Crustaceana 87 (2)

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1 Crustaceana 87 (2) A REVIEW OF THE COLLECTIONS OF THE INFRAORDERS THALASSINIDEA LATREILLE, 1831 AND CALLIANASSIDEA DANA, 1852 (DECAPODA, PLEOCYEMATA) LODGED IN THREE GERMAN MUSEUMS, WITH REVISED KEYS TO THE GENERA AND SPECIES BY KATSUSHI SAKAI 1,3 ) and MICHAEL TÜRKAY 2,4 ) 1 ) Shikoku University, , Tokushima, Japan 2 ) Senckenberg Forschungsinstitut und Naturmuseum, Senckenberganlage 25, D Frankfurt am Main, Germany ABSTRACT The collections of the infraorders Thalassinidea Latreille, 1831 and Callianassidea Dana, 1852 logded in three German Museums, viz., Zoologische Staatssammlung, Munich (ZSM), Zoologisches Museum der Universität Hamburg, Hamburg (ZMH), and Museum für Naturkunde, Berlin (ZMB) were examined. The results of the present study indicate that the material of these collections comprises 10 families, 31 genera, including one new genus, and 73 species, three of which are new to science. ZUSAMMENFASSUNG Die Sammlungen der Infraordnungen Thalassinidea Latreille, 1831 und Callianassidea Dana, 1852 dreier deutscher Museen, Zoologische Staatssammlung, München (ZSM), Zoologisches Museum der Universität Hamburg (ZMH) und Museum für Naturkunde, Berlin (ZMB), wurden studiert. Die Ergebnisse der vorliegenden Studie zeigen daß das Material dieser Sammlungen aus 10 Familien, 31 Gattungen (einschließlich einer neuen) und 73 Arten (einschließlich drei neuen) besteht. INTRODUCTION The infraorder Thalassinidea Latreille, 1831 was earlier considered to be a diphyletic taxon that could be separated into two sections, Thalassinida Latreille, 1831 and Callianassida Dana, 1852 (cf. Sakai & Sawada, 2006) by the difference in morphology of the gastric mill. In the present paper, however, those two sections 3 ) ksakai@mf.pikara.ne.jp 4 ) Michael.Tuerkay@senckenberg.de Koninklijke Brill NV, Leiden, 2014 DOI: /

2 130 KATSUSHI SAKAI & MICHAEL TÜRKAY are both assigned to a new rank as infraorders of the suborder Pleocyemata Burkenroad, 1963 (cf. Felgenhauer & Abele, 1983: 295), viz., Thalassinidea Latreille, 1831 and Callianassidea Dana, Hence, they are now assigned to the same taxonomic rank as the infraorders Astacidea Latreille, 1802, Nephropidea Dana, 1852, and Palinura Latreille, 1802 (cf. also Sakai & Sawada, 2006), as well as the infraorders Anomura MacLeay, 1838 and Brachyura Latreille, The Thalassinidea Latreille, 1831 include three families, characterized as follows: (1) the median tooth of the urocardiac ossicle is triangular in shape, protruding downwards and concave posteriorly, bearing a longitudinal median carina with a row of fine, transverse septa on the posterior surface; (2) the P2 is subchelate or simple; and the Callianassidea Dana, 1852 include 19 families, in which (1) the median tooth of the urocardiac ossicle is simple and smooth on the posterior surface, and (2) the P2 is chelate (Sakai, 2005a; Sakai & Sawada, 2006). In the present study, the specimens of those taxa lodged in the three German museums were examined and identified, and it was found that they comprise 10 families, 31 genera, including one new genus, and 73 species, three of which are new to science. [ Two families in the Thalassinidea (Laomediidae De Haan, 1849; Upogebiidae Borradaile, 1903), and 8 families in the Callianassidea (Axiidae Huxley, 1879; Callianideidae Kossmann, 1880; Callianassidae Dana, 1852; Calocarididae Ortmann, 1891; Ctenochelidae Manning & Felder, 1991; Meticonaxiidae Sakai, 1992; Strahlaxiidae Poore, 1994; Eucalliacidae Manning & Felder, 1991 (sensu Sakai, 2011))] (cf. Sakai, 2011). All material examined is presented below, with comments as necessary and provided with (additional) descriptions as required. MATERIAL AND METHODS The collections of the infraorders Thalassinidea and Callianassidea lodged in the three major German museums, viz., the Zoologische Staatssammlung, Munich (ZSM), the Zoologisches Museum der Universität Hamburg, Hamburg (ZMH), and the Museum für Naturkunde, Berlin (ZMB) were examined. A few specimens from the collection of the Senckenberg Research Institute and Natural History Museum, Frankfurt am Main (SMF), so far omitted from our former paper (Sakai & Türkay, 2012) have also been included. RESULTS AND DISCUSSION De Saint Laurent (1979: 28) proposed her subclassification of the infraorder Thalassinidea Latreille, 1831 as follows: [in translation] Thalassinidea are separated into two sections by the chela-form of the P1-2, viz., Infraorder Axiidea

3 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 131 Huxley, 1879 with chelate P1-2, and Infraorder Gebiidea De Saint Laurent, 1979 with simple P1-2. The name Gebiidea, including the families Upogebiidae, Laomediidae, and Thalassinidae, seems to have been derived from the earlier rejected family name Gebiidae (ICZN, opinion 434), and, though the Code (ICZN) has no bearing on names above the level of the family group (with as highest rank superfamily), Gebiidea should, in the opinion of the present authors, not have been used, but rather Upogebiidea, derived from the family name Upogebiidae Borradaile, 1903, might have been preferable. However, the family name Upogebiidae was published later than the name Thalassinidae. Besides, the infraorder name Gebiidea De Saint Laurent, 1979b was proposed much later than the name Thalassinidea Latreille, 1831, so that the name Gebiidea De Saint Laurent, 1979 is considered nomenclaturally inappropriate, while instead Thalassinidea Latreille, 1831 would be desirable as the representative name for this group. The infraorder Axiidea De Saint Laurent, 1979, including, inter alia, Axiidae Huxley, 1879, Callianassidae Dana, 1852, Callianideidae Kossman, 1880, Ctenochelidae Manning & Felder, 1991, Micheleidae Sakai, 1992, and Strahlaxiidae Poore, 1994, seems derived from the family name Axiidae Huxley, Yet the name Axiidea is, in our opinion, not nomenclaturally desirable for representing this group, because the name Axiidae Huxley, 1879 was introduced later than the name Callianassidae Dana, 1852, which makes the infraorder name Axiidea De Saint Laurent, 1979b less appropriate, whereas the name Callianassidea Dana, 1852 would be desirable as the representative name for this group. As a consequence, Thalassinidea Latreille, 1831 and Callianassidea Dana, 1852 are herein adopted as the representative names for the groups also known as the infraorders Gebiidea De Saint Laurent, 1979 and Axiidea De Saint Laurent, De Saint Laurent s (1979) taxon names were used by Poore (1994), Tudge et al. (2000), and Robles et al. (2009), and this practice was recently followed by Dworschak et al. (2012). Though we do not want to create any confusion at the level of the infraorders of Pleocyemata, we think a well-considered and proper choice of names, as we make herein, is to be preferred. The present taxa of the infraorders Thalassinidea and Callianassidea are based on two monographs (Sakai, 2006, 2011), and reference is made to those two papers for a full account of the families, genera, and species included in these infraorders, which were still acknowledged as sections by then. The present results of the examination and identification of the specimens from the three German museums to be classified in these taxa are listed below. SYSTEMATICS Infraorder THALASSINIDEA Latreille, 1831 [sensu Sakai, 2011]

