with row of 10-13 spines; flagellum with short setae 1 flagellar article in length or less. Mandible (Figure 24D) with incisor process consisting of several irregu- larly-shaped teeth; second (Figure 24A) and third maxillipeds each with slender, long exopod about 10 times as long as broad. Epistome (Figure 24C) with median region strongly produced anteriorly (somewhat pyramid-shaped with ven- tral face concave); terminating in small, often inconspicuous blunt spine. Chelipeds markedly dissimilar. Right cheliped (Figure 24E, F) sparsely setose, with some iridescence on surfaces of merus, carpus and chela; chela with dorsal surface smooth or with few scattered small spines or tubercles; palm with dorsolateral margin at most weakly delimited by row of FIGURE 25 Paragiopagurus insolitus lemaitre, 1997, paratype male (2.0 mm) USNm 276037, indonesia, KarUbar stn DW 28, 448-467 m. A, left second maxilliped, internal view. B, right second pereopod, lateral view. C, dactyl of same, mesial view. D, propodus and dactyl of left fourth pereopod, lateral view. E, dactyl and distal end of propodus of same, lateral view. F, left second gonopod, anterior view. Scales bar 0.25 mm for a, D, F; 0.5 mm for b, C; 0.2 mm for e. (adapted from lemaitre 1996). F E B-D A AFE B CD rafael lemaitre368 small spines or tubercles, mesial face rounded; dactyl set at moderately oblique angle to longitudinal axis of palm; carpus with dorsomesial margin delimited in distal half by row of small spines. Left cheliped (Figure 24G) usually weakly calci- fied laterally on carpus and merus; palm unarmed, with scattered setae; carpus with small dorsodistal spine. Ambulatory legs (Figure 25B, C) with dactyls each having dorsal and distal dorsomesial rows of long setae, and 5-8 minute spinules on ventromesial margin. Anterior lobe of sternite XII (of third pereopods) subsemicircular, unarmed, setose. Fourth pereo- pod (Figure 25D, E) with dactyl terminating in blunt corneous claw entirely masked by long, stiff plumose setae arising near base of claw; propodal rasp consisting of 1 row of ovate scales. Uropods and telson markedly asymmetrical. Telson (Figure 24H) lacking transverse suture; posterior lobes separated by wide and shallow unarmed U-shaped cleft, terminal margins of lobes armed with long, often strongly curved corneous spines. Males lacking first gonopods; with paired second gonopods (Figure 25F) having distal segments flat, long setae distally and on anterodistal face, with or without rudimentary exopod. Females lacking vestigial second right pleopod. COLOUR — Unknown. HABITAT — Unknown, probably gastropod shells. REMARKS — As summarized in Lemaitre (1997), P. insolitus is a singularly distinctive parapagurid species. Unique cha- racters among the parapagurids in P. insolitus include the incisor process of the mandible that consists of several irregu- larly-shaped teeth; the long and slender exopods (about 10 times as long as broad) of the second and third maxillipeds; the pyramidal shape of the epistome; and the specialized setal arrangement on the dactyl of the fourth pereopod, with the distal end of the claw almost entirely masked by long plumose setae (Figure 25D, E). The long ocular long acicles of P. insolitus were also noted by Lemaitre (1997) as a differentiating feature; however, similar long acicles have been found in P. orthotenes n. sp. Paragiopagurus orthotenes n. sp. Figures 26-29 Paragiopagurus sp. 1 – mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, New Caledonia, SMIB 3, stn CP 4, 24°54.00’S, 168°21.50’E, 530 m, 20.05.1987, male 3.4 mm (MNHN-IU-2012-717). Paratypes: Solomon Islands. SALOMON 1: stn DW 1770, 8°19.6’S, 160°38.7’E, 453-542 m, 28.09.2000, 1 ovigerous female 4.3 mm (MNHN-IU-2008-15876). – stn CP 1859, 9°32.6’S, 160°37.3’E, 283-305 m, 07.10.2000, 1 female 2.4 mm (MNHN-IU-2008-15877). Tonga. BORDAU 2: stn DW 1584, 18°36’S, 174°01’W, 439 m, 13.06.2000, 1 male 3.1 mm, 1 ovigerous female 3.4 mm (MNHN-IU-2008-15875). New Caledonia. BIOCAL: stn DW 66, 24°55.43’S, 168°21.67’E, 505- 515 m, 03.09.1985, 6 males 2.1-4.2 mm, 3 ovigerous female 2.4-3.2 mm (MNHN-IU-2012-708), 2 males 1.3, 1.4 mm, 2 fe- males 1.3, 3.1 mm, 2 juveniles 0.8, 1.4 mm (USNM 1184049). – stn CP 67, 24°55.44’S, 168°21.55’E, 500-510 m, 03.09.1985, 1 ovigerous female 2.4 mm (MNHN-IU-2012-707). – MUSORSTOM 4: stn DW 230, 22°52.50’S, 167°11.80’E, 390-420 m, 30.09.1985, 1 male 2.2 mm (MNHN-IU-2012-711). – MUSORSTOM 5: stn DW 301, 22°06.90’S, 159°24.60’E, 487-610 m, 12.10.1986, 2 males 1.7, 1.9 mm, 2 female 1.8, 2.2 mm (MNHN-IU-2012-713). – stn DC 379, 19°53.20’S, 158°39.50’E, 370- 400 m, 20.10.1986, 1 female 2.5 mm (MNHN-IU-2012-712). – CHALCAL 2: stn CC 2, 24°55.48’S, 168°21.29’E, 500-610 m, 28.10.1986, 1 male 2.1 mm, 1 ovigerous female 3.2 mm (MNHN-IU-2012-709). – stn DW 72, 24°54.50’S, 168°22.30’E, 527 m, 28.10.1986, 8 males 2.2-3.2 mm, 7 females 2.1-2.9 mm (USNM 1184050), 6 males 1.4-3.1 mm, 1 ovigerous female 2.2 mm (MNHN-IU-2012-710). – SMIB 3: stn DW 3, 24°55.00’S, 168°21.70’E, 513 m, 20.05.1987, 7 males 3.0-3.9 mm, 4 ovigerous females 2.7-4.3 mm, 7 females 2.2-3.1 mm (USNM 1184051). – stn DW 5, 24°54.90’S, 168°21.60’E, 502 m, 21.05.1987, 3 males 2.2-3.3 mm, 1 ovigerous female 2.5 mm (MNHN-IU-2012-718). – stn DW 6, 24°56.40’S, 168°21.20’E, ParagioPagurus: wordldwide review and new species 369 505 m, 21.05.1987, 4 males 2.8-3.1 mm (MNHN-IU-2012-719) . – stn DW 7, 24°54.60’S, 168°21.30’E, 505 m, 21.05.1987, 5 males 2.8-3.3 mm, 2 females 2.4, 3.0 mm (MNHN-IU-2012-720). – stn DW 8, 24°45.20’S, 168°08.00’E, 233 m, 21.05.1987, 1 male 2.9 mm (MNHN-IU-2012-721). – MUSORSTOM 6: stn DW 428, 20°23.54’S, 166°12.57’E, 420 m, 17.02.1989, 1 female 2.1 mm (MNHN-IU-2012-714). – stn DW 457, 21°00.42’E, 167°28.71’E, 353 m, 20.02.1989, 1 female 2.2 mm (MNHN-IU-2012-715). – stn DW 460, 21°01.72’S, 167°31.45’E, 420 m, 20.02.1989, 2 males 1.8, 3.6 mm, 1 female 2.7 mm (USNM 1184052). – stn DW 487, 21°23.30’S, 167°46.40’E, 500 m, 23.02.1989, 1  male 2.6  mm (MNHN-IU-2012-716). – SMIB 4: stn DW 55, 23°21.40’S, 168°04.50’E, 260 m, 09.03.1989, 1 female 3.0 mm (MNHN-IU-2012-722). – SMIB 5: stn DW 91, 22°18.40’S, 168°41.10’E, 340 m, 11.09.1989, 1 female 1.6 mm (MNHN-IU-2012-723). – BERYX 11: stn DW 38, 23°37.53’S, 167°39.42’E, 550-690 m, 19.10.1992, 2 males 2.6, 3.4 mm (MNHN-IU-2008-15849). – SMIB 8: stn DW 147, 24°54.90’S, 168°21.85’E, 508-532 m, 27.01.1993, 3 males 3.1-3.2 mm (MNHN-IU-2008-15850). – stn DW 148, 24°55.15’S, 168°21.62’E, 510 m, 27.01.1993, 1 male 2.7 mm (MNHN-IU-2008-15851). – stn DW 149, 24°54.94’S, 168°21.82’E, 508- 510 m, 27.01.1993, 2 males 1.6, 2.4 mm, 1 female 2.9 mm (MNHN-IU-2008-15852). – stn DW 150, 24°54.30’S, 168°22.23’E, 519-530 m, 27.01.1993, 1 male 3.7 mm (MNHN-IU-2008-15853), 5 males 2.5-3.2 mm, 3 females 2.7-3.0 mm (MNHN- IU-2008-15854). – stn DW 152, 24°54.35’S, 168°22.23’E, 514-530 m, 27.01.1993, 8 males 2.1-4.0 mm, 4 females 2.7-2.8 m (MNHN-IU-2008-15855), 1 male 2.5 mm (MNHN-IU-2008-15856), 3 males 2.5-3.3 mm, 5 females 2.2-2.5 mm (MNHN- IU-2008-15857), 2 males 2.2, 2.7 mm (MNHN-IU-2008-15858). – stn DW 160, 24°46.12’S, 168°08.12’E, 280-282  m, 28.01.1993, 1 female 1.6 mm (MNHN-IU-2008-15859). – stn DW 165, 24°47.64’S, 168°09.61’E, 372-600 m, 28.01.1993, 1 male 1.8 mm (MNHN-IU-2008-15860). – stn DW 198, 22°51.59’S, 167°12.44’E, 414-430 m, 01.02.1993, 1 female 2.4 mm (MNHN-IU-2008-15861). – BATHUS 2: stn DW 723, 22°50.21’S, 167°26.84’E, 430-433 m, 11.05.1993, 1 male 2.0 mm (MNHN-IU-2008-15839). – stn DW 729, 22°52.42’S, 167°11.90’E, 349-400 m, 12.05.1993, 1  ovigerous female 2.7 mm (MNHN-IU-2008-15840). – stn DW 730, 23°02.56’S, 166°58.30’E, 350-400 m, 12.05.1993, 2 males 1.9, 2.4 mm, 1 female 1.9 mm (MNHN-IU-2008-15841). – BATHUS 3: stn CP 811, 23°41.42’S, 168°15.50’E, 383-408 m, 28.11.1993, 6 males 2.2- 3.1 mm, 1 female 3.1 mm (MNHN-IU-2008-15842). – stn CP 813, 23°45.23’S, 168°16.57’E, 410-415 m, 28.11.1993, 1 male 3.2 mm (MNHN-IU-2008-15843). – stn DW 817, 23°42.38’S, 168°15.5’E, 405-410 m, 28.11.1993, 1 male 1.9 mm, 2 females 2.3, 2.6 mm (MNHN-IU-2008-15844). – stn DW 818, 23°43.89’S, 168°16.32’E, 394 m, 28.11.1993, 6 males 2.8-3.5 mm, 6 females 1.6-3.1 mm (MNHN-IU-2008-15845). – stn DW 829, 23°21.37’S, 168°01.84’E, 386-390 m, 29.11.1993, 8 males 1.7-3.2 mm, 3 females 2.0-2.9 mm, 12 spms in housing (MNHN-IU-2008-15846). – stn DW 838, 23°00.81’S, 166°55.87’E, 400-402 m, 30.11.1993, 1 female 2.3 mm (MNHN-IU-2008-15847). – BATHUS 4: stn DW 940, 18°59.53’S, 163°25.90’E, 305 m, 08.08.1994, 1 ovigerous female 1.3 mm (MNHN-IU-2008-15848). – SMIB 10: stn DW 203, 24°56’S, 168°22’E, 508- 502 m, 10.01.1995, 1 male 2.7 mm (MNHN-IU-2008-15863). – stn DW 205, 24°57’S, 168°21’E, 517-559 m, 10.01.1995, 25 males 2.0-3.7 mm, 2 ovigerous females 2.2, 2.7 mm, 8 females 2.0-2.8 mm (MNHN-IU-2008-15862). – stn DW 207, 24°57’S, 168°21’E, 508-553 m, 10.01.1995, 8 males 2.7-3.7 mm (MNHN-IU-2008-15864). – stn DW 208, 24°49’S, 168°09’E, 270 m, 10.01.1995, 1 male 2.5 mm (MNHN-IU-2008-1586), 1 male 2.7 mm (MNHN-IU-2008-15866). – stn DW 215, 24°56’S, 168°21’E, 508-553 m, 11.01.1995, 1 ovigerous female 2.7 mm (MNHN-IU-2008-15867). – stn DW 216, 24°56’S, 168°21’E, 512-541 m, 11.01.1995, 10 males 2.4-4.0 mm, 1 female 3.1 mm (MNHN-IU-2008-15868). – LITHIST: stn DW 07, 24°55.4’S, 168°21.6’W, 504-530 m, 11.08.1999, 2 males 2.8, 2.9 mm (MNHN-IU-2008-15869). – stn CP 08, 24°54.2’S, 168°21.3’W, 540 m, 11.08.1999, 1 male 2.9 mm, 1 ovigerous female 2.6 mm (MNHN-IU-2008-15870). – stn DW 11, 24°46.7’S, 168°08.3’W, 254-283 m, 11.08.1999, 4 males 4.0-4.1 mm, 2 females 1.7, 1.8 mm (MNHN-IU-2008-15871). – stn DW 13, 23°45.0’S, 168°16.7’W, 400 m, 12.08.1999, 1 ovigerous female 2.1 mm (MNHN-IU-2008-15872). – stn CP 15, 23°40.4’S, 168°15.0’W, 389-404 m, 12.08.1999, 2 ovigerous females 2.9, 3.5 mm, 1 female 4.0 mm (MNHN-IU-2008-15873). – NOR- FOLK 1: stn DW 1692, 24°56’S, 168°21’E, 507-967 m, 23.06.2001, 1 male 2.8 mm (MNHN-IU-2008-15874). – NORFOLK 2: stn DW 2081, 25°54.40’S, 168°21.64’E, 500-505 m, 28.10.2003, 2 males 1.8, 2.1 mm (MNHN-IU-2008-1607). – stn DW 2108, 23°46.52’S, 168°17.12’E, 403-440 m, 31.10.2003, 1 male 2.4 mm (MNHN-IU-2008-16074). – stn DW 2137, 23°01.18’S, 168°22.70’E, 547-560 m, 03.11.2003, 1 male 1.7 mm (MNHN-IU-2008-16075). – EBISCO: stn DW 2525, 22°46.650’S, 159°23.106’E, 408-410 m, 09.10.2005, 1 male 1.6 mm, 5 males 1.4-1.6 mm MNHN-IU-2012-1122. – stn DW 2565, 20°22.31’S, 158°40.51’E, 414-419 m, 13.10.2005, 1 male 2.3 mm MNHN-IU-2012-1123. – stn DW 2578, 20°19.785’S, 158°39.714’E, rafael lemaitre370 14.10.2005, 5 males 2.0-2.6 mm, 2 females 1.8, 2.4 mm MNHN-IU-2012-1124. – stn DW 2606, 19°36.805’S, 158°42.062’E, 442-443 m, 18.10.2005, 2 males 2.4, 2.6 mm, 1 ovigerous female 2.4 mmm, 3 females 2.3-2.5 mm MNHN-IU-2012-1125. – stn DW 2610, 19°33.147’S, 158°40.618’E, 486-494 m, 19.10.2005, 4 males 2.7, 2.8 mm MNHN-IU-2012-1126. – stn DW 2613, 19°36.516’S, 158°42.131’E, 519-522 m, 19.10.2005, 2 males 2.5, 3.4 mm MNHN-IU-2012-1127. – stn DW 2626, 21°04.488’S, 160°48.791’E, 728-739 m, 21.10.2005, 1 male 2.4 mm MNHN-IU-2012-1128. FIGURE 26 Paragiopagurus orthotenes n. sp., holotype male (3.4 mm) mNHN-iU-2012-717, New Caledonia, Smib 3 stn CP 4, 530 m. A, shield and cephalic appendages. B, right antennal peduncle and anterior portion of branchiostegite, lateral view. C, anterior lobe of sternite Xii (of third pereopods), ventral view. D, telson, dorsal view. E, second left pleopod (gonopod) and pleura, lateral view. Scale bars 1 mm for a; 0.25 mm for b-e. D, E B, C A ED C AB ParagioPagurus: wordldwide review and new species 371 FIGURE 27 Paragiopagurus orthotenes n. sp., left mouthparts, internal view. A-F, paratype male (3.1 mm) mNHN-iU-2012-719, New Caledonia, Smib 3 stn DW 6, 505 m. G, holotype male (3.4 mm) mNHN-iU-2012-717, New Caledonia, Smib 3 stn CP 4, 530 m. A, mandible. B, maxillule. C, maxilla. D, first maxilliped. E, second maxilliped. F, third maxilliped. G, basis and ischium of third maxilliped with crista dentata. Scale bars 0.25 mm. E, F A-D, G GFD B ECA rafael lemaitre372 MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — Western Pacific: Solomon Islands, Tonga, and New Caledonia region. Depth: 233 to 967 m. DESCRIPTION — Eleven pairs of biserial gills, with lamellae undivided distally. Shield (Figure 26A) approximately as broad as long; dorsal surface weakly calcified along midline, just posteriorly to rostrum, and anterior and anterolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections bluntly subtriangular. Anterolateral margins sloping. Ventrolateral margins with small spine on one or both sides. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, with longitudinal row of short setae on dorsal surface. Corneas slightly dilated. Ocular acicles subtriangular, about 0.6 times length of ocular peduncles, each terminating in long and strong, simple spine reaching to midlength of ocular peduncle; mesial margins nearly parallel; separated basally by about half width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment; ventral flagellum with about 6 articles. Ultimate segment about 2.0 times as long as penultimate, with scattered setae dorsally. Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 3 or 4 spines, and strong spine proximally. Antennal peduncles (Figure 26B) slightly exceeding distal margins of corneas. Fifth segment unarmed, with few setae on lateral and mesial margins. Fourth segment with strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong spine reaching beyond midpoint of acicle; mesial margin with spine on dorsolateral distal angle. First segment with lateral surface unarmed; ventromesial angle produced, with row of 4 or 5 spines laterally. Antennal acicles nearly straight (dorsal view), reaching to distal margins of corneas, each terminating in strong spine; mesial margin setose, armed with row of 9 to 11 strong spines. Flagellum reaching to about distal end of right cheliped; with short (< 1 article in length) setae throughout, and at least distally also with long setae (4-8 articles in length) every 4 to 12 articles. Mandible (Figure 27A) with 3-segmented palp. Maxillule (Figure 27B) with external lobe of endopod moderately developed, internal lobe with long seta. Maxilla (Figure 27C) with endopod slightly exceeding distal margin of scaphognathite. First maxilliped (Figure 27D) with endopod exceeding distal margin of exopod. Second maxilliped (Figure 27E) without distinguishing characters. Third maxilliped (Figure 27F, G) with distal 3 segments about twice to 2.5 as long as wide; crista dentata with about 11-14 corneous-tipped teeth, proximal 2 or 3 teeth distinctly larger than others; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine short, blunt or sharp. Chelipeds markedly dissimilar. Right cheliped (Figure 28A-D) massive, with chela, carpus and merus sparsely setose; chela somewhat dorsoventrally flattened, operculate; proportions of palm and carpus affected by sexual dimorphism, varying from approximately as long as broad to slightly longer than broad in large males (sl > 3.0 mm) or slightly broader than long in females. Fingers curving ventromesially, terminating in small, usually blunt corneous claw; cutting edges with 3 or 4 large calcareous teeth. Dactyl subequal in length to palm, set at strongly oblique angle to longitudinal axis of palm when closed; dorsal surface with scattered small tubercles or spines; mesial margin well defined by row of spines (most distal corneous-tipped); ventromesial face concave, ventral face smooth. Fixed finger broad at base, dorsal surface with scattered small tubercles or spines. Palm with dorsal surface covered with well spaced small tubercles or spines; lateral margin well delimited by row of spines; dorsal surface with scattered well-spaced, small sharp or blunt spines; dorsomesial margin delimited by row of small spines; mesial face rounded, with well-spaced small spines or tubercles; dorsal surface covered with well-spaced, small spines or tubercles; ventral surface smooth. Carpus covered with small spines or tubercles on dorsal and dorsolateral surfaces; ventrolateral margin well defined by row of spines; dorsodistal and ventromesial distal margins each with row of small spines. Merus with lateral