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Centrophorus uyato (Rafinesque, 1810) Little Gulper Shark Synonymy. Dalatias nocturnus Rafinesque, 1810: 11, pl. 14, fig. 3 (Type locality: Sicily, Italy)— Cuvier, 1818: 455; Cuvier, 1829: 392; Cuvier, 1837: 246; Swainson, 1838: 129, 160; Swainson, 1839: 313; Duméril, 1865: 436; Jordan & Evermann, 1917: 77; Bigelow & Schroeder, 1948: 500. Spinax uyatus — Bonaparte, 1834: 49, fig. 2 (Italy); Bonaparte, 1846: 16 (Mediterranean); Böhlke, 1984: 158. Acanthias uyatus — Müller & Henle, 1839: 85 (Mediterranean); Gray, 1851: vii, 71 (Mediterranean); Steindachner, 1864: 27; Duméril, 1865: 439 (Algerian coast, Mediterranean); Barbosa du Bocage & de Brito Capello, 1866: 7, 21; Günther, 1870: 419 (Mediterranean); Canestrini, 1872: 40; Steindachner, 1875: 466; Moreau, 1881: 346 (Mediterranean); Réguis, 1882: 72; Rochebrune, 1883: 47 (Senegal and Gambia); Hilgendorf, 1884: 518; Duncan, 1891: 35 (Mediterranean); Moreau, 1892: 39; Seeley, 1895: 35 (Mediterranean); Parona, 1898: 38 (Ligurian Sea); Duncker, 1914: 291; de Buen, 1916: 303, figs (Mediterranean coast of Morocco); Landolt, 1947: 353. Centrophorus granulosus — Müller & Henle, 1839: 89, pl. 33 (Sicily, Italy); Barbosa du Bocage & de Brito Capello, 1864: 261 (Portugal); Vinciguerra, 1883: 18 (482) (Gulf of Genoa); Goode & Bean, 1896: 12, pl. 3, fig. 11 (W Atlantic, Mediterranean and adjacent areas); Boutan, 1926: 1 (Algeria); Dieuzeide, 1928a: 15, figs (Algeria); Dieuzeide, 1928b: 1314 (Algeria?); Andr & Canal, 1929: 511 (Algeria); Arcidiacono, 1931: 609 (Gulf of Squillace, Italy); Gruvel, 1931: 74 (State of Syria [Syria, Lebanon, Israel]); Ranzi, 1932: 240 (Naples, Italy); Belloc, 1934: 146, fig. (Western Sahara, Morocco, Canary Islands and Madeira); Ranzi, 1934: 343, 370 (Naples, Italy); Fowler, 1936: 73 (Mediterranean); Fowler, 1941 (in part): 231 (Mediterranean, inc. Italy); Šoljan, 1948: 66, figs (Adriatic Sea); Bigelow et al., 1955: 6 (Gulf of Mexico); Kirinčić & Lepetić, 1955: 24 (southern Adriatic Sea); Tortonese, 1956: 176, figs 94-95 (Italy); Cadenat, 1959: 748, fig. 1b (West Africa); Maurin, 1968: 82 (Morocco to Algeria); Maurin & Bonnet, 1970: 147 (Canary Islands to Cape Verde); Karrer, 1975: 64 (Namibia); Karlovac, 1976: 601, fig. 4 (Adriatic Sea); Guitart, 1979: 67, fig. 46 (Cuba); Bouchet et al., 1982: 577 (Tunisia); Zupanovic & El-Buni, 1982: 111 (Libya); Uyeno et al., 1983: 62, figs (Suriname); Compagno, 1984a: 37, figs (Atlantic and Indo-West Pacific); Gilat & Gelman, 1984: 263 (Levant Sea, Israel); McEachran & Branstetter, 1984: 130, fig. (NE Atlantic and Mediterranean); Muñoz-Chapuli, 1984: 9 (NE Atlantic);? Quéro, 1984: 43, fig.; Capapé, 1985: 97, fig. 1–7 (Eastern Atlantic); Jardas, 1985: 50 (Adriatic Sea); Anon, 1986: 93, fig. 22 (Atlantic Ocean); Bass et al., 1986 (in part): 50, fig. 5.1 (Walvis Bay and Mozambique); Golani, 1986: 23, fig. 2 (Levantine Sea, Israel); Muñoz-Chapuli & Ramos, 1989: 65, figs 1a, 3a, 4b and c, 5a, 6a, 7a (NE Atlantic and Mediterranean); Compagno et al., 1991: 54 (Hondeklip Bay to Namibia); Benli et al., 1993: 133, figs 1 and 2 (Sea of Marmara); Pisanty & Golani, 1995: 388 (Levantine Sea, Israel); Lanfranco, 1996: 6, pl. 4 (Malta); Aldebert, 1997: 284 (Gulf of Lion, France); Guallart, 1998: 1, figs (Balearic Sea); Bello, 1999: 69 (Adriatic Sea); Hernández-Hamón & Núñez, 1998: 107 (Pacific Columbia); Ungaro et al., 1999: 180 (Albania); Capapé et al., 2000: 129 (Languedoc coast, France); Bertrand et al., 2000: 5 (Gibraltar to Aegean Sea, Mediterranean); Golani & Pisanty, 2000: 71 (Levantine Sea, Israel); Baino et al., 2001: 234 (Alboran Island to Aegean Sea, Mediterranean); Guallart & Vicent, 2001: 135, fig. 4 (Balearic Sea, Spain); Bilecenoglu et al., 2002: 16 (Turkey); Schembri et al., 2003: 76, fig. 3c (Malta); McLaughlin & Morrissey, 2004: 481, fig. 3 (Cayman Trench, Jamaica); Moreno García, 2004: 214 (in part, not figs) (Mediterranean Sea); Sion et al., 2004: 155 (Ionian Sea, Greece); Golani, 2005: 11 (Levantine Sea, Israel); Lteif, 2015: 16, figs 9a, 14, (Lebanon); Serena, 2005: 27, figs, pl. I, 8 (Mediterranean Sea); Serét, 2005: 21 (Libya); Bessho, 2006: 28, figs 27–30, 33–35 (Japan, Namibia); Golani et al., 2006: 38, fig. (eastern Mediterranean); Hadjichristophorou, 2006: 163 (Cyprus); Megalofonou & Chatzispyrou, 2006: 67, fig. 4 (Crete, Greece); Mejía-Falla et al., 2007: 116 (Colombia); Psomadakis et al., 2009: 200 (Gulf of Naples, Italy); D’Onghia et al., 2010: 401 (Ionian Sea, Greece); Lipej & Dulĉić, 2010: 10 (Adriatic Sea); Damalas & Vassilopoulou, 2011: 145 (Aegean Sea, Greece); Colloca & Lelli, 2012: 12 (Lebanon); Costa et al., 2012: 7 (Portugal); Guijarro et al., 2012: 89 (Balearic