4 132 KATSUSHI SAKAI & MICHAEL TÜRKAY Superfamily THALASSINOIDEA Latreille, 1831 Family LAOMEDIIDAE Borradaile, 1903 Genus Jaxea Nardo, 1847 Jaxea nocturna Nardo, 1847 Genus Laomedia De Haan, 1841 Laomedia astacina De Haan, 1841 Family UPOGEBIIDAE Borradaile, 1903 Subfamily UPOGEBIINAE Borradaile, 1903 Genus Pomatogebia Williams & Ngoc-Ho, 1990 Pomatogebia operculata (Schmitt, 1924) Genus Upogebia Leach, 1814 A. The East Atlantic-Mediterranean species Upogebia capensis (Krauss, 1843) Upogebia deltaura (Leach, 1815) Upogebia nitida (A. Milne-Edwards, 1868) Upogebia poensis De Saint Laurent & Ngoc-Ho, 1979 Upogebia pusilla (Petagna, 1792) Upogebia talismani Bouvier, 1915 B. The West Atlantic species Upogebia australis Thatje & Gerdes, 2000 Upogebia brasiliensis Holthuis, 1956 Upogebia jamaicensis Thistle, 1973 Upogebia omissa Gomes Corrêa, 1968 Upogebia vasquezi Ngoc-Ho, 1989 C. The East Pacific species Upogebia affinis (Say, 1818) Upogebia dawsoni Williams, 1986 Upogebia galapagensis Williams, 1986 D. The Indo-West Pacific species Upogebia africana (Ortmann, 1894) Upogebia ancylodactyla De Man, 1905 Upogebia barbata (Strahl, 1862) Upogebia brucei Sakai, 1975 Upogebia carinicauda (Stimpson, 1860) Upogebia darwinii (Miers, 1884) Upogebia digitina (Sakai, 1975) Upogebia fijiensis Sakai, 1982 Upogebia foresti Ngoc-Ho, 1989 Upogebia issaeffi (Balss, 1913) Upogebia longicauda Sakai, 1975 Upogebia major (De Haan, 1841) Upogebia pseudochelata Tattersall, 1921 Upogebia savignyi (Strahl, 1862) Upogebia spongium Sakai, 1975 Upogebia yokoyai Makarov, 1938 Genus Wolffogebia Sakai, 1982 Wolffogebia phuketensis Sakai, 1982 Infraorder CALLIANASSIDEA Dana, 1852 Superfamily AXIOIDEA Huxley, 1879 Family AXIIDAE Huxley, 1879

5 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 133 Genus Allaxius Sakai & De Saint Laurent, 1989 Allaxius aethiopicus (Nobili, 1904) Genus Axiopsis Borradaile, 1903 Axiopsis serratifrons (A. Milne-Edwards, 1873) Genus Balssaxius Sakai, 2011 Balssaxius habereri (Balss, 1913) Genus Boasaxius Sakai, 2011 Boasaxius princeps (Boas, 1880) Genus Newzealandaxius Sakai, 2011 Newzealandaxius novaezealandiae (Borradaile, 1916) Genus Scytoleptus Gerstaecker, 1856 Scytoleptus serripes Gerstaecker, 1856 Family CALLIANIDEIDAE Kossmann, 1880 Genus Callianidea H. Milne Edwards, 1837 Callianidea typa H. Milne Edwards, 1837 Family CALOCARIDIDAE Ortmann, 1891 Genus Calocaris Bell, 1846 Calocaris macandreae Bell, 1846 Family METICONAXIIDAE Sakai, 1992 Genus Meticonaxius De Man, 1905 Meticonaxius coeca (Balss, 1921) Family STRAHLAXIIDAE Poore, 1994 Genus Neaxius Borradaile, 1903 Neaxius glyptocercus (Von Martens, 1869) Genus Strahlaxius Sakai & De Saint Laurent, 1989 Strahlaxius plectrorhynchus (Strahl, 1862) Superfamily CALLIANASSOIDEA Dana, 1852 Family CALLIANASSIDAE Dana, 1852 Subfamily CALLIANASSINAE Dana, 1852 [sensu Sakai, 2011] Genus Callianassa Leach, 1814 [sensu Sakai, 2011] Callianassa subterranea (Montagu, 1808) Genus Cheramus Bate, 1888 [sensu Sakai, 2011] Cheramus pacificus (Guzmán & Thatje, 2003) Genus Gilvossius Manning & Felder, 1992 [sensu Sakai, 2011] Gilvossius bouvieri (Nobili, 1904) Gilvossius candidus (Olivi, 1792) Gilvossius gravieri (Nobili, 1905) Gilvossius tyrrhenus (Petagna, 1792) Gilvossius uncinatus (H. Milne Edwards, 1837) Genus Notiax Manning & Felder, 1991 [sensu Sakai, 2011] Notiax brachyophthalma (A. Milne-Edwards, 1870) Notiax ceramica (Fulton & Grant, 1906) Genus Trypaea Dana, 1852 [sensu Sakai, 2011] Trypaea biformis (Biffar, 1971) Trypaea japonica (Ortmann, 1891) Trypaea joculatrix (De Man, 1905) Trypaea rotundocula sp. nov. Trypaea santarita (Thatje, 2000) Genus Balsscallichirus Sakai, 2011 Balsscallichirus guineensis (De Man, 1928)

6 134 KATSUSHI SAKAI & MICHAEL TÜRKAY Genus Callichirus Stimpson, 1866 Callichirus kraussi (Stebbing, 1900) Genus Glypturus Stimpson, 1866 Glypturus armatus (A. Milne-Edwards, 1870) Glypturus longiventris (A. Milne-Edwards, 1870) Genus Lepidophthalmoides Sakai, 2011 Lepidophthalmoides rosae (Nobili, 1904) Lepidophthalmoides siriboia (Felder & Rodrigues, 1993) Genus Lepidophthalmus Holmes, 1904 Lepidophthalmus turneranus (White, 1861) Genus Neocallichirus Sakai, 1988 Neocallichirus indicus (De Man, 1905) Neocallichirus moluccensis (De Man, 1905) Neocallichirus mucronatus (Strahl, 1862) Neocallichirus pola sp. nov. Family CTENOCHELIDAE Manning & Felder, 1991 Genus Ctenocheles Kishinouye, 1926 Ctenocheles balssi Kishinouye, 1926 Family EUCALLIACIDAE Manning & Felder, 1991 [sensu Sakai, 2011] Subfamily EUCALLIACINAE Manning & Felder, 1991 [sensu Sakai, 2011] Genus Calliaxina Ngoc-Ho, 2003 [sensu Sakai, 2011] Calliaxina panglaoensis (Dworschak, 2006) Genus Calliaxiopsis gen. nov. Calliaxiopsis madagassa sp. nov. Abbreviations. CL, carapace length; TL, total length measured with thread from tip of rostrum to end of telson; P, pereiopod; Plp, pleopod; SAM, South African Museum, Cape Town. TAXONOMY Infraorder THALASSINIDEA Latreille, 1831 Superfamily THALASSINOIDEA Latreille, 1831 Family LAOMEDIIDAE Borradaile, 1903 For details on this family, see also Sakai & Türkay (2012: 755). Genus Jaxea Nardo, 1847 Jaxea nocturna Nardo, 1847 Cancer nocturnus Chiereghin, 1818: 38, fig. 48. Jaxea nocturna Nardo, 1847: 4; Ngoc-Ho, 2003: , figs. 22, 23; Sakai & Türkay, 2012: 756, fig. 12; Dworschak et al., 2012: 192. Calliaxis adriatica Heller, 1856: 717; 1863: 208. Material examined. ZSM 7312, 2 males (TL/CL, 28.0/10.1, 28.0/9.4 mm), 1 female (TL/CL, 22.0/7.7 mm), 1 ovig. female (TL/CL, 25.0/9.6 mm), eastern Mediterranean Sea, sta. Me/5-50, N E, m, beam trawl, F.S. (= R/V) Me-