and ventral surfaces nearly smooth; ventromesial distal margin with row of spines. Ischium unarmed except for setae on dorsal margin and ventral surface; coxa with 2 or 3 spines on distolateral and distomesial margins, and ventromesial row of long setae. ParagioPagurus: wordldwide review and new species 373 FIGURE 28 Paragiopagurus orthotenes n. sp. A-C, E, holotype male (3.4 mm) mNHN-iU-2012-717, New Caledonia, Smib 3 stn CP 4, 530 m. D, paratype ovigerous female (4.3 mm) USNm 1184051, New Caledonia, Smib 3 stn DW 3, 513 m. A, D, right cheliped, dorsal view. B, chela of same, mesial view. C, same chela, lateral view. E, left cheliped, dorsal view. Scale bar 1 mm. ADE BC rafael lemaitre374 FIGURE 29 Paragiopagurus orthotenes n. sp., holotype male (3.4 mm) mNHN-iU-2012-717, New Caledonia, Smib 3 stn CP 4, 530 m. A, left second pereopod, lateral view. B, dactyl of same, mesial view. C, left third pereopod, lateral view. D, dactyl of same, mesial view. E, merus of left third pereopod, mesial view. F, propodus and dactyl of left fourth pereopod, lateral view. G, propodus and dactyl of left fifth pereopod, lateral view. Scale bars 1 mm for a-e; 0.25 mm for F, G. F, G A-E C F G E D BA ParagioPagurus: wordldwide review and new species 375 Left cheliped (Figure 28E) well calcified. Fingers terminating in sharp corneous claws, often gaping when closed; dorsal and ventral surfaces unarmed except for tufts of setae; cutting edges of dactyl and fixed finger with row of closely-set small corneous teeth. Dactyl longer than palm. Palm with scattered setae and tubercles on dorsal surface, with dorsomesial row of small blunt spines; dorsolateral and ventral faces smooth except for scattered setae. Carpus with bristle-like setae and small blunt spines on dorsal margin, and dorsodistal spine; mesial and ventral faces smooth. Merus unarmed except for dorsal and ventral setae, and 2 or 3 spines on ventrolateral distal margin. Ischium unarmed except for dorsal and ventral setae. Coxa with spine on distolateral and distomesial margin, and ventromesial row of long setae. Ambulatory legs (Figure 29A-E) similar from right to left, at most reaching to about tip of extended right cheliped, or in large males (sl > 3.0 mm) frequently reaching only to about midlength of right chela. Dactyl long, approximately 1.8 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, and ventromesial row of 5-14 corneous spines; lateral and mesial faces with shallow, longitudinal sulcus on proximal half (deeper on mesial face). Propodus with setae on dorsal margin, and sparse setae on ventrolateral margin. Carpus with small dorsodistal spine, and setae on dorsal margin. Merus with setae on dorsal and ventral margins; merus of third pereopods with row of few small spines on dorsal margin (more visible in mesial view, Figure 29E). Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods) usually with strong sharp or blunt subterminal spine (Figure 26C). Fourth pereopod (Figure 29F) subchelate. Dactyl elongate, broadly curved, terminating in sharp corneous claw; with ventrolateral row of small corneous spinules. Propodus longer than wide, rasp consisting of 1 row of rounded scales at least distally. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 29G) chelate. Propodal rasp extending to mid-length of segment. Uropods and telson (Figure 26D) markedly asymmetrical. Telson lacking lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 broadly rounded projections by wide, shallow cleft; rounded projections each armed with row of corneous spines, those on left projection with 1-3 distinctly stronger spines. Male lacking first gonopods; with rudimentary, bud-like, unpaired left second gonopod (Figure 24E). Female lacking rudimentary second right pleopod. COLOUR — Unknown. HABITAT — Gastropod shells. REMARKS — This new species is similar to P. insolitus in the shape and size of the ocular acicles. In both species the acicles terminate in a long, slender spine directed straight forward, and extend to mid-length or more of the ocular peduncles. The males of both species are also similar in lacking first gonopods. However, that is where the similarity ends. As noted under P. insolitus, that species is quite unique in a number of characters. Aside from those characters, P. orthotenes n. sp. differs from P. insolitus in having an operculate shaped right chela, the shape and armature of the telson (less asymme- trical and more strongly armed in P. orthotenes n. sp.), and in males, the presence of only an unpaired, rudimentary left second gonopod. The proportions of the segments of the right cheliped in P. orthotenes n. sp. vary according to sex and size. In large females (sl > 3.5 mm), the palm and carpus, in particular, are distinctly broader than in males of comparable size. In large females, the palm can be up to 1.7 times as broad as long, and the carpus about as broad as long (Figure 28D); whereas the chela and carpus in males is about 1.2 times as broad as long, and the carpus is distinctly longer (up to 1.3 times) than broad (Figure 28A). Also, the ventromesial face of the palm in large females is more expanded distally than in males. ETYMOLOGY — The specific name is from the Greek orthotenes, stretched out, straight, and refers to the shape of the ocular acicles. rafael lemaitre376 Paragiopagurus oxychelos n. sp. Figures 1H , 30-33 Paragiopagurus sp. 2 – mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, New Caledonia, SMIB 5, stn DW 73, 23°41.40’S, 168°00.60’E, 240 m, 07.09.1989, male 2.8 mm (MNHN-IU-2012-733). Paratypes: Taiwan. TAIWAN 2000: stn CP 212, 24°34.60’N, 122°05.84’E, 223-260 m, 26.08.2003, 1 male 3.5 mm (NTOU A01347). New Caledonia. N.O. “Vauban”: stn DR15, 22°49’S, 167°12’E, 380  m, 10.04.1978, 1 female 2.4 mm (MNHN-IU-2012-732). – MUSORSTOM 4: stn DW 181, 18°57.20’S, 163°22.40’E, 355 m, 18.09.1985, 1 male 1.4  mm, 1 female 2.0 mm (MNHN-IU-2012-726). – stn DW 212, 22°47.40’S, 167°10.50’E, 375- 380 m, 28.09.1985, 2 males 2.2, 2.7 mm, 1 ovigerous female 2.2 mm, 1 female 3.4 mm (USNM 1184053). – stn DW 230, 22°52.50’S, 167°11.80’E, 390-420 m, 30.09.1985, 1 female 2.1 mm (MNHN-IU-2012-727). – stn DW 234, 22°15.50’S, 167°08.30’E, 350-365 m, 02.10.1985, 1 male 1.4 mm, 1 ovigerous female 2.2 mm (MNHN-IU-2012-728), 1 female 2.2 mm (USNM 1184054). – MUSORSTOM 5: stn DW 301, 22°06.90’S, 159°24.60’E, 487-610 m, 12.10.1986, 1 ovige- rous female 1.7 mm (MNHN-IU-2012-730), 3 males 1.8-3.3 mm, 2 ovigerous females 2.4, 2.8 mm (USNM 1184055). – stn DW 378, 19°53.74’S, 158°38.30’E, 355 m, 20.10.1986, 1 female 2.7 mm (MNHN-IU-2012-729). – CHALCAL 2: stn DW 81, 23°19.60’S, 168°03.40’E, 311  m, 31.10.1986, 1 male 1.4 mm (MNHN-IU-2012-725). – MUSORSTOM 6: stn DW 478, 21°08.96’S, 167°54.28’E, 400 m, 22.02.1989, 1 male 2.2 mm (MNHN-IU-2012-731).– Banc Alis: 250 m, 09.1989, 1 male 3.9 mm, 1 female 2.8 mm (MNHN-IU-2012-724). –SMIB 5: stn DW 73, 23°41.40’S, 168°00.60’E, 240 m, 07.09.1989, 1 male 2.7 mm (MNHN-IU-2012-734). – stn DW 91, 22°18.40’S, 168°41.10’E, 340 m, 11.09.1989, 2 males 1.7, 2.0 mm (MNHN-IU-2012-735). – SMIB 6: stn DW 125, 18°57.40’S, 163°23.50’E, 350 m, 03.03.1990, 3 males 2.2-2.7 mm (MNHN-IU-2012-736). – SMIB 8: stn DW 174, 23°39.94’S, 168°00.55’E, 235-240 m, 29.01.1993, 1 male 2.4 mm (MNHN- IU-2008-15829). – stn DW 182, 23°19.28’S, 168°04.82’E, 314-330 m, 31.01.1993, 1 male 1.3 mm (MNHN-IU-2008-15830). – stn DW 189, 23°17.60’S, 168°05.51’E, 400-402 m, 31.01.1993, 2 males 1.9, 2.1 mm (MNHN-IU-2008-15831). – stn DW 197, 22°51.27’S, 168°12.54’E, 414-436 m, 01.02.1993, 1 male 2.5 mm (MNHN-IU-2008-15832). – stn DW 198, 22°51.59’S, 167°12.44’E, 414-430 m, 01.02.1993, 1 male 2.6 mm, 1 ovigerous female 2.7 mm, 1 female 2.7 mm (MNHN- IU-2008-15833). – BATHUS 2: stn DW 729, 22°52.42’S, 167°11.90’E, 349-400 m, 12.05.1993, 3 males 2.0-2.7 mm (MNHN- IU-2008-15822). – BATHUS 3: stn CP 811, 23°41.42’S, 168°15.50’E, 383-408 m, 28.11.1993, 4 males 2.7-3.1 mm, 1 ovi- gerous female 3.0 mm, 1 female 2.9 mm (MNHN-IU-2008-15823). – stn DW 817, 23°42.38’S, 168°15.51’E, 405-410 m, 28.11.1993, 3 females 3.0-3.2 mm (MNHN-IU-2008-15824). – stn DW 818, 23°43.89’S, 168°16.32’E, 394 m, 28.11.1993, 3 males 2.7-3.0 mm, 1 female 2.7 mm (MNHN-IU-2008-15825). – stn DW 830, 23°19.75’S, 168°01.45’E, 361-365 m, 29.11.1993, 2 males 1.6, 3.0 mm (MNHN-IU-2008-15826). – BATHUS 4: stn DW 925, 18°54.55’S, 163°23.75’E, 370- 405 m, 07.08.1994, 1 male 2.1 mm (MNHN-IU-2008-15827). – stn DW 940, 18°59.53’S, 163°25.90’E, 305 m, 08.08.1994, 1 ovigerous female 2.0 mm, 1 female 1.9 mm (MNHN-IU-2008-15828). – LITHIST: stn CP 16, 23°43.2’S, 168°16.2’W, 379-391 m, 12.08.1999, 1  female 3.0 mm (MNHN-IU-2008-15834). – NORFOLK 1: stn DW 1709, 23°42’S, 168°16’E, 380-389 m, 25.06.2001, 1 ovigerous female 3.3 mm (MNHN-IU-2008-15801). – stn CP 1716, 23°22’S, 168°03’E, 266- 276  m, 26.06.2001, 1 male 2.8  mm (MNHN-IU-2008-15802). – NORFOLK 2: stn CP 2139, 23°00.68’S, 168 22.60’E, FIGURE 30 Paragiopagurus oxychelos n. sp. A, paratype female (2.1 mm) mNHN-iU-2012-727, New Caledonia, mUSOrStOm 4 stn DW 230, 390-420 m. B-E, holotype (2.8 mm) mNHN-iU-2012-733, New Caledonia, Smib 5 stn DW 73, 240 m. F, paratype male (2.7 mm) USNm 1184053, New Caledonia, mUSOrStOm 4 stn DW 212, 375-380 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages. C, right antennal peduncle and anterior portion of branchiostegite, lateral view. D, telson, dorsal view. E, first gonopods, anterior portion of abdomen, and coxae and sternite XiV (of fifth pereopods), ventral view. F, left second pleopod, lateral view. Scale bars 0.25 mm for a, C, D-F; 0.5 mm for b. ParagioPagurus: wordldwide review and new species 377 C B A, D-F F E B D CA rafael lemaitre378 372-373  m, 03.11.2003, 1 male 2.8 mm MNHN-IU-2012-1129. – EBISCO: stn DW 2508, 24°42.17’S, 159°43.610’E, 304-350 m, 07.10.2005, 1 male 1.8 mm MNHN-IU-2012-1130. – stn DW 2520, 24°06.960’S, 159°41.270’E, 350-400 m, 08.10.2005, 1 male 2.9 mm MNHN-IU-2012-1131. – stn DW 2525, 22°46.650’S, 159°23.106’E, 408-410 m, 09.10.2005, 1 male 2.1  mm, 1 ovigerous female 2.0 mm, 1 female 2.3 mm MNHN-IU-2012-1132. – stn  DW 2527, 22°45.380’S, 159°22.370’E, 330-340 m, 09.10.2005, 1 male 2.3 mm MNHN-IU-2012-1133. – stn DW 2530, 22°46.911’S, 159°22.810’E, 338-343 m, 09.10.2005, 1 male 1.8 mm MNHN-IU-2012-1134. – stn DW 2533, 22°17.205’S, 159°27.480’E, 36-370 m, 10.10.2005, 2 males 2.3, 3.3 mm MNHN-IU-2012-1135. – stn DW 2537, 22°18.459’S, 159°29.226’ E, 990 m, 10.10.2005, 1  male 2.4 mm MNHN-IU-2012-1136. French Polynesia. BENTHAUS: stn DW 1973, 23°23.49’S, 150°43.87’W, 200- 350 m, 20.11.2002, 1 ovigerous female 3.4 mm (MNHN-IU-2008-15838). Marquesas Islands. MUSORSTOM 9: stn DW 1145, 09°19.0’S, 140°06.3’W, 150-180 m, 22.08.1997, 3 males 0.9-2.3 mm (MNHN-IU-2008-15835). – stn DW 1197, 08°57.4’S, 140°01.9’W, 277-372 m, 27.08.1997, 1 female 1.9 mm (MNHN-IU-2008-15836). – stn DW 1288, 08°53.9’S, 139°38.0’W, 200-220 m, 08.09.1997, 2 males 1.8, 1.9 mm, 2 ovigerous female 1.8, 1.9 mm (MNHN-IU-2008-15837). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — Western Pacific, from Taiwan and New Caledonia region; and South Pacific, from the Marquesas Islands, and French Polynesia. Depth: 150 to 610 m. DESCRIPTION — Eleven pairs of biserial gills (Figure 30A), with lamellae undivided distally. Shield (Figure 30B) approxi- mately as broad as long; dorsal surface weakly calcified along midline, just posteriorly to rostrum, and anterior and ante- rolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections bluntly subtriangular, or with small terminal spine. Anterolateral margins sloping. Ventrolateral margins usually without spine. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, each with longitudinal row of setae on dorsal surface. Cornea weakly dilated. Ocular acicles subtriangular, about 0.25 times as long as ocular peduncles, each terminating in strong simple spine; separated basally by about half width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by 0.75 or more length of ultimate segment; ventral flagellum with about 6 articles. Ultimate segment about twice as long as penultimate, with scattered setae dorsally. Basal segment with strong ventromesial spine; lateral face with usually unarmed distal subrectangular lobe, and strong spine proximally. Antennal peduncles (Figure 30C) slightly exceeding distal margins of corneas. Fifth segment unarmed, with few, short bristle-like setae on dorsolateral distal angle, and few setae on mesial margin. Fourth segment with strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong simple spine extending to about midlength of acicle; mesial margin with small spine on dorsolateral distal angle. First segment with lateral surface armed with small spine; ventromesial angle produced, with row of 3-5 spines laterally. Antennal acicles nearly straight or weakly curved outwards (dorsal view), reaching or almost reaching distal margins of corneas, and terminating in strong spine; mesial margin setose, with row of 8-12 spines. Flagellum exceeding distal end of right cheliped; with short setae < 1 article in length, and long setae about 4 articles in length every 5-9 articles. FIGURE 31 Paragiopagurus oxychelos n. sp., paratype female (2.1 mm) mNHN-iU-2012-727, New Caledonia, mUSOrStOm 4 stn DW 230, 390-420 m, left mouthparts, internal view. A, mandible. B, maxillule. C, maxilla. D, first maxilliped. E, second maxilliped. F, third maxilliped. G, basis and ischium of third maxilliped with crista dentata. Scale bar 0.25 mm. ParagioPagurus: wordldwide review and new species 379 E G F CB DA rafael lemaitre380 Mandible (Figure 31A) with 3-segmented palp. Maxillule (Figure 31B) with external lobe of endopod slender, moderately developed, internal lobe with 1 long seta. Maxilla (Figure 31C) with endopod slightly exceeding distal margin of scaphognathite. First maxilliped (Figure 31D) with endopod exceeding distal margin of exopod. Second maxilliped (Figure 31E) without distinguishing characters. Third maxilliped (Figure 31F, G) with dactyl about 2.5 times as long as broad, carpus and propodus about 3 times as long as broad; crista dentata with about 11-14 small teeth, proximal teeth often distinctly larger than distal ones; coxa and basis each with small mesial tooth. Sternite of third maxillipeds with small spine on each side of midline. Epistomial spine short, sharp or blunt, occasionally absent. Chelipeds markedly dissimilar. Right cheliped (Figure 32A-E) massive; chela somewhat dorsoventrally flattened, operculate, with scattered setae or tufts of setae dorsally. Fingers slightly curving ventromesially, each terminating in small, usually blunt corneous claw; cutting edges with row of calcareous teeth, that of dactyl also sometimes with short row of small, blunt corneous spines distally; dorsal faces with scattered small sharp or blunt spines; ventral faces naked. Dactyl slightly longer than mesial length of palm, set at strongly oblique angle relative to longitudinal axis of palm; dorsal surface with scattered small tubercles or spines; dorsomesial margin serrate (Figure 32D), drawn out as sharp, thin somewhat translucent ridge with short transverse lines suggesting spine fusion, and with minute and often corneous-tipped teeth; ventromesial face concave. Fixed finger broad at base; dorsal surface with scattered small tubercles or spines. Palm broader than long; dorsal surface with scattered well-spaced, small sharp or blunt spines; lateral margin sharp, well delimited by row of small, closely set and often corneous-tipped spines; dorsomesial margin delimited by irregular row of small blunt spines; mesial face rounded, unarmed or with scattered spines or tubercles; ventral surface naked, smooth, or with scattered small tubercles. Carpus with several irregular rows of small spines on dorsal margin; dorsodistal margin with row of small blunt spines; dorsomesial surface with few long setae, and scattered small spines; ventromesial distal margin with row of spines; ventrolateral margin unarmed except for row of small blunt spines anteriorly; ventrolateral distal angle frequently with strong, ventrally directed protuberance armed with few small spines (Figure 32B, E). Merus with few setae on dorsal margin, and row of setae near dorsodistal margin; ventromesial margin with row of spines. Ischium unarmed but with setae on dorsal margin and ventral surface; coxa with row of small blunt spines on ventrodistal margin, and ventromesial row of long setae. Left cheliped (Figure 32F) usually well calcified. Fingers terminating in sharp corneous claws; dorsal and ventral surfaces unarmed except for few tufts of setae; cutting edge of dactyl with row of closely-set small corneous spinules, cutting edge of fixed finger with row of small calcareous teeth. Dactyl about as long as palm. Palm unarmed but with tufts of setae dorsally. Carpus with setae on dorsal margin, dorsodistal spine, and dorsolateral distal spine. Merus with few setae dorsally, and row of spines on ventrolateral distal margin. Ischium with setae dorsally and ventrally. Coxa with ventrodistal margin unarmed or with row of small spines, and ventromesial row of long setae. Ambulatory legs (Figure 33A-D) similar from right to left, usually exceeding right cheliped by about 0.3 length of dactyl of legs when fully extended. Dactyl approximately 1.8 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, ventromesial margin armed with irregular row of 7-10 corneous spines; mesial face with shallow, longitudinal sulcus on proximal half. Propodus with setae or tufts of setae on dorsal margin. Carpus with small dorsodistal spine, and short setae or tufts of setae on dorsal margin. Merus with setae on dorsal margin; merus of second pereopod unarmed ventrodistally but with row of setae. Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods; Figure 33G) setose, usually lacking spine or at most with minute subterminal spine. FIGURE 32 Paragiopagurus oxychelos n. sp. A-D, F, holotype (2.8 mm) mNHN-iU-2012-733, New Caledonia, Smib 5 stn DW 73, 240 m. E, paratype female (2.1 mm) mNHN-iU-2012-727, New Caledonia, mUSOrStOm 4 stn DW 230, 390-420 m. A, right cheliped, dorsal view. B, carpus and chela of same, lateral view. C, chela of same, mesial view. D, mesial margin of dactyl of right cheliped, ventral view (stippled area indicates semi-transparent, thin margin). E, carpus of right cheliped, lateral view. F, left cheliped, lateral view. Scale bars 1 mm for a-C, F, e; 0.25 mm for D. ParagioPagurus: wordldwide review and new species 381 E D A-C, F C D E AF B rafael lemaitre382 Fourth pereopod (Figure 33E) subchelate. Dactyl broadly curved, terminating in sharp corneous claw, longer and more slender in females than in males; with ventrolateral row of small corneous spinules. Propodus longer than wide, rasp consisting of 1 row (at least distally) of rounded scales. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 33F) chelate. Propodal rasp extending posteriorly to mid-length of segment. Uropods and telson (Figure 30D) markedly asymmetrical. Telson lacking lateral indentations, dorsal surface with scattered setae; posterior margin separated into 2 lobes by shallow U-shaped cleft; terminal margins of lobes armed with row of often curved, corneous spines. Male with paired first gonopods, and unpaired left second gonopod; first gonopod (Figure 28E) with distal lobe subtriangular, and concave mesial face; second left pleopod (Figure 28F) uniramous, bud like, or occasionally absent. Female lacking rudimentary second right pleopod. FIGURE 33 Paragiopagurus oxychelos n. sp., holotype (2.8 mm) mNHN-iU-2012-733, New Caledonia, Smib 5 stn DW 73, 240 m. A, left second pereopod, lateral view. B, dactyl of same, mesial view. C, third left pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of left fourth pereopod, lateral view. F, propodus and dactyl of left fifth pereopod, lateral view. G, anterior lobe of sternite Xii (of third pereopods), ventral view. Scale bars 1 mm for a-D; 0.25 mm for e-G. G E, F A-D F E BD G C A ParagioPagurus: wordldwide review and new species 383 COLOUR (Figure 1H) — Shield orangish medially, faint yellow laterally. Ocular peduncles light violet proximally, with red- dish stripe dorsally. Antennular peduncles semi-transparent, with light yellow or orange tint on ultimate and penultimate segments dorsally. Antennal peduncles orange. Right cheliped light; dorsolateral and dorsomesial margins light yellow. Left cheliped orange except for dorsal red stripe on merus and carpus, and light violet on mesial face of merus and car- pus; chela mostly light yellow. Meri, carpi, propodi and dactyls of ambulatory legs with dorsolateral red stripe, light violet dorsal margin and lateral faces, and orange ventral margins. HABITAT — Gastropod shells. REMARKS — This new species is the only one in Paragiopagurus in which males have paired first gonopods and rudimen- tary, unpaired left second gonopod. The right chela is also unique in that the dactyl has a sharp, finely cristate dorsomesial margin that is somewhat drawn out and is somewhat transparent (Figure 32D). The dorsolateral margin of the right chela is also sharp and cristate, although usually not as sharp as the dorsomesial margin of the dactyl. ETYMOLOGY — The specific name is a compound using the Greek adjective oxys, sharp, and the noun chele, claw, and refers to the sharp margins of the right chela (lateral margin of palm and fixed finger, and mesial margin of dactyl) that distinguish this new species. Paragiopagurus fasciatus Lemaitre & Poupin, 2003 Figures 1i, 34 Paragiopagurus fasciatus lemaitre & Poupin, 2003: 2, figs 1-4 (type locality: austral islands French Polynesia). — mclaughlin et al., 2010: 39, pl. 19e. TYPE MATERIAL — Holotype, French Polynesia, Austral islands, Neilson reef, BENTHAUS, stn CAS 1916, 27°00.3’S, 146°03.6’W, 180 m, 11.11.2002, male 9.9 mm (MNHN-IU-2008-15806). Paratypes, French Polynesia. BENTHAUS: stn  CAS  1878, 27°52.3’S, 143°31.7’W, 129-122 m, 05.11.2002, 1 ovigerous female 9.6 mm (MNHN-IU-2008-15803). – stn DW 1881, 27°54.6’S, 143°28.5’W, 112-121 m, 06.11.2002, 1 juvenile male 3.3 mm (MNHN-IU-2008-15804). – stn DW 1915, 27°03.4’S, 146°03.9’W, 120-200 m, 11.11.2002, 1 ovigerous female 8.6 mm, (USNM 1016947). – stn CP 1918, 27°03.4’S, 146°04.0’W, 130-140 m, 11.11.2002, 1 female 6.5 mm (MNHN-IU-2008-15807). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — South Pacific: Austral islands, French Polynesia. Depth: 112 to 200 m. DIAGNOSIS — Gill lamellae undivided or weakly divided distally. Antennal peduncles and acicles at most reaching mid- point of corneas. Shield (Figure 34A) about as broad as long; dorsal surface weakly calcified medially and anteriorly; ros- trum broadly rounded, with short mid-dorsal ridge sometimes narrowing and extending posteriorly for about one-third of shield; lateral projections subtriangular, with terminal spine. Ocular peduncles more than half length of shield; corneas slightly dilated. Ocular acicles subtriangular with mesial margins nearly parallel, each terminating in strong, simple spine. Maxillule with internal lobe usually with 4 long terminal setae. Epistomial spine short, straight. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment. Antennal peduncles (Figure 34B) not exceeding distal margins of corneas, at most reaching to about midpoint of corneas; fourth segment with strong spine on dorsolateral distal angle; second segment with dorsolateral distal angle produced, terminating in strong, simple or multi- fid spine (sometimes with 1 or 2 small spines dorsally) reaching to about midpoint of acicle; acicle weakly curved outward (dorsal view), reaching at most to midpoint of cornea, terminating in strong spine, mesial margin armed with row of 3 rafael lemaitre384 I H G F B A, C, D, E GH F I A DEB C ParagioPagurus: wordldwide review and new species 385 (juveniles, sl < 3.5 mm) to 7 (adults) strong spines; flagellum with few short (<1 article in length) setae throughout, and at least distally with long setae (1-3 articles in length) every 15-20 articles. Right chela (Figure 34C, D) dorsoventrally flattened, somewhat operculate; dorsal surface with numerous blunt to sharp spines or tubercles. Left cheliped (Figure 34E) with dorsal margin of carpus armed with row of 8-10 spines (less in juveniles sl < 3.5 mm) and strong dorsodistal spine. Ambulatory legs (Figure 34F, G) each with merus, carpus and propodus unarmed except for short, bristle-like setae on dorsal margins, and dorsodistal spine on carpus; dactyl with dorsal row of long, bristle-like setae and dorsomesial row of corneous spinules. Fourth pereopod (Figure 34H) with propodal rasp consisting of 1 row of ovate scales at least dis- tally. Uropods and telson (Figure 34I) markedly asymmetrical. Telson with weak or obsolete transverse suture, posterior margin separated by narrow, V-shaped shallow cleft, into rounded projections armed with short corneous spines (more numerous in females), some often ventrally curved; left anterior ventrolateral margin with long setae in males, or with long setae mixed with row of slender, corneous spinules in females. Males with well developed paired first and second gonopods; second gonopod often with rudimentary exopod. Female with rudimentary second right pleopod. COLOUR (Figure 1i) — “Shield white, with pair of broad, submedian red stripes; lateral margins red. Posterior carapace whitish to pale red, with red stripes over cardiobranchial grooves. Ocular peduncles with white dorsal surface, and red ventral and lateral surfaces (peduncles slightly twisted in Figure 49I, showing part of lateral surfaces in red, and dorsal surface in white). Antennular and antennal peduncles whitish-pink; flagella pale yellow. Right cheliped with chela and carpus mostly cream or white, with white spines; chela also with well-spaced small red spots; palm with dorsal surface iridescent; carpus cream or light brown, with white spines, and iridescent dorsal surface. Left cheliped white; chela with well-spaced red spots; carpus with dorsolateral, dorsomesial, and ventrolateral red stripes, and scattered red spots; merus with dorsolateral and dorsomesial red stripes. Ambulatory legs white with narrow dorsolateral and ventrolateral red stripes; merus, carpus and propodus with dorsolateral, dorsomesial, and ventrolateral red stripes; dactyl with light red dorsolateral stripe. Fourth and fifth pereopods with continuous red stripe on lateral faces of merus, carpus, and pro- podus, and on dactyl of fourth pereopod. Abdomen purple or pinkish, with red spots on tergites, and dorsal surface of telson” (Lemaitre & Poupin 2003: 9). HABITAT — Gastropod shells. REMARKS (see also P. laperousei n. sp.) — As discussed by Lemaitre & Poupin (2003), P. fasciatus is most similar to two other species also distributed in French Polynesia, P. bougainvillei and P. wallisi, and a new species from the northern Hawaiian Islands, P. laperousei n. sp. Individuals of these four species can grow to relatively large sizes (up to sl 12.5 mm). The strong armature of the carpus of the left cheliped (Figure 34E) immediately sets P. fasciatus apart from the other three similar species. The striped color pattern of the shield and ambulatory legs of P. fasciatus is strikingly different (Figure 1I) from all other Paragiopagurus species. FIGURE 34 Paragiopagurus fasciatus lemaitre & Poupin, 2003, holotype male (9.9 mm) mNHN-iU-2008-15806, French Polynesia, beNtHaUS stn DW 1916, 180 m. A, shield and cephalic appendages, dorsal view. B, right antennal peduncle, lateral view. C, carpus and chela of right cheliped, dorsal view. D, chela of same, lateral view. E, left cheliped, dorsal view. F, right second pereopod, lateral view. G, dactyl of same, mesial view. H, propodus and dactyl of left fourth pereopod, lateral view. I , telson, dorsal view. Scale bars 10.0 mm for a, C-G; 5.0 mm for b; 1 mm for H, i. (adapted from lemaitre & Poupin 2003). rafael lemaitre386 Paragiopagurus diogenes (Whitelegge, 1900) Figures 15a, b, 35a, 36 restricted synonymy (for other synonyms see lemaitre, 1996: 208): Sympagurus diogenes Whitelegge, 1900: 172, pl. 34-fig. 3 (lectotype locality: New South Wales, autralia). — lemaitre, 1989: 37; 1994: 412, fig. 27g. Parapagurus diogenes – de Saint laurent, 1972: 108. — takeda, 1982: 66, fig. 196. — miyake, 1975: 117, 2 unnumbered photos; 1982: 119, fig. 2; 1990: 117, 2 unnumbered photos. — baba et al., 1986: 197, fig. 145. Paragiopagurus diogenes – lemaitre, 1996: 208, figs. 14d, e, 24; 2000: 219, fig 70. — Spiridonov & Zhadan, 1999: 627. — Forest & mclay, 2001: 702, fig. 11. — Davie, 2002: 90. — Poore, 2004: 282, fig. 81e. — asakura, et al., 2006: 219, fig. 4. — mclaughlin et al., 2007: 323, 3 unnumbered figs. — Wang, 2008: 759. — mclaughlin et al., 2010: 39. — Webber, et al., 2010: 226. [not] Sympagurus arcuatus diogenes – Hale, 1941: 279. [see remarKS] [not] Paragiopagurus diogenes – Crist et al., 2009: 27, unumbered fig. [= P. laperousei n. sp.] [not] Paragiopagurus cf. diogenes – mclaughlin et al., 2010: 103, fig. 19D. [= P. laperousei n. sp.] TYPE MATERIAL — Lectotype (subsequent designation by Lemaitre, 1996), Australia, New South Wales, 3 km E of Port Hacking, Thetis, stn 35, 34°03.5’S, 151°12.5’E, 40-69 m, 10.03.1898, female 7.9 mm (AM G2379). Paralectotypes, Austra- lia, New South Wales. Thetis: stn 48, 34°27’S, 151°04’E, 101-102 m, 18.03.1898, 2 males 3.9, 4.4 mm (AM G2380). MATERIAL EXAMINED — the type material (see above). Madagascar. N.O. “Vauban”: stn P4-Ch. 52, 15°21.0’S, 46°12.5’e, 150 m, 08.11.1972, 3 males 4.4-6.3 mm, 6 females 4.1-4.9 mm (mNHN-Pg 5786). – stn P4-Ch. 53, 15°21.7’S, 46°12.6’e, 90-130 m, 08.11.1972, 1 male 6.3 mm. (mNHN-Pg 5787). Japan. tosa bay: 130 m, 04.06.1985, 1 male 8.5 mm (USNm). Taiwan. taiwan Strait, Formosa bank: 60 m, 02.month not specified.1972, 1 male 5.7 mm (California academy of Sciences 046660). Philippines. mUSOrStOm 1: stn CP 27, 13°59.8’N, 120°18.6’e, 192-188 m, 22.03.1976, 1 male 9.3 mm (mNHN-Pg 6841). – mUSOrStOm 2: stn CP 01, 14°00’S, 120°19’e, 188-198 m, 20.11.1980, 1 ovigerous female 10.2 mm (mNHN-iU-2008-16068). – stn CP 02, 14°01’S, 120°17’e, 184-186 m, 20.11.1980, 1 male 10.1 mm (mNHN-iU-2008-16069). – stn CP 17, 14°00’S, 120°17’e, 174-193 m, 22.11.1980, 1 ovigerous female 10.7 mm (mNHN-iU-2008-16070). – stn CP 35, 13°28’S, 121°12’e, 160-198 m, 24.11.1980, 1 male 11.1 mm, 1 female 7.4 mm mNHN-iU-2012-1137. – stn CP 51, 13°59’S, 120°16’e, 170-187 m, 27.11.1980, 1 male 9.9 mm, 1 ovigerous female 10.0 mm (mNHN-iU-2008-16071). – stn CP 69, 14°05’S, 120°02’e, 180 m, 30.11.1980, 1 ovigerous female 9.7 mm mNHN-iU-2012-1138. Vanuatu. VOlSmar: stn DW 42, 22°17.00’S, 168°41.50’e, 400 m, 08.06.1989, 1 male 7.8 mm (mNHN-iU-2012-706). – mUSOrStOm 8: stn CP 963, 20°20,10’S, 169°49,08’e, 400-440 m, 21.09.1994, 1 ovigerous female 8.4 mm (mNHN-Pg 6831). Chesterfield Islands. CHalCal 1: stn DW 64, 22°11.50’S, 159°15.40’e, 305 m, 27.07.1984, 1 male 5.7 mm (mNHN-Pg 6830). – mUSOrStOm 1: no station data, 1986, 1 ovigerous female 6.9 mm (USNm). New Caledonia. mUSOrStOm 4: stn DW 183, 19°01.80’S, 163°29.5’e, 275 m, 18.09.1985, 1 male 5.1 mm (USNm). – stn CC 201, 18°55.80’S, 163°13.80’e, 500 m, 20.09.1985, 1 juvenile male 4.5 mm (mNHN-iU-2012-696). – mUSOrStOm 5: stn DW 263, 25°21.30’S, 159°46.44’e, 150-225 m, 08.10.1986, 1 female 7.8 mm, 2 ovigerous females 6.3, 6.7  mm (mNHN-iU-2012-697), 1 juvenile male 2.6 mm mNHN-iU-2012-1139, 2 ovigerous females 6.3, 6.7 mm, 1 female 4.8 mm (USNm). – stn DW 265, 25°21.10’S, 159°45.20’e, 190-260 m, 08.10.1986, 1 juvenile 2.7 mm (USNm). – stn DW 266, 25°20.20’S, 159°45.70’e, 240 m, 08.10.1986, 1 male 4.8 mm, 1 ovigerous female 6.4 mm (USNm). – stn DW 270, 24°48.85’S, 159°34.13’e, 223 m, 09.10.1986,1 ovigerous females 5.8 mm (USNm). – stn DW 291, 23°07.70’S, 159°28.40’e, 300 m, 11.10.1986, 1 male 5.5 mm (USNm). – stn DW 299, 22°47.20’S, 159°23.7’e, 360-390 m, 11.10.1986, FIGURE 35 Coloration (in life or fresh). A, Paragiopagurus diogenes (Whitelegge, 1900), female (6.2 mm) NtOU, taiwan, Gushan fishing port, Kaohsiung County (from mclaughlin et al. 2007: 323, unnumbered photo.). B-D, P. laperousei n. sp., Hawaiian islands, French Frigate Shoals, baited traps, 215-228 m: B, holotype male (4.9 mm) UF 12304 (from mclaughlin et al. 2010: fig. 19D, as P. cf. diogenes); C, paratype ovigerous female (6.6 mm) UF 12303; D, not sexed or measured. E, P. wallisi (lemaitre, 1994), French Polynesia, austral islands. F, P. bougainvillei (lemaitre, 1994), French Polynesia, tuamotu. G, H, P. ventilatus lemaitre, 2004, taiwan, Gueishan island, Yilan County, Vent stn 9, 200 m: G, male (NtOU, from mclaughlin et al. 2007: 320, unnumbered photo.); H, ovigerous female (NtOU, from mclaughlin et al. 2007: 320, unnumbered photo.). I, P. pilimanus (a. milne-edwards, 1880), female (3.9 mm) USNm 1190206, off Curaçao, Caribbean Sea, CUraSUb stn 12-03, 144.8-182.8 m. (Photograps: a, G, H, t.