Islands, Spain); Güven et al., 2012: 278 (Antalya Bay, Turkey); Iglésias, 2013: 38, pl. 16 (France and Cape Verde); Carneiro et al., 2014: 13 (Portugal); Farrugio & Soldo, 2014: 33 (Sicily, Italy and Tunisia); Veríssimo et al., 2014: 6 (Gulf of Mexico); Goren & Galil, 2015: 510 (Levant Sea, Israel); Barría et al., 2015a: 226 (Catalan Sea, Spain and Gulf of Lions, France); Barría et al., 2015b: 114 (Catalan Sea, Spain and Gulf of Lions, France); Carpenter & De Angelis, 2016: 1170, figs (Eastern Atlantic); Ramírez-Amaro et al., 2016: 639 (western Mediterranean); Cariani et al., 2017: 5 (Mediterranean);? Haroun et al., 2017: 84 (Egypt); Gajić, 2019: 101, fig. 14 (Croatia, Montenegro and Albania); Bariche & Fricke, 2020: 17, fig. 12 (Lebanon). ? Acanthias nigrescens Nardo, 1860: 70, 96 (Type locality: Venice, Italy). Entoxychirus uyatus — Gill, 1862: 498; Whitley, 1934: 199 (Australia). Acanthias ujatus — Döderlein, 1878: 30 (Sicily, Italy); Döderlein, 1881: 92 (Italy). Centrophorus uyatus — Goode & Bean, 1896: 508; Garman, 1906: 204; Garman, 1913: 9, 196. Squalus uyatus — Garman, 1899: 28. Squalus uyato — Garman, 1906: 204. Centrophorus bragancae Regan, 1906: 438 (Type locality: Cezimbra, Portugal)— Regan, 1908: 53 (coast of Portugal); Strand, 1908: 83. Centrophorus uyato — Fowler, 1936: 72, fig. 21 (Mediterranean and eastern Atlantic); Tortonese, 1938: 310 (Mediterranean); Poll 1951: 64, figs 33–34 (Angola and Namibia); Bigelow et al., 1953: 227, fig. 4 (Gulf of Mexico; Nice, Mediterranean Sea); Aksiray, 1954?: 233 (Turkish Seas); Bigelow et al., 1955: 5, 9; Springer & Bullis, 1956: 42 (Gulf of Mexico); Bigelow & Schroeder, 1957: 54, 66, 69, 72, 79–84, fig. 8e (Gulf of Mexico); Springer & Garrick, 1964: 81, 91; Krefft & Tortonese, 1973: 39 (NE Atlantic and Mediterranean); Bass et al., 1976: 31, figs 22, 24E, 24F, pl. 7 (southern Mozambique); Bridger, 1978: 26 (west of Ireland and Britain); Compagno, 1981: 8 [sharks] (Eastern Central Atlantic); Castro, 1983: 54, figs (Gulf of Mexico); Allu et al., 1984: 125 (Namibia); Compagno, 1984a: 45, figs (Atlantic, Indo-West Pacific); Compagno, 1984b: 9 [sharks] (Western Indian Ocean); McEachran & Branstetter, 1984: 132, fig. (NE Atlantic and Mediterranean); Lloris, 1986: 96, fig 24 (Namibia); Turón et al., 1986: 63 (Namibia); Fischer et al., 1987: 823 (Mediterranean); Compagno, 1988b: 603 (Comoro Islands); Compagno et al., 1989: 24, fig. (Hondeklip Bay, South Africa to Namibia);? Clark & Kristof, 1990: 277, fig. 9 (Caribbean); Springer, 1990: 11 (Gulf of Mexico, Mediterranean, and Natal, South Africa); Applegate et al., 1993: 35 (Atlantic Mexican waters); Gomon et al., 1994: 92, 94, figs 30, 31 (southern Australia); Last & Stevens, 1994: 60, figs, pl. 4 (fig. 8.5) (southern Australia); Meriç, 1995: 192 (Sea of Marmara, Turkey); Perry et al., 1995: 139 (Gulf of Mexico); Reiner, 1996: 22, fig. (Cape Verde); Bonfil, 1997: 105 (Veracruz, Mexico); Joseph, 1999: [unpaginated] (Sri Lanka); Cervigón & Alcalá, 1999 (Venezuela); Clarke, 2000: 377 (Rockall Trough, NE Atlantic); Baino et al., 2001: 234 (Alboran Island to Aegean Sea, Mediterranean); Graham et al., 2001: 551 (south-eastern Australia); Yearsley et al., 2001: 35, 360 (southern Australia); Bilecenoglu et al., 2002: 17 (Turkey); Daley et al., 2002: 53 (southern Australia); Ali & Saad, 2003: 58, fig. (Syria); Schembri et al., 2003: 77, fig. 3d (Malta); Serena, 2005: 28, figs, pl. I, 9 (Mediterranean Sea, NE Atlantic, Western Indian, Gulf of Mexico and Taiwan); Meriç et al., 2007: 31 (Turkey); White, 2008: 87, figs (southern Australia); Scacco et al., 2010: 39, fig. 3f (Mediterranean Sea); Castro, 2011: 81, figs 16a-e (north-western Atlantic); Davenport et al.: 2011: 557 (north-western Atlantic, USA); White et al., 2013: 36, fig. 2, 15, 16 (Atlantic); Veríssimo et al., 2014: 6, fig. 5 (Gulf of Mexico); Hipes, 2015: 1, fig. 11 (Gulf of Mexico, USA); Wienerroither et al., 2015: 834, fig. 2 (northern Norway); Farrag, et al., 2016: 481, fig. 2a (Egypt); Driggers et al., 2017: 52 (Gulf of Mexico); Haroun et al., 2017: 84 (Egypt); Lteif et al., 2017: 1491 (Lebanon); Biscoito et al., 2018: 471, fig. 7 (Madeira, Portugal); Carneiro et al., 2019: 36 (Portugal); Ehemann et al., 2019: 4 (Venezuela); Fernando et al., 2019: 231, figs 5e, 18d-f (Mutur, Sri Lanka); Follesa et al., 2019: 85 (Mediterranean); Psomadakis et al., 2019: 162, figs, pl. X (fig. 70) (Myanmar); Iglésias, 2020: 46, pl. 21 (France and Cape Verde); Ebert & Dando, 2021: 217, figs (NE Atlantic and Mediterranean); Kousteni et al., 2021: 1, figs 2 and 3 (Cypriot waters). Centrophorus machiquensis Maul, 1955: 5, figs 13–16 (Type locality: Madeira)— Krefft & Tortonese, 1973: 39 (Madeira); Ali & Saad, 2003: 