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Etmopterus samadiae, new species Papuan Lanternshark (Figures 1���10; Table 1) Etmopterus sp.��� Fricke et al., 2014: 14 (Madang) Holotype. NTUM 10078 (tissue accession GN 17184), adult male 265 mm TL, east of Malmal Passage, Madang, Papua New Guinea, 05��07' S, 145��50' E, 527���539 m depth, 30 Nov 2012. Paratypes. 9 specimens: ASIZ P.73777, adult male 230 mm TL, ASIZ P.73778, female 188 mm TL, ASIZ P.73765, pregnant female 277 mm TL, off Lae, Huon Gulf, Morobe Province, Papua New Guinea, 06��51.841' S, 147��04.672' E, 395���406 m depth, 22 Aug 2010; NTUM 10313 (tissue accession GN 17195), female 269 mm TL, northern Cape King William, Morobe Province, Papua New Guinea, 06��00' S, 147��38' E, 785 m depth, 10 Dec 2012; NTUM 10314 (tissue accession GN 17197), female 258 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05��22' S, 145��48' E, 420���490 m depth, 14 Dec 2012; NTUM 10315 (tissue accession GN 17198), female 154 mm TL, Astrolabe Bay, Madang, Papua New Guinea, 05��22' S, 145��48' E, 340���385 m depth, 14 Dec 2012; NTUM 10316 (3 specimens; tissue accessions GN 17210���2), female 177 mm TL, subadult male 201 mm TL, female 228 mm TL, west of Kairiru Island, East Sepik, Papua New Guinea, 03��19' S, 143��27' E, 422���425 m depth, 19 Dec 2012. Diagnosis. Etmopterus samadiae is a relatively small, slender, species of linear���denticled Etmopterus that can be separated from its closest congeners within the E. lucifer clade by a combination of characteristics including the length of its anterior flank markings being slightly shorter than its posterior branch, long caudal base marking, and irregular and variable number of black, horizontal, dash-like marks on sides of body. The new species is morphologically and genetically (based on the NADH 2 marker) closest to E. brachyurus, but differs from this species in having a shorter posterior caudal marking (2.8���4.4 vs. 4.2���6.1% TL), a longer caudal base marking (10.6���14.1 vs. 7.0���7.8% TL), and flank marking with a slightly shorter posterior branch (9.1���11.2 vs. 11.4���12.6% TL). Description. Values expressed as a percentage of total length (TL) for the holotype, followed by the range of values for 9 paratypes (Table 1). Body fusiform, trunk sub-cylindrical (Fig. 1), width 1.1 (0.7���1.7) in trunk height; head sub-conical, long, 21.3 (20.6���23.7)% TL, slightly depressed, height 0.7 (0.5���0.8) times width. Snout moderately long, conical in lateral view, in dorsal view triangular���shaped becoming rounded at snout���tip (Fig. 3), head width 8.2 (9.8���11.8)% TL. Eyes oval-shape, large, orbit length 3.8 (3.0���3.7) in head and 2.6 (2.0���3.3) times orbit height; orbits with anterior and posterior notches; moderately spaced, inter���orbital space 1.2 (1.2���1.5) in width of head and orbit length 1.2 (1.1���1.4) times in inter���orbital distance. Spiracles small, semi-circular, greatest diameter 0.9 (1.4���2.5)% TL, 6.1 (2.7���4.0) times orbit length, distance to eye 2.9 (1.4���2.6)% TL, eye���spiracle length 0.7 (1.0���1.9) in orbit height. Nostrils large, oblique, length almost equal to internarial width, less than orbit length; anterior nasal flap well developed, triangular, anterior tip extending across nasal opening, length 1.0 (0.5���0.9) times spiracle length. Gill openings small, narrow, slightly oblique, in horizontal series, subequal in height, inter-gill length 4.9 (3.0���4.7)% TL. Mouth broad, length 3.4 (3.3���4.8) times in width, slightly arched, width 0.8 (0.7���0.9) times preoral length. Teeth dissimilar in upper and lower jaw (Fig. 4); upper teeth multicuspid in three functional series, functional teeth in lower jaw unicuspid in single series; multicuspid upper teeth small, upright, with strong central cusp flanked by 2 or 3 lateral cusplets on each side, decreasing in size distally; teeth in lower jaw fused into single row, blade-like, cusp oblique. Tooth count in first row of upper jaw 33 (27���28) and in first row of lower jaw 35 (28���31). First dorsal fin small, rounded at apex, length of first dorsal fin 8.6 (8.5���9.9)% TL, origin just anterior to pectoral-fin free rear tip; fin base insertion well anterior of pelvic-fin origin; pre���first dorsal fin length 1.3 (1.4���1.8) times inter���dorsal distance; first dorsal���fin spine straight, short, 1.6 (1.3���2.0) times height of first dorsal fin, located over pectoral fin rear margin. Second dorsal fin conspicuously larger, more erect than first dorsal fin, length of first dorsal fin 0.6 (0.6���0.8) times second dorsal fin, height of first dorsal fin 0.4 (0.5���0.7) times second dorsal fin; apex sub���angular, posterior margin concave, free rear tip elongated, length 13.2 (12.2���13.6)% TL, pre���second dorsal length 2.5 (2.6���3.1) times inter���dorsal distance; second dorsal���fin spine large, height slightly taller than fin, curved near tip towards fin apex; origin posterior to insertion of pelvic fins, over pelvic fin free rear tips. Interspace between first and second dorsal fins 1.0 (0.8���1.0) times pre���pectoral length. Pectoral fins relatively large, length 9.4 (10.1���11.3)% TL, subangular at free rear tips, base 2.0 (1.7���2.1) times in anterior margin, posterior margin nearly straight. Caudal peduncle relatively long, dorsal-caudal space 16.1 (13.6���15.3)% TL, height slightly greater than width, rounded, and tapering posteriorly. Caudal fin elongate, subequal to head length, terminal lobe distinct; length of lower preventral caudal fin margin less than one-half upper caudal fin margin. E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Total length (mm) 265 154 277 256 172.0 343.0 224 350 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 ......continued on the next page E. samadiae n.sp. E. fusus E. evansi E. brachyurus Holotype Paratypes NTUM n = 4 n = 4 Min. Max. 10318 Min. Max. Min. Max. Dermal denticles hook-like, posteriorly directed rearwards; organized in distinct rows laterally, characteristic of E. lucifer clade members (Fig. 5, 6). Distance between lateral rows mostly consistent along length, decreasing only very slightly towards caudal peduncle. Flank mark area denticles more dense and pointed ventrally. Luminescent markings distinct, intricate (Fig. 7 D); ventral head surface markings blackish, starting from almost at snout tip, extending to level of nostrils and orbits at just below level of anterior notch, then extending straight from just below posterior eye notch towards gill slits, weakly demarcated from belly marking by a weak band of transverse dermal folds across throat extending from below lower edges of first three gill openings on either side. Head dorsal surface photophore pattern as follows: a single midline along back originating at level of first gill slits extending posteriorly to caudal���fin origin; scattered photophores on paler fontanelle area of central head and also on paler area above orbits (as short dashes in some paratypes). Belly marking originates behind mouth on posterior portion of transverse dermal folds and extends ventrally along pectoral fin bases extending upwards to level with fin origin and upper edges of gill slits, and posteriorly to pelvic fin bases; ventral surface of pectoral fin very dark along lower base and upper base where ceratotrichia originate, bisected by distinct lighter to white band or blotch forming a two���prong fork pattern; margin on lateral trunk (pectoral���pelvic space) very clearly defined, line extending from rear margin of pectoral���fin base nearly to pelvic���fin insertion except for paler area at pelvic-fin origin; dark ventral belly surface continuous onto caudal peduncle to about lower caudal fin origin. Flank markings well defined (Fig. 7 d), consisting of both an anterior and posterior branch; anterior branch relatively short, length 10.3 (8.2���10.5)% TL, slender, nearly straight, extending well anterior to pelvic���fin origin; posterior branch straight, slightly thicker, width at level of base end of second dorsal fin 0.6 (0.5���0.8)% TL, slightly longer than anterior branch (shorter than anterior branch in paratype ASIZ P.73777), length 11.2 (9.1��� 10.8)% TL, extending to just anterior to second dorsal-fin free rear tip but well posterior to second dorsal-fin insertion; base of flank marking wide, origin slightly posterior to pelvic-fin insertion. Ventral