7 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 135 teor, 06.i.1987, det. M. Türkay; ZSM 73/1, 1 male (TL/CL, 61.8/21.8 mm), Adriatic Sea; ZSM A , 1 male (TL/CL, 50.0/16.0 mm, lacking rostrum), 1 female (TL/CL, 56.0/20.0 mm), Trieste, Italy. Distribution. Irish Sea, Atlantic Ocean; Mediterranean Sea, m, sandy mud. Genus Laomedia De Haan, 1841 Laomedia astacina De Haan, 1841 Laomedia astacina De Haan, 1841: 165, pl. 35 fig. 8; Sakai, 1962: 27-34, pls. 5-7 (figs. 1-24); Ngoc- Ho, 1997: 732, fig. 1; Dworschak et al., 2012: 192. Material examined. ZSM 74/1, 1 female (TL/CL, 72.0/23.4 mm), Japan, Nagasaki, ded. Müller-Beeck, leg. Doflein, (probably meant as: 19.iv.1905), det. H. Balss (mentioned in Balss, 1914: 88). Distribution. Japan; Korea; China; Taiwan, in shallow brackish water (Ngoc- Ho, 1997). Family UPOGEBIIDAE Borradaile, 1903 Subfamily UPOGEBIINAE Borradaile, 1903 Genus Pomatogebia Williams & Ngoc-Ho, 1990 Pomatogebia operculata (Schmitt, 1924) Upogebia (Gebiopsis) operculata Schmitt, 1924: 91, pl. 5 figs Gebiopsis hartmeyeri Balss, 1924: 178, figs Pomatogebia operculata Williams & Ngoc-Ho, 1990: 614, fig. 1a-c; Williams, 1993: 10, fig. 5; Sakai, 2006: 34-36, fig. 9A; Felder et al., 2009: 185; Dworschak et al., 2012: 192. Material examined. ZMB 20282, 1 female (TL/CL, 17.0/5.0 mm) [holotype of Gebiopsis hartmeyeri Balss, 1924], St. Thomas, Savanna passage, U.S.-Virgin Islands, leg. Kükenthal & Hartmeyer. Distribution. Florida; Panama; Colombia; Brazil. Genus Upogebia Leach, 1814 A. The East Atlantic-Mediterranean species Species included. Upogebia aristata Le Loeuff & Intès, 1974; U. capensis (Krauss, 1843) [= U. subspinosa (Stimpson, 1860)]; U. crosnieri Le Loeuff & Intès, 1974; U. deltaura (Leach, 1815) [= U. littoralis Sars, 1884; U. sp. γ De Man, 1928]; U. demani De Saint Laurent & Le Loeuff, 1979; U. furcata (Aurivillius, 1898); U. mediterranea Noël, 1992; U. nitida (A. Milne-Edwards, 1868); U. poensis De Saint Laurent & Ngoc-Ho, 1979; U. pusilla (Petagna, 1792) [= U. gracilipes De Man, 1927; U. tipica (Nardo, 1869)]; U. senegalensis Ngoc- Ho, 2001; U. stellata Montagu, 1808; U. talismani Bouvier, 1915.

8 136 KATSUSHI SAKAI & MICHAEL TÜRKAY KEY TO THE EAST ATLANTIC-MEDITERRANEAN SPECIES OF THE GENUS Upogebia 1. Anterolateralmarginofcarapaceunarmed...2 Anterolateralmarginofcarapacearmedwithspine A2segment3unarmedventrally...3 A2segment3armedventrallywithdistalspine Cervicalgroovesmoothlaterally...U. demani Cervicalgroovespinoselaterally...U. furcata 4. P1palmunarmeddorsally...U. nitida P1 palm armed with subdistal spine or a row of distinct spines dorsally P1Palmwitharowofdistinctspinesdorsally...U. crosnieri P1 palm with subdistal spine dorsally P2merusunarmeddorsally...U. senegalensis P2 merus armed with subdistal spine dorsally Abdominal somite 1 terminating ventrally in acute angle....u. deltaura Abdominal somite 1 terminating ventrally in obtuse angle......u. mediterranea 8. Infrarostralspinepresent...9 Infrarostralspineabsent Lateralridgesofgastricregionprotrudingforward;P1palmunarmedventrally......U. poensis Lateral ridges of gastric region not protruding forward; P1 palm armed ventrally with large spineposteriortofixedfinger...u. talismani 10. P1palmarmedventrallywithspinesposteriortofixedfinger...U. aristata P1palmunarmedventrally Rostrum about as long as wide; P1-3 coxae each bearing a spine U. capensis Rostrumtwiceaslongaswide Dactylusarmeddorsallywithconicalproximaltooth...U. stellata Dactylusunarmeddorsally...U. pusilla (= U. tipica) The following are species lodged in the German museums. Upogebia capensis (Krauss, 1843) Gebia major var. capensis Krauss, 1843: 54; Ortmann, 1891: 54. Gebia subspinosa Stimpson, 1860: 91; Ortmann, 1893: 49. [Type locality: Simon s Bay, South Africa, 15 m.] Gebia capensis Ortmann, 1893: 49. Upogebia capensis Stebbing, 1900: 45; Sakai, 2005b: 42; Nel & Branch, 2013: Material examined. ZMH K 8395, 3 males (TL/CL, 37.0/ /23.0 mm), 2 females (TL/CL, 62.0/19.0, 50.0/18.0 mm), 1 ovig. female (TL/CL, 86.0/26.0 mm), S.W. Africa, Lüderitzbucht [Namibia, Luderitz Bay], intertidal beach, Hamb. Deutsch.-S.W. Afr. Studienreise 1911, vii.1911, leg. W. Michaelsen, det. H. Balss [cf. Balss, 1916, as U. capensis Krauss], rev. Sakai, x.1977; ZSM A , 2 males (TL/CL, 70.0/24.0, 58.0/20.0 mm), 2 females (TL/CL, 59.0/19.0, 58.0/18.0 mm), 1 ovig. female (TL/CL, 68.0/25.0 mm), Africa, Namibia, Lüderitzbucht, leg. L. Schultze, vii.1903; ZMB 17944, 5 males (TL/CL, 28.0/ /5.5 mm); 3 females (TL/CL, 20.0/ /4.5 mm), Deutsche Südpolar Exped , Simons Bay, Kapland [= Cape Province], det. Sakai as U. subspinosa, x.1977; as U. capensis, 16.vii.2012; ZMB 8733, 1 female