57, fig. (Syria); Biscoito et al., 2018: 470, fig. 6 (Madeira, Portugal); Almeida & Biscoito, 2019: 99 (Canary Islands and Madeira); Carneiro et al., 2019: 36 (Portugal). Centrophorus ujato — Tortonese, 1956: 178, fig. 96 (Genova, Italy); Bini, 1967: 97, fig. (Italy); Sara, 1968: 1, figs 1–3 (west of Sicily); FAO, 1971: no pagination (Mediterranean Sea). Centrophorus spp. (granulosus group)— Forster et al, 1970 (in part): 388 (Western Indian Ocean). Centrophorus (forme) uyato-machiquensis — Cadenat & Blache, 1981: 58, figs. 36, 37 and 40 (NE Atlantic). Centrophorus spp. — Peyronel et al., 1984: 643 (Bay of Ajaccio, Corsica, France). Centrophorus cf. harrissoni (Undescribed gulper shark #2)— Kiraly et al., 2003: 2 (Puerto Rico, US Virgin Islands, Virginia, North Carolina and Florida Straits to Dry Tortugas). Centrophorus bragance — Hernández-Hamón & Núñez, 1998: 108 (as questionable synonym of C. granulosus). Centrophorus sp. (uyato ?)— Morón et al., 1998: 144 (Beruwela, Sri Lanka). Centrophorus sp. cf. uyato — Saad, et al., 2004: 430 (Syria). ? Centrophorus sp. (non uyato)— Serét, 2005: 21 (Libya). Centrophorus zeehaani White, Ebert & Compagno, 2008: 1, figs 8-10 (Type locality: South Australia)— Last & Stevens, 2009: 68, pl. 9.7, figs (southern Australia); Pethybridge et al., 2010: 1369 (Tasmania and Great Australian Bight, Australia); Pethybridge et al., 2011: 2743 (Tasmania and Victoria, Australia); Graham & Daley, 2011: 583 (southern Australia); Daley et al., 2012: 708 (southern Australia); White et al., 2013: 41 (southern Australia); Daley et al., 2015:127 (southern Australia); Wienerroither et al., 2015: 834, fig. 2 (northern Norway); Bineesh et al., 2016: 461 (Kollam, India). Centrophorus cf. uyato — McLaughlin & Morrissey, 2005: 1185, figs 2, 3 (Cayman Trench, Jamaica); Veríssimo et al., 2014: 7 (Gulf of Mexico); Serena et al., 2020: 502, 509 (Mediterranean); Bellodi et al., 2022: 2 (Mediterranean Sea). Centrophorus zeehani — Daley et al., 2012: fig. 2 (misspelling; southern Australia); Bineesh et al., 2016: 466 (misspelling; Kollam, India). Centrophorus ‘uyato’ — White et al., 2017: 86 (Eastern Atlantic); Almeida & Biscoito, 2019: 100 (Mediterranean Sea, Canary Islands and Madeira). Centrophorus cf. granulosus — Follesa et al., 2019: 85 (Mediterranean); FAO, 2018: unpaginated, fig. (Mediterranean Sea). ? Centrophorus granulosus — Parenti, 2019: 102 (Sicily). Material examined. Neotype: BMNH 2021.10.4.1 (eviscerated; GenBank accession ON167716), female 983 mm TL, between Gorgona and Capraia islands, Ligurian Sea, 43°19.8′ N, 9°56.1′ E, 180 m, 20 Dec. 2012. Other specimens: Australia: AMS I 44310 –001 (paratype of Centrophorus zeehaani; GenBank accession ON167706), adult male 826 mm TL, southwest of Coffin Bay, South Australia, 35°14′ S, 134°29′ E, 360–600 m, 28 July 2005; CSIRO CA 4104, adult male 843 mm TL, east of Gabo Island, Victoria, 37°40′ S, 150°15′ E, 504–508 m, 4 May 1984; CSIRO H 866–02, immature male 456 mm TL, CSIRO H 867–01, female 439 mm TL, east of Jervis Bay, New South Wales, 34°58′ S, 151°09′ E, 490–576 m, 10 Sep. 1986; CSIRO H 2268–02, adult male 800 mm TL, west of Bunbury, Western Australia, 33°03′ S, 114°25′ E, 701 m, 10 Feb. 1989; CSIRO H 6307–01 (skeletal parts), female 1027 mm TL, 12 July 2004, east of Flinders Island, Tasmania, ~ 40° S, ~ 149° E, 350–430 m; CSIRO H 6309–01 (skeletal parts; GenBank accession ON167708), adult male 865 mm TL, CSIRO H 6309–02 (skeletal parts), adult male 876 mm TL, CSIRO H 6309–04 (skeletal parts), adult male 906 mm TL, east of Flinders Island, Tasmania, ~ 40° S, ~ 149° E, 400–450 m, 1 Aug. 2004; CSIRO H 6310–04 (skeletal parts), female 970 mm TL, northeast of Flinders Island, Tasmania, 39°04′ S, 148°39′ E, 500–680 m, 24 Jul. 1986; CSIRO H 6311–01 (skeletal parts), female 655 mm TL, east of St. Helens, Tasmania, 41°27′ S, 148°44′ E, 850–860 m, 5 Jun. 1987; CSIRO H 6500-02, adult male 862 mm TL, east of Flinders Island, Tasmania, 40°15′ S, 148°45′ E, 329–512 m, 21 Aug. 2003; CSIRO H 6503–02 (skeletal parts), female 991 mm TL, CSIRO H 6503–03 (skeletal parts; GenBank accession ON167709), female 1023 mm TL, CSIRO H 6503–04 (skeletal parts; GenBank accession ON167710), female 987 mm TL, CSIRO H 6503–05 (skeletal parts), female 957 mm TL, northeast of Flinders Island, Tasmania, 39°20′ S, 148°45′ E, 370–420 m, 7 Apr. 2003; CSIRO H 6504–02, adult male 854 mm TL, CSIRO H 6504–03, female 817 mm TL, CSIRO H 6504–04, juvenile male 666 mm TL, CSIRO H 6504–05, adult male 861 mm TL, east of Jervis Bay, New South Wales, 35°12′ S, 150°58′ E, 320–500 m, July to Aug. 2003; CSIRO H 6628–01 (paratype of Centrophorus zeehaani), immature male 506 mm TL, CSIRO H 6628–02 (paratype of Centrophorus