caudal base marking distinct, short, length about equal to base of flank marking, not extending as a saddle on to caudal peduncle, anterior finger very short, posterior finger long, total length (including base) 11.4 (10.6���14.1)% TL. No central caudal marking. Posterior caudal fin marking very narrow, its length 3.2 (2.8���4.4) % TL. Vertebral counts: total counts 80 (80���86), total precaudal counts 55 (55���59); monospondylous 34 (36���37); diplospondylous precaudal 21 (19���22); caudal 25 (25���27). Coloration. In life, greyish to silvery black dorsally and laterally on body, becoming dark black ventrally; transition between lateral and ventral surfaces well demarcated by a paler lateral stripe below the flank markings in most specimens (less distinct in holotype and some paratypes). Dorsal midline with a broad, pale stripe originating just posterior to midpoint of inter-spiracle space, extending to first dorsal fin origin, continuing between dorsal fins, and from second dorsal fin insertion to upper caudal fin origin (Fig. 2); most prominent anterior to first dorsal fin and between dorsal fins, less prominent along upper caudal peduncle; a single row of dark photophores extending along middle of white dorsal stripe. Body with variable number and arrangement of short, horizontal, dash-like black markings (Fig. 8); most numerous in paratype NTUM 10314 (Fig. 8). Pectoral and pelvic fins dark at base and along anterior fin edge, becoming translucent to white on remainder of fins. Dorsal fins dark at base and along anterior edges, becoming translucent to white on remainder of fins. Black lateral flank markings demarcated by surrounding lighter colored lateral flanks (not sharply demarcated in most preserved specimens). Paler colored flank area on lateral surface between pectoral and pelvic fins sharply demarcates lateral and ventral surfaces; ventral surface black. Caudal fin with a distinct, large dark blotch centrally, occupying the area between the posterior finger of the caudal base marking and the upper caudal marking; posterior margin dark. Ventral surface mostly dark black around mouth, belly, and with a dark stripe between pelvic fin insertions and caudal origin; snout not distinctly paler than mouth. After preservation coloration similar but with paler markings often less obvious and coloration becoming a light or dark brown; dorsal median pale stripe and whitish flank area and black lateral flank markings less prominent but still distinct in most cases; dark blotch on central caudal fin obvious and darker pores extending laterally on body still clearly visible under microscope. Molecular analysis. The analysis of the NADH 2 data confirms that Etmopterus samadiae belongs in the E. lucifer clade and represents a monophyletic lineage that is distinct from, but most closely related to Etmopterus brachyurus (Fig. 9). It should be noted that this inference is based on a single mitochondrial marker. Inclusion of multiple nuclear markers could affect the presented inference. Size. Specimens examined ranged in size from 154 to 277 mm TL. A 277 mm TL female (ASIZ P.73765) was found to be pregnant (embryo caudal fin visible in cloaca) and two males 230 (ASIZ P.73777) and 265 mm TL (NTUM 10078) were determined to be mature. Distribution. The new species is known from off the northern Papua New Guinea mainland, from west of Kairiru Island in East Sepik Province to off Lae in the Huon Gulf (Morobe Province) and at a depth range of 340 to 785 m (Fig. 10). Etymology. The species is named after Dr Sarah Samadi (MNHN) who was one of the key principal investigators of the 2010 and 2012 MNHN expeditions in Papua New Guinea from which all the type specimens were obtained. The proposed English common name is Papuan Lanternshark., Published as part of White, William T., Ebert, David A., Mana, Ralph R. & Corrigan, Shannon, 2017, Etmopterus samadiae n. sp., a new lanternshark (Squaliformes: Etmopteridae) from Papua New Guinea, pp. 339-354 in Zootaxa 4244 (3) on pages 340-349, DOI: 10.11646/zootaxa.4244.3.3, http://zenodo.org/record/427411, {"references":["Fricke, R., Allen, G. R., Andrefouet, S., Chen, W. - J., Hamel, M. A., Laboute, P., Mana, R., Hui, T. H. & Uyeno, D. (2014) Checklist of the marine and estuarine fishes of Madang District, Papua New Guinea, western Pacific Ocean, with 820 new records. Zootaxa, 3834 (1), 1 - 247."]}

White, William T., Kawauchi, Junro, Corrigan, Shannon, Rochel, Elisabeth, Naylor, Gavin J. P. (2015): Redescription of the eagle rays Myliobatis hamlyni Ogilby, 1911 and M. tobijei Bleeker, 1854 (Myliobatiformes: Myliobatidae) from the East Indo-West Pacific. Zootaxa 3948 (3): 521-548, DOI: http://dx.doi.org/10.11646/zootaxa.3948.3.7

Myliobatis tobijei Bleeker, 1854 (Figs. 8 ���15, 17 b; Table 2) Myliobates aquila (not Linnaeus): Temminck & Schlegel, 1850: 310, pl. 142 (Japan); Boeseman, 1947: 228 (Japan); Tokida & Kobayashi, 1967: 180 (Japan). Myliobatis aquila (not Linnaeus): Bleeker, 1853: 22 (Japan); G��nther, 1880: 63 (Yokohama, Japan); Pietschmann, 1908: 638 (Japan). Myliobatis tobijei Bleeker, 1854: 425 (Nagasaki, Japan): Bleeker, 1855: 130 (Japan); Bleeker, 1859: 270 (Nagasaki, Japan); Bleeker, 1860: 10 (Nagasaki, Japan); Martens, 1876: 410 (Yokohama, Japan); Jordan & Snyder, 1900: 338 (Tokyo, Japan); Jordan & Snyder, 1901: 43 (Yokohama and Nagasaki, Japan); Jordan & Fowler, 1903: 663 (Tokyo, Japan); Jordan, 1905: 557 (Japan); Garman, 1913: 433 (Japan); Jordan & Metz, 1913: 6 (Fusan[=Busan]); Fowler, 1929: 507 (Japan); Fowler (1930): 186 (China); Schmidt, 1931: 15 (Nagasaki, Japan);? Fang & Wang, 1932: 277, fig. 27 (Chefoo [=Yantai], China);? Wang, 1933: 113 (Ningpo[=Ningbo], China); Masuda et al., 1984: 16, pl. 19 -D (Japan); Amaoka et al., 1989: 257 (Hokkaido, Japan); Nishida, 1990: 4, figs 9 C, 16 A, 19 D, 28 A, 38 H, 40 B, 57 B (Japan); Miya et al., 1994 a: 111 (Central Japan); Miya et al., 1994 b: 120 (Chitose, Japan); Miya et al., 1995: 198 (Choshi, Japan); Kitamura et al., 1996: 340 (Tokyo, Japan); Shinohara et al., 1998: 108 (Nagasaki, Japan); Compagno & Last, 1999: 1513, 1519, fig. (northwestern Pacific); Nakabo, 2002: 185, figs (Honshu, Shikoku and Kyushu, Japan); Yamaguchi, 2002: 29 (Ariake Bay, Japan); Senou et al., 2006: 404 (Sagami Bay, Japan); Matsuura et al., 2009: 15 (Aomori, Iwate, Miyagi and Fukushima, Japan); Abe et al., 2012: 190 (Japan); Grigorov & Orlov, 2013: 928 (Sea of Okhotsk); Nakabo, 2013: 229, figs (Japan). Myliobatis cornuta G��nther, 1870: 490 (Japan): Ishikawa & Matsuura, 1897: 60 (Matsushima, Japan); Philippi, 1892: 8 (Japan); Tokida & Kobayashi, 1967: 180 (Japan). Myliobatis cornutus Jordan & Snyder, 1901: 43 (Nagasaki, Japan). Myliobatis tobijaei: Luther, 1909: 152 (Yokohama, Japan). Aetobatis tobijei: Jordan et al., 1913: 30 (Hakodate to Nagasaki). Holorhinus hamlyni: Fowler, 1941: 460 (Japan). Holorhinus tobijei: Fowler, 1941: 463 (Japan, Korea,? China); Okada, 1955: 34, fig. (southern Japan); Okada & Mori, 1958: 35, fig. 33 (Mie Prefecture, Japan); Katayama & Fujioka, 1958: 1149 (Yamaguchi Prefecture, Japan); Dotsu & Tomiyama, 1967: 6 (Nagasaki, Japan); Ueno, 1971: 70 (southern Shakotan Peninsula, western Cape Erimo, coast of Soya and Kitami districts, and near Sendai Bay, Japan); Shiogaki & Dotsu, 1973: 15 (Nomozaki, Japan). Aetobatus tobijei: Kamohara, 1950: 19 (Tosa and Kishu, Japan); Kuroda, 1951: 317 (Hokkaido, Tokyo Bay, Suruga Bay, Kii Peninsula, Shikoku, Kyushu, Tsushima in Japan; Korea?); Kamohara, 1952: 11 (Kochi, Mimase, Urado and Susaki, Japan); Mori, 1952: 27 (Fusan[=Busan] and Quelpart[=Jeju] Island, Korea). Myliobatis tobiyei: Kamohara, 1958: 8 (Hokkaido to southern Japan). Myriobatis tobijei: Kamohara, 1964: 10 (Kochi Prefecture, Japan); Yamaguchi, 2004: 44, fig. 3 (Ariake Bay, Japan). Holotype. RMNH 7461 (largest of 2; holotype of Myliobatis tobijei), juvenile female 236 mm DW (405 mm TL), Nagasaki, Japan. Other specimens. (53 specimens): BMNH 1862.11. 1.74 (holotype of Myliobatis cornuta), adult male 429 mm DW, Japan; BMNH 1878.4. 54.86, 2 specimens, Japan; BMNH 1879.5. 14.423, subadult male, Japan; BMNH 1925.3. 13.2, 2 juvenile males 272 mm DW and 276 mm DW, China; BMNH 1937.7. 