-Y. Chan; b, C, G. Paulay; D, S. middleton © 2006; e, F, J. Poupin; i, b. b. brown © 2012). ParagioPagurus: wordldwide review and new species 387 A H I G F E D CB rafael lemaitre388 1  male 7.2  mm (mNHN-iU-2012-699), 1 ovigerous female 6.1 mm (USNm), 1 female 4.6 mm (mNHN-iU-2012-698). – stn DW 301, 22°06.90’S, 159°24.60’e, 487-610 m, 12.10.1986, 1 male 7.5 mm, 1 ovigerous female 5.3 mm (mNHN-iU-2012-700). – stn DW 304, 22°10.34’S, 159°25.51’e, 385- 420 m, 12.10.1986, 1 ovigerous female 7.2 mm (mNHN-iU-2012-701). – stn 311, 320 m, 1 male 3.0 mm (mNHN-iU-2012-702). – stn  CP  316, 22°25.13’S, 159°24.00’e, 330 m, 13.10.1986, 1 female 7.5 mm (USNm). – CHalCal 2: stn CH 5, 24°44.0’S, 168°08.5’e, 223 m, 27.10.1986, 1 male 9.7 mm (mNHN-iU-2012-694). – stn CP 18, 24°42.85’S, 168°09.73’e, 271 m, 27.10.1986, 1 juvenile male 1.7 mm (USNm). – stn DW 71, 24°42.26’S, 168°09.52’e, 230 m, 27.10.1986, 1 female 7.2 mm (USNm). – stn DW 83, 23°20.30’S, 168°05.50’e, 200 m, 31.10.1986, 2 males 5.2, 7.6 mm (mNHN- iU-2012-695). – Smib 4: stn DW 41, 24°44.00’S, 168°08.60’e, 235 m, 08.03.1989, 1 male 9.6 mm (USNm). – stn DW 43, 24°46.60’S, 168°08.80’e, 235- 245 m, 08.03.1989, 1 male 9.8 mm (mNHN-iU-2012-703). – stn DW 44, 24°46.00’S, 168°08.20’e, 300 m, 08.03.1989, 1 male 10.0 mm (USNm). – Smib 6: stn DW 116, 18°59.30’S, 163°26.20’e, 300 m, 02.03.1990, 1 juvenile male 3.4 mm (mNHN-iU-2012-705). – Smib 8: stn DW 154, 24°45.39’S, 168°08.41’e, 235-252 m, 28.01.1993, 1 female 3.4 mm (mNHN-Pg 6832). – stn CP 162, 24°46.99’S, 168°08.76’e, 254-264 m, 28.01.1993, 1 male 9.2 mm (mNHN-Pg 6833). – stn DW 170, 23°41.23’S, 168°00.56’e, 241-244 m, 29.01.1993, 1 male 4.0 mm, 1 female 5.6 mm (mNHN-Pg 6834). – stn DW 175, 23°41.13’S, 168°00.38’e, 235-240 m, 29.01.1993, 1 female 5.4 mm (mNHN-Pg 6835). – stn DW 177, 23°39.06’S, 168°00.05’e, 320-370 m, 29.01.1993, 1 male 4.2 mm (mNHN-Pg 6836). – batHUS 4: stn CP 905, 19°02.45’S, 163°15.65’e, 294-296 m, 04.08.1994, 1 male 9.4 mm (mNHN-Pg 6827). – stn CP 939, 18°58.18’S, 163°25.37’e, 304-320 m, 08.08.1994, 2 ovigerous females 6.5, 7.1 mm, 1 female 3.7 mm (mNHN-Pg 6828). – stn CP 905, 19°02.45’S, 163°15.65’e, 294-296 m, 04.08.94, 1 male 9.4 mm mNHN-iU-2012-1140. – stn CP 939, 18°58.18’S, 163°25.37’e, 304-320 m, 08.08.94, 2 ovigerous females 6.5, 7.1 mm, 1 female 3.7 mm mNHN-iU-2012-1141. – stn DW 940, 18°59.53’S, 163°25.90’e, 305 m, 08.08.1994, 1 male 7.4 mm, 2 spms in housing (mNHN-Pg 6823). – stn DW 942, 19°04.26’S, 163°27.36’e, 270-264 m, 08.08.1994, 1 male 6.0 mm (mNHN-Pg 6829). – Smib 10: stn DW 208, 24°49’S, 168°09’e, 270 m, 10.01.1995, 1 male 4.2 mm (mNHN-Pg 6838), 1 female 8.7 mm (mNHN-Pg 6837). – stn DW 209, 24°49’S, 168°09’e, 329-560 m, 10.01.1995, 1 female 9.5 mm (mNHN-Pg 6839). – stn DW 210, 24°49’S, 168°09’e, 308-510 m, 10.01.1995, 1 male 8.1 mm (mNHN-Pg 6840). – NOrFOlK 2: stn DW 2093, 24°43,90’S, 168°08,70’e, 230 m, 29.10.2003, 1 female 4.4 mm (mNHN-iU-2008-16072). – ebiSCO: stn CP 2494, 24°45’S, 159°42’e, 348-354 m, 06.10.2005, 1 ovigerous female 8.8 mm mNHN-iU-2012-1142. – stn CP 2499, 24°53’S, 159°52’e, 286- 529 m, 07.10.2005, 1 male 8.7 mm mNHN-iU-2012-1143. – stn DW 2504, 24°48’S, 159°46’e, 390-600 m, 07.10.2005, 2 males 7.3, 8.3 mm mNHN- iU-2012-1144. – stn CP 2505, 24°45’S, 159°43’e, 328-463 m, 07.10.2005, 1 male 10.4 mm mNHN-iU-2012-1145. – stn DW 2508, 24°41’S, 159°43’e, 304-350 m, 07.10.2005, 2 males 3.3, 7.1 mm mNHN-iU-2012-1146. – stn DW 2509, 24°08’S, 159°35’e, 265 m, 08.10.2005, 1 male 3.0 mm mNHN- iU-2012-1147. – stn DW 2513, 24°06’S, 159°42’e, 280-500 m, 08.10.2005, 1 ovigerous female 6.4 mm mNHN-iU-2012-1148. – stn DW 2530, 22°48’S, 159°23’e, 338-343 m, 09.10.2005, 1 male 4.8 mm mNHN-iU-2012-1149. – stn DW 2532, 22°15’S, 159°27’e, 350 m, 10.10.2005, 1 male 9.5 mm mNHN- iU-2012-1150. – stn DW 2533, 22°18’S, 159°28’e, 360-370 m, 10.10.2005, 1 female 4.6 mm mNHN-iU-2012-1151. – stn DW 2534, 22°17’S, 159°28’e, 390-430 m, 10.10.2005, 3 males 4.9-8.3 mm, 1 female 5.7 mm mNHN-iU-2012-1152. – stn CP 2539, 22°19’S, 159°25’e, 315-320 m, 10.10.2005, 1 male 3.8 mm mNHN-iU-2012-1153. – stn CP 2541, 22°18’S, 159°28’e, 360-369 m, 10.10.2005, 1 male 5.8 mm mNHN-iU-2012-1154. Australia. 38 km NNe of montague island, 164 m, 09.1926, 1 male 12.4 mm (am P9312). Hawaiian Islands. NOaa/NmFS “townsend Cromwell”: stn 33-8, 21°00.3’N, 156°45.4’W, 223-238 m, 30.10.1967, 2 males 4.0, 4.3 mm, 3 females 3.7-4.0 mm (bPbm S11007). – stn tC 33-10, 21°00.2’N, 156°46.3’W, 210-219 m, 31.10.1967, 1 ovigerous female 4.5 mm (bPbm S10960). – stn 33-15, 21°03.2’N, 156°43.5’W, 241-254 m, 31.10.1967, 1 male 4.3 mm, 1 ovigerous fe- male 5.2 mm (bPbm S11002). – stn tC 33-18, 21°03.7’N, 156°43.7’W, 245 m, 01.11.1967, 1 male 5.5. mm, 1 ovigerousg female 5.1 mm (bPbm S10948). – stn tC 33-26, 20°43.1’N, 156°51.3’W, 221-232 m, 04.11.1967, 1 male 6.6 mm (bPbm S10991), 5 males 3.4-5.1 mm, 1 female 3.7 mm, (bPbm S11009). – stn 35-2, 20°58.0’N, 156°46.0’W, 201 m, 28.03.1968, 1 male 7.9 mm, 1 ovigerous female 6.4 mm (bPbm S10997). – stn tC 35-33, 21°09.7’N, 157°29.8’W, 183 m, 07.04.1968, 3 males 3.4-3.9 mm, 5 females 2.5-4.9 mm (bPbm S11008), 8 males 2.4-4.6 mm, 2 females 3.0, 4.0 mm (bPbm S11008). – stn tC 35-34, 21°09.7’N, 157°29.8’W, 08.04.1968, 2 males 3.6, 4.3 mm, 8 females 2.4-4.2 mm (bPbm), 1 damaged spm (bPbm S10996). – stn tC 35-35, 21°09.7’N, 157°29.8’W, 183 m, 08.04.1968, 1 male 3.9 mm, 1 ovigerous female 4.3 mm (bPbm S11010). – stn tC 36-8, 21°01.2’N, 156°44.1’W, 27.04.1968, 29 males 3.7-8.2 mm, 7 ovigerous females 3.7-6.4 mm, 4 females 3.4-4.8 mm (bPbm). – stn tC 36-9, 21°01.7’N, 156°44.3’W, FIGURE 36 Paragiopagurus diogenes (Whitelegge, 1900). A-G, I-K, male (8.1 mm) am P39441, New South Wales, australia. H, F (4.0 mm) am P40389, New South Wales, australia. A, shield and cephalic appendages, dorsal view. B, right antennal peduncle, lateral view. C, left cheliped, dorsal view. D, right second pereopod, lateral view. E, right third pereopod, lateral view; F, dactyl of same, mesial view. G, propodus and dactyl of male left fourth pereopod, lateral view. H, propodus and dactyl of F left fourth pereopod, lateral view. I, telson, dorsal view. J, male left first gonopod, mesial view. K, male left second gonopod, anterior view. Scale bars 2 mm for a; 1 mm for b, G, i-K; 3 mm for C-F; 0.5 mm for H. (adapted from lemaitre 1996). ParagioPagurus: wordldwide review and new species 389 H C-F B, G, I-K A G H I B C K J F E A D rafael lemaitre390 223 m, 27.04.1968, 1 male 6.4 mm (bPbm), 19 males 3.0-6.9 mm, 3 ovigerous females 4.5-4.8 mm, 6 females 4.2-8.1 mm (bPbm S11018), 9 males 3.7-5.4 mm, 3 ovigerous females 4.2-5.7 mm, 7 females 3.1-5.7 mm (bPbm S10954). – stn tC 36-10, 20°57.2’N, 156°47.1’W, 219 m, 28.04.1968, 1 male 8.5 mm, 2 ovigerous females 4.5, 6.4 mm (bPbm S10993), 5 males 5.4-6.7 mm, 1 ovigerous female 3.9 mm, 1 female 4.2 mm (bPbm). – stn tC 36-11, 21°04.0’N, 156°44.1’W, 221 m, 28.04.1968, 7 males 3.4-8.8 mm, 5 ovigerous females 3.7-8.9 mm, 1 female 4.8 mm (bPbm), 1 female 4.8 mm (bPbm S11005). – stn tC 36-12, 20°57.9’N, 156°47.4’W, 219 m, 28.04.1968, 1 male 3.2 mm (bPbm S11012), 3 males 3.9-5.4 mm, 1 ovigerous female 4.3 mm (bPbm S10999). – stn tC 36-17, 21°37.8’N, 158°03.5’W, 101 m, 03.05.1968, 1 male 3.0 mm (bPbm S11014). – stn tC 36-21, 21°09.7’N, 157°29.0’W, 183 m, 04.05.1968, 1 male 2.7 mm (bPbm S11013), 3 males 3.6-4.5 mm (bPbm S11006). – stn tC 40-2, 21°09.9’N, 157°24.1’W, 183 m, 6.11.1968, 8 males 3.3-6.3 mm, 3 ovigerous females 2.5-5.2 mm, 1 female 3.7 mm (bPbm). – stn tC 40-12, 21°09.8’N, 157°30.1’W, 183 m, 08.11.1968, 2 males 4.2, 6.1 mm (bPbm S10950). – stn tC 40-14, 21°09.7’N, 157°24.6’W, 183 m, 08.11.1968, 9 males 2.7-5.7 mm, 2 ovigerous females 2.5, 4.5 mm, 3 fe- males 2.4-2.7 mm (bPbm). – stn tC 40-15, 21°0.9.8’N, 157°30.2’W, 181 m, 08.11.1968, 2 males 3.1, 4.9 mm (bPbm S10945). – stn tC 40-28, 21°09.7’N, 157°29.8’W, 185 m, 11.11.1968, 4 males 4.8-6.1 mm, 1 female 4.8 mm (bPbm S10952). – stn tC 40-48, 20°57.3’N, 156°47.5’W, 218 m, 17.11.1968, 1 male 6.3 mm (bPbm S10951). – stn tC 40-50, 21°01.7’N, 156°43.2’W, 210 m, 17.11.1968, 3 males 3.7-7.6 mm, 2 ovigerous females 6.7, 7.5 mm, 4 females 3.6-7.5 mm (bPbm). – stn tC 40-53, 20°57.5’N, 156°47.7’W, 219 m, 17.11.1968, 1 male 6.7 mm, 1 ovigerous female 3.6 mm, 1 female 5.2 mm (bPbm S11001).– bUrCH: stn 80051, 21°16.9’N, 157°55.0’W, 183 m, 30.08.1980, 2 males 3.1, 4.2 mm, 1 female 3.1 mm (bPbm S10990). For additional material examined, including specimens illustrated or photographed in Figures 33 and 34a,b see lemaitre (1996). DISTRIBUTION — Western Indian Ocean: from Madagascar. Western and central Pacific: Japan, Taiwan, South China Sea, Philippines, Vanuatu, Chesterfield Islands, New Caledonia region, Australia, New Zealand, and Hawaiian Islands. Depth: 4 to 695 m. DIAGNOSIS. — Gill lamellae distally divided. Shield (Figure 36A) as broad as long; dorsal surface weakly calcified medially, and with low blister-like tubercles; rostrum broadly rounded, with short mid-dorsal ridge; lateral projections subtrian- gular, with terminal spine. Ocular peduncles more than half length of shield. Corneas slightly dilated. Ocular acicles sub- triangular, terminating in strong spine usually directed anteromesially. Epistome with strong, straight spine. Antennular peduncles exceeding distal margin of corneas by nearly entire length of ultimate segment. Antennal peduncles (Figure 36B) exceeding distal margin of corneas by at most 0.20 length of fifth segment; third segment with strong ventromesial distal spine; acicle curved outward (in dorsal view), not exceeding distal margin of cornea, and armed mesially with 4-7 strong spines; flagellum articles with short setae < 1 flagellar article, interspersed with long setae every 5 to 15 articles. Chelipeds markedly dissimilar, glabrous or at most with scattered short setae, carpi and chelae usually iridescent dorsally. Right cheliped (Figure 15A, B) operculate, with numerous small tubercles or spines on dorsal surfaces of carpus and chela (tubercles or spines decreasing in size and number with increased size of individuals); fingers curved ventromesially; dactyl set at strongly oblique angle to longitudinal axis of palm, with ventromesial face concave; palm with dorsolateral margin well delimited by row of blunt to sharp spines, mesial face rounded. Left cheliped (Figure 36C) with dorsolateral face of carpus frequently weakly calcified; chela with dorsomesial row of few small spines; carpus with irregular row of small spines dorsally. Ambulatory legs (Figure 36D-F) unarmed except for dorsodistal spine on carpi; dactyls each with ventromesial row of about 8-14 corneous spines. Anterior lobe of sternite XII (of third pereopods) unarmed, or rarely with small blunt marginal spine. Fourth pereopod (Figure 36G, H) with dactyl terminating in sharp corneous claw (longer and more slender in females than in males); propodal rasp consisting of 1 row of ovate scales. Uropods and telson (Figure 36I) markedly asymmetrical; telson with weak transverse suture; dorsal surface usually with low, blister-like tubercles; posterior lobes separated by narrow U-shaped cleft, terminal margin of lobes armed with numerous corneous spines (of- ten strongly curved on left lobe). Male with paired first and second gonopods well developed; first gonopod (Figure 36J) with concave distal lobe; second gonopod (Figure 36K) with distal segment setose on distomesial face, and row of short bristle-like setae on lateral margin. Females occasionally with paired first pleopods; with vestigial second right pleopod. COLOUR (Figure 35A). — “In fresh specimens, general color orange or reddish (in preservative, color fades to cream white except on chelipeds where patterns remain for considerable time). Chelipeds with fingers cream white; dorsal surface of carpus and chela orange or reddish, iridescent, fading to white laterally. Walking legs orange or reddish, fading to cream ParagioPagurus: wordldwide review and new species 391 white towards the dactyls. Carpi of chelipeds and walking legs each with dark red band proximally. Merus of right cheli- ped with dark red stripe dorsomesially. Carapace and chelae with small, scattered red spots” (Lemaitre 1996: 211). HABITAT — Gastropod shells. REMARKS (see also P. laperousei n. sp.) — As discussed by Lemaitre (1996: 211), Hale (1941) considered Whitelegge’s (1900) Sympagurus diogenes (= Paragiopagurus diogenes) a subspecies of S. arcuatus A. Milne-Edwards & Bouvier, 1893 (=  Oncopagurus gracilis (Henderson, 1888), an Atlantic species). However, even from Hale’s brief description of the dense pilosity of his specimens from Tasmania and Macquaire Island, it is clear that his speicmens are not P. diogenes but instead of another, undetermined species. Paragiopagurus laperousei n. sp. Figures 35b-D, 37, 38, 39a-C, 40, 41 Paragiopagurus diogenes – Crist et al., 2009: 27, unumbered fig. [not Whitelegge, 1900] Paragiopagurus cf. diogenes – mclaughlin et al., 2010: 103, fig. 19D. [not Whitelegge, 1900] “anemone crab” – Knowlton, 2010: photos on inside title page and backcover. TYPE MATERIAL — Holotype, Northwestern (Leeward) Hawaiian Islands, French Frigate Shoals, 23°88.73’N, 166°13.57’W, baited traps, 215-228 m, 25.10.2006, male 4.9 mm (UF 12304). Paratype, Northwestern (Leeward) Hawaiian Islands. French Frigate Shoals, 23°88.73’N, 166°13.57’W, baited traps, 215-228 m, 25.10.2006, 1 ovigerous female 6.6 mm (UF 12303). MATERIAL EXAMINED — the type material (see above). DISTRIBUTION — Known so far known only from the types collected in the northern Hawaiian Islands. Depth: 215-228 m. DESCRIPTION — Eleven pairs of biserial, phyllobranchiate gills, with lamellae undivided (Figure 37A). Shield (Figure 37B) approximately as broad as long; dorsal surface weakly calcified along midline, just posteriorly to rostrum, and anterior and anterolateral margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins concave. Lateral projections bluntly to acutely subtriangular. Anterolateral margins sloping. Ventrolateral mar- gins without spine. Posterior margin broadly rounded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, at most with scattered setae. Corneas slightly dilated. Ocular acicles subtriangular, about 0.3 times length of ocular peduncles, each terminating in long and strong, simple spine directed slightly inward, and with tuft of setae dorsally; separated basally by about half width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment; ventral flagellum with about 9 articles. Ultimate segment about twice as long as penultimate, with scattered setae dorsally. Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 3 or 4 small spines, and strong spine proximally. Antennal peduncles (Figure 37C) slightly exceeding distal margins of corneas. Fifth segment unarmed, with few setae on lateral and mesial margins. Fourth segment with strong spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong spine reaching to about midpoint of acicle; mesial margin with spine on dorsolateral distal angle. First segment with lateral surface armed with small spine; ventromesial angle produced, with row of 2-4 strong spines laterally. Antennal acicles slightly sinuous (dorsal view), reaching at most to about proximal margins of corneas, each terminating in strong spine; mesial margins rafael lemaitre392 FIGURE 37 Paragiopagurus laperousei n. sp., holotype male (4.9 mm) UF 12304, Hawaiian islands, French Frigate Shoals, 215-228 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages. C, right antennal peduncle and anterior portion of branchiostegite, lateral view. D, left cheliped, dorsal view. E, male left first gonopod, mesial view. F, male left second gonopod, anterior view. Scale bar 0.25 mm for a, e, F; 1 mm for b, D; 0.5 mm for C. FE B A D C ParagioPagurus: wordldwide review and new species 393 FIGURE 38 Paragiopagurus laperousei n. sp., holotype male (4.9 mm) UF 12304, Hawaiian islands, French Frigate Shoals, 215-228 m, left mouthparts, internal view. A, mandible. B, maxillule. C, maxilla. D, first maxilliped. E, second maxilliped. F, third maxilliped. Scale bars 0.3 mm for a-D; 0.5 mm for e, F. E, F A-D E DBA C F rafael lemaitre394 setose, each armed with row of 5-8 strong spines. Flagellum reaching beyond distal end of right cheliped; with scattered short (<1 article in length) setae throughout. Mandible (Figure 38A) with 3-segmented palp. Maxillule (Figure 38B) with external lobe of endopod obsolete, internal lobe with 3 long setae. Maxilla (Figure 38C) with endopod not exceeding distal margin of scaphognathite. First maxilliped (Figure 38D) with endopod exceeding distal margin of exopod. Second maxilliped (Figure 38E) without distinguishing characters. Third maxilliped (Figure 38F) with distal 3 segments about twice to 2.5 times as long as wide; crista dentata with about 8-10 corneous-tipped teeth, proximal teeth slightly larger than others; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine short and sharp. Chelipeds markedly dissimilar. Right cheliped (Figure 39A-C) massive, with chela, carpus and merus at most with scattered short setae; chela somewhat dorsoventrally flattened, operculate; chela longer than broad. Fingers curving ventromesially, terminating in small, usually blunt corneous claw; cutting edges with 2 or 3 large calcareous teeth interspersed with smaller ones; dorsal faces with strong, mostly blunt spines. Dactyl slightly longer than palm, set at strongly oblique angle to longitudinal axis of palm when closed; mesial margin well defined by row of spines; ventromesial face concave, ventral face smooth. Fixed finger broad at base. Palm with dorsal surface covered with small, well spaced spines or tubercles; lateral margin well delimited by row of spines; mesial face rounded, with well-spaced small spines or tubercles; ventral surface smooth. Carpus with dorsal surfaces covered with small, well-spaced spines or tubercles similar to dorsal surface of palm; ventrolateral margin rounded, with small spines or tubercles; dorsodistal margin with row of small spines; ventromesial distal margin with row of spines, 1 or 2 stronger. Merus with surfaces nearly smooth, unarmed except for dorsodistal spine, and row of small spines on ventromesial distal margin. Ischium unarmed except for setae on dorsal margin. Coxa unarmed except for ventromesial row of long setae. Left cheliped (Figure 37D ) well calcified. Fingers terminating in sharp corneous, inwardly curved claws; dorsal and ventral surfaces unarmed except for tufts of setae; cutting edge of dactyl with row of closely-set, small corneous teeth, cutting edge of fixed finger with row of small calcareous teeth. Dactyl about as long as palm. Palm with scattered small tubercles and tufts of setae on dorsal surface; dorsomesial margin with irregular row of tufts of setae. Carpus with long setae and row of small spines on dorsal margin, and strong dorsodistal spine; dorsolateral surface with scattered small tubercles or spines; mesial and ventral faces smooth. Merus unarmed except for few setae on dorsal and ventral surfaces, and 2 or 3 spines on ventrolateral distal margin. Ischium unarmed except for scattered dorsal and ventral setae. Coxa unarmed except for ventromesial row of long setae. Ambulatory legs (Figure 40A-D) similar from right to left, reaching to about tip or slightly beyond tip of extended right cheliped. Dactyl long, approximately 1.8 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of bristle-like setae, and ventromesial row of 8-18 corneous spines. Merus, carpus and propodus with scattered short setae on dorsal margins. Carpus with small dorsodistal spine. Ischium and coxa unarmed; with ventromesial setae. Anterior lobe of sternite XII (of third pereopods) rounded, setose (Figure 40E). Fourth pereopod (Figure 41A, B) subchelate. Dactyl elongate, broadly curved, terminating in sharp corneous claw that is short in males, and slender and long (more half length of dactyl) in females; with ventrolateral row of small corneous spinules. Propodus longer than wide, rasp consisting of 1 row of rounded scales at least distally. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 41C) chelate. Propodal rasp extending to mid-length of segment. Uropods (Figure 41F, G) and telson (Figure 41E) markedly asymmetrical. Telson lacking or at most obsolete lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 broadly rounded projections by narrow, shallow U-shaped cleft; rounded projections each armed with row of corneous spines, those on left projection with some strong, curved spines; armature of rounded projections varying with sex, being stronger and with more numerous spines in females than in males. Male with paired first and second gonopods well developed (Figure 37E, F). Female with rudimentary second right pleopod. ParagioPagurus: wordldwide review and new species 395 COLOUR (Figure 35b-D) — Shield mostly whitish with orangish tint (darker orangish tint on each side medially). Ocular peduncle light purple fading near cornea. Antennular peduncle purple, with whitish flagellum. Antennal peduncle and flagellum whitish. Chelipeds with fingers cream white; dorsal surface of carpus and chela dark orange or reddish dor- somesially, with iridescent area fading to white dorsolaterally. Ambulatory legs whitish except for orangish tips of dactyls, scattered small orangish spots dorsally on propodi, and orangish dorsal surface and ventral margins of carpi and ventro- distal surface of meri. HABITAT — Gastropod shell with small actinian attached on shell opening and positioned under hermit crab body (Figure 35D). REMARKS — A careful examination of the specimen used in the colour photograph listed as Paragiopagurus cf. diogenes in McLaughlin et al. (2010: fig. 19D) showed that it represents this new species, P. laperousei n. sp. Paragiopagurus laperousei n. sp. is most similar to three species that occur in French Polynesia, P. fasciatus, P. bougainvillei, and P. wallisi, and to a lesser extent a species broadly distributed in the Indo-Pacific, P. diogenes. The colorations of these five species, however, are strikingly different, and can easily be used to separate them (Figures 1I, 35A-D, E, F). There is some similarity in the general coloration of the chelipeds of P. laperousei n. sp. and P. diogenes; however, otherwise the FIGURE 39 Carpus and chela of right cheliped. A-C, Paragiopagurus laperousei n. sp., holotype male (4.9 mm) UF 12304, Hawaiian islands, French Frigate Shoals, 215- 228 m. D, P. bougainvillei (lemaitre, 1994), paratype male (11.8 mm) USNm 265394, French Polynesia, tuamotu archipelago, 280 m. E, P. wallisi (lemaitre, 1994), paratype male (11.4 mm) USNm 265393, French Polynesia, tuamotu archipelago. A, D, E, dorsal view. B, lateral view. C, mesial view. Scale bars 3 mm for a; 2 mm for b, C; 5 mm for D, e. ED C A B rafael lemaitre396 FIGURE 40 Paragiopagurus laperousei n. sp., holotype male (4.9 mm) UF 12304, Hawaiian islands, French Frigate Shoals, 215-228 m. A, second right pereopod, lateral view. B, dactyl of same, mesial view. C, third right pereopod, lateral view. D, dactyl of same, mesial view. E, anterior lobe of sternite Xii (of third pereopods), ventral view. Scale bars 2 mm for a-D; 0.5 mm for e. E A-D D E C B A ParagioPagurus: wordldwide review and new species 397 FIGURE 41 Paragiopagurus laperousei n. sp. A, C, D, F, G, holotype male (4.9 mm) UF 12304, Hawaiian islands, French Frigate Shoals, 215-228 m. B, E, paratype ovigerous female (6.6 mm) UF 12303, Hawaiian islands, French Frigate Shoals, 215-228 m. A, B, propodus and dactyl of right (a) and left (b) fourth pereopods, lateral view. C, propodus and dactyl of right fifth pereopod, lateral view. D, E, telson, dorsal view. F, G, left (F) and right (G) exopod of uropods, dorsal view. Scale bars 0.5 mm for a-C; 1 mm for D-G. D-G B A-C BA C E GDF rafael lemaitre398 color of the ocular peduncles (light purple in P. laperousei n. sp., reddish in P. diogenes) and ambulatory legs (white in P. laperousei n. sp., light orange with dark red band proximally on each carpus in P. diogenes), is markedly different in these two species. The ocular peduncles, shield, and ambulatory legs have dark red stripes in P. fasciatus; the carpi of the left cheliped and ambulatory legs have a dark spot on the lateral face distally, and the ambulatory legs are overall yellowish with numerous, small red spots in P. bougainvillei; the right cheliped is reddish with white tubercles, and the ambulatory legs are light pink in P. wallisi. Other than coloration, these five species can be distinguished primarily by the armature of the chelipeds, although the armature of the telson and shape of the gill lamellae can be useful as well. The cheliped armature is strongest in P. fasciatus (Figure 34C-E), particularly on the left cheliped, and weakest in P. diogenes (Figures 36C, 15A, B). This difference in armature can be used to separate these two species from each other as well as from P. laperousei n. sp., P. bougainvillei, and P. wallisi. The cheliped armature is intermediate in strength between P. fasciatus and P. diogenes, in P. laperousei n. sp. (Figures 37D, 39A), P. bougainvillei (Figures 39D), and P. wallisi (Figures 39E). This new species is distinguished from P. bougainvillei and P. wallisi by subtle characters, such as the shape of the gill lamella (undivided distally in the new species, divided distally in P. bougainvillei and P. wallisi), the shape of the spine or tubercles of the right cheliped (weak and well spaced in the new species, distinct and mammilliform in P. bougainvillei and P. wallisi). A photograph (Figure 35D) of a hermit crab collected from French Frigate Shoals in the northwestern Hawaiian Islands, identified as Paragiopagurus diogenes or by the common names “blue-eyed hermit crab” or “anemone crab”, has been prominently used in various books and reports from the Census of Marine Life project (e.g., Crist et al. 2009; Knowlton 2010). Comparison of this photograph with others of specimens examined of P. laperousei n. sp., have shown that the Census of Marine Life photograph is actually of a specimen of this new species. ETYMOLOGY — The name of this new species honors the French explorer Jean François de Galaup, comte de Lapérouse (23 August 1741-1788?), whose famous expedition left Botany Bay, Australia in 1788 and vanished in Oceania. The name French Frigate Shoals, where specimens of this new species were collected and photographed, also commemorates this famous explorer who nearly lost two frigates while attempting to navigate these shoals in 1786. Furthermore, the specific name is appropriate given that this new species is morphologically closest to two congenerics, Paragiopagurus bougain- villei and P. wallisi, that are also named for famous explorers. Paragiopagurus wallisi (Lemaitre, 1994) Figures 35e, 39e, 42 Sympagurus wallisi lemaitre, 1994: 392, figs 13-15, 27h-i, 28f (type locality : tuamotu, French Polynesia). — Poupin, 1996a: 20, pl. 9-fig. g; 1996b: 96. — Parin et al., 1997: 163. Paragiopagurus wallisi – lemaitre, 1996: 207. — Zhadan, 1997: 69. — mclaughlin et al., 2010: 39, pl. 19G. TYPE MATERIAL — Holotype, French Polynesia, Tuamotu, Vanavana, stn 331, 20°45.7’S, 139°10.1’W, 240 m, 28.10.1990, male 11.0 mm (MNHN-Pg 5144). For paratypes from French Polynesia, including specimen photographed in Figures 35E and 42A, see Lemaitre (1994 as Sympagurus wallisi). MATERIAL EXAMINED — the type material (see above). French Polynesia. beNtHaUS: stn CaS 1931, 24°39,08’S, 146°01,58’W, 200 m, 13.11.2002, 1 female 11.1 mm (mNHN-Pg 6915). DISTRIBUTION — Known so far only from the Tuamotu Archipelago, French Polynesia, and Nazca and Sala-y-Gómez Ridges. Depth: 200 to 280 m. ParagioPagurus: wordldwide review and new species 399 FIGURE 42 Paragiopagurus wallisi (lemaitre, 1994). A, paratype male (11.5 mm) USNm 265393, French Polynesia, tuamotu archipelago. B-G, I, holotype male (11.0 mm) mNHN-Pg 5144, French Polynesia, tuamotu archipelago, stn 331, 240 m. H, paratype female (11.4 mm) USNm 265393, French Polynesia, tuamotu archipelago, stn 231. A, lamella of posterior arthrobranch. B, shield and cephalic appendages, dorsal view. C, right antennular peduncle and anterior portion of branchiostegite, lateral view. D, left cheliped, dorsal view. E, third left pereopod, lateral view. F, dactyl of same, mesial view. G, H, propodus and dactyl of left fourth pereopod in male (G) and female (H). I, telson, dorsal view. Scale bars 1 mm for a; 5 mm for b, D; 2 mm for C; 6 mm for e, F; 3 mm for G-i. E-I C B, D A E A B F G HID C rafael lemaitre400 DIAGNOSIS — Gill lamellae weakly divided distally (Figure 42A). Shield (Figure 42B) about as broad as long; dorsal surface usually weakly calcified medially and anteriorly on half or more of surface; rostrum broadly rounded, often obsolete, with short, low mid-dorsal ridge; lateral projections subtriangular, with small terminal spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, terminating in strong spine; corneas slightly dilated. Epistomial spine straight, strong. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment. Antennal peduncles (Figure 42C) exceeding distal margins of corneas by about 0.3 length of fifth segment; fourth seg- ment with strong spine on dorsolateral distal angle; second segment with dorsolateral distal angle produced, terminating in strong, occasionally bifid spine; acicle slightly curved outward (in dorsal view), usually not reaching to distal margin of cornea, terminating in strong spine, and with mesial margin armed with 2-6 strong spines; flagellum with short setae. Chelipeds strongly dissimilar. Right cheliped (Figure 39E) massive, operculate, with scattered short setae, dorsal surface of palm and carpus with some iridescence medially; chela longer than broad; fingers curved ventromesially, armature on dorsal surfaces stronger than on rest of chela, covered with strong, mammilliform tubercles or spines; dactyl set at stron- gly oblique angle to longitudinal axis of palm, with ventromesial face concave; palm about as long as broad, dorsal surface covered with well-spaced tubercles or spines, dorsolateral margin well delimited by row of blunt to sharp spines, mesial face rounded and with strong spines or tubercles; carpus with spines or tubercles on dorsal surface, dorsodistal margin with row of sharp spines. Left cheliped (Figure 42D) usually well calcified, dorsal surfaces of carpus and palm with some iridescence medially; palm with scattered setae and small tubercles or spines on dorsal surface; carpus with scattered small spines on dorsomedial surface, row of small spines on dorsal margin, and strong dorsodistal spine. Ambulatory legs (Figure 42E, F) with dactyls each having ventromesial row of about 15 corneous spinules, and dorsal and dorsome- sial rows of long bristle-like setae; carpus with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) usually unarmed or occasionally with small spines. Fourth pereopod (Figure 42G, H) with merus, carpus and propodus with low, blister-like tubercles on lateral faces; dactyl terminating in sharp, corneous claw (longer and more slender in females than in males); propodal rasp consisting of 1 row of ovate scales. Uropods and telson strongly asymmetrical. Tel- son (Figure 42I) with weak transverse suture separating anterior and posterior lobes; dorsal surface with low, blister-like tubercles; posterior lobes separated by narrow V-shaped, median cleft; terminal margins armed with curved corneous spines. Male with paired first and second gonopods well developed; first gonopod with concave distal lobe; second gono- pod with low, blister-like tubercles on posterior face of distal and basal segments, distal segment setose on distomesial face and bristle-like setae on mid-lateral margin. Female with vestigial right second pleopod. COLOUR — “Shield orange-brown medially, cream yellow elsewhere. Ocular peduncles amber, cornea