zeehaani; GenBank accession ON167705), immature male 645 mm TL, CSIRO H 6628–03 (paratype of Centrophorus zeehaani), adult male 875 mm TL, CSIRO H 6628–04 (paratype of Centrophorus zeehaani), adult male 910 mm TL, CSIRO H 6628–05 (holotype of Centrophorus zeehaani), adult male 893 mm TL, CSIRO H 6628–06 (paratype of Centrophorus zeehaani), adult male 852 mm TL, CSIRO H 6628–07 (paratype of Centrophorus zeehaani), adult male 906 mm TL, NMV A 29736 –001 (paratype of Centrophorus zeehaani), adult male 820 mm TL, southwest of Coffin Bay, South Australia, 35°14′ S, 134°29′ E, 360–600 m, 28 July 2005; CSIRO unreg. (DB 02/181), Great Australian Bight, adult male 857 mm TL; CSIRO unreg. (LJVC 880517), female 840 mm TL, unknown location; PMH095-11 (jaws only), female 104 cm TL, Albany, Western Australia. Eastern Atlantic (including Mediterranean): AMNH 78267, female 922 mm TL, AMNH 78269, female 937 mm TL, AMNH 78271, female 1016 mm TL, AMNH 78273, adult male 872 mm TL, AMNH 78277, adult male 895 mm TL, AMNH 78279, adult male 890 mm TL, between Tenerife and Gran Canaria, Canary Islands, Spain, 3 Oct. 1986; AMNH 78280, female 996 mm TL, AMNH 78282, female 1059 mm TL, AMNH 78283, female 1056 mm TL, AMNH 78284, adult male 832 mm TL, AMNH 78285, female 921 mm TL, AMNH 78286, female 1004 mm TL, AMNH 78291, female 1034 mm TL, AMNH 78292, female 980 mm TL, AMNH 78294, adult male 891 mm TL, between Tenerife and Gran Canaria, Canary Islands, Spain, 4 Oct. 1986; BMNH 1862.4.22.29, female 449 mm TL, Madeira, Portugal; BMNH 1864.7.18.1, female 894 mm TL, Madeira, Portugal; BMNH 1904.11.30.9- 10 (2 specimens), female 1036 mm TL, adult male 888 mm TL, west of Faro, Portugal, 662 m depth; BMNH 1904.11.30.11 (paralectotype of Centrophorus bragancae), juvenile male 488 mm TL, off Sesimbra, Portugal, 841 m depth; BMNH 1904.11.30.12 (lectotype of Centrophorus bragancae), female 467.5 mm TL, off Sesimbra, Portugal, 505 m depth; BMNH 2013.9.20.34, female 875 mm TL, BMNH 2013.9.20.35, female 930 mm TL, BMNH 2013.9.20.36, female 935 mm TL, BMNH 2013.9.20.37, female 731 mm TL, BMNH 2013.9.20.38, adult male 843 mm TL, BMNH 2013.9.20.39, adult male 865 mm TL, northeast Atlantic; CSIRO H 7471-01 (GenBank accession ON167715), adult male 853 mm TL, east of Corsica, France, 42°05.8′ N, 9°44.5′ E, 550–565 m, 30 May 2012; CSIRO H 7472-01, juvenile male 440 mm TL, southeast of Corsica, France, 41°47.9′ N, 9°30.5′ E, 466–480 m, 31 May 2012; ERB 1288, male 770 mm TL, Concarneau fish-market, France, 9 Sep. 2000; ERB 0763, male, 850 mm TL, Concarneau, France, 13 May 2009; HUJ 10885, female 760 mm TL, Hadera, Israel, 13 Sep. 1982; HUJ 11339, adult male 796 mm TL, Haifa Bay, Israel, 1 Mar. 1983; HUJ 17029, female 476 mm TL, Haifa, Israel, 16 Mar. 1983; HUMZ 151304, juvenile male 578 mm TL; HUMZ 151306, female 978 mm TL, Namibia; HUJ 21135 (GenBank accession ON167718), female 753 mm TL, Haifa fishing port, Israel, 29 July 2013; MNHN-IC 1905- 0568, juvenile male 454 mm TL, Portugal, 460 m depth, June 1903; MNHN-IC 1969-0269, juvenile male 499 mm TL, southwest of Monrovia, Liberia, 6°08′ N, 10°56′ W, 400 m depth, 27 Apr. 1964; MNHN-IC 0000-1224, female 537 mm TL, Naples, Italy; MNHN-IC 2005-0169, adult male 892 mm TL, west of Ireland, 28 Apr. 2005; NMW 15009, female 889 mm TL, Nice; NMW 63020, adult male 880 mm TL, Nice, France, 1902, Published as part of White, William T., Guallart, Javier, Ebert, David A., Naylor, Gavin J. P., Mo, Ana Veríssi-, Cotton, Charles F., Harris, Mark, Serena, Fabrizio & Iglésias, Samuel P., 2022, Revision of the genus Centrophorus (Squaliformes: Centrophoridae): Part 3 - Redescription of Centrophorus uyato (Rafinesque) with a discussion of its complicated nomenclatural history, pp. 1-51 in Zootaxa 5155 (1) on pages 15-41, DOI: 10.11646/zootaxa.5155.1.1, http://zenodo.org/record/6669082, {"references": ["Rafinesque, C. S. (1810) Caratteri di alcuni nuovi generi e nuove specie di animali e piante della Sicilia, con varie osservazioni sopra i medisimi. Sanfilippo, Palermo, 1, 3 - 69. https: // doi. org / 10.5962 / bhl. title. 104418", "Cuvier, G. (1818) Dictionnaire des sciences naturelles, dans lequel on traite meithodiquement des diffeirens e \u02c6 tres de la nature, consideireis soit en eux-me \u02c6 mes, d'apreIs l'eitat actuel de nos connoissances, soit relativement aI l'utilitei qu'en peuvent retirer la meidecine, l'agriculture, le commerce et les artes. 