17.18 and 19, 2 embryos, male 159 mm DW, female 158 mm DW, Japan; FNU unregistered, dried jaws only from adult specimen, Ariake Bay, Japan; HUMZ 74436, adult male 619 mm DW (1257 mm TL), Japan; HUMZ 105905, juvenile male 266 mm DW, HUMZ 105906, juvenile male 277 mm DW, off Itado, Shimoda, Shizuoka Prefecture, Japan, 19 Nov. 1985; HUMZ 105937, juvenile male 304 mm DW, Shimoda Bay, Shimoda City, Shizuoka Prefecture, Japan, 18 Nov. 1985; HUMZ 105969, juvenile male 286 mm DW, HUMZ 105983, juvenile male 268 mm DW (480 mm TL), off Tatado, Shimoda City, Shizuoka Prefecture, Japan, 20 Nov. 1985; HUMZ 107419, female 342 mm DW, HUMZ 107420, female 396 mm DW (592 mm TL), HUMZ 107421, female 304 mm DW, HUMZ 107422, juvenile male 323 mm DW (536 mm TL), HUMZ 107423, female 356 mm DW, Shimoda Bay, Shimoda City, Shizuoka Prefecture, Japan, Sep. 1985; HUMZ 107437, female 295 mm DW (493 mm TL), Japan; HUMZ 107438, juvenile male 278 mm DW, HUMZ 107439, adult male 472 mm DW (~ 715 mm TL), Japan; HUMZ 107440, female 324 mm DW, Shimoda Bay, Shimoda City, Shizuoka Prefecture, 24 Nov. 1985; HUMZ 107454, juvenile male 296 mm DW, off Shirahama, Shimoda City, Shizuoka Prefecture, Japan, 1986; HUMZ 107455, juvenile male 254 mm DW (476 mm TL), HUMZ 107456, juvenile male 245 mm DW, off Shirahama, Shimoda City, Shizuoka Prefecture, Japan, 27 Jan. 1986; HUMZ 107457, female 404 mm DW (666 mm TL), off Shirahama, Shimoda City, Shizuoka Prefecture, Japan, 30 Jan. 1986; HUMZ 107458, female 299 mm DW, presumably Japan; HUMZ 109718, adult male 480 mm DW, Japan; HUMZ 109719, juvenile male 323 mm DW, HUMZ 109720, 308 mm DW (548 mm TL), HUMZ 109721, female 348 mm DW, HUMZ 109722, female 296 mm DW, HUMZ 109723, female 317 mm DW, HUMZ 109724, female 214 mm DW, Japan; HUMZ 109725, juvenile male 248 mm DW (469 mm TL), HUMZ 109726, juvenile male 303 mm DW, HUMZ 109727, juvenile male 317 mm DW, HUMZ 109728, subadult male 410 mm DW (742 mm TL), HUMZ 109729, female 312 mm DW, Japan; HUMZ 111032, female 257 mm DW, off Shirahama, Izu Peninsula, Shizuoka Prefecture, Japan, 3 Mar. 1986; HUMZ 114481, female 266 mm DW, presumably Japan; NSMT-P 53341, juvenile female 243.5 mm DW (462 mm TL), Tachibana Bay, Japan, 32 �� 35.64 ��� N, 129 �� 55.37 ��� E, 60 m depth, 9 Sep. 1996; NSMT-P 61770, juvenile male 248 mm DW (464 mm TL), Miho Peninsula, Honshu, Japan, 24 Apr. 1978; NSMT-P 65303, female 301 mm DW (534 mm TL), Totoro fishing port, Nobeoka City, Japan; NSMT-P 74603, juvenile female 264 mm DW (478.5 mm TL), Onahama, Iwaki City, Fukushima Prefecture, Honshu, Japan, 1931���1935; NSMT-P 75070, juvenile male 289 mm DW (561 mm TL), off Miyako, Shimohei County, Iwate Prefecture, Honshu, Japan, 6 or 7 Jan. 1938; NSMT-P 92524, female 261 mm DW (510 mm TL), Honshu, Japan; NSMT-P SK 871, juvenile female 287 mm DW (510 mm TL), eastern Honshu, Japan, 15 Oct. 1957; NAG002 (not retained), juvenile female 324 mm DW (605 mm TL), NAG003 (not retained), juvenile female 441 mm DW, NAG004 (not retained), pregnant female 656 mm DW, Nagasaki, Japan, June 2013. Diagnosis. A small Myliobatis (attaining about 665 mm DW) with the following combination of characters: dorsal surfaces yellowish brown, usually with variable, irregular dark blotches; tail with a very weak ventral skin fold; stinging spine(s) relatively long (longest spine 10.3���18.6 % DW); interorbital space shallowly concave in adult males; anterior margins of pectoral fins moderately convex; cranial fontanelle (visible in dorsal view without dissection) relatively narrow with mostly straight lateral margins; claspers of adult males 6.6���10.2 % DW; predorsal length 66.6���78.8 % DW; teeth in 7 rows in each jaw, with a broad median row flanked by three smaller rows on each side; pectoral-fin radials 79���85 (excluding rostral propterygial radials anterior of eyes); total vertebral centra (including synarcual) 115���124; males mature by 429 mm DW. Description. Disc diamond-shaped, broad, moderately long, width about 1.70 (1.53���1.71) times disc length; anterior projection 4.09 (3.61���4.32) in disc length; axis of greatest width of disc just posterior to scapular region, over anterior abdominal cavity, its horizontal distance from snout tip 1.48 (1.29���1.56) times in distance from tip of snout to pectoral-fin insertion; moderately deep, greatest thickness above scapular region and posterior head, thickness 8.62 (7.45���9.63) in disc width; without denticles or thorns; a short, bony ridge on midline above scapular region. Pectoral fins very large, wing-like, triangular, not or slightly falcate; anterior margin weakly to moderately convex, moderately convex distally; apex moderately rounded, pectoral angle 64 (58.5���67)��; posterior margin shallowly concave; free rear tip angular; inner margin moderately convex; length of anterior margin 47.7 (46.2��� 51.2)% DW, 1.11 (1.05���1.20) times its base length, inner margin 5.06 (5.30���7.97) in its base; origin below and just posterior to margin of eye; apex located at about level with pectoral mid-base; insertion just posterior to pelvic-fin origin, well anterior to dorsal-fin origin; free rear tip partly overlapping pelvic-fin anterior margin. Head pronounced, relatively low, short and broad; projecting well anterior to pectoral-fin origins; subquadrangular in cross-section at pectoral-fin origin; cranial region of head very broadly rounded to almost truncate in dorsoventral view; snout abruptly convex anterior of eyes, becoming deeply concave at origin of rostral lobe; moderately to slightly convex ventrally; ventral head length 27.3 (26.0��� 31.2)% DW, 1.58 (1.28���1.70) times width at pectoral-fin origins, 4.18 (3.38���5.30) times preorbital length (horizontal), 2.52 (2.32���3.30) times interorbital width; preoral snout length (1.00��� 1.41) times mouth width, 1.64 (1.35���1.84) times internarial width, 0.63 (0.55���0.84) times distance between first gill slits; head width at pectoral-fin origin 17.2 (17.7���21.9)% DW, 1.81 (1.63���2.21) times its height. Rostral lobe fleshy, broad, short (slightly longer in adult males); broadly rounded in dorsoventral view with a broadly rounded apex; narrowly rounded in lateral view; dorsal surface with a roughly T-shaped patch of pores medially; its length 5.2 (3.8���7.8)% DW, 5.22 (4.02���7.38) in head length, its width 1.14 (1.14���1.35) in head width at pectoral-fin origin. Interorbital space broad, shallowly convex (moderately convex in adult males) but with a broad medial depression over the cranial fontanelle, without ridges, denticles or thorns; interorbital width 10.8 (9.1���12.1)% DW, 1.56 (1.49���2.01) times orbit length, 0.66 (0.56���0.67) times head width at mid-eye; margins of cranial fontanelle narrowest posteriorly, gradually widening anteriorly, without a rapid change in degree of widening. Eyes moderately large, oval, lateral on head (eyes not visible in dorsal view), angling very slightly inwards anteriorly, diameter 2.04 (1.63���2.75) in spiracle length, 5.35 (4.98���7.47) in head width at pectoral-fin origin; orbits pronounced (very pronounced in adult males) and visible above upper margin of head; a small bony horn present above anterior quarter of each orbit in adult males. Spiracles moderately large, elliptical to slit-like, situated almost entirely laterally on head (not or only slightly visible in dorsal view), just posterior to orbit and above pectoral-fin origin, length 6.6 (6.1���7.5)% DW, 4.87 (2.80���5.70) times width; upper margins with a fleshy fold which is mostly nearly straight but angled inwards near posterior margin of spiracle. Nostrils narrowly oval with a narrow, fleshy oronasal groove; anterior nasal fold thin, membranous, internal; posterior nasal fold larger, fleshy, extending about two thirds of nostril width from its lateral margin; internarial space 0.98 (0.77���1.07) in prenasal length, 1.44 (1.30���1.84) times nostril length. Nasal curtain large, broad, elongate, width (1.58���1.98) times length; lateral margin weakly concave with a small lateral protuberance at widest point; posterior margin weakly concave, not divided by a medial notch, bordered by a long, curtain-like fringe which follows contour of lower jaw; apices subangular to moderately rounded; a patch of minute pores present near central posterior margin; apex and posterolateral margin recessible within oronasal groove; a small, low fleshy protuberance present on mid ventrolateral margin. Mouth moderately large, broad, transverse, located ventrally, width (8.2���10.3)% DW, (0.71 ���1.00) times preoral length, (1.89���2.30) in