9 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 137 (TL/CL, 19.0/5.5 mm, lacking telson), in stomach of scylline sharks, Deutsche Südpolar Exped , Simons Bay, Kapland [= Cape Province], leg. Fritsch, det. Sakai as U. subspinosa, x.1977; as U. capensis, 16.vii Type locality. Saldanha Bay, South Africa. Distribution. South Africa (Angola to Mossel Bay). Upogebia deltaura (Leach, 1815) Gebia deltaura Leach, 1815: 342, figs Upogebia deltaura Schellenberg, 1928: 77, fig. 58; Sakai, 2006: 43; Dworschak et al., 2012: 194. Material examined. ZSM 88/2, 1 male (TL/CL, 45.0/14.5 mm), 1 ovig. female (TL/CL, 56.0/6.2 mm), Naples, det. H. Balss; ZSM 88/1, 1 female (TL/CL, 46.0/14.0 mm), Monaco, iii.1908; ZSM 88/3, 1 female (TL/CL, 21.0/6.2 mm), Alexandria, Egypt, sta. 50, 9 fathoms (= 16.2 m), leg. Steuer, 1933; ZMH K 38295, 1 female (TL/CL, 43.0/14.0 mm), Kristineberg, Gullmarsfjorden [Gullmarfjord], 05.ix.1897; ZMB 27273, 1 female (TL/CL, 41.0/11.8 mm), German Bight, 32 m, fine silt, leg. & det. Thatje; ZMB 1576, 1 female (TL/CL, 16.0/5.0 mm), data unknown. Distribution. N. Atlantic Ocean (Norway to Channel Islands); Mediterranean Sea (Spain, Adriatic Sea, Aegean Sea, Cyprus, Haifa Bay). Upogebia nitida (A. Milne-Edwards, 1868) Gebiopsis nitidus A. Milne-Edwards, 1868: 63, pl. 18 figs Upogebia nitida Le Loeuff & Intès, 1974: 58, figs. 20a-o, 20bis; De Saint Laurent & Le Loeuff, 1979: 37 (key), 38, fig. 2a-i; Ngoc-Ho, 2005: 115; Sakai, 2006: 47. Material examined. ZMB 1572, 1 female (TL/CL, 40.0/10.0 mm), included in specimens of U. pusilla (det. as Gebia littoralis), Mediterranean Sea. Remarks. The material examined (ZMB 1572) was found in a collection of U. pusilla and had been identified as such. If the locality is correct, which cannot be proven with certainty, this would be the first record from the Mediterranean Sea. However, this record might remain doubtful, as the species has never been collected in that region yet. Distribution. Atlantic coast (Mauritania to Togo);?Mediterranean Sea [doubtful record]. Upogebia poensis De Saint Laurent & Ngoc-Ho, 1979 (fig. 1) Gebicula Hupferi Balss, 1916: 35, figs [partim]. Upogebia poensis De Saint Laurent & Ngoc-Ho, 1979: 65, figs ; De Saint Laurent & Le Loeuff, 1979: 37 (key), 44, fig. 6; Sakai, 2006: 48. Material examined. ZMH K (ex. K 8404), holotype, female (TL/CL, 23.0/6.8 mm); paratypes, 2 females (TL/CL, 14.0/4.9, 21.0/6.1 mm); paratype, 1 ovig. female (TL/CL, 17.0/5.8 mm),

10 138 KATSUSHI SAKAI & MICHAEL TÜRKAY Fig. 1. Upogebia poensis De Saint Laurent & Ngoc-Ho, A, carapace, lateral view; B, gastric region of carapace, lateral view; C, gastric region of carapace, dorsal view; D, P1 on right side, lateral view. A-D, ZMH K (ex. K 8404), holotype female (TL/CL, 23.0/6.8 mm), Equatorial Guinea, Gulf of Guinea, Bioko [= Fernando Poo]. Scale bar, A-D, 1 mm. Equatorial Guinea, Gulf of Guinea, Bioko [= Fernando Poo], N E, 5 fathoms [= 9m], leg. Capt. Hupfer, det. de St. Laurent & Ngoc-Ho, Diagnosis. Small species measuring less than 23.0 mm in total length. Rostrum triangular, slightly longer than wide, bearing 1-3 lateral teeth (fig. 1A, B) on each side and 1-2 ventrorostral teeth. Eyestalks shorter than rostrum. A1 peduncle shorter than A2 peduncle. A2 penultimate segment with two ventral spines. Carapace setose and usually provided with anterolateral spine on each side, though anterolateral spine on left side and 2 anterolateral spines on right side in holotype; lateral ridges of gastric region protruding forward and sharply pointed anteriorly, and armed with a row of teeth (fig. 1C); cervical groove located at ca. anterior 0.6 of carapace and provided irregularly with some spinules.

11 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 139 P1 subchelate (fig. 1D); palm twice as long as wide, bearing 8 dorsal teeth and a few ventral teeth; fixed finger armed with a few teeth on prehensile margin. P2 carpus with subdistal tooth dorsally. Telson subsquare, bearing transverse carina at proximal 0.25 of dorsal surface, but no median spine on slightly convex posterior margin. Uropodal endopod broadened. Remarks. De Saint Laurent & Ngoc-Ho (1979) designated four females chosen from among the type specimens of Gebicula Hupferi Balss, 1916 (now Upogebia talismani Bouvier, 1915) as holotype (CL/TL, 6.5/19.0 mm) and paratypes (CL/TL, 6.4/18.0, 6.2/15.0, 4.5/13.0 mm) of Upogebia poensis De Saint Laurent & Ngoc-Ho, However, it has turned out in the re-examination reported herein that the present measurements of these four females are not in accordance with those by De Saint Laurent & Ngoc-Ho, which suggests that there is some confusion about these female type specimens handled by De Saint Laurent & Ngoc-Ho (1979), because the measurements in the present re-examination are as follows; CL/TL, 6.8/23.0, 6.1/21.0, 4.9/14.0, 5.8/17.0 (ovig.) mm, and two out of these four female type specimens are provided with two ventrorostral spines, and the other two are with one ventrorostral spine. So, one of the former two female specimens, measuring TL/CL, 23.0/6.8 mm is considered to be the holotype figured by De Saint Laurent & Ngoc-Ho (1979), who apparently did not produce a correct figure of the anterolateral teeth of the carapace, because in this holotype specimen the carapace bears two anterolateral teeth on the right side (fig. 1A, B) unlike in fig. 26 by De Saint Laurent & Ngoc-Ho (1979), in which the carapace bears one anterolateral tooth on the right side, and one anterolateral tooth on the left side. The other specimen of these (TL/CL, 21.0/6.1 mm) with two ventrorostral spines (fig. 1B) is considered to be a paratype, because the dissected mouth parts preserved together with the body in a tube were figured as a paratype by De Saint Laurent & Ngoc-Ho (1979: 68, figs ). As a result of the present re-examination of their type specimens, and comparison with the results of De Saint Laurent & Ngoc-Ho s description and figures (1979), it appears that the female holotype (the largest one) was incorrectly measured and figured as mentioned above. The present species, U. poensis, is well separable from Balss s specimens of U. hupferi Balss, 1916 (= U. talismani Bouvier, 1915), because those two species are different from each other as presented in table I. Distribution. So far only the type locality, Equatorial Guinea, Gulf of Guinea, Bioko [= Fernando Poo], N E. Upogebia pusilla (Petagna, 1792) Astacus pusillus Petagna, 1792: 418, pl. 5 fig. 5. [Type locality: Naples: Habitat in nostri maris arena.]