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GENUS † PSEUDORHINOBATOS Zoobank registration: urn:lsid:zoobank.org:act: 75367FF8-9DCA-406F-B257-8A34595CE1BF Type species: Trygonorhina dezignii Heckel, 1853. Etymology: After the Ancient Greek ψεῦδος (pseudos), false, and Rhinobatos, one of the living rhinobatid genera, therefore remarking their apparent similarity; gender masculine. Diagnosis: A rhinobatid guitarfish characterized by a wedge-shaped pectoral disc, longer (44.8% TL) than wide (38.5% TL); head length (from snout to scapulocoracoid) 29.8% TL; tail length (from pelvicfin origin to the posteriormost tip) 56% TL; pelvic-fin length 17.8% TL; snout to pelvic-fin origin 44.8% TL: well-developed lateral skin folds extending along the lateral margins of tail; rostral cartilage about 65% of neurocranial length; no horn-like processes on nasal capsules; 42 nasal lamellae; nuchal cartilages absent; anteriormost synarcual centrum located near the midlength of the synarcual; 118–138 vertebral centra, of which 18 monospondylous, and 100–120 diplospondylous; about 20 rib pairs; propterygia extending close to the margins of the pectoral disc; first propterygial segment not reaching the level of nasal capsules; propterygial radials extending as far as nasal capsule level; 60–62 pectoral-fin radials, of which 30–32 propterygial, seven to nine mesopterygial, one to two neopterygial, 20–22 metapterygial; about 21 pelvic-fin radials; closely arranged, small dermal denticles forming a continuous and regular covering on the ventral side of the body; denticle crown smooth, rhomboidal or lozenge in shape; thorns completely covering the dorsal side of the body; thorns of globular or arrow shape with prominent ridges; teeth extremely small (up to 750 µm); crown low, not globular; lingual visor slightly convex; regular central lingual uvula with apices rounded and not enlarged; two very incipient, not divergent, lateral lingual uvulae; labial apron absent; transverse keel separating the crown into labial and lingual faces, and forming a wide obtuse angle in lingual view; enameloid surface completely smooth; holaulacorhizid root bilobed, wide, displaced lingually, not broader or higher than the crown; root lobes triangular in shape in basal view, with regular, not undulated margins; lobes separated by a marked and deep furrow exhibiting a single central nutritive foramen; two additional foramina on lingual root face; low collar on upper part of the root stem., Published as part of Marramà, Giuseppe, Carnevale, Giorgio, Naylor, Gavin J. P., Varese, Massimo, Giusberti, Luca & Kriwet, Jürgen, 2021, Anatomy, taxonomy and phylogeny of the Eocene guitarfishes from the Bolca Lagerstätten, Italy, provide new insights into the relationships of the Rhinopristiformes (Elasmobranchii: Batomorphii) in Zoological Journal of the Linnean Society 192, {"references":["Heckel MJ. 1853. Bericht uber die vom Herrn Cavaliere Achille de Zigno hier angelangte. Sammlung fossiler Fische. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Mathematisch-Naturwissenschaftliche Klasse 11: 122 - 138."]}

Background: Modern sequencing technologies should make the assembly of the relatively small mitochondrial genomes an easy undertaking. However, few tools exist that address mitochondrial assembly directly. Results: As part of the Vertebrate Genomes Project (VGP) we develop mitoVGP, a fully automated pipeline for similarity-based identification of mitochondrial reads and de novo assembly of mitochondrial genomes that incorporates both long (> 10 kbp, PacBio or Nanopore) and short (100–300 bp, Illumina) reads. Our pipeline leads to successful complete mitogenome assemblies of 100 vertebrate species of the VGP. We observe that tissue type and library size selection have considerable impact on mitogenome sequencing and assembly. Comparing our assemblies to purportedly complete reference mitogenomes based on short-read sequencing, we identify errors, missing sequences, and incomplete genes in those references, particularly in repetitive regions. Our assemblies also identify novel gene region duplications. The presence of repeats and duplications in over half of the species herein assembled indicates that their occurrence is a principle of mitochondrial structure rather than an exception, shedding new light on mitochondrial genome evolution and organization. Conclusions: Our results indicate that even in the “simple” case of vertebrate mitogenomes the completeness of many currently available reference sequences can be further improved, and caution should be exercised before claiming the complete assembly of a mitogenome, particularly from short reads alone.