head width at pectoral-fin origin; margin of lower jaw slightly concave laterally and moderately convex medially, not indented at symphysis; not strongly protrusible, one series of anterior teeth of lower jaw sometimes visible when mouth closed; skin on chin and at margin of lower jaw fleshy, strongly furrowed, papillate. Teeth in 7 rows in each jaw, coalesced to form plates; middle series of teeth in both jaws broad and hexagonal, flanked by three rows of much smaller, similarly-sized, diamond-shaped teeth on each side arranged in a pavement-like fashion; median tooth row about 5 times width of outer tooth rows; upper tooth plate length slightly wider than long (based on FFNU unregistered, see Fig. 15 a); lower jaw tooth plate length about 1.4 times its width, its width about two thirds mouth width, in 11 series (based on FFNU unregistered, see Fig. 15 b). Gill openings small, elongated S-shaped, forming a weakly fringed lobe laterally; length of first gill slit 1.15 (1.19���1.73) times length of fifth gill slit, (3.45���5.42) in mouth width; distance between first gill slits 2.62 (2.01��� 2.55) times internarial space, 0.59 (0.49���0.64) times ventral head length; distance between fifth gill slits 1.42 (1.03���1.41) times internarial distance, 0.32 (0.25���0.35) times ventral head length. Pelvic fins relatively large, moderately broad, subquadrangular, anterior margin slightly concave, apex subangular, posterior margin slightly convex with scalloped edge, free rear tip very broadly rounded to subangular, inner margin slightly convex; extending well beyond pectoral-fin free tips; pelvic-fin length 16.9 (14.5���19.6)% DW, 1.23 (0.91���1.23) times width across fin bases, inner margin 6.2 (6.2 ���9.0)% DW. Claspers of adult males (n= 4) moderately long, moderately broad, not tapering distally except near tip, apex bluntly pointed, outer length (6.5��� 10.2)% DW. Dorsal fin small, slightly raked back, almost semicircular, its origin well posterior to pelvic-fin free rear tips by about an eye diameter or more; anterior margin moderately convex; apex broadly rounded, just anterior to or opposite insertion of fin; posterior margin moderately convex; free rear tip subangular, inner margin short, straight; predorsal length 1.42 (1.27���1.50) in disc width, fin length 6.6 (5.5���8.2)% DW, height 0.40 (0.37���0.52) times its length, inner margin 16.93 (4.75���12.73) in fin length. Tail very long, slender, whip-like, its length (from cloaca origin) 1.18 (0.94���1.48) times disc width; tapering gradually at base to stinging spine, and gradually becoming more whip-like beyond sting; base moderately compressed, suboval in cross section at pelvic-fin insertion, tail width at pelvic insertion 1.29 (1.29���1.85) times height; almost quadrangular in cross section near origin of stinging spine, width 0.97 (1.02���1.44) times height at first spine origin; a very weak, low dorsal skin fold present beyond sting; a distinct, low ventral skin fold present; a lateral skin fold present on either side from about level of mid-pelvic inner margin to below base of first stinging spine; a weak groove on dorsal surface of tail immediately posterior to base of stinging-spine(s), partially housing spines. Stinging spines 1���2, very elongate, slender, moderately broad-based, strongly tapered, mostly serrated laterally except for basal portion; distance from sting base to pectoral-fin insertion 25.7 (20.9���28.7)% DW; longest stinging spine 14.6 (10.3���18.6)% DW, 2.21 (1.59���2.90) times dorsal-fin length. Vertebral centra total (including synarcual) 115���124 (n= 7); total (excluding synarcual) 110���117 (n= 7); monospondylous (including synarcual) 36���37 (n= 9); monospondylous (excluding synarcual) 29���31 (n= 9); predorsal diplospondylous 34���44 (n= 9); post-dorsal diplospondylous 41 ���ca. 47 (n= 7). Total pectoral-fin radials (excluding rostral propterygial radials anterior of eyes) 79���85 (n= 12); propterygium (posterior of eyes) 20���22 (n= 7), mesopterygium 16���21 (n= 7), metapterygium 42���45 (n= 6). Pelvic-fin radials: 1 (2���3 fused elements) + 17��� 18 (n= 5 males); 1 (2 fused elements) + 24 (n= 1 female). Holotype Other specimens (n= 25) Min. Max. Mean ......continued on the next page Holotype Other specimens (n= 25) Min. Max. Mean Width of fifth gill slit 1.4 1.3 2.1 1.5 Distance from edge of disc to first gill slit 8.3 7.0 8.2 7.6 Distance between first gill slits 16.0 14.9 17.8 16.4 Distance between fifth gill slits 8.7 7.6 9.9 8.7 Tail at axil of pelvic fins (width) 5.4 5.2 6.5 5.9 Tail at axil of pelvic fins (height) 4.2 3.4 4.7 3.9 Tail at origin of stinging spine(s) (width) 1.9 2.0 3.0 2.5 Tail at origin of stinging spine(s) (height) 1.9 1.6 2.5 2.1 Pectoral-fin insertion to spine origin (horiz.) 25.7 20.9 28.7 24.0 Length of first stinging spine 14.6 4.4 18.6 12.6 Length of second stinging spine ��� 10.3 12.7 11.8 Pectoral-fin insertion to dorsal-fin origin (horiz.) 16.6 14.4 20.7 16.7 Dorsal-fin length 6.6 5.5 8.2 6.7 Dorsal-fin anterior margin 5.7 4.6 6.3 5.3 Dorsal-fin height 2.6 2.3 3.3 2.9 Dorsal-fin posterior margin 2.6 1.9 3.3 2.8 Dorsal-fin inner margin 0.4 0.4 1.4 1.0 Snout to anterior cloaca 53.8 51.0 62.8 54.7 Lower jaw to anterior cloaca 44.2 41.2 51.8 44.6 Cloaca anterior to tail tip 117.8 93.6 147.7 124.8 Cloaca anterior to stinging spine 27.3 20.3 26.5 23.7 Width across pelvic fin bases 13.8 14.1 17.9 16.1 Greatest span of pelvic fins 24.7 16.5 34.2 27.8 Pelvic-fin length 16.9 14.5 19.6 17.1 Pelvic-fin anterior margin 15.0 12.6 16.6 14.5 Pelvic-fin base 12.2 10.4 14.3 11.9 Pelvic-fin posterior margin 10.4 8.8 15.3 12.0 Pelvic-fin inner margin 6.2 6.2 9.0 7.3 Clasper outer length ��� 5.2 10.2 7.9 Clasper base width ��� 1.9 3.4 2.9 Colour. When fresh: Dorsal surface yellowish brown; irregular shaped and sized dark brown blotches variably present on dorsal surface of disc; blotches vary from completely absent (Fig. 9), to only several present, to densely covering dorsal surface (Fig. 11 a, b); dark (dorsal) and pale (ventral) surfaces well demarcated (waterline) along edge of pectoral fins and rostral lobe; paler laterally below eye. Tail dark dorsally; anterior third slightly paler ventrally with a mottled, diffuse waterline on mid-lateral margin; paler ventral coloration often just visible in dorsal view at base of tail; uniformly dark posteriorly. Ventral surface mostly whitish; pectoral fin apices, and anterior and posterior margins darkish brown, with junction between brown areas and whitish ventral colour strongly mottled; distal quarter of pelvic fins brownish; rostral lobe and nasal curtain white. Size. Males and females examined in this study (excluding embryos which were 158 and 159 mm DW) ranged in size from 245���619 and 214���656 mm DW, respectively. Five males between 429 and 619 mm DW were mature and a 410 mm DW male was adolescent, while 18 males between 245 and 323 mm DW were immature. A female of 656 mm DW was pregnant. Yamaguchi (2004) reported on an abnormal female specimen which was 665 mm, Published as part of White, William T., Kawauchi, Junro, Corrigan, Shannon, Rochel, Elisabeth & Naylor, Gavin J. P., 2015, Redescription of the eagle rays Myliobatis hamlyni Ogilby, 1911 and M. tobijei Bleeker, 1854 (Myliobatiformes: Myliobatidae) from the East Indo-West Pacific, pp. 521-548 in Zootaxa 3948 (3) on pages 532-542, DOI: 10.11646/zootaxa.3948.3.7, http://zenodo.org/record/242908, {"references":["Bleeker, P. (1854) Faunae ichthyologicae japonicae. Species Novae. Natuurkundig Tijdschrift voor Nederlandsch Indie, 6, 395 - 426.","Temminck, C. J. & Schelgel, H. (1850) Pisces. 