12 140 KATSUSHI SAKAI & MICHAEL TÜRKAY TABLE I Comparison of features between Upogebia poensis De Saint Laurent & Ngoc-Ho, 1979 and Upogebia talismani Bouvier, 1915 Upogebia poensis Upogebia talismani Rostrum Armed with 1-2 ventral spines Armed with 1-4 ventral spines P1 palm Armed with 8 teeth dorsally and a few teeth ventrally Armed with sharp teeth dorsally and 1 strong spine followed proximally by 1 small spine ventrally; armed with 6-7 sharp teeth subdorsally and 2 longitudinal rows of 6-7 interspaced teeth on mesial surface P1 dactylus Stout and smoothly narrowing distally Slender and abruptly narrowing at Telson Slightly convex on posterior margin without median spine middle of prehensile margin Slightly concave on posterior margin without median spine Upogebia pusilla Holthuis, 1947: 321, fig. 1(5); Sakai, 2006: 48-59, fig. 10, tabs. 1-2; Dworschak et al., 2012: 194. Material examined. ZMB 1572, 2 males (TL/CL, 30.0/9.3, 32.0/10.3 mm), 2 females (TL/CL, 30.0/10.0, 54.0/15.9 mm), Mediterranean Sea; ZMB 1574, 1 male (TL/CL, 40.0/12.4 mm), Mediterranean Sea; ZMB 1575, 1 male (TL/CL, 25.0/6.70 mm), Messina, Italy, leg. Haeckel; ZMH K 8394, 1 male (TL/CL, 13.0/4.0 mm), Nice, France; ZSM A , 1 male (TL/CL, 43.0/15.0 mm), 2 females (TL/CL, 37.0/11.0, 38.0/13.0 mm), Mediterranean Sea, det. H. Balss, 1914; ZSM A , 4 males (TL/CL, 12.0/ /11.0 mm), 4 females (TL/CL, 18.0/ /8.0 mm), probably Adriatic Sea, det. H. Balss as U. litoralis; ZMH K 27264, 1 male (TL/CL, 23.0/7.0 mm), 3 females (TL/CL, 22.0/ /17.0 mm), Venezia, Italy, 1954, leg. Caspers, det. Gruner; ZSM A , 1 female (TL/CL, 47.0/13.6 mm), Rovinj, Istria, Croatia, salty mud flat, leg. H. Geiselbrecht, 28.viii.2010, det. H. Geiselbrecht, viii.2010; ZSM A , 2 males (TL/CL, 38.0/12.4, 35.0/11.5 mm), 1 female (TL/CL, 40.0/13.5 mm), Rovinj, Istria, Croatia, salty mud flat, leg. R. Melzer, R. Meyer, S. Friedrich, 19.iii.2003, det. R. Melzer; ZMH K 8408, 2 females (TL/CL, 40.0/13.0, 46.0/15.0 mm), Naples, Italy; ZMH K 8407, 2 females (TL/CL, 29.0/11.0, 36.0/13.0 mm), Sicily, ded. A. Krohn; ZSM A , 1 female (CL, 8.0 mm, lacking abd. 5- telson), off Al-Iskandariyya [= Alexandria], Egypt, sta. 15, 19.x.1933, leg. Steuer; ZSM A , 1 male (TL/CL, 19.0/7.0 mm), off Al-Iskandariyya [= Alexandria], Egypt, sta. 53, 26.x.1933, leg. Steuer; ZMH K 27402, 7 males (TL/CL, 35.0/ /25.0 mm), 5 females (TL/CL, 48.0/ /19.0 mm), 2 male carapaces (CL, 18.0, 14.0 mm), Black Sea, Bay of Varna, Bulgaria, leg. Caspers, det. Panning. Distribution. Atlantic Ocean (Mauritania to Bay of Biscay); Mediterranean Sea; Black Sea; Suez Canal. Upogebia talismani Bouvier, 1915 (fig. 2) Upogebia Talismani Bouvier, 1915: 184; De Man, 1927: 56. Gebicula Hupferi Balss, 1916 (partim): 35, figs Upogebia (Upogebia) Talismani De Man, 1928a: 24 (list), 36, 38, 47 (key).

13 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 141 Upogebia talismani Buchanan, 1958: 24, 28; De Saint Laurent, 1971: 1259, figs. 1-3; Le Loeuff & Intès, 1974: 52, fig. 15d-n; Beaubrun, 1979: 79; De Saint Laurent & Le Loeuff, 1979: 45, fig. 7; Števčič, 1979: 128; Thessalou-Legaki, 1986: 183; Koukouras et al., 1992: 223 (list); 1993: 195; Falciai & Minervini, 1996: 149, 3 figs.; Sakai, 2006: 60. Gebiacantha talismani Ngoc-Ho, 1989a: 118, 121; Noël, 1992: 82; Froglia, 1995: 8; d Udekem d Acoz, 1995: 61; García Raso, 1996: 738; Pancucci-Papadopoulou et al., 1999: 393; d Udekem d Acoz, 1999: 156; González-Gordillo et al., 2001: 279; Markham, 2001, tabs. 1-2; Türkay, 2001: 289; Ngoc-Ho, 2003: 506, figs. 24, 25; Dworschak et al., 2012: 192. Material examined. ZMH K 8402, 1 male (TL/CL, 17.0/7.0 mm by Balss, 1916; TL/CL, 19.0/6.50 mm by present measurement), [lectotype of Gebicula hupferi Balss, 1916], Equatorial Guinea, Bata, N E, 6 fathoms [= 10.8 m], mud, leg. Capt. Hupfer; ZMH K 8403, 1 female (TL/CL, 30.0/11.0 mm) [paralectotype of Gebicula hupferi Balss, 1916], Gulf of Guinea, Bioko [= Fernando Poo], N E, leg. Capt. Hupfer, rev. De Saint Laurent & Ngoc-Ho, 1979; ZMH K 8404, 1 male (TL/CL, 36.0/11.3 mm) [paralectotype of Gebicula hupferi Balss, 1916], 2 damaged specimens, Equatorial Guinea, Bioko [= Fernando Poo], Gulf of Guinea, leg. Capt. Hupfer; ZSM A , 1 male (TL/CL, 26.0/8.0 mm), 1 female (TL/CL, 25.0/8.1 mm), Gulf of Guinea, Bioko [= Fernando Poo], N E, leg. Capt. Hupfer. Remarks. Balss (1916: 35, fig. 11) described Gebicula hupferi, based on specimens from Bioko and Bata, Equatorial Guinea. Later, De Saint Laurent & Ngoc-Ho (1979: 65) synonymized Gebicula hupferi Balss, 1916 with Upogebia talismani Bouvier, 1915, after examining Balss s syntypes, from among which De Saint Laurent & Ngoc-Ho chose four females (ZMH 31904) including one ovigerous, and assigned them to a new species, Upogebia poensis. In U. talismani the shape of the rostrum in dorsal view differs between males and females; it is broader in females than in males (fig. 2C, D). It is known that U. talismani is closely similar to U. lifuensis (Ngoc-Ho, 1994) from Loyalty Island, N.W. Australia, in the shape of the rostrum, the spines on the anterolateral region and margin of the carapace, as well as the spines on the lateral margin of the cervical groove, the morphology of the A2 peduncle, the P1 propodus, and the telson, but differs in that the P1 dactylus is abruptly narrowing at the middle of the prehensile margin (fig. 2F, G) (vs. smoothly narrowing distally in U. lifuensis). Distribution. Mediterranean Sea: Libya (De Saint Laurent), Aegean Sea (Thessalou-Legaki); Atlantic Ocean, Morocco (Bouvier), Ivory coast (Le Loeuff & Intès) to Congo (Balss). B. The West Atlantic species Species included. Upogebia acanthura (Coêlho & Ramos, 1973); U. aestuari Williams, 1993; U. affinis (Say, 1818); U. annae Thistle, 1973; U. aquilina Williams, 1993; U. australis Thatje & Gerdes, 2000; U. bermudensis Williams, 1993; U. brasiliensis Holthuis, 1956; U. careospina Williams, 1993; U. casis Williams, 1993; U. corallifora Williams & Scott, 1989; U. felderi Williams, 1993; U. inomissa Williams, 1993; U. jamaicensis Thistle, 1973; U. marina Coêlho

14 142 KATSUSHI SAKAI & MICHAEL TÜRKAY Fig. 2. Upogebia talismani Bouvier, A, whole body of male, lateral view; B, P1-5 of same male, lateral view; C, carapace with rostrum of male, dorsal view; D, same of female, dorsal view; E, carapace of female, lateral view; F, G, P1 carpus, palm, and dactylus of male, mesial view; H, abdominal somite 6, telson and uropod on left side of male, dorsal view. A, B, C, F, H, ZSM A , male (TL/CL, 26.0/8.0 mm); D, E, female (TL/CL, 25.0/8.1 mm); G, ZMH K 8404, paralectotype male (TL/CL, 36.0/11.3 mm); Equatorial Guinea, Bioko [= Fernando Poo], Gulf of Guinea. Scale bar, A-H, 1 mm.