black centrally. Cephalic appendages and ocular acicles whitish. Right cheliped with white tubercles and spines on reddish dark-pink background (bases of tubercles and spines darker than remaining surfaces); mesial face of palm and entire dorsal surface of carpus with darker colouration than other surfaces of cheliped. Left cheliped with chela red-spotted (spots at bases of tufts of setae) on whitish background; carpus with spines on reddish background. Second to fifth pereopods more or less uniformly pinkish.” (Lemaitre 1994: 396). HABITAT — Found living in gastropod shells with a small unidentified actinian growing on the shell. REMARKS — See REMARKS under P. laperousei n. sp. and P. bougainvillea. Paragiopagurus bougainvillei (Lemaitre, 1994) Figures 35F, 39D, 43 Sympagurus bougainvillei lemaitre, 1994: 397, figs 16-19, 27j, 28g (type locality: tuamotu, French Polynesia). — Poupin, 1996a: 20, pl. 9- fig. b; 1996b: 96. Paragiopagurus bougainvillei – lemaitre, 1996: 207. — mclaughlin et al., 2010: 39, pl. 19C. — Poupin et al., 2012 : 51. ParagioPagurus: wordldwide review and new species 401 TYPE MATERIAL — Holotype, French Polynesia, Tuamotu, Makemo, stn 308, 16°34.5’S, 143°39.9’W, 280 m, 07.10.1990, male 11.9 mm (MNHN-Pg 5148). Paratypes: French Polynesia. Tuamotu, Makemo: stn 308, 16°34.5’S, 143°39.9’W, 280 m, 07.10.1990, 2 males 11.8, 12.2 mm (USNM 265394). Marquesas Islands. Tahuata: stn 300, 09°54.5’S, 139°07.9’W, 190 m, 01.09.1990, 1 male 11.7 mm (MNHN-Pg 5149). MATERIAL EXAMINED — the type material (see above). marquesas islands. mUSOrStOm 9: stn DW 1235, 09°41.8’S, 139°03.5’W, 105-285 m, 31.08.1997, 1 male 10.7 mm (mNHN-Pg 6414). – stn DW 1288.8°53.9’S, 139°38.0’W, 200-220 m, 08.09.1997, 1 male 11.6 mm (mNHN-Pg 6413). For additional material examined see lemaitre (1994, as Sympagurus bougainvillei). DISTRIBUTION — South Central Pacific: so far known from the Marquesas Islands, and Tuamotu Archipelago, in French Polynesia. Depth: 105 to 285 m. DIAGNOSIS — Gill lamellae weakly divided distally (Figure 43A). Shield (Figure 43B) about as broad as long; dorsal surface usually weakly calcified on half or more of surface; rostrum broadly rounded, with short, low mid-dorsal ridge; anterior margins weakly concave; lateral projections subtriangular, with small subterminal spine. Ocular peduncles more than half length of shield; ocular acicles subtriangular, each terminating in strong spine; corneas slightly dilated. Epistomial spine absent. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment. Antennal peduncles (Figure 43C) slightly exceeding distal margins of corneas; third segment with strong ventromesial distal spine; second segment with dorsolateral distal angle produced, terminating in strong, usually bifid spine (occasionally with additional small spines dorsally); acicles reaching distal margins of corneas, mesial margin each armed with 6-8 strong spines; flagellum with short setae. Chelipeds strongly dissimilar. Right cheliped (Figure 39D) massive, operculate; chela longer than broad, sparsely setose, dorsal surface covered with well-spaced, small spines; fingers curved ventromesially; dactyl set at strongly oblique angle to longitudinal axis of palm, with ventromesial face concave; palm about as long as broad or slightly longer than broad in large specimens (sl > 11.0 mm), with dorsolateral margin well delimited by row of spines, mesial face rounded and with well-spaced spines or tubercles; carpus with small spines or tubercles on dorsal and ventral surfaces. Left cheliped (Figure 43D) usually well calcified; palm with dorsomesial row of small spines, and dorso- lateral face with few small tubercles or spines; carpus with dense setae dorsally, and strong dorsodistal spine. Ambulatory legs (Figure 43E, F) with dactyls each having ventromesial row of about 14 corneous spines, and dorsal and dorsomesial rows of long bristle-like setae; carpi each with small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) usually unarmed. Fourth pereopod (Figure 43G) with dactyl terminating in sharp corneous claw; propodal rasp consis- ting of 1 row of ovate scales. Uropopds and telson strongly asymmetrical. Telson (Figure 43H) with weak transverse suture separating anterior and posterior lobes; dorsal surface with inconspicuous low, blister-like tubercles; posterior lobes separated by narrow and shallow V-shaped median cleft, terminal margins armed with strongly curved corneous spines. Male with paired first and second gonopods; first gonopod with concave distal lobe; second gonopod with low, blister-like tubercles on posterior face of distal and basal segments, with distal segment setose on distomesial face and bristle-like setae on mid-lateral margin. Female with vestigial right second pleopod. COLOUR (Figure 35F) — Shield whitish to light yellow, except for light orange region anteriorly on each side of midline; with orange-red spots more numerous on calcified surfaces. Ocular peduncles light orange, cornea with light blue tinge. Cephalic appendages and ocular acicles whitish or light orange. Right cheliped with chela and merus whitish and with small orange spots; carpus orange with large dark orange-red spot on mesial face distally. Left cheliped with chela, car- pus, and merus whitish with small orange spots; carpus with large dark orange spot on lateral and mesial faces distally. Second to fifth pereopods whitish with small orange spots; carpi with large, dark orange-red spot on lateral and mesial faces distally. Abdominal tergites and telson with small orange spots. HABITAT — Found in gastropod shells with an unidentified actinian growing on the shells. rafael lemaitre402 FIGURE 43 Paragiopagurus bougainvillei (lemaitre, 1994). A, paratype male (11.8 mm) USNm 265394, French Polynesia, tuamotu archipelago, stn 308, 280 m. B-H, holotype male (11.9 mm) mNHN-Pg 5148, French Polynesia, tuamotu archipelago, stn 308, 280 m. A, lamella of posterior arthrobranch. B, shield and cephalic appendages, dorsal view. C, left antennal peduncle and anterior margin of branchiostegite, lateral view. D, left cheliped, dorsal view. E, left third pereopod, lateral view. F, dactyl of same, mesial view. G, propodus and dactyl of left fourth pereopod, lateral view. H, telson dorsal view. Scales bar 1 mm for a, 5 mm for b, D, F; 3 mm for C, e, G, H. A B, D C, H E, F G F H B D G C A E ParagioPagurus: wordldwide review and new species 403 REMARKS (see also P. laperousei n. sp.) — Morphologically, P. bougainvillei most closely resembles P. wallisi and P.  laperousei n. sp. As previously mentioned, these three species differ markedly in colouration (Figure 35D, E, F). If colour is not available, P. bougainvillei can be separated from P. wallisi by the shape and strength of the spines on the distal lobes of the telson. The lobes of the telson bear several large spines that are abruptly curved ventromesially in P.  bougainvillei, whereas the spines are small and weakly curved in P. wallisi. The separation of P. bougainvillei from P. laperousei n. sp. can be done by the shape of the gill lamella (distally divided in P. bougainvillei, undivided in P.  laperousei n. sp.) , and the shape of the tubercles on the fingers of the right cheliped (mammilliform in P. bougainvillei, weak and well spaced in P. laperousei n. sp.). Paragiopagurus ventilatus Lemaitre, 2004 Figures 35G, H, 44 Paragiopagurus ventilatus Lemaitre, 2004c: 326, figs 1-5 (type locality: Taiwan); 2006a: 435, figs 1, 2. — McLaughlin et al., 2007: 321, 2 unnumbered figs; 2010: 39, pl. 19F. — Wang, 2008: 760. — Komai et al., 2010: 2, figs 1-5. — Tsang et al., 2011: 5, figs 2, 4 (phylogenetic trees). TYPE MATERIAL — Holotype, Taiwan, TAIWAN 2001, stn CP 113, 24°50.8’N, 121°59.9’E, 281 m, 21.05.2001, male 5.6 mm (NTOU A00438). Paratypes, Taiwan. TAIWAN 2001: stn CP 81, 24°50.5’N, 121°59.9’E, 205 m, 08.05.2001, 1 male 5.6 mm (MNHN-IU-2008-15796). – stn CP 113, 2 males 5.4, 5.8 mm, 1 ovigerous female 3.8 mm (MNHN-IU-2008-15797), 1 male 5.4 mm, 2 ovigerous females 4.2, 4.4 mm (USNM 1068936), 6 male 4.5-5.7 mm, 1 ovigerous female 4.9 mm (NTOU A00439). – stn CP 114, 24°51.0’N, 121°51.3’E, 128-250 m, 21.05.2001, 1 male 5.1 mm (MNHN-IU-2008-15798). MATERIAL EXAMINED — The type material (see above). For additional material see McLaughlin et al. (2007). DISTRIBUTION — Known from the northeastern coast of Taiwan (Lemaitre 2004) and the Mariana Trough (Komai et al. 2010). Depth: 128 to 410 m. DIAGNOSIS — Gill lamellae undivided distally. Shield (Figure 44A) approximately as broad as long; dorsal surface weakly calcified along midline and sometimes anterior portions; rostrum broadly rounded, with short mid-dorsal ridge; lateral projections subtriangular, with small terminal spine. Ocular peduncles more than half length of shield; corneas slight- ly dilated. Ocular acicles (Figure 44B) subtriangular, terminating in strong, simple spine (rarely bifid) with transverse striae. Antennular peduncles exceeding distal margins of corneas by nearly entire length of ultimate segment. Antennal peduncles (Figure 44C) reaching to about distal margin of corneas; fourth segment unarmed, lacking spine on dorsola- teral distal angle; second segment with dorsolateral distal angle produced, terminating in strong, simple spine reaching slightly beyond midpoint of acicle; acicle nearly straight or weakly curved outward (dorsal view), reaching to distal mar- gin of cornea, terminating in strong spine, mesial margin armed with row of 4-6 spines (rarely 8 or 9) diminishing in size distally; flagellum with short setae < 1 to 2 flagellar articles in length. Maxillule with internal lobe with long terminal seta. Third maxilliped stout; with basis, ischium merus, carpus, propodus, and dactyl about same length (measured along lateral margin); outer face of basis, and ischium, with long, dense, plumose bacteriophore setae denser on larger male specimens (sl > 5.0 mm); crista dentata with about 16 corneous-tipped teeth. Epistome usually with small, straight simple to trifid epistomial spine, or rarely unarmed. Chelipeds markedly dissimilar; proportions and armature of right strongly affected by size and sex (see Remarks). Right cheliped about 3 times as long as sl in females (Figure 44D), or up to 6.9 times as long as sl in males; chela somewhat dorsoventrally flattened in females, less so in males, dorsal and ventral surfaces with numerous blunt to sharp spines or tubercles, most with tuft of setae near base anteriorly; fingers slightly curved ventromesially; dactyl shorter (males) or longer (females) than mesial margin of palm, set at weakly oblique angle to longitudinal axis of palm, ventromesial face concave; palm longer than broad in males or broader than long in females, rafael lemaitre404 G E, F D C B A F E A G D B C ParagioPagurus: wordldwide review and new species 405 lateral margin more distinctly delimited by spines or tubercles in females than in males, mesial face rounded, dorsomesial margin distinctly delimited in females by row of spines or tubercles; carpus armed similarly to chela, length/width ratio ranging from 1.2 in females to as much as 2.7 in males; dorsodistal margin with row of strong (females) or weak (males) spines; ventrolateral margin well delimited by spines (females) or not delimited (males); mesial surface rounded, with small tubercles, ventromesial distal margin with row of small blunt spines; merus armature and setation similar to that of carpus, with mesial and ventral surfaces with dense, long plumose bacteriophore setae, more dense in large males; ischium with dense, long plumose bacteriophore setae on distomesial and dorsal surfaces. Left cheliped (Figure 44A) well calcified; palm with tufts of setae and small tubercles on dorsal surfaces; carpus with irregular rows of small spines or tubercles obscured by tufts of setae on dorsal surface, and 1 or 2 strong dorsodistal spines; merus with dense, long bacteriophore setae on proximomesial and ventral surfaces; ischium usually with dense, long bacteriophore setae on dis- tomesial and dorsal surfaces. Ambulatory legs (Figure 44A, E, F) with dactyls having tufts of long setae on dorsal margins and 18-40 corneous spines (more numerous on third pereopod) arranged irregularly on ventromesial surface; propodi, carpi, and meri with tufts or short transverse rows of setae on all surfaces; carpi each with small dorsodistal spine; meri with ventrodistal row of small spines or tubercles (second pereopod), or lacking tubercles (third pereopod), ventral margin with long bacteriophore setae proximally, more dense on second pereopod than on third; ischia with long bac- teriophore setae on ventral margins. Anterior lobe of sternite XII (third pereopods) subsemicircular, setose, with strong subterminal simple or more often, bifid spine. Fourth pereopod with propodus longer than high, rasp formed of 1 row (at least distally) of ovate scales. Uropods and telson markedly asymmetrical. Telson (Figure 44G) with weak or obsolete transverse suture; posterior margin separated by broad, shallow cleft, into rounded projections armed with relatively few short, corneous spines, some often ventrally curved. Male lacking first gonopods; with unpaired left second gonopod rudimentary, uniramous. Female with rudimentary second right pleopod. COLOUR (Figure 35G, H) — “In life, shield and cephalic appendages cream with some faded pink areas. Chelipeds and ambulatory legs red or pink; propodi and dactyls of ambulatory legs each with faded pink or white stripe on lateral face” (Lemaitre 2004c: 331). HABITAT — Has been found living in gastropod shells of the buccinid genus Siphonalia, and siboglonid worm tubes of Lamellibrachia satsuma. Paragiopagurus ventilatus is believed to be associated with chemosynthetically-based vent eco- systems, although it probably does not live exclusively in such vent systems (Lemaitre 2004). REMARKS — As described by Lemaitre (2004c), morphological variations related to sex and size affect primarily the right cheliped in this species. The elongation of the right cheliped is particularly pronounced in specimens sl > 5.0 mm. The right chela in females (Figure 44D) is more dorsoventrally flattened than in males, the former having the dorsolateral and dorsomesial margins of the chela well defined by rows of spines, whereas in the latter the dorsolateral and dorsomesial margins are weakly defined by low tubercles. The bacteriophore setae on the outer surfaces of bases and ischia of the third maxillipeds, the mesial surfaces and ventral margins of the ischia and meri of the chelipeds, and the ventral margins of ischia and meri of the second and third pereopods, are more dense in large males (sl > 5.0 mm). Additional observations on variations were provided by Komai et al. (2010), who also noted that the bacteriophore setae are absent in juveniles. FIGURE 44 Paragiopagurus ventilatus lemaitre, 2004. A-C, E-G, holotype male (5.6 mm) NtOU a00438, taiwan, taiWaN stn CP 113, 281 m. D, paratype ovigerous female (4.9 mm) NtOU a00439, taiwan, taiWaN stn CP 113, 281 m. A, entire specimen except for abdomen, and carpi, propodi and dactyl of left second and third pereopods. B, ocular acicles and rostrum, dorsal view. C, right antennal peduncle, lateral view. D, right cheliped, dorsal view. E, F, dactyls of right second (e) and right third (F) pereopods, mesial view. G, telson, dorsal view. Scale bars 2 mm for a; 0.5 mm for b, C, G; 1 mm for D-F. rafael lemaitre406 Paragiopagurus ventilatus can be separated from other species of Paragiopagurus by the presence of bacteriophore setae on the third maxillipeds, chelipeds, and second and third pereopods in subadult to adult; the striae on the dorsal surface of the spines of the ocular acicles; the irregular rows of spines on the ventromesial margins of the dactyls of second and third pereopods; and the bifid spine on the anterior lobe of sternite XII (of third pereopods). There are seven other Paragiopagurus species in which males lack first gonopods and have unpaired rudimentary left second gonopod like in P. ventilatus: P. acutus, P. bicarinatus, P. hirsutus, P. hobbiti, P. orthotenes n. sp., P. trilineatus n. sp., and P. ruticheles. Paragiopagurus hobbiti (Macpherson, 1983) Figure 45 Parapagurus hobbiti macpherson, 1983a: 472, figs 1-3 (type locality: Valdivia bank); 1984: 86. Sympagurus hobbiti – lemaitre, 1989: 37; 1990: 231, fig. 8; 1994: 412. Paragiopagurus hobbiti – lemaitre, 1996: 207. — mclaughlin et al., 2010: 39. TYPE MATERIAL — Holotype, Valdivia Bank, off coast of Namibia, Valdivia 1, stn 7, 15, 18, 23 (all with same locality data), 26°11’S, 06°12’E, 234-242 m, 21.05.1982, male 2.2 mm (ICMD 665/1991). Paratypes, Valdivia Bank. Valdivia 1: stn P-7, 4  males 2.5-3.0 mm, 4 ovigerous female 1.8-2.2 mm (ICMD 670/1991), 1 male 2.2 mm, 1 ovigerous female 2.5 mm (USNM 240165). MATERIAL EXAMINED — the type material (see above). For additional material see macpherson (1984). DISTRIBUTION — Southeastern Atlantic: known so far only from the Valvivia Bank, off the coast of Namibia, at a depth of about 250 m. DIAGNOSIS — Gill lamellae undivided. Shield (Figure 45A) about as long as broad; dorsal surface weakly calcified me- dially and anteriorly; rostrum broadly rounded, with low dorsal ridge; lateral projections subtriangular, often terminating in small spine. Ocular peduncles about 0.75 length of shield; ocular acicles subtriangular, each terminating in strong, forwardly directed spine. Antennular peduncles exceeding distal margins of corneas by about 0.75 length of ultimate segment. Antennal peduncles (Figure 45B) usually not exceeding distal margins of corneas; fourth segment armed with strong spine on dorsolateral distal angle; first segment with small spine on lateral face distally; acicle not exceeding distal margin of cornea, terminating in strong spine, mesial margin armed with 6 or 7 spines. Epistomial spine short, straight. Right cheliped (Figure 45C, D) with carpus and chela armed dorsally with scattered small spines or low tubercles; fingers weakly curved ventromesially; palm with mesial face strongly sloping; dorsolatereal margin more or less well delimited by row of tubercles or spines. Left cheliped (Figure 45E) usually well calcified; chela with dorsomesial row of small spines; carpus with dorsal row of small spines and dorsodistal spine. Ambulatory legs (Figure 45F, G) with dactyls each having ventromesial row of 6-12 minute spinules, and dorsal and distal dorsomesial rows of setae; carpi each with dorsal row of 4-7 small spines and small dorsodistal spine. Anterior lobe of sternite XII (of third pereopods) armed with 1 spine. Fourth pereopod (Figure 45H) with propodal rasp consisting of 1 row of ovate scales. Uropods strongly asymmetrical. Telson (Figure 45I) moderately asymmetrical; with terminal margin divided into unequal lobes by broad, shallow U-shaped cleft, lobes armed with few corneous spines (larger, and more strongly curved on left than on right lobe). Male lacking first gonopods, and with unpaired vestigial second left gonopod. Female lacking second right pleopod; ovigerous with 23-31 eggs about 0.6-0.8 mm in diameter. COLOUR — “White to pale cream, eyes black” (Macpherson 1983a: 475). ParagioPagurus: wordldwide review and new species 407 HABITAT — Found occupying gastropod shells, and living inside siliceous sponges, Pachastrella monolifera Schmidt, 1868 (Macpherson 1984). REMARKS — Subsequently to Macpherson’s (1983a) description of this species, additional material was reported by Mac- pherson (1984) from the type locality, Valdivia Bank, all from stn P-15. FIGURE 45 Paragiopagurus hobbiti (macpherson, 1983), paratype male (2.2 mm) USNm 240165, off Namibia, Valdivia bank, ValDiVia 1 stn P-1. A, shield and cephalic appendages. B, right antennal peduncle, lateral view. C, right cheliped. D, chela of same, mesial view. E, left cheliped. F, right second pereopod, lateral view. G, dactyl of same, mesial view. H, propodus and dactyl of left fourth pereopod, lateral view. I, telson, dorsal view. Scale bars 1 mm for a; 0.5 mm for b, H, i; 2 mm for C-e; 1 mm for F, G. (adapted in part from lemaitre 1990). B, H, I F, G E C, D A A G F CE B H I D rafael lemaitre408 Paragiopagurus pilimanus (A. Milne-Edwards, 1880) Figures 35i, 46 [For a complete synonymy see lemaitre (1989: 46)] Eupagurus pilimanus a. milne-edwards, 1880: 43 (in part; lectotype locality: Guadeloupe). Sympagurus pilimanus – a. milne-edwards & bouvier, 1893: 63, pl. 5-figs 8-20. — lemaitre, 1989: 37, 46, figs 20-23; 1994: 412. — boschi, 2000: 128. Parapagurus pilimanus – de Saint laurent, 1972: 105. — Williams et al., 1989: 32. — Gaytán Caballero & escobar briones, 2008 : 339. Paragiopagurus pilimanus – lemaitre, 1996: 207. — mclaughlin et al., 2005: 246; 2010: 39. — Campos et al., 2005: 140, figs 101, 102. — Felder et al., 2009: 1071. — martínez Campos et al., 2012 : 240. TYPE MATERIAL — Lectotype, western Atlantic, United States Coast Survey Steamer “Blake”, stn 167, 16°09.40’N, 61°29.25’W, 315 m, 29.01.1879, male 9.4 mm (MCZ 4013). Paralectotype, western Atlantic, United States Coast Survey Steamer “Blake”: stn 148, 17°17.12’N, 62°46.43’W, 374 m, 14.01.1879, 1 female 7.0 mm (MCZ 6329) MATERIAL EXAMINED — the type material (see above). Caribbean Sea. Off Curaçao Sea aquarium, Willemstaad, CUraSUb: stn 12-03, 12°05.078’N, 68°53.788’W, 144.8-182.8 m, 05.28.2012, 1 female 3.9 mm (USNm 1190206, colour voucher number CUri 12039). For additional mate- rial examined, including specimens illustrated in Figure 44b-F, J, see lemaitre (1989, under Sympagurus pilimanus). DISTRIBUTION —Western Atlantic: from Bermuda, the Bahamas, Straits of Florida and Gulf of Mexico, to the southeas- tern Caribbean. Depth: 36-2034 m. DIAGNOSIS — Gill lamellae undivided distally. Shield (Figure 46A) about as long as broad; dorsal surface usually weakly calcified medially on posterior half, with numerous, low blister-like tubercles; rostrum rounded, with long, often subdi- vided mid-dorsal ridge; lateral projections broadly subtriangular, unarmed. Ocular peduncles more than half length of shield; ocular acicles subtriangular, each terminating in strong spine; corneas slightly dilated. Maxillule with internal lobe of endopod bearing up to 6 long setae. Epistome unarmed. Antennular peduncles exceeding distal margins of corneas by less than half length of ultimate segment. Antennal peduncles usually not exceeding distal margins of cornea; second segment with dorsolateral distal angle produced, terminating in strong simple spine reaching to midpoint of antennal acicle; acicle not exceeding distal margin of cornea, mesial margin with 3 or 4 strong spines; flagellum with series of 2 or 3 long setae every 10-20 flagellar articles. Chelipeds markedly dissimilar, surfaces with moderately dense to dense setae. Right cheliped (Figure 46B, C) massive, with palm about as broader as long, fingers weakly curved ventromesially, dactyl set at weakly oblique angle to longitudinal axis of palm; palm with several irregular longitudinal rows of weak to strong spines on dorsal surface, mesial and lateral faces rounded and armed with weak to strong spines, ventral face unarmed or with scattered small tubercles; carpus with several irregular longitudinal rows of spines or tubercles on dorsal surface. Left cheliped (Figure 46D) well calcified, or occasionally carpus with weakly calcified lateral face; with 3 or more strong spines dorsally on carpus, including often strong dorsodistal spine. Ambulatory legs (Figure 46E, F) with dactyls each armed ventromesially with 6-18 minute or small spinules, and dorsal and dorsomesial rows of long setae; carpi each with 1-3 dorsodistal spines. Anterior lobe of sternite XII (of third pereopods) setose, armed with 1 strong subterminal spine. Fourth pereopod (Figure 46G) with propodal rasp consisting of 2 or 3 regular or irregular rows of ovate scales. Uropods and telson (Figure 46H) markedly asymmetrical, with low, blister-like tubercles on dorsal surfaces; telson with weak transverse suture separating anterior and posterior lobes, latter separated by shallow, V-shaped sinus, lobes with weak and strong, often ventrally curved, corneous spines on terminal margins. Males with paired first and second gonopods well developed; distal lobe of first gonopod (Figure 46I) with concave mesial face; second gonopod (Figure 46J) frequently with rudimentary exopod; small individuasl (sl < 5.0 mm) commonly with rudimentary first pleopods and poorly deve- loped second pleopods. Females with vestigial right second pleopod. ParagioPagurus: wordldwide review and new species 409 FIGURE 46 Paragiopagurus pilimanus (a. milne-edwards, 1880). A, G-I, lectotype male (9.4 mm) mCZ 4013, Guadeloupe, blake stn 167, 315 m. B-F, female (7.6 mm) Umml, Straits of Florida, GerDa stn 637. J, male (8.3 mm) Umml 32:4418, Straits of Florida, GerDa stn 929. A, shield and cephalic appendages, dorsal view. B, right cheliped, dorsal view. C, same, mesial view. D, left cheliped, dorsolateral view. E, left second pereopod, lateral view. F, dactyl of same, mesial. G, propodus and dactyl of left fourth pereopod, lateral view. H, telson, dorsal view. I, left first gonopod, mesial view. J, left second gonopod, anterior view. Scale bars 5 mm for a; 2 mm for b-F; 3 mm for G, H; 1 mm for i, J. (adapted in part from lemaitre 1990). I, J H G B-F A D C BHIJ E G FA rafael lemaitre410 COLOUR (Figure 35I) — Antennular and antennal peduncles mostly white, or faintly yellow distally on ultimate antennular and fifth antennal segments, and flagella. Ocular peduncles white proximally, and dark orange distally. Epistomial region light orange. Right chelipeds with merus and chela light yellow with white spines; left cheliped mostly white or with faint yellow on mesial face of merus and carpus. Second and third pereopods white except for light orange tinge on mesial faces of meri and carpi. HABITAT — Gastropod shells. REMARKS — A description of the morphological variations of this exclusively western Atlantic species, and taxonomic remarks, can be found in Lemaitre (1989). Paragiopagurus tuberculosus (de Saint Laurent, 1972) Figures 47, 48 Parapagurus tuberculosus de Saint Laurent, 1972: 118, figs 11, 25 (type locality: NW of Laysan, Hawaiian Islands). Sympagurus tuberculosus – Lemaitre, 1989: 37; 1994: 412. Paragiopagurus tuberculosus – Lemaitre, 1996: 207. — McLaughlin et al., 2005: 246; 2010: 39. — Castro, 2011: 30. TYPE MATERIAL — Holotype, Hawaiian Islands, NW of Laysan, Albatross, stn 3938, 25°52’17”N, 171°46’35”W, 270-300 m, 16.05.1902, female 3.6 mm (damaged) (USNM 168312). MATERIAL EXAMINED — The type material (see above). Hawaiian Islands. Mamala Bay, Oahu, BURCH: stn 80008, 21°15.4’N, 157°51.8’W, 402 m, 03.02.1980, 1 male 1.8 mm (BPBM S10984). – NOAA/NMFS “Townsend Cromwell”: stn TC 81-01, no detail station data, 369 m, 1 male 5.5 mm (BPBM S10995). Marquesas Islands. MUSORSTOM 9: stn DW 1146, 9°18.8’S, 140°06.2’W, 200 m, 22.08.1997, 5 males 2.0-3.9 mm, 3 ovigerous females 3.5, 3.8 mm, 2 females 2.8, 3.6 mm (MNHN-Pg 6902). – stn DW 1148, 9°18.9’S, 140°06.3’W, 300 m, 22.08.1997, 1 male 6.0 mm (MNHN-Pg 6904), 2 ovigerous females 3.8, 4.6 mm (MNHN-Pg 6903). – stn CP 1175, 8°45.1’S, 140°16.2’W, 300 m, 25.08.1997, 1 male 1.5 mm (MNHN-Pg 6905), 1 male 6.0 mm (MNHN-Pg 6906). – stn CP 1229, 9°44.3’S, 138°51.2’W, 310-320 m, 30.08.1997, 1 ovigerous female 3.6 mm (MNHN-Pg 6907). – stn CP 1268, 7°55.8’S, 140°42.6’W, 285-320 m, 04.09.1997, 1 male 3.9 mm, 2 ovigerous females 3.6, 4.5 mm, 3 females 3.0-3.9 mm (MNHN-Pg 6908). – stn DW 1281, 7°47.8’S, 140°20.8’W, 450-455 m, 07.09.1997, 1 male 3.6 mm, 3 females 4.3-5.2 mm (MNHN-Pg 6909). – stn CP 1282, 7°51.7’S, 140°30.6’W, 416-460 m, 07.09.1997, 4 males 5.1-5.4 mm, 4 females 4.0-4.6 mm (MNHN-Pg 6910). – stn DW 1287, 7°54.5’S, 140°40.2’W, 163-245 m, 07.09.1997, 5 males 1.9-5.6 mm, 2 females 3.3, 4.0 mm (MNHN-Pg 6911), 1 male 4.3 mm (MNHN-Pg 6913), 1 female 2.4 mm (MNHN-Pg 6912). – stn CP 1306, 8°55.2’S, 140°14.8’W, 283-448 m, 10.09.1997, 1 male 5.2 mm (MNHN-Pg 6914). DISTRIBUTION — Central Pacific: Hawaiian and Marquesas Islands. Depth: 163 to 460 m. REDESCRIPTION — Gill lamellae (Figure 47A) divided distally. Shield (Figure 47B) about as broad as long or slightly broa- der than long; dorsal surface weakly calcified along midline and anterior margins; with scattered short setae. Rostrum broadly rounded, with short mid-dorsal ridge. Anterior margins weakly concave. Lateral projections subtriangular, with small terminal spine. Anterolateral margins sloping. Ventrolateral margins without spine. Posterior margin broadly roun- ded. Anterodistal margin of branchiostegite rounded, unarmed, setose. Ocular peduncles more than half length of shield, each with longitudinal row of few, short setae on dorsal surface. Corneas dilated. Ocular acicles subtriangular, each terminating in strong simple spine; separated basally by about width of 1 acicle. Antennular peduncles exceeding distal margins of corneas by about 0.75 to 0.5 length of ultimate segment; ventral flagellum with about 8-10 articles. Ultimate segment about twice as long as penultimate, with scattered setae dorsally. ParagioPagurus: wordldwide review and new species 411 FIGURE 47 Paragiopagurus tuberculosus (de Saint laurent, 1972). A, male (4.8 mm) mNHN-Pg 6910, marquesas islands, mUSOrStOm 9 stn CP 1282, 416-460 m. B, C, G-I, male (3.9 mm) mNHN-Pg 6902, marquesas islands, mUSOrStOm 9 stn DW 1146, 200 m. D, male (6.0 mm) mNHN-Pg 6904, marquesas islands, mUSOrStOm 9 stn DW 1148, 300 m. A, lamella of posterior arthrobranch. b, shield and cephalic appendages. C, right antennal peduncle, lateral view. D, right cheliped, dorsal view. E, merus of same, mesial view. F, carpus and chela of right cheliped, dorsal view (from de Saint laurent 1972). G, left cheliped, dorsal view. H, left first gonopod, mesial view. I, left second gonopod, anterior view. Scale bars 0.25 mm for a; 2 mm for b, D, F; 1 mm for C, G, H, i; 3 mm for e. H, I G F E D C B A B A GH DI C EF rafael lemaitre412 Basal segment with strong ventromesial spine; lateral face with distal subrectangular lobe armed with 1 or 2 spines, and strong spine proximally. Antennal peduncles (Figure 47C) at most reaching to distal margins of corneas. Fifth segment unarmed, with few, short bristle-like setae on dorsolateral distal angle, and mesial margin. Fourth segment with 1 small spine on dorsolateral distal angle. Third segment with strong ventromesial distal spine. Second segment with dorsolateral distal angle produced, terminating in strong simple or multifid spine extending for about 0.3 length of acicle; mesial margin with small spine on dorsolateral distal angle. First segment with lateral surface armed with minute spine; ventromesial angle produced, with row of 3-5 spines laterally. Antennal acicle nearly straight or weakly sinuous (in dorsal view), at most reaching distal margin of cornea, and terminating in strong spine; mesial margin setose, with row of 7-10 strong spines diminishing in strength distally. Flagellum exceeding distal end of right cheliped; with few short setae less than 2 flagellar articles in length. G E, F A-D G F DC B E A ParagioPagurus: wordldwide review and new species 413 Mandible with 3-segmented palp. Maxillule with external lobe of endopod slender, moderately developed, internal lobe with long seta. Maxilla with endopod slightly exceeding distal margin of scaphognathite. First maxilliped with endopod exceeding distal margin of exopod. Second maxilliped without distinguishing characters. Third maxilliped with dactyl, carpus and propodus about 2.5 as long as wide; crista dentata with about 14 small teeth; coxa and basis each with small mesial tooth. Sternite IX (of third maxillipeds) with small spine on each side of midline. Epistomial spine short, straight. Chelipeds markedly dissimilar. Right cheliped (Figure 47D-F) elongate, much more so in large males (sl > 5.5 mm). Fingers straight or nearly so, each terminating in small, usually blunt, internally curved tip with corneous claw; dorsal surfaces with few setae; cutting edges each with row of calcareous teeth, that of dactyl also with short row of small corneous spines distally. Dactyl longer than length to palm, set at weakly oblique angle to longitudinal axis of palm; dorsal surface with numerous small spines on proximal half; ventral face with few small spines or tubercles; ventromesial