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Myliobatis hamlyni Ogilby, 1911 (Figs. 1 ���8, 17 a; Table 1) Myliobatis hamlyni Ogilby, 1911: 40 (Queensland, Australia): Ogilby, 1916: 89 (Queensland, Australia); McCulloch, 1929: 29 (Queensland, Australia); Whitley, 1939: 258, fig. 18 (Queensland, Australia); Whitley, 1940: 223, fig. 256 (Queensland, Australia); Last & Stevens, 1994: 454, pl. 80, figs (northern Australia); Compagno & Last, 1999: 1513, 1519, fig. (Australia); Johnson, 1999: 718 (Moreton Bay, Queensland); Kyne et al., 2005: 325 (Queensland, Australia); Schwartz, 2008: 39 (Australia); Last & Stevens, 2009: 472, pl. 86, fig. 52.5 (northern Australia); Johnson, 2010: 307 (Queensland, Australia). ? Aetobatis tobijei: Snyder, 1912: 489 (Naha, Okinawa, Japan). Myliobatis tobijei: Chen, 1948: 11, fig. 9 (Taiwan); Chen & Chung, 1971: 44 (Taiwan);? Okamura & Kitajima, 1984: 73, fig. 40 (Okinawa Trough); Chen & Yu, 1986: 162 (Taiwan); Nakaya & Shirai, 1992: 41 (Okinawa Trough); Chen & Joung, 1993: 90, pl. 13 (fig. 5) (Taiwan); Shao et al., 1993: 126 (Taiwan);? Yano, 1999: 356 (Ryukyu Islands);? Randall & Lim, 2000: 111 (South China Sea); Chen, 2004: 25 (Taiwan); White et al., 2006: 300, fig. (Indonesia); White & Dharmadi, 2007: 1815 (Indonesia); Shao et al., 2008: 237 (Taiwan); Ward et al., 2008: p. 3 of Accessory Publication 1 (Indonesia); Shen & Wu, 2011: 104, fig. (Taiwan); Naylor et al., 2012: 82, fig. 62 (Taiwan, Philippines); Ebert et al., 2013: 367 (Taiwan); Straube et al., 2013: 276 (Taiwan). Holorhinus hamlyni: Fowler, 1941: 460 (Queensland, Australia). Myliobatis tobiyei: Kamohara, 1958: 8 (Formosa [= Taiwan]). Myliobatis cf. tobijei: Compagno et al., 2005: 79 (Philippines). ? Myliobatis sp.: Obara et al., 2009: 19, 20 (Tokyo Submarine Canyon)���uncertain. Aetobatus tobijei: Kuroda, 1951: 317 (Formosa [= Taiwan]). Holotype. QM I. 1567, juvenile male 270 mm DW (~ 474 mm TL), Moreton Island, Queensland, Australia, 27 ��02��� S, 153 �� 28 ��� E. Other specimens. Australia (5 specimens): AMS I 23718 -003, (1000 mm TL), off Coffs Harbour, New South Wales, Australia, 30 �� 23 ��� S, 153 �� 24 ��� E, 229 m depth, 25 Jul. 1981; CSIRO CA 2878, female 456 mm DW (907 mm TL), north of Forestier Island, Western Australia, 18 �� 31 ��� S, 118 ��09��� E, 200���201 m depth, 2 Apr. 1982; CSIRO H 2578 -01, adult male 704 mm DW (1351 mm TL), southwest of Shark Bay, Western Australia, 26 �� 45 ��� S, 112 �� 36.6 ��� E, 346���367 m depth, 31 Jan. 1991; CSIRO H 3860 -01, female 596 mm DW (1006 mm TL), east of Swansea, New South Wales, 33 ��07��� S, 151 �� 56 ��� E, 117���121 m depth, 9 Nov. 1994; QM I 33318, female 429 mm DW (943 mm TL), Swains Reef, Queensland, Australia, 20 �� 40 ��� S, 151 �� 20 ��� E, 210���220 m depth, 7 Aug. 1999. Indonesia (3 specimens): CSIRO H 5866 -02 (Genbank accession EU 398924), female 838 mm DW (1663 mm TL), Kedonganan fish market, Jimbaran Bay, Bali, 24 Mar. 2002; CSIRO H 5889 - 36, adult male 697 mm DW, Kedonganan fish market, Jimbaran Bay, Bali, Jul. 2002; MZB 15442, adult male 651 mm DW, Tanjung Luar fish market, Lombok, 11 Oct. 2004. Philippines (3 specimens): BRU 165, female 540 mm DW, BRU 166, subadult male 479 mm DW (1064 mm TL), Cadiz Pala-Pala, caught off Samal Island, 6 Apr. 2000; JPAG 329, female 663 mm DW, Pasil Fish Port II, Cebu City, Philippines, 13 Apr. 2000. Taiwan (6 specimens): AMNH 258303 (tissue accession GN 9949), female 395 mm DW, CSIRO H 6296 -06, female embryo 192 mm DW, Da-xi fish market, northeastern Taiwan, 25 May 2005; NMMBP 6308, Da-xi fish market, northeastern Taiwan, 30 Apr. 2002; NMMBP 15686, female 417 mm DW (849 mm TL), Da-xi fish market, northeastern Taiwan; NMMBP 16227 (tissue accession GN 9950), subadult male 478 mm DW (952 mm TL), Da-xi fish market, northeastern Taiwan, 13 Mar. 2012; NMMBP 18796, female 401 mm DW (802 mm TL), Taiwan; Japan (1 specimen): OCA-P 20030411 - 6, off Kunigami, Okinawa, 20 m depth, Apr. 2003. Diagnosis. A medium-sized Myliobatis (attaining at least 1142 mm DW) with the following combination of characters: dorsal surfaces uniformly purplish brown to greenish brown, without dark spots or blotches; tail with a distinct, low ventral skin fold; stinging spine(s) relatively long (longest spine 13.1���15.9 % DW); interorbital space moderately concave in adult males; anterior margins of pectoral fin straight to slightly convex; cranial fontanelle (visible in dorsal view without dissection) relatively wide with slightly convex lateral margins; claspers of adult males 5.5���5.7 % DW; predorsal length 63.6���68.6 % DW; teeth in 7 rows in each jaw, with a broad median row flanked by three smaller rows on each side; pectoral-fin radials 85���92 (excluding rostral propterygial radials anterior of eyes); total vertebral centra (including synarcual) 114���119; males mature at about 650 mm DW. Description. Disc diamond-shaped, broad, moderately long, width about 1.64 in holotype (1.61���1.76 in other specimens) times disc length; anterior projection 4.62 (3.50���4.17) in disc length; axis of greatest width of disc just posterior to scapular region, over anterior abdominal cavity, its horizontal distance from snout tip 1.38 (1.31���1.50) times in distance from tip of snout to pectoral-fin insertion; moderately deep, greatest thickness above scapular region and posterior head, thickness 7.89 (8.55���10.15) in disc width; without denticles or thorns; a short, bony ridge on midline above scapular region. Pectoral fins very large, wing-like, triangular, weakly falcate; anterior margin nearly straight to weakly convex, moderately convex distally; apex narrowly rounded, pectoral angle 61 (57���62)��; posterior margin shallowly concave; free rear tip angular; inner margin moderately convex; length of anterior margin 49.6 (46.9���49.8)% DW, 1.13 (1.11���1.20) times its base length, inner margin 6.36 (5.82���7.78) in its base; origin below posterior margin of eye; apex located at slightly posterior to pectoral mid-base; insertion posterior to pelvic-fin origin by about half eye diameter, well anterior to dorsal-fin origin; free rear tip overlapping about half or nearly entire pelvic-fin anterior margin. Head pronounced, relatively low, short and broad; projecting well anterior to pectoral-fin origins; subquadrangular in cross-section at pectoral-fin origin; cranial region of head very truncate in dorsoventral view; snout abruptly convex anterior of eyes, becoming deeply concave at origin of rostral lobe; weakly to moderately convex ventrally; ventral head length 28.1 (26.3���31.4)% DW, 1.40 (1.34���1.64) times width at pectoral-fin origins, 4.86 (3.62���6.16) times preorbital length (horizontal), 2.78 (2.64���3.39) times interorbital width; preoral snout length 1.25 (0.98���1.53) times mouth width, 1.48 (1.36���1.92) times internarial width, 0.73 (0.61���0.98) times distance between first gill slits; head width at pectoral-fin origin 20.1 (18.5���20.1)% DW, 1.98 (1.73���2.24) times its height. Rostral lobe fleshy, broad, short (slightly longer in adult males); broadly rounded to broadly parabolic in dorsoventral view with a bluntly pointed apex; narrowly rounded in lateral view; dorsal surface with a roughly Tshaped patch of pores medially; its length 5.0 (3.9���7.6)% DW, 5.04 (3.87���7.58) in head length, its width 1.19 (1.20���1.39) in head width at pectoral-fin origin. Interorbital space broad, weakly concave, with a broad medial depression over the cranial fontanelle, without ridges, denticles or thorns; interorbital width 10.1 (8.7���10.4)% DW, 1.59 (1.41���1.98) times orbit length, 0.53 (0.53���0.61) times head width at mid-eye; margins of cranial fontanelle narrowest posteriorly, gradually widening anteriorly, then rapidly becoming its widest near the front of head. Eyes relatively large, oval, lateral on head (eyes not visible in dorsal view), angling very slightly inwards anteriorly, diameter 1.88 (1.59���3.02) in spiracle length, 4.97 (5.05���7.71) in head width at pectoral-fin origin; orbits pronounced and visible above upper margin of head; a small, distinct bony protuberance present above anterior quarter of each orbit in adult males. Spiracles moderately large, elliptical to slit-like, situated almost entirely laterally on head (not or only slightly visible in dorsal view), just posterior to orbit and above pectoral-fin origin, length 7.6 (6.2���7.6)% DW, 3.40 (4.09���7.56) times width; upper margins with a fleshy fold which is mostly nearly straight but angled inwards near posterior margin of spiracle. Nostrils narrowly oval with a narrow, fleshy oronasal groove; anterior nasal fold thin, membranous, internal; posterior nasal fold larger, fleshy, extending about half of nostril width from its lateral margin; internarial space 0.79 (0.74���1.19) in prenasal length, 1.58 (1.39���2.03) times nostril length. Nasal curtain large, broad, elongate, width 1.63 (1.71���1.98) times length; lateral margin weakly concave with a small lateral protuberance at widest point; posterior margin weakly concave to straight, without a medial notch, bordered by a long, curtain-like fringe which follows