15 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 143 & Ramos, 1973; U. molipollex Williams, 1993; U. noronhensis Fausto-Filho, 1969; U. omissa Gomes Corrêa, 1968; U. omissago Williams, 1993; U. paraffinis Williams, 1993; U. pillsbury Williams, 1993; U. spinistipula Williams & Heard, 1991; U. vasquezi Ngoc-Ho, [Revised after Sakai, 2006: 61.] A West Atlantic species, U. australis Thatje & Gerdes, 2000, not included in the previous key to the species in the revision of the Upogebiidae (cf. Sakai, 2006), is added to this revised key in the present study. KEY TO THE WEST ATLANTIC SPECIES OF THE GENUS Upogebia 1. Infrarostralspinepresent...2 Infrarostralspineabsent Lateral ridges of gastric region protruding forward as moderate spines Lateralridgesofgastricregionprotrudingdistinctlyforward Rostrum with small infrarostral spines; anterolateral margin of carapace with strong ocular spine...u. felderi Rostrum with 3 strong infrarostral spines; anterolateral margin of carapace with ocular spine andsomeadditionalones...u. bermudensis 4. Anterolateralmarginofcarapacewithsomespinules...U. pillsbury Anterolateralmarginofcarapacewithsinglepostocularspine DorsalmarginofP1palmsmoothbetweenproximalspineanddistalone...U. affinis DorsalmarginofP1palmdenticulate P1 dactylus with denticulate carina on mesial surface; A2 penultimate segment unarmed ventrally...u. paraffinis P1 dactylus without denticulate carina on mesial surface; A2 penultimate segment armed withspinesventrally...u. spinistipula 7. Anterolateral margin of carapace without postocular spine Anterolateralmarginofcarapacewithone(ormore)postocularspine(s) P1 merus with subdistal tooth dorsally u. australis P1 merus without subdistal tooth dorsally u. acanthura 9. Anterolateral margin of carapace usually with 4 or more postocular spines.. U. jamaicensis Anterolateral margin of carapace usually with single postocular spine, occasionally 2 spines A2segment3armedwithdistoventralspine...11 A2segment3unarmed Telson square and convex posteriorly; eyestalks with distoventral spine U. omissago Telson subsquare and concave posteriorly; eyestalks without distoventral spine u. vasquezi 12. P1 dactylus armed with conical tooth subterminally on prehensile margin P1dactylusunarmedonprehensilemargin P1palmdenticulatedorsally...U. omissa P1palmnotdenticulatedorsally P2merusunarmedventrally...U. brasiliensis P2 merus armed with single subproximal spine ventrally P3merusarmedwith3spinesventrally...U. inomissa

16 144 KATSUSHI SAKAI & MICHAEL TÜRKAY P3merusunarmedventrally P1meruswitharowofspinulesventrally...U. careospina P1merusarmedwithfewspinulesorunarmedventrally...U. marina 17. P2 merus with strong subproximal spine ventrally P2 merus without strong subproximal spine ventrally P2 carpus armed with dorsal spine and ventral one on distal margin P2carpusunarmedondistalmargin P3merusarmedwith3strongspinesventrally...U. aestuari P3merusarmedwithsomespinulesventrally...U. corallifora 20. P1palmwith3rowsofspinulesonmesialsurface...U. noronhensis P1 palm with a subdorsal row of spinules on mesial surface....u. aquilina 21. P1 palm without terminal spine dorsally u. molipollex P1palmarmedwithterminalspinedorsally P1carpuswitharowofspinulesonlateralcarina...U. casis P1 carpus without a row of spinules on lateral carina....u. annae The following are species lodged in the German museums. Upogebia australis Thatje & Gerdes, 2000 Upogebia australis Thatje & Gerdes, 2000: 232, figs Material examined. ZMB 27251, holotype, male (TL/CL, 40.0/12.1 mm), Chile, Beagle Channel, S W, 255 m, box corer, x.1994, leg. R/V Victor Hensen. Distribution. So far only the type locality, Chile, Beagle Channel, S W, 255 m. Upogebia brasiliensis Holthuis, 1956 Upogebia brasiliensis Holthuis, 1956: 175, figs Upogebia (Upogebia) brasiliensis Coêlho, 1971: 231. Material examined. ZMH K-27275, paratypes, 2 males (TL/CL, 20.0/6.0, 20.0/7.0 mm), 3 females (TL/CL, 18.0/ /8.0 mm), Cananeia, S. Brazil, vi.1954, leg. S. Gerlach; ZMH K 8418, 1 male (TL/CL, 34.0/10.0 mm, lacking chelipeds), Tutoia, W. Brazil, 04.vii.1912, leg. W. Schwinghammer. Distribution. Belize to Brazil. Upogebia jamaicensis Thistle, 1973 Upogebia jamaicensis Thistle, 1973: 16, fig. 4; Williams, 1986: 10 (key); Williams, 1993: 48, fig. 21; Sakai, 2006: 66. Material examined. ZMB 6048, 1 male (TL/CL, 46.0/14.1 mm), Cuba. Distribution. Jamaica; Panama to Colombia.

17 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS 145 Upogebia omissa Gomes Corrêa, 1968 Upogebia omissa Gomes Corrêa, 1968: 98, figs. 1-15, 28, 29. Upogebia omissago Williams, 1993: 57, fig. 25; Sakai, 2006: 68; Felder et al., 2009: 185. Material examined. ZMH K 8422, 1 female (TL/CL, 27.0/8.0 mm), Amapala, Honduras, 22.x.1906, leg. R. Paessler, ded. R. Paessler, 25.ii Distribution. U.S.A.: Horn Island, Mississippi (Williams, 1993), Vero Beach, Seminole shores, Jim Island, Sawyer Key and Andrews Bay, Florida (Williams, 1993). Upogebia vasquezi Ngoc-Ho, 1989 Upogebia affinis [non Upogebia affinis Say, 1818] Schmitt, 1936: 375. Upogebia vasquezi Ngoc-Ho, 1989b: 866, figs. 1-2; Markham et al., 1990: 424; Williams, 1993: 67, figs Material examined. SMF 25817, 1 male (TL/CL, 18.0/5.1 mm), 1 ovig. female (TL/CL, 21.0/5.5 mm), Bahia Ballena, Golfo de Nicoya, Costa Rica, GN-TBB-20, N W, 20 m, KG-1, 04.ii.1944, leg. R/V Victor Hensen ; ZMB 3307, 2 males (TL/CL, 39.0/11.4, 27.0/7.5 mm), 2 females (TL/CL, 45.0/11.8, 29.0/7.9 mm), Costa Rica. Distribution. Florida; Bahamas; Mexico; Costa Rica; Panama; Colombia to Brazil. C. The East Pacific species Species included. Upogebia acanthops Williams, 1986; U. affinis (Say, 1818); U. baldwini Williams, 1997; U. burkenroadi Williams, 1986; U. cortesi Williams & Vargas, 2000; U. dawsoni Williams, 1986; U. galapagensis Williams, 1986; U. hertwecki Sakai & Türkay, 2012; U. jonesi Williams, 1986; U. lepta Williams, 1986; U. longipollex (Streets, 1871); U. maccraryae Williams, 1986; U. macginitieorum Williams, 1986; U. onychion Williams, 1986; U. pugettensis (Dana, 1852a); U. ramphula Williams, 1986; U. schmitti Williams, 1986; U. spinigera (Smith, 1871) [= U. sturgisae Boone, 1931; U. rostrospinosa Bott, 1955]; U. tenuipollex Williams, 1986; U. thistlei Williams, 1986; U. vargasae Williams, 1997; U. veleronis Williams, In the present study, the key is revised by adding another species, Upogebia hertwecki Sakai & Türkay, 2012, which was not included in the key in the revision of the Upogebiidae (cf. Sakai, 2006). KEY TO THE EAST PACIFIC SPECIES OF THE GENUS Upogebia 1. Infrarostralspinepresent...U. affinis Infrarostralspineabsent Anterolateral margin of carapace without postocular spine