Hemitrygon yemenensis sp. nov. (Figs. 1–5, Table 1) Holotype. NMW 60783, adult male 223 mm DW, Gischin, Yemen, collected 1902. Paratype. NMW 99841, female 206 mm DW, collected with holotype. Diagnosis. A small species of Hemitrygon (attaining at least 22 cm DW) with the following combination of characters: disc width subequal to length; snout elongate, tip narrowly pointed, angle 107–110°, length 2.1–2.3 times interorbital width; preoral length 2.6–2.7 times mouth width; internasal distance 1.8–1.9 in prenasal length; body and tail mostly naked, denticles confined largely to head; single continuous median row (broken in paratype) of 8–18 small to large thorns on disc (two scapular thorns on each side of adult male), 4–13 similar thorns on tail before caudal sting (more thorns on male than female); tail moderately elongate, slender, whip-like beyond sting, width 0.5–1.2 times its depth, postcloacal tail length 1.9–2.2 times precloacal length; ventral cutaneous fold long, very slender, its length 1.9–2.0 in DW, height 0.3–0.4 in tail depth at its midlength; dorsal skin fold elongate, low, much shorter than ventral fold; distance from cloaca to sting 1.5–1.6 in precloacal length; pectoral-fin radials ~115; total vertebral centra 125–126, monospondylous vertebrae (including all synarcual) 38–39. Description. Disc quadrangular, bluntly angular anteriorly and slightly produced; length subequal to width, width 0.99 times length in adult male holotype (0.98 in female paratype); axis of greatest width of disc slightly forward of scapular region, its distance from snout tip 1.83 (1.89) times in distance from tip of snout to pectoral-fin insertion; body moderately robust, thickness 7.1 (7.4) times in disc width, raised slightly above cranium (marginally more so in scapular region); anterior margin of disc concave anteriorly, straight medially, strongly and evenly convex just before pectoral-fin apex; apex broadly rounded; posterior margin weakly convex; free rear tip broadly rounded. Pelvic fins weakly triangular, anterior and posterior margins almost straight, apices narrowly rounded, free rear tip broadly rounded; relatively large, length 24.3% (25.3%) DW; 1.49 (1.66) times width across fin bases. Tail moderately elongate, slender, with a long, very low ventral skin fold and a shorter and lower dorsal fold (modified somewhat through partial desiccation); postcloacal tail 2.22 (1.88) times precloacal length; width at axil of pelvic fin 1.31 (1.63) times its depth; tapering gradually and evenly to sting base (missing in holotype but position evident as a scar; sting broken at base in paratype); narrowly oval in cross-section near origin of ventral skin fold, width 1.03 (1.25) times height at fold origin; tapering evenly in dorsoventral view below sting scar; very slender, whip-like beyond sting scar, becoming progressively more compressed toward tail tip; subquadrangular in cross-section above mid ventral fold, depth 1.36 (1.16) times width; at end of fold suboval, weakly compressed, depth 1.37 (1.07) times width; filamentous towards tail apex. Dorsal skin fold reduced to low, elongate ridge, length 77 (36) times its height, 1.37 (2.03) in snout length, 2.68 (3.99) in length of ventral fold; its height 1.91 (1.04) in height of mid ventral fold; origin near likely position of undamaged sting apex. Ventral skin fold long, very narrow, length 2.03 (1.87) in disc width, 3.82 (3.05) in post cloacal tail; commencing almost below sting origin, origin 1.1% (1.0%) before sting origin; depth at quarter length 0.43 (0.27), at mid length 0.39 (0.30), at three quarter 0.32 (0.29) in adjacent tail height; distance from cloaca to sting origin 1.54 (1.62) in precloacal length; length of tail beyond ventral fold 0.65 (1.00) in fold length, 2.48 (3.04) in tail length. Lateral line on ventral surface distinct. Snout elongate, subtriangular; apex lobe-like, narrowly and bluntly pointed; angle 107 o (110 o); preoral snout length 2.58 (2.66) times mouth width, 2.34 (2.48) times internarial distance, 1.16 (1.23) times distance between first gill slits; direct preorbital snout length 2.05 (2.31) times interorbital length; snout to maximum disc width 1.97 (2.10) in DW; interorbital space broad, almost flat; eyes small, almost lateral, barely protruding in both types, ventral margin partly covered by thin skin fold; orbit not elevated above disc, diameter 1.00 (0.91) in spiracle length, eye diameter 1.58 (1.35) in spiracle length; inter-eye distance 3.81 (3.31) times eye diameter. Spiracles subrectangular to suboval, enlarged, opening dorsolaterally. Nostril elongate, suboval, directed posterolaterally; anterior margin fleshy; anterior nasal fold internal, broad, membranous; weak oronasal groove present; internasal distance 1.78 (1.93) in prenasal length, 2.51 (2.84) times nostril length. Nasal curtain skirt-like, relatively broad, short, width 1.74 (1.70) times length; not bilobed; surface flat, smooth, without longitudinal medial groove, not covered with minute pores; apex partly recessible within lateral margin