contour of lower jaw; apices rounded; a patch of minute pores present near central posterior margin; apex and posterolateral margin recessible within oronasal groove; a small, low fleshy protuberance present on mid ventrolateral margin. Holotype Other specimens (n= 12) Min. Max. Mean ......continued on the next page Holotype Other specimens (n= 12) Min. Max. Mean Mouth moderately large, broad, transverse, located ventrally, width 8.8 (9.1 ���10.0)% DW, 0.80 (0.66���1.02) times preoral length, 2.28 (1.92���2.14) in head width at pectoral-fin origin; margin of lower jaw slightly concave laterally and moderately convex medially, not indented at symphysis; not strongly protrusible, one series of anterior teeth of lower jaw visible when mouth closed in some specimens; skin on chin and at margin of lower jaw fleshy, strongly furrowed, papillate. Teeth in 7 rows in each jaw, coalesced to form plates; middle series of teeth in both jaws broad and hexagonal, flanked by three rows of much smaller, similarly-sized, diamond-shaped teeth on each side arranged in a pavement-like fashion; median tooth row about 5 times width of outer tooth rows; upper tooth plate length almost equal to its width, in about 9 series (based on CSIRO H 5889 - 36, Fig. 7 a); lower jaw tooth plate length about 1.6 times its width, its width almost two thirds mouth width, in 13 series (based on adult male CSIRO H 5889 - 36, Fig. 7 b). Gill openings small, elongated S-shaped, forming a weakly fringed lobe laterally; length of first gill slit 1.53 (1.12���1.59) times length of fifth gill slit, 4.50 (3.83���5.79) in mouth width; distance between first gill slits 2.03 (1.95���2.33) times internarial space, 0.54 (0.47���0.61) times ventral head length; distance between fifth gill slits 1.12 (0.95���1.13) times internarial distance, 0.30 (0.23���0.31) times ventral head length. Pelvic fins relatively large, moderately broad (broadest in adult females), subquadrangular, anterior margin nearly straight to very slightly convex, apex subangular, posterior margin mostly straight with scalloped edge, free rear tip very broadly rounded to subangular, inner margin slightly convex; extending well beyond pectoral-fin free tips; pelvic-fin length 15.3 (14.3 ���16.0)% DW, 1.06 (0.94���1.12) times width across fin bases, inner margin 6.7 (6.1��� 7.7)% DW. Claspers of adult males (n= 2) moderately long, moderately broad, not tapering distally except near tip, apex bluntly pointed, outer length (5.5���5.7)% DW. Dorsal fin small, raked back, its origin just to well posterior to pelvic-fin free rear tips; anterior margin moderately convex; apex broadly rounded, just anterior to insertion of fin; posterior margin moderately convex; free rear tip subangular, inner margin short, straight; predorsal length 1.54 (1.46���1.57) in disc width, fin length 6.6 (5.8���7.3)% DW, height 0.55 (0.44���0.56) times its length, inner margin 4.70 (3.96���8.26) in fin length. Tail very long, slender, whip-like, its length (from cloaca origin) 1.24 (1.19���1.70) times disc width; tapering gradually to about mid-length of stinging spine, and gradually becoming more whip-like beyond sting; base compressed, oval in cross section at pelvic-fin insertion, tail width at pelvic insertion 1.44 (1.15���1.85) times height; almost quadrangular to hexangular in cross section near origin of stinging spine, width (1.02���1.37) times height at first spine origin; no dorsal skin fold (rarely a very weak fold evident); a weak, very low ventral skin fold usually present; a lateral skin fold present on either side from about level of mid-pelvic inner margin to below base of first stinging spine; a weak groove on dorsal surface of tail immediately posterior to base of stinging-spine(s), partially housing spines. Stinging spines 1���2, very elongate, slender, moderately broad-based, strongly tapered, mostly serrated laterally except near its base; distance from sting base to pectoral-fin insertion (18.1���23.1)% DW; longest stinging spine (13.1���15.9)% DW, (2.26) times dorsal-fin length. Vertebral centra total (including synarcual) 114���119 (n= 9); total (excluding synarcual) 108���114 (n= 9); monospondylous (including synarcual) 30���38 (n= 13); monospondylous (excluding synarcual) 24���32 (n= 13); predorsal diplospondylous 37���48 (n= 10); post-dorsal diplospondylous 36���45 (n= 9). Total pectoral-fin radials (excluding propterygial radials anterior of eyes) 85���92 (n= 12); rostral propterygium (anterior of eyes) ~ 8���11 (n= 7); propterygium (posterior of eyes) 20���23 (n= 12), mesopterygium 18���22 (n= 13), metapterygium 42���49 (n= 12). Pelvic-fin radials: 1 (2 fused elements) + 17 ���~ 19 (n= 5 males); 1 (2���3 fused elements) + 21 ���~ 24 (n= 8 females). Colour. When fresh: Dorsal surface uniformly purplish brown to greenish brown; no spots or blotches; dark (dorsal) and pale (ventral) surfaces well demarcated (waterline) along edge of pectoral fins and rostral lobe; slightly paler laterally below eye. Tail dark dorsally; anterior third paler ventrally with a mottled, diffuse waterline on lower lateral margin; uniformly dark posteriorly. Ventral surface mostly whitish; pectoral fin apices and most of posterior margins darkish brown, with junction between brown areas and whitish ventral colour strongly mottled; distal third of pelvic fins brownish; rostral lobe and nasal curtain white. Size. Specimens examined in this study ranged in size from 270���838 mm DW (up to 1663 mm TL). Two males of 478 and 479 mm DW were adolescent, while two of 697 and 704 mm DW were adult. Compagno et al. (2005) also reported two additional adolescent males of 557 and 571 mm DW. An additional six adult males recorded at the Tanjung Luar and Kedonganan fish landing sites in Indonesia ranged from 651���800 mm DW and 9 females ranged from 600���1142 mm DW. Distribution. Patchy distribution in the East Indo ���West Pacific (Fig. 8). In Australia, known from several locations off eastern Australia (Swain Reefs and the type locality Cape Moreton in Queensland; and off Swansea and Coffs Harbour in New South Wales) and off Western Australia (off Forestier Island and Shark Bay) in depths of 120��� 350 m. In Indonesia, recorded in the landings at the Tanjung Luar fishing port in eastern Lombok and the Kedonganan fish market in southern Bali. Depth of capture unknown but landed with Mustelus and Squalus spp., presumably on the outer continental shelf and upper slope; not landed with shallower water species such as Rhinobatos, Gymnura and dasyatids. In the Philippines, reported from the Pasil fish market in Cebu City and Palapala fishing port in Cadiz City (Compagno et al., 2005). Recorded in the South China Sea by Randall & Lim (2000), but specimens needed to confirm their identity. In Taiwan, recorded from the Da-xi fish market in Yilan; no depth information available. In Japan, one specimen was taken from off Okinawa. Records from Ryukyu Islands and Okinawa Trough (Okamura & Kitajima, 1984; Yano, 1999) possibly refer to this species but this requires verification. Specimens recorded off the Tokyo Submarine Canyon by Obara et al. (2009), as Myliobatis sp., between 150 and 500 m depth may also refer to this species, but this needs to be confirmed. Deepwater records of M. tobijei from Japan may refer to this species., Published as part of White, William T., Kawauchi, Junro, Corrigan, Shannon, Rochel, Elisabeth & Naylor, Gavin J. P., 2015, Redescription of the eagle rays Myliobatis hamlyni Ogilby, 1911 and M. tobijei Bleeker, 1854 (Myliobatiformes: Myliobatidae) from the East Indo-West Pacific, pp. 521-548 in Zootaxa 3948 (3) on pages 523-531, DOI: 10.11646/zootaxa.3948.3.7, http://zenodo.org/record/242908, {"references":["Ogilby, J. D. (1911) Descriptions of new or insufficiently described fishes from Queensland waters. Annals of the Queensland Museum, 10, 36 - 58.","Ogilby, J. D. (1916) Check-list of the cephalochordates, selachians, and fishes of Queensland. Memoirs of the Queensland Museum, 5, 70 - 98.","McCulloch, A. R. (1929) A checklist of the fishes recorded from Australia. Part I. Australian Museum Memoir, 5 (1), 1 - 144. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.473","Whitley, G. P. (1939) Taxonomic notes on sharks and rays. The Australian Zoologist, 9, 227 - 262.","Whitley, G. P. (1940) The fishes of Australia. Part 1. The sharks, rays, devil fishes and other primitive fishes of Australia and New Zealand. Royal Zoological Society of New South Wales, Sydney, 230 pp.","Last, P. R. & Stevens, J. D. (1994) Sharks and Rays of Australia. CSIRO Publishing, Hobart, 513 pp.","Compagno, L. J. V. & Last, P. R. (1999) Myliobatidae: eagle rays. In: Carpenter, K. E. & Niem, V. H. (Eds.). The living marine resources of the Western Central Pacific. FAO species identification guide for fishery purposes. Volume 3; Batoid fishes, chimaeras and bony fishes part 1 (Elopidae to Linophrynidae). Food and Agricultural Organization (FAO), Rome, pp. 1511 - 1519","Johnson, J. W. (1999) Annotated checklist of the fishes of Moreton Bay, Queensland, Australia. Memoirs of the Queensland Museum, 43, 709 - 762.","Kyne, P. M., Johnson, J., Courtney, A. J. & Bennett, M. B. (2005) New biogeographic information on Queensland chondrichthyans. Memoirs of the Queensland Museum, 50, 321 - 327.","Schwartz, F. J. (2008) A survey of the tail spine characteristics of stingrays frequenting Indo-Pacific ocean areas between the International Date Line and the Chagos-Maldive Archipelago waters. Journal of the North Carolina Academy of Science, 124 (2), 27 - 45.","Last, P. R. & Stevens, J. D. (2009) Sharks and Rays of Australia. 2 nd Edition. CSIRO Publishing, Melbourne, 644 pp.","Johnson, J. W. (2010) Fishes of the Moreton Bay Marine Park and adjacent continental shelf waters, Queensland, Australia. Memoirs of the Queensland Museum, 54, 299 - 353.","Snyder, J. O. (1912) The fishes of Okinawa, one of the Riu Kiu Islands. Proceedings of the United States National Museum, 42, 487 - 519. http: // dx. doi. org / 10.5479 / si. 00963801.42 - 1913.487","Chen, J. T. - F. (1948) Notes on the fish-fauna of Taiwan in the collections of the Taiwan Museum. I. Some records of Platosomeae from Taiwan, with description of a new species of Dasyatis. Quarterly Journal of the Taiwan Museum, 1, 1 - 14. [Taipei]","Chen, J. T. - F. & Chung, I. - H. (1971) A review of rays and skates or Batoidea of Taiwan. Tunghai University, Biological Bulletin 40, Ichthyology Series, 8, 1 - 53.","Okamura, O. & Kitajima, T. (Eds.) (1984) Fishes of the Okinawa Trough and the adjacent waters. Vol. 1. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, 414 pp.","Chen, J. T. - F. & Yu, M. - J. (1986) A synopsis of the vertebrates of Taiwan. Vol. 1. Taiwan Commercial Press, Taipei, 442 pp.","Nakaya, K. & Shirai, S. (1992) Fauna and zoogeography of deep-benthic chondrichthyan fishes around the Japanese Archipelago. Japanese Journal of Ichthyology, 39 (1), 37 - 48.","Chen, C. - T. & Joung, S. - J. (1993) Chondrichthyes. In: Shen, S. - C., Lee, S. - C., Shao, K. - T., Mok, H. - K., Chen, C. - T. & Chen, C. - H. (Eds.), Fishes of Taiwan. Department of Zoology, National Taiwan University, Taipei, pp. 1 - 960.","Shao, K. - T., Chen, J. - P., Kao, P. - H. & Wu, C. - Y. (1993) Fish fauna and their geographic distribution along the western coast of Taiwan. Acta Zoologica Taiwanica, 4, 113 - 140.","Yano, K. (1999) Chondrichthyans of the Ryuku Islands, Japan. In: Seret, B. & Sire, J. Y. (Eds.), Proceedings of the 5 th Indo- Pacific Fish Conference, Noumea. Societe Francaise d'Ichtyologie, Paris, pp. 351 - 365.","Randall, J. E. & Lim, K. K. P. (2000) A checklist of the fishes of the South China Sea. The Raffles Bulletin of Zoology, Supplment 8, 569 - 667.","Chen, C. - T. (2004) Checklist of the fishes of Penghu. FRI Special Publication No. 4, 175 pp.","White, W. T., Last, P. R., Stevens, J. D., Yearsley, G. K. & Dharmadi, F. (2006) Economically Important Sharks and Rays of Indonesia. ACIAR Monograph Series, No 124, ACIAR Publishing, Canberra, 329 pp.","White, W. T. & Dharmadi (2007) Species and size compositions and reproductive biology of rays (Chondrichthyes, Batoidea) caught in target and non-target fisheries in eastern Indonesia. Journal of Fish Biology, 70, 1809 - 1837. http: // dx. doi. org / 10.1111 / j. 1095 - 8649.2007.01458. x","Shao, K. - T., Ho, H. - C., Lin, P. - L., Lee, P. - F., Lee, M. - Y., Tsai, C. - Y., Liao, Y. - C., Lin, Y. - C., Chen, J. - P. & Yeh, H. - M. (2008) A checklist of the fishes of southern Taiwan, northern South China Sea. The Raffles Bulletin of Zoology, Supplement 19, 233 - 271.","Ward, R. D., Holmes, B. H., White, W. T. & Last, P. R. (2008) DNA barcoding Australasian chondrichthyans: results and potential uses in conservation. Marine and Freshwater Research, 59, 57 - 71. http: // dx. doi. org / 10.1071 / MF 07148","Shen, S. - C. & Wu, K. - Y. (2011) Fishes of Taiwan. National Museum of Marine Biology and Aquarium, Pingtung, 896 pp.","Naylor, G. J. P., Caira, J. N., Jensen, K., Rosana, K. A. M., White, W. T. & Last, P. R. (2012) A DNA sequence-based approach to the identification of shark and ray species and its implications for global elasmobranch diversity and parasitology. Bulletin of the American Natural History Museum, 367, 1 - 263. http: // dx. doi. org / 10.1206 / 754.1","Ebert, D. A., White, W. T., Ho, H. - C., Last, P. R., Nakaya, K., Seret, B., Straube, N., Naylor, G. J. P. & de Carvalho, M. R. (2013) An annotated checklist of the chondrichthyans of Taiwan. Zootaxa, 3752 (1), 279 - 386. http: // dx. doi. org / 10.11646 / zootaxa. 3752.1.17","Fowler, H. W. (1941) The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi obtained by United States Bureau of Fisheries Steamer Albatross in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bulletin of the United States National Museum, Series 100, 13, 1 - 879.","Kamohara, T. (1958) A catalogue of fishes of Kochi Prefecture (Province Tosa), Japan. Reports of the USA Marine Biological Station, 5 (1), 1 - 76.","Compagno, L. J. V., Last, P. R., Stevens, J. D. & Alava, M. N. R. (2005) Checklist of Philippine Chondrichthyes. CSIRO Marine Laboratories Report 243, Hobart, 103 pp.","Obara, G., Jo, K., Yamagami, K., Kojima, T. & Taniuchi, T. (2009) Species composition of chondrichthyans in the Tokyo Submarine Canyon. Report of Japanese Society for Elasmobranch Studies, 44, 8 - 20.","Kuroda, N. (1951) A nominal list with distribution of the fishes of Suruga Bay, inclusive of the freshwater species found near the coast. Japanese Journal of Ichthyology, 1 (5), 314 - 338.","Holotype. QM I. 1567, juvenile male 270 mm DW (~ 474 mm TL), Moreton Island, Queensland, Australia, 27 ° 02 ' S, 153 ° 28 ' E."]}

Documentation of conducted analyses. (PDF 1710Â kb)

Last, Peter R., Corrigan, Shannon, Naylor, Gavin (2014): Rhinobatos whitei, a new shovelnose ray (Batoidea: Rhinobatidae) from the Philippine Archipelago. Zootaxa 3872 (1): 31-47, DOI: http://dx.doi.org/10.11646/zootaxa.3872.1.3

Rhinobatos whitei sp. nov. [English Name: Philippine Guitarfish] (Figs 1���8, Tab. 1) Rhinobatos schlegelii: Casto de Elera, 1895, 619; Manila, Luzon. Fowler, 1941, 308; listing of Casto de Elera���s record without new material. Compagno & Last, 1999, 1429. Western Central Pacific, Philippines (misidentifications)? Rhinobatos cf. schlegelii M��ller & Henle, 1841: Compagno et al., 2005. Holotype. PNM 15189 (formerly SUML F 1086 and JPAG 079), adolescent male 597 mm TL, Dipolog City Public Market, collected J.A. Gaudiano, 10 April 1999. Paratypes. 10 specimens, Suba, Pasil and Pasil Fish Port I, Cebu City; Dipolog City Public Market; Palapala Fish Port, Bacolod City; Palapala Fish Port, Cadiz City; Punta Miray, Baliangao; Silliman Beach, Dumaguete City, PNM 10318 (formerly SUML JPAG 277), female, 620 mm TL, 6 April 2000; CSIRO H 7542 ���01 (formerly SUML JPAG 300), mature male, 641 mm TL, 10 April 2000; CSIRO H 7542 ���02 (formerly SUML JPAG 301), female, 494 mm TL, 10 April 2000; CSIRO H 7542 ���03 (formerly SUML JPAG 306, mature male, 556 mm TL, 10 April 2000; CSIRO H 7542 ���04 (formerly SUML JPAG 308), mature male, 636 mm TL, 10 April 2000; SUML JPAG 309, immature male, 471 mm TL, 10 April 2000; CSIRO H 7542 ���05 (formerly SUML JPAG 310), mature male, 617 mm TL, April 10, 2000; SUML BRU 144, immature male, 317 mm TL, 28 March 2000; SUML MMLM 0 0 1, female, 764 mm TL, 1 March 1999; SUML F 1 (formerly MMLM 012), female, 686 mm TL, 1 April 1999. Other material. 