18 146 KATSUSHI SAKAI & MICHAEL TÜRKAY Anterolateralmarginofcarapacewithpostocularspine(s) P1 dactylus with strong conical subdistal tooth on prehensile margin; P1 merus armed with subdistal spine dorsally u. tenuipollex P1 dactylus without subdistal conical tooth on prehensile margin; P1 merus unarmed dorsally P1 chelate; dactylus and fixed finger finely and evenly toothed on prehensile margin u. ramphula P1subchelate P1 robust; merus armed with denticles ventrally; carpus armed with more than 1 strong spine dorsally; fixed finger armed with distinct, stout median tooth on prehensile margin u. macginitieorum P1 slender; merus unarmed ventrally; carpus unarmed dorsally; fixed finger unarmed on prehensilemargin...u. cortesi 6. P1 dactylus with subdistal conical tooth dorsally u. vargasae P1 dactylus without subdistal conical tooth dorsally P1 dactylus denticulate on whole prehensile margin, usually bearing subdistal conical tooth...8 P1dactylussmoothondistalhalfofprehensilemargin P2 merus armed with single subproximal spine ventrally....9 P2merusunarmedventrally Telsonwithdistincttransversecarina...10 Telsonwithindistincttransversecarina Telson distinctly wider than long, bearing denticulate transverse carina located at middle of dorsalsurface...u. baldwini Telson slightly wider than long, bearing denticulate transverse carina located at proximal 1/3 ofdorsalsurface...u. spinigera 11. Tailfanspinuloseondorsalsurface...U. longipollex Tailfannotspinuloseondorsalsurface P2carpusunarmeddorsally...U. acanthops P2 carpus armed with subdistal spine dorsally u. thistlei 13. P2merusunarmeddorsally...U. burkenroadi P2 merus armed with subdistal spine dorsally P2 carpus armed with subdistal spine dorsally u. veleronis P2carpusunarmeddorsally...U. maccraryae 15. Telsonwithdistinctdenticulatetransversecarinadorsally...U. onychion Telson with or without indistinct transverse carina dorsally Rostrumelongate,twiceaslongaswide...U. lepta Rostrum triangular P1fixedfingerarmedwithstouttooth,ordenticulateonprehensilemargin...18 P1fixedfingersmoothonprehensilemargin P1 fixed finger armed with stout tooth on prehensile margin U. pugettensis P1fixedfingerdenticulateonproximalhalfofprehensilemargin P1dactylusarmedwithproximaltoothonprehensilemargin...U. hertwecki P1 dactylus armed with obsolescent teeth on proximal half of prehensile margin, occasionallyfolloweddistallybymediantooth...u. dawsoni

19 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS Cervicalgroovespinoselaterally...U. schmitti Cervicalgrooveunarmed P1carpusarmedwithdistalspinedorsally...U. jonesi P1carpusdenticulatedorsally...U. galapagensis The following are species lodged in the German museums. Upogebia affinis (Say, 1818) Gebia affinis Say, 1818: 241; Dworschak, 2012: 194. Upogebia affinis Stebbing, 1893: 185; Sakai, 2006: 63. Material examined. ZMH K 8406, 1 male, damaged; 1 female, damaged, locality unknown. Distribution. U.S.A. (Georgia, Massachussetts, southern Texas, Florida); Colombia. Upogebia dawsoni Williams, 1986 Upogebia dawsoni Williams, 1986: 10 (key), 14, fig. 5; Hendrickx, 1995: 390 (list); 2013: 14, Sakai, 2006: 72. Material examined. ZMB 6096, 3 females (TL/CL, 22.0/ /6.7 mm), California, leg. Forster. Distribution. Mexico; Costa Rica to Panama. Upogebia galapagensis Williams, 1986 Upogebia galapagensis Williams, 1986: 10 (key), 17, fig. 6; Hendrickx, 1995: 390 (list). Material examined. ZMB 2824, 1 male (TL/CL, 57.0/16.3 mm); 2 females (55.0/14.9, 60.0/17.0 mm, abdomen divided into two pieces between abd. somite 2 and 3), leg. Jagor, California. Distribution. Galapagos Islands (Williams, 1986); California. D. The Indo-West Pacific species Species included. Upogbia acanthochela Sakai, 1967; U. acarinicauda Sakai, 2006; U. acutispina De Saint Laurent & Ngoc-Ho, 1979; U. africana Ortmann, 1894; U. albengai (Ngoc-Ho, 2005); U. allobrancha Ngoc-Ho, 1991; U. allspachi Sakai, 2006; U. amboinensis (De Man, 1888); U. anacantha Ngoc-Ho, 1994; U. ancylodactyla De Man, 1905; U. arabica (Ngoc-Ho, 1989); U. assisi Barnard, 1947; U. balmaorum Ngoc-Ho, 1990; U. balssi De Man, 1927; U. barbata (Strahl, 1862a) [= partim U. intermedia (De Man, 1887)]; U. baweana Tirmizi & Kazmi, 1979; U. borradailei Sakai, 1982; U. bowerbankii (Miers, 1884) [= U. (Calliadne) octoceras var. australiensis De Man, 1927]; U. brucei Sakai, 1975; U. cargadensis Borradaile, 1910; U. carinicauda (Stimpson, 1860) [= partim