of oronasal groove; lateral margin concave (possibly distorted through preservation; paratype almost straight), smooth edged; posterior margin not strongly fringed, weakly concave, not following contour of lower jaw, well removed from symphysis of lower jaw when mouth closed. Jaws asymmetric with teeth visible when mouth closed. Upper jaw strongly arched (teeth highly visible), symphysial part of jaw projecting ventrally; lower jaw strongly convex with truncate apex, broad band of symphysial teeth visible when mouth closed; lateral grooves deep (most evident on right side of holotype), almost straight, extending from nostril to well below lower jaw, longer than nasal curtain length. Lower jaw not projecting forward when mouth open, mouth not protrusible; skin on chin very fleshy, corrugate (more so in less dehydrated paratype); jaws of types not prised apart to reveal oral papillae. Teeth of adult male holotype rather large, variable in shape; those in mid lateral part of upper jaw almost plate-like, their crowns more than twice size of those either side; those at symphysis of upper jaw much smaller, upright, crowns with well-developed slender cusps; cusps much shorter in those teeth posterolaterally; crowns on symphysial teeth in lower jaw somewhat globular, slightly large than those at symphysis of upper jaw; those toward angle of lower jaw concealed; teeth not close-set in either jaw, in straight to semi-oblique rows, not obviously arranged quincuncially; rows in upper jaw ~31 (counted from photograph). Teeth of female paratype smaller than adult male, more similar in size and shape, more closely arranged, quincuncial. Gill openings S-shaped; length of first gill slit 1.36 (1.32) times length of fifth gill slit, 3.64 (3.38) times in mouth width; distance between first gill slits 2.02 (2.01) times internasal distance, 0.43 (0.41) times ventral head length; distance between fifth gill slits 1.31 (1.24) times internasal distance, 0.28 (0.25) times ventral head length. Squamation. Disc and tail with well-developed thorns and an otherwise largely naked disc; holotype with small, widely spaced stellate, dermal denticles on raised part of head; denticles sparse elsewhere, scattered few on post sting tail. Mid-dorsal thorn series in holotype in a single continuous, closely spaced row from nuchal region to sting base; 18 thorns on disc (8 on head, 10 on posterior disc), length 2.0– 6.3 mm, width 1.1 to 2.8 mm; 13 thorns on tail before sting scar, length 3.1–7.9 mm, width 2.1–2.8 mm; thorns very well developed, with rectangular bases and long, semi-erect, pungent lanceolate crowns; two similar thorns on each side of scapulocoracoid; thorns on tail much larger and more widely spaced than those on disc. Squamation of paratype less well developed; fewer dermal denticles on head (~ 12 in orbito-spiracular region); median thorn row less well developed, discontinuous, 8 thorns on disc (4 on head, 4 on posterior disc), length 2.8–5.5 mm, width 1.5–2.5 mm; 4 thorns on tail before sting base, length 6.2–7.2 mm, width 2.5– 2.5 mm. Both type specimens lacking an intact sting; distance from sting base to pectoral-fin insertion 49.4% (49.6%) DW; distance from cloaca to sting base 0.54 (0.53) in disc length. Clasper of adult moderately depressed, robust basally, convex distally and tapering to a blunt or narrowly rounded point; post cloacal length 31.0% DW. Total pectoral radials ~115 left side only (~115 right side only); propterygials ~51 (~47), mesopterygials 16–17 (17–18) and metapterygials ~48 left side only (~51 right side only). Total pelvic radials in right side of paratype 1 + ~26. Total vertebral segments (including first synarcual centra) 125 (126); all synarcual and monospondylous centra 38 (39); total diplospondylous centra 87 (87). Colouration. No information on live colour. In preservative (holotype): Uniformly brownish on dorsal surface, lightest around orbit, orbital membrane darker brown; thorns typical paler than surrounding skin, obvious. Ventral surface yellowish brown (blotching likely to be due to preservation and partial desiccation. Paratype: Dorsal surface pale brownish, lighter than holotype (possible artefact of preservation); scapular region, orbit and interorbit palest. Ventral surface almost uniformly white, paler than dorsal surface. Size.