face concave proximally. Fixed finger armed similarly to dactyl on dorsal and ventral faces. Palm proportions varying with sex from about 0.8 as broad as long in females, to about 1.4 as long as broad in males; dorsal surface densely covered with short, fine plumose setae, armed with blunt tubercles or spines or with scattered small spines or tubercles; dorsolateral and dorsomesial faces rounded, with small spines or tubercles; ventral surface sparsely setose, with scattered small spines or tubercles. Carpus with dorsal surface densely covered with short, fine plumose setae; dorsal, lateral, and mesial surfaces armed with numerous small spines or tubercles; lateral face rounded, mesial face strongly sloping; ventromesial and ventrolateral margins each with row of blunt or sharp spines. Merus with scattered small tubercles on dorsal margin; lateral surface nearly smooth; ventromesial and ventrolateral margins each with row of blunt spines; ventral face with small spines or tubercles and long, dense, mostly plumose and bristle-like setae. Ischium unarmed except for setae on dorsal margin and ventral surface; coxa with row of small tubercles or spines on ventrodistal margin, and ventromesial row of long setae. Left cheliped (Figure 47G) well calcified, or with weakly calcified dorsal surface of carpus. Fingers terminating in sharp corneous claws; dorsal and ventral surfaces unarmed except for tufts of setae; cutting edge of dactyl with row of closely- set or fused small corneous spinules, cutting edge of fixed finger with small calcareous teeth interspersed with small corneous spinules. Dactyl longer than palm. Palm with row of small blunt or sharp spines and long setae on dorsomesial margin; dorsal and dorsolateral surfaces with scattered tufts of setae. Carpus with dorsal margin armed with row of small spines, dorsodistal spine, and long setae. Merus with setae on dorsal margin, and row of small spines on ventrolateral and ventromesial distal margins. Ischium unarmed except for dorsal and ventral setae. Coxa with ventrodistal margin unarmed or with small spine mesially, and ventromesial row of long setae. Ambulatory legs (Figure 48A-D) similar from right to left, slightly exceeding right cheliped when fully extended. Dactyl approximately 1.5 times as long as propodus, terminating in sharp corneous claw; with dorsomesial and dorsolateral rows of setae, ventromesial margin armed with row of about 10-12 corneous spinules; mesial faces with shallow, longitudinal sulcus on proximal half. Propodus with short, bristle-like setae on dorsal margin; ventrodistal margin with scattered setae distally. Carpus with small dorsodistal spine, and short, bristle-like setae on dorsal margin. Merus unarmed except for short setae on dorsal margin; ventral margin unarmed or with scattered setae. Ischium and coxa unarmed except for short setae on dorsal margin and ventromesially. Anterior lobe of sternite XII (of third pereopods) setose, with small terminal spine. FIGURE 48 Paragiopagurus tuberculosus (de Saint laurent, 1972), male (cl 3.9 mm) mNHN-Pg 6902, marquesas islands, mUSOrStOm 9 stn DW 1146, 200 m. A, second right pereopod, lateral view. B, dactyl of same, mesial view. C, third right pereopod, lateral view. D, dactyl of same, mesial view. E, propodus and dactyl of fourth left pereopod, lateral view. F, propodus and dactyl of fifth left pereopod, lateral view. G, telson, dorsal view. Scale bars 2 mm for a-D; 0.5 mm for e, F; 1 mm for G. rafael lemaitre414 Fourth pereopod (Figure 48E) subchelate. Dactyl broadly curved, terminating in corneous claw; with ventrolateral row of small corneous spinules. Propodus longer than high, rasp consisting of 2 rows (at least distally) of ovate scales. Carpus with long setae on dorsal margin. Merus with rows of long setae on dorsal and ventral margins. Fifth pereopod (Figure 48F) chelate. Propodal rasp extending posteriorly to or beyond mid-length of segment. Uropods and telson markedly asymmetrical. Telson (Figure 48G) without or at most with weak lateral indentations, dorsal surface with scattered setae; posterior margins separated into 2 broadly rounded and moderately unequal projections by shallow U-shaped cleft; rounded projections each armed with row of often strong and curved, corneous spines. Male with paired first and second gonopods well developed; first gonopod (Figure 47H) with distal lobe subtriangular, and concave mesial face; second gonopod (Figure 47I) with distal segment setose on distomesial face, and with short, bristle-like setae on mid-lateral margin. Female lacking vestigial right second pleopod. COLOUR — Recently preserved specimens have the right cheliped orangish with white spines or tubercles. HABITAT — Found in gastropod shells, frequently completely covered by an unidentified actinian. A young male, sl 1.5 mm (MNHN-Pg 6905), was found inhabiting a piece of hollow wood. REMARKS — Previously, this species was known exclusively from the damaged, dismembered, and incomplete female holotype. In her description of this species, de Saint Laurent (1972) did mention under “Distribution” a lot from Albatross stn 4101, with only a right cheliped, presumably of this species; however, that lot has not been found. During this study a good number of specimens of P. tuberculosus were identified from various expeditions in the Hawaiian and Marquesas Islands, and has provided the opportunity to redescribed this species. Paragiopagurus tuberculosus stands out among its congeners by having stouter ocular peduncles and more dilated corneas than any other species in the genus. As is generally the case in many parapagurids, there is considerable sexual dimorphism in the shape of the right cheliped in P. tuberculosus. The shape of the right cheliped in this species is more elongated than in other congeners. The right cheliped tends to be more elongated and slender in males than in females of comparable size. In large males (sl > 5.5 mm), the chela and carpus can be as much as 2.7 times along as broad. The overall shape of the right chela is ovate in females, with lateral and mesial margins broadly arched, whereas the shape tends to be subrectangular in large males, with lateral and mesial margins nearly straight. Armature of spines and tubercles also varies with size and sex, being stronger and denser in females and males of comparable size, than in large males. DISTRIBUTION OF PARAGIOPAGURUS SPECIES The rich collection of specimens of Paragiopagurus species used for the present study has provided sufficient data to present a picture of the general vertical and horizontal distribution patterns of the species in this genus. However, it is important to consider that there are still vast areas with underwater features (e.g., seamounts, submerged ridges) and habitats of the world oceans that have not been adequately sampled or not sampled at all. Furthermore, the type of sampling gear used during deep-sea expeditions most often consists of trawls or dredges, or less frequently traps, and these can sample only certain types of bottoms and habitats. Thus, the information on the existence and distribution of species of this group of parapagurids is biased in favor of individuals that can be captured by these types of gear or those living in a narrow range of habitats. BATHYMETRIC (Figure 49) — The 24 species of Paragiopagurus occur almost exclusively on the continental shelf and slope, from 36 to 1000 m. A total of 12 species (50%) have a range that includes the Continental Shelf (< 200 m), and 10 species (42% %) occur exclusively on the Continental Slope (200-1000 m). Five species have been found beyond 1000 m on the Continental Rise: P. bicarinatus, P. hirsutus, P. pilimanus, P. ruticheles, and P. schnauzer. One of these five species, P. pili- manus, has the widest bathymetric range (36-2034 m) of all Parapagurus species; however, the finding of this species at the minimum and maximum depth of its range is a rare occurrence. ParagioPagurus: wordldwide review and new species 415 GEOGRAPHIC (table 1, Figure 50) — As expected, the highest concentration of Paragiopagurus species is found in the large and geologically older Indo-West Pacific region, where 15 species occur. Of those, 12 are found in the western Pacific but not in the Indian Ocean. The three species from the Indian Ocean, P. boletifer, P. diogenes and P. ruticheles, are known exclusively from the western side of that Ocean around Madagascar, although all three are also distributed in the western and central Pacific, and southeastern South Pacific (French Polynesia, and Nazca and Sala-y-Gómez Ridges). Seven of those 15 Indo-West Pacific species are known from the western Pacific, in an area that includes Japan, South China Sea, and Philippines; and 12 from an area in the South Pacific that encompasses the Solomon Islands to Samoa Islands, New Zealand, eastern and southern Australia, and Tasmania. So far, no species of Paragio- pagurus have been reported from the Indian Ocean other than from the western side, although this may simply be a result of poor sampling efforts. Surprisingly, nearly as many species (13) occur in the Central and eastern South Pacific from Hawaii, French Polynesia, and Nazca and Sala-y-Gómez Ridges, as in the western Pacific (15). Six species are known from Hawaii, eight from French Polynesia and the Marquesas Islands, and four from the Nazca and Sala-y-Gómez Ridges. Of those 13, it is striking that eight (62%) species are found exclusively in the Central and eastern South Pacific, whereas five (38%) are shared with the Indo-West Pacific region. It appears that the vast expanse between the faunistically rich western Pacific tropical region and French Polynesia does not represent a dispersal barrier for at least some Paragiopagurus species presumably have been able to reach the Hawaiian Islands, French Polynesia, and the Nazca FIGURE 49 bathymetric distribution of Paragiopagurus species from the world, with maximum and minimum depths indicated for each. P. ac ut us P. bo let ife r P. bi ca rin at us P. pa cifi cu s P. ox yc he lo s n . s p. P. or th ot en es n . s p. P. m ac ro ce ru s P. la pe ro us ei n. sp . P. in so lit us P. ho bb iti P. hi rsu tu s P. fa sc ia tu s P. di og en es P. bo ug ai nv ille i P. ru go su s P. pi lim an us P. ru tic he les P. sc hn au ze r P. sc ul pt oc he la P. tri lin ea tu s n . s p. P. tu be rcu lo su s n . s p. P. um bo na tu s n . s p. P. ve nt ila tu s P. wa llis i 200 280 0 500 1000 1500 2000 de pt h (m ) 128 281381 650 163 460 360 516 218 400 161 780 274 1070 85 806 105 285 40 695 112 200 221 1063 250 100 708 215 228 128 280 233 967 150 610 366 2034 360 221 100 1440 1200 1665 36 rafael lemaitre416 and Sala-y-Gómez Ridges. The presence of a high percentage of species endemic to the Central and eastern South Pacific, suggests that significant speciation among parapagurids has also occurred there. The Atlantic Ocean is poor in species of Paragiopagurus, with only four species occurring in this ocean. In the western Atlantic there is only one known species, P. pilimanus, distributed from Bermuda and off the eastern coast of the United States to the southeastern Caribbean. The eastern Atlantic has three species, P. hobbiti, P. macrocerus and P. ruticheles; the former two are known only from the southeastern Atlantic off western Africa, whereas the latter ranges broadly to the eastern Indian Ocean, western and Central Pacific, and southeastern South Pacific. No species of Paragiopagurus are known from the eastern Pacific. As previously mentioned in the systematic account, P. ruticheles has a puzzling and broad, disjunct distribution (Table 1). This species has been found in the eastern Atlantic, western Indian Ocean, off many of the tropical western Pacific islands, the central Pacific (Hawaiian Islands), and the eastern South Pacific (French Polynesia, Nazca and Sala-y-Gómez Ridges). The species, however, is absent from the eastern Pacific and western Atlantic. Individuals from widely separate TABLE 1 Summary of geographical distribution of Paragiopagurus species from the world. INDO-wEST PACIFIC CENTRAL + SOUTH PACIFIC In di an O ce an Ja pa n Ta iw an So ut h Ch in a Se a Ph ili pp in es M ar ia na Tr ou gh In do ne sia So lo m on W al lis & F ut un a Fi jii Va nu at u To ng a Ch es te rfi el d N ew C al ed on ia Au st ra lia N ew Z ea la nd H aw ai ia n Isl an d Jo hn st on A to ll M ar qu es as Fr en ch P ol yn es ia N az ca /S al a y G om ez Ea st er n Pa ci fic W es te rn A tla nt ic Ea st er n At la nt ic P. acutus • • • • • • • • • • P. bicarinatus • • • • • • • • • P. boletifer • • • • • • • • • • P. bougainvillei • • P. diogenes • • • • • • • • • • • P. fasciatus • P. hirsutus • • • • • • • • • • • P. hobbiti • P. insolitus • • P. laperousei n. sp. • P. macrocerus • P. orthotenes n. sp. • • • P. oxychelos n. sp. • • • • P. pacificus • P. pilimanus • P. rugosus • • P. ruticheles • • • • • • • • • • • P. schnauzer • P. sculptochela • P. trilineatus n. sp. • • • • • P. tuberculosus n. sp. • • P. umbonatus n. sp. • • • P. ventilatus • • • P. wallisi • • TOTAL SPECIES 3 4 7 5 4 1 2 7 2 3 7 7 2 11 4 2 6 2 4 7 4 0 1 3 ParagioPagurus: wordldwide review and new species 417 localities of this morphologically unique species are indistinguishable, even in colouration. One other parapagurid species, Strobopagurus gracilipes (A. Milne-Edwards, 1891), also has a similar disjunct distribution (Lemaitre 2004). Whether this disjunct distribution reflects the remnants of populations of widely distributed species during the geologic past, or can be attributed to an unexplained dispersal ability of these species, is difficult to ascertain at this time. ACKNOwLEDGEMENTS This study was inspired and made possible primarily by the vast and species rich collections gathered during French expeditions over the last four decades, and placed at my disposal during many visits to the MNHN in Paris, and through loans. Among the French collaborators, I am once again indebted to the leadership of A. Crosnier (formerly at MNHN) and P. Bouchet (MNHN), both of whom made sure that collection and data were perfectly organized to study them, facilitated working visits to their laboratories, and the latter for agreeing to deposit some duplicate materials at the USNM. I am also grateful at MNHN to D. Guinot, R. Cléva, Laure Corbari, and P. Maestrati, for their kind help in many museum matters, access to collections, and station data. Numerous other colleagues from other museums or institutions also helped in many unselfish ways, sharing photographs, data and/or information, and to them I extend my gratitude; they include: A. Asakura (Kobe University, Japan), K. Baba (formerly Kumamoto University, Japan), N. Budaeva (P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia), B. L. Burch (formerly BPBM), FIGURE 50 Generalized distributions of Paragiopagurus species from the world. 90° 70° 60° 40° 20° 60° 40° 20° 0° 60° 30° 0° 30° 60° 90° 120° 150° 180° 150° 120° 90° 90° 60° 30° 0° 30° 60° 90° 120° 150° 180° 150° 120° 90° 70° 60° 40° 20° 60° 40° 20° 0° 1 3 3 7 6 0 8 4 12 15 rafael lemaitre418 P. Castro (California State Polytechnic University, USA), T.-Y. Chan (NTOU), V. N. Ivanenko, T. Komai (Natural History Museum & Institute, Chiba, Japan), E. Macpherson (Centro de Estudios Avanzados de Blanes, Blanes, Spain), the late P. A. McLaughlin (Shannon Point Marine Center, Western Washington University, USA), T. N. Molodtsova (P. P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia), B. Richer de Forges (formerly IRD), G. Paulay (FLMNH), and J. Poupin (Institut de Recherche de l'École Navale, Brest, France). I am most thankful to photographers S. Middleton and B. B. Brown (Substation Curaçao), for giving permissions to use colour photographs of P. laperousei n. sp., and P. pilimanus, respectively. Figures 44B-F of P. pilimanus were made by B. Stolen. As usual, the help of Rose Gulledge in preparing the electronic versions of all figures, graphs, and table, as well as assistance in dealing with museum cataloguing, is gratefully acknowledged. This paper is posthumously dedicated to Patsy A. 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