20 148 KATSUSHI SAKAI & MICHAEL TÜRKAY U. intermedia (De Man, 1887)]; U. ceratophora De Man, 1905; U. dampieri (Ngoc-Ho, 2007); U. darwinii (Miers, 1884) [= U. hexaceras (Ortmann, 1894); U. octoceras Nobili, 1904]; U. digitina (Sakai, 1975); U. dromana Poore & Griffin, 1979; U. edentata Lin et al., 2001; U. edulis Ngoc-Ho & Chan, 1992; U. fijiensis Sakai, 1982; U. foresti Ngoc-Ho, 1989; U. kempi Sankolli, 1972; U. hirtifrons (White, 1847); U. holthuisi Sakai, 1982; U. imperfecta Sakai, 1982; U. iriomotensis Sakai & Hirano, 2006; U. issaeffi (Balss, 1913); U. kuekenthali Sakai, 1982; U. laemanu Ngoc-Ho, 1990; U. lagonensis (Ngoc-Ho, 1989a); U. laurentae (Ngoc-Ho,1989a);U. lenzrichtersi Sakai, 1982; U. lifuensis (Ngoc-Ho,1994b);U. lincolni Ngoc-Ho, 1977; U. longicauda Sakai, 1975; U. major (De Haan, 1839); U. miyakei Sakai, 1967; U. monoceros De Man, 1905; U. mortenseni Sakai, 2006; U. multispinosa (Ngoc-Ho, 1994b); U. narutensis Sakai, 1986; U. neglecta De Man, 1927; U. osiridis Nobili, 1904; U. ovalis Ngoc-Ho, 1991; U. plantae Sakai, 1982; U. poorei (Ngoc-Ho, 1994); U. priochela Sakai, 1993; U. pseudochelata Tattersall, 1921; U. pugnax De Man, 1905; U. quddusiae Tirmizi & Nasima Ghani, 1978; U. reunionensis (Ngoc-Ho, 1989); U. richeri (Ngoc-Ho, 1989); U. rupicola Komai, 2005; U. saigusai Sakai & Hirano, 2006; U. saintlaurentae Ngoc- Ho, 2007; U. sakaii Ngoc-Ho, 1994; U. savignyi (Strahl, 1862) [= U. isodactyla (Ortmann, 1891); U. rhadames Nobili, 1904]; U. seychellensis Sakai, 1982; U. shenchiajuii Yü, 1931; U. snelliusi Ngoc-Ho, 1989b; U. spinidactylus Sakai & Hirano, 2006; U. spinifrons (Haswell, 1881) [= U. nobilii Sakai & Türkay, 1995]; U. spinimana Ngoc-Ho, 1994; U. spongium Sakai, 1975; U. srilankaensis Sakai, 2006; U. stenorhyncha Ngoc-Ho, 1991; U. takaoensis Sakai & Türkay, 1995; U. tractabilis Hale, 1941; U. wuhsienweni Yü, 1931; U. yokoyai Makarov, 1938; U. sp. Borradaile, 1904 ; U. sp. Edmondson, 1944; U. sp. β De Man, ( U. sp. Borradaile, 1904 is not included in the key.) In the present key to the Indo-West Pacific species, the following 6 species: U. foresti Ngoc-Ho, 1989; U. miyakei Sakai, 1967; U. rupicola Komai, 2005; U. quddusiae Tirmizi & Nasima Ghani, 1978; U. edentata Lin et al., 2001; and U. shenchiajuii Yü, 1931, which were not included in the key in the revision of the Upogebiidae (cf. Sakai, 2006), and another 5 species: U. dampieri (Ngoc-Ho, 2007); U. saigusai Sakai & Hirano, 2006; U. iriomotensis Sakai & Hirano, 2006; U. spinidactylus Sakai & Hirano, 2006; U. saintlaurentae Ngoc-Ho, 2007, which were newly described after the publication of the revision (Sakai, 2006), are added. U. foresti Ngoc-Ho, 1989, earlier synonymized with Upogebia kempi Sankolli, 1972, is added to the revised key as a good species. KEY TO THE INDO-WEST PACIFIC SPECIES OF THE GENUS Upogebia 1. Lateralridgesofgastricregionnotprotrudingforward...2 Lateralridgesofgastricregiondistinctlyprotrudingforward...36

21 THALASSINIDEA AND CALLIANASSIDEA IN THREE GERMAN MUSEUMS Infrarostralspinespresent...3 Infrarostralspinesabsent Singleinfrarostralspinepresent...4 Pleuralinfrarostralspinespresent P1 palm without a row of spines mesially u. monoceros P1palmwitharowofspinesmesially Telsonslightlyconcaveposteriorly...U. dampieri Telsondistinctlyconcaveposteriorly A2segment3withventrodistalspineandsegment4with4ventralspines......U. reunionensis A2 segment 3 with ventrodistal spine and segment 4 with 1 ventral spine... U. ceratophora 7. Twoinfrarostralspinespresent...8 Morethantwoinfrarostralspinespresent Cervical groove with spinules; A2 penultimate segment with 1 distal spine ventrally u. lagonensis Cervical groove without spinules but with a single hepatic spine; A2 penultimate segment with3spinesventrally P1carpuswith2strongdistalspinesonmesialsurface...U. arabica P1carpuswith3strongdistalspinesonmesialsurface Rostrum elongate; 2 infrarostral spines arranged close to each other and located distally, distal one smaller than subdistal....u. acanthochela Rostrum rounded; 2 strong infrarostral spines arranged apart from each other, subdistally anddistally...u. poorei 11. Infrarostral spines arranged regularly along ventral margin from median to distal Infrarostralspinesarrangedverticallyatdistalend,formingasetof3spines Mesial surface of P1 palm without rows of spines....u. albengai MesialsurfaceofP1palmwithrowsofspines Threeinfrarostralspinespresent...U. priochela Fourinfrarostralspinespresent P1dactylusnotcarinatedorsomesially...U. lifuensis P1dactyluscarinatedorsomesially...U. plantae 15. Mesial surface of P1 palm without rows of spines u. laurentae MesialsurfaceofP1palmwithrowsofspines Proximal knob on lateral margin of uropodal endopod indiscernible u. acutispina Proximal knob on lateral margin of uropodal endopod distinct P1palmwithventralspinesposteriortofixedfinger...U. richeri P1 palm without ventral spines posterior to fixed finger u. multispinosa 18. P1chelate;anterolateralmarginofcarapacearmedwithocularspineorunarmed...19 P1subchelate;anterolateralmarginofcarapacearmedwithocularspine Anterolateralmarginofcarapacearmedwithocularspine...20 Anterolateralmarginofcarapaceunarmed Epistomebidentatedistally...U. mortenseni Epistome rounded distally P1palmdenticulateondorsalmargin...U. rupicola P1palmsmoothondorsalmargin...U. iriomotensis

22 150 KATSUSHI SAKAI & MICHAEL TÜRKAY 22. Rostrumwith2frontalteeth...23 Rostrumwith4frontalteeth Rostrum elongate and narrowly triangular, bearing 4 frontal teeth U. srilankaensis Rostrum broadly triangular in dorsal view P1merusunarmedonventralmargin...U. amboinensis P1merusarmedwithtuberclesorspinesonventralmargin P1meruswithtuberclesonventralmargin...U. holthuisi P1meruswithdistinctspinesonventralmargin...U. fijiensis 26. Rostrum wider than long and rounded anteriorly u. quddusiae Rostrum longer than wide and rounded anteriorly P1fixedfingerunarmedonprehensilemargin...U. barbata P1fixedfingerarmedwithdenticlesonprehensilemargin P1palmarmedwithtoothondistalmargin...U. baweana P1palmunarmedondistalmargin...U. ancylodactyla 29. Telsonwith2transversecarinae...U. dromana TelsonwithU-shapedcarina P1palmunarmedventrallyposteriortofixedfinger...31 P1palmarmedwithventraltoothposteriortofixedfinger Rostrum elongate and rounded anteriorly u. kempi Rostrum short and rounded anteriorly Rostrumwith2teethlaterally...U. foresti Rostrumwith3teethlaterally...U. balssi 33. P1 dactylus denticulate on prehensile margin, bearing a denticulate row medially on mesial surface...u. saintlaurentae P1 dactylus unarmed on prehensile margin, bearing no denticulate row medially on mesial surface P1 fixed finger narrow proximally and slender by deep concavity between fixed finger and palm, bearing 2 proximal denticles on prehensile margin U. edentata P1 fixed finger broad proximally and triangular Epistomeunarmed...U. sakaii Epistomewithterminalspine...U. pugnax 36. Infrarostralspinepresent...37 Infrarostralspineabsent Uropodal endopod without proximal knob on lateral margin Uropodal endopod with proximal knob on lateral margin Anterolateral margin of carapace with ocular spine; 2 short infrarostral spines located ventrodistally;telsonlackingtransversecarina...u. imperfecta Anterolateral margin of carapace with distinct ocular spine and 4 spinules; infrarostral spine locatedventrodistally;telsonbearingstrongtransversecarina...u. snelliusi 39. P1palmarmedwithlargeventralspineposteriortofixedfinger...40 P1palmunarmedventrallyposteriortofixedfinger P1fixedfingerwithtubercleonprehensilemargininmale...U. wuhsienweni P1fixedfingersmoothonprehensilemargininmale...U. edulis 41. P1 palm without translucent ridges mesially in male....u. narutensis P1palmwithsometranslucentridgesmesiallyinmale...42