— Male holotype mature at 223 mm DW; stage of development of female paratype 206 mm DW unknown. Distribution. Type material reported as collected from Gischin, assumed to be Qishn in eastern Yemen, on the Arabian Sea coast (Fig. 6). Etymology. Epithet derived from the country of collection. Vernacular name: Heins’ stingray, after Marie and Wilhelm Hein who collected the type material shortly before Wilhelm’s death, and who also collected the holotype of the rare shark Carcharhinus leiodon. Remarks. The dasyatid genus Hemitrygon Müller & Henle 1838 presently comprises several small to medium-sized stingray species distributed in shallow marine, estuarine and freshwater environments. The new species H. yemenensis is currently known only from its collection locality (off eastern Yemen in the northwestern Indian Ocean), whereas almost all species of the genus Hemitrygon are largely restricted to the western Pacific (H. bennetti and H. parvonigra also occur in the eastern Indian Ocean) (Last et al., 2016). The extent of this disjunct distribution is unusual for a small-bodied coastal batoid, so additional scrutiny of the provenance of these specimens was required. However, we believe the collection locality of Gischin (Qishn) in Yemen is most likely correct for a number of reasons. Firstly, Wilhelm and Marie Hein undertook a well-documented expedition from Vienna to South Arabia from December 1901 – April 1902, most of which was spent in Gischin, although they stopped in Aden and Mukalla (Fig. 6). In Gischin they researched Mehri, a Modern South Arabian language that is unique to eastern Yemen and southern Oman. Wilhelm Hein is not known to have visited the western Pacific region (i.e. the core distribution of Hemitrygon) during his lifetime, and died in 1903, the year after the expedition to Gischin (Müller 1909, Klein-Franke 2006). Secondly, listings of NMW holdings of reptiles and teleost fish collected by the Heins on the same expedition appear to be consistent with the fauna of southern Arabia (ABMM, unpubl. data). Finally, the only other elasmobranch that the Heins collected in Gischin, the smoothtooth blacktip shark C. leiodon, has been recorded in eastern Yemen and the adjacent waters of southern Oman, despite having a highly limited known distribution (Henderson & Reeve, 2011; Moore et al., 2013), providing further evidence that the collection locality of H. yemenensis is likely correct. Hemitrygon yemenensis sp. nov. most closely resembles H. akajei (Müller & Henle, 1841) from the western North Pacific, H. bennetti (Müller & Henle, 1841) from the Indo-West Pacific and the Bay of Bengal, and H. fluviorum (Ogilby, 1908) from Australian seas, than any other 10 members of the genus treated recently in a guide to rays of the world (Last et al., 2016). These species all have well-developed thorns and tubercles in a median row on the disc and tail, and a short thorn row on each shoulder of adult males. However the disc shape of H. yemenensis is characteristically marked by a longer and more narrowly pointed snout than these species (and any other member of the genus), and the disc’s length is shorter than its width for all species of Hemitrygon other than H. yemenensis. Compared to the most similar species, H. bennetti, it has a shorter tail (cloaca origin to tail tip 163–188% DW in H. yemenensis vs. 209–258% DW in H. bennetti, n=6), longer disc (length 101–102% DW vs. 91–97% DW), head (length 51–52% DW vs. 44–49% DW) and snout (direct length 25–27% DW vs. 22–24% DW), slightly narrower interorbit (width 11.8–12.3% DW vs. 12.7–14.0% DW), larger eye and orbit (eye diameter 4.6–4.7% DW vs. 3.1–3.7% DW); longer proportions around the head of H. yemenensis are reflected by longer prenasal and preoral lengths. Although the elasmobranch fauna of Yemen has been poorly documented, at least fourteen other species of the family Dasyatidae either occur or are likely to occur there (Last et al., 2016). Of these, H. yemenensis is most similar in morphology to Brevitrygon walga (Müller & Henle, 1841), Maculabatis species (e.g. M. ambigua Last, Bogorodsky & Alpermann, 2016), and Pateobatis jenkinsii (Annandale, 1909) (Last et al., 2016) but unlike all of these species has dorsal and ventral folds on the tail. Hemitrygon yemenensis can also be distinguished from these species by a combination of small size of adult males, squamation (notably an absence of a denticle band, and the arrangement of thorns on disc, tail, and shoulder in adult males), and tail morphology (i.e. length, thickness). Although no fresh material was available for this study, colouration of live or freshly caught H. yemenensis may also be an important distinguishing field character and should be documented at the earliest opportunity. It is noteworthy that nearly 120 years have elapsed since the type specimens of H. yemenensis were collected, and the species has not been recorded nor additional material collected since. It remains to be seen if this apparent rarity is simply a reflection of a paucity of sampling effort near the type locality or a change in its conservation status.