11 specimens. SUML RSE 0 0 5, mature male, 705 mm TL, 4 March 1999; SUML JPAG 047a, male, 556 mm TL, 21 March 1999; SUML JPAG 278, female, 690 mm TL, 6 April 2000; SUML JPAG 307, immature male, 497 mm TL, 10 April 2000; SUML JPAG 311, female, 473 mm TL, 10 April 2000; SUML JPAG 0 43, mature male, 643 mm TL, 21 March 1999; SUML JPAG 0 45, female, 661 mm TL, 21 March 1999; SUML BRU 0 71, female, 820 mm TL, 15 April 1999; SUML BRU 0 72, female, 820 mm TL, 15 April 1999; SUML BRU 0 73, mature male, 720 mm TL, 15 April 1999; SUML BRU 0 74, female, 844 mm TL, 15 April 1999. Diagnosis. A species of the genus Rhinobatos distinguished by the following combination of adult characters: wedge-shaped disc with an moderately concave anterior snout margin, width 31���35 % TL, disc length 1.3���1.4 times width; snout length 3.0��� 3.4 times interspiracular distance, 3.9���5.2 times interorbital width; medium-sized orbit, 1.4���1.7 times spiracle length; weakly oblique nostrils, their length 1.3���1.6 times internarial distance; preoral length 6.8���7.8 times internarial distance; anterior nasal flaps penetrating slightly into internarial space but well separated at their insertion; posterior nasal flaps broad; large spiracles with two folds, outermost fold distinctly longer than inner fold; ridges of rostral cartilage almost parallel, converging slightly anteriorly but not constricted medially; anterior cartilage subtriangular; distance between fifth gill slits 3.0��� 3.4 times in ventral head length; prebranchial sensory pore patch obvious, extending to just behind first gill slit; postscapular sensory canal obvious, with exposed lateral pores, grooved; thorn patches on supraorbit, nuchal and scapular regions, and dorsal midline rudimentary, not conspicuous; pelvic-fin inner margin longer than its base in males, subequal in females; interdorsal distance 2.6���3.2 times length of first dorsal-fin base; dorsal caudal margin 2.1���2.6 times preventral margin; upper jaw with ca. 65���92 tooth rows; 173���179 post-synarcual centra; 50���53 nasal lamellae; dorsal surface of adults brownish, covered with diffuse orange and dusky blotches, and indistinct pale spots; melanophores pronounced on dorsal surface, and dorsal fins largely pale anteriorly, darker posteriorly with a blackish blotch above the free rear tip when fresh. Description. Disc broadly wedge shaped, bluntly angular anteriorly and mildly bottlenose shaped, margin concave near tip then almost straight to its apex; snout angle before eyes 56 o (56���60 o in 4 paratypes); outer margins broadly rounded, more narrowly rounded distally; females slightly broader than males, length 1.37 in holotype (1.31���1.37 in adult male paratypes, 1.27���1.30 in adult-sized females) times width. Pelvic fins elongate, shortbased, base length 0.84 (0.85���1.06 in all paratypes) of inner margin length; total length 2.12 (1.81���2.29) times their base length, 2.69 (2.15���2.58) times width; anterior margin double concave, strongly concave anteriorly, apex broadly rounded, posterior margin noticeably convex. Tail elongate, slender anteriorly, tapering gradually; in crosssection nearly flat below, rounded slightly above; tail length from anterior cloaca 1.38 (1.28���1.46) times precloacal length, 1.35 (1.21���1.44) times disc length, 6.26 (5.90���6.10, 4.98���6.44) times its width at pelvic-fin insertions; tail width sexually dimorphic anteriorly, 2.09 (2.16���2.20, 2.28���2.51) times depth at pelvic-fin insertions, 2.52 (2.40���2.50, 2.39���2.73) at first dorsal-fin origin, 2.03 (1.90���2.12, 1.93���2.17) at second dorsal-fin origin. Dermal fold lateral on tail, originating forward of free rear tip of pelvic fin, reaching just behind ventral caudal-fin origin; fold moderately well developed, maximum width at about at end of interdorsal space; width about half width of Head elongate, its ventral length 29.2 (27.4���31.3)% TL; snout moderately long and bluntly pointed; preoral length 3.41 (2.92���3.31) times mouth width, 7.43 (7.26���7.79, 7.11���7.52) times internarial distance, 1.45 (1.21���1.46) times dorsal caudal-fin margin, 5.44 (5.45���6.44, 4.60���5.48) times distance from nostril to margin of disc; preorbital snout length (direct) 3.40 (2.95���3.31) times interspiracular length, 4.69 (3.90���4.80) times orbit diameter, 5.22 (3.94���5.01) times interorbital width; interorbital space weakly concave to flat, rather broad; eye dorsolateral, not elevated or protruding, orbit relatively small, diameter 1.58 (1.35���1.68) times spiracle length, 1.11 (0.82���1.21) times interorbital distance. Spiracles narrowly bean-shaped, relatively large; two strongly compressed folds on posterior margin, length of innermost spiracular fold 0.8 (0.7���0.9) of outer fold length; distance between bases of folds 0.7 (0.5���0.8) length of shortest fold. Nostril moderately large, oblique, nasal flaps well developed; anterior aperture suboval, width slightly exceeding length; nostril length 3.22 (2.93���3.49) times anterior aperture width, 1.35 (1.30���1.57) times anterior nasal-flap base 0.94 (1.14���1.18, 0.93���1.05) times distance from nostril to edge of disc, 1.29 (1.30���1.61) times internarial distance. Anterior nasal flap moderate with long, slender process anteriorly; flap base length 1.86 (1.90���2.09, 1.28���1.70) times its width at process, 2.38 (2.03���2.47) times anterior aperture width; inserted by about length of longest spiracular fold into internarial space; distance between insertions of flaps 3.38 (3.27���3.88) in greatest distance across nostrils anteriorly, 1.05 (1.00��� 1.09) in minimum internarial distance; process of flap about twice as long as wide, bluntly pointed distally, abutting posterolateral nasal flap and determining hind margin of anterior aperture. Posterolateral nasal flap lobe-like, broadest medially, length 4.99 (3.06���6.27) times width; originating just behind lateral extremity of anterior nasal aperture, extending postero-medially as a free fold to about level of insertion of anterior nasal flap and almost to inner edge of posterior nasal flap. Posterior nasal flap strongly lobe-like, base length 2.22 (1.85���2.74) times its width, its innermost edge well short of innermost margin of nostril; width 0.98 (0.80���1.07) times anterior aperture width, 1.65 (1.02���2.19) times posterolateral nasal-flap width. Mouth width 1.69 (1.50���1.79) times nostril length, 7.06 (6.61���7.26) in precloacal length; positioned over hind margin of orbit. Upper jaw weakly convex, upper lip arched slightly; lower lip pronounced, separated from oral groove by ridges of strongly corrugated skin; deep, short lateral grooves around corners of mouth. Teeth very small, rhomboidal, quincuncial; crowns blunt or with weak posterior cusps; cusps most prominent near jaw angle; ca. 78 (65���92) rows in upper jaw; upper and lower jaw teeth similar in shape and size, similar between sexes, slightly larger toward symphysis. Gill openings s-shaped, fifth less so; length of third gill slit 2.52 (2.51���2.90, 2.00��� 2.34) in nostril length, 6.15 (5.83���6.41, 5.31���5.76) in distance between fifth gill slits; distance between first gill slits 1.42 (1.41���1.51) times distance between fifth gill slits; distance between fifth gill slits 3.14 (3.17���3.73) times internarial distance, 1.44 (1.42���1.61) times mouth width, 0.29 (0.29���0.34) of ventral head length. Dorsal fins variable in shape and height, upright, broad to falcate; apices narrowly rounded to almost angular; anterior margins weakly convex, more extremely convex in some individuals; posterior margins weakly to deeply concave; free rear tips almost forming right angle, not produced; first dorsal fin marginally taller than second, length of first 1.05 (0.84���1.02) times its height, base length 1.38 (1.58���1.81) times inner margin length; second dorsal-fin length 1.15 (1.02���1.28, 0.95���1.12) times its height, base length 2.06 (1.96���2.54) times inner margin length. First dorsal fin well behind pelvic-fin rear tip, interspace 0.78 (0.74���0.96) times interdorsal distance; interdorsal space moderate, 2.12 (1.79���1.98, 1.68���1.97) times second dorsal-fin height, 3.19 (2.63���2.91) times base of first dorsal fin, 1.43 (1.33���1.41, 1.11���1.37) times tail width at origin of first dorsal fin, 1.71 (1.64���1.81) times interspace between second dorsal-fin insertion and upper origin of caudal fin. Caudal fin medium-sized, deep; dorsal caudal margin 2.28 (2.05���2.57) times preventral margin length. Dermal denticles of holotype mostly minute, narrowly arrow-shaped with prominent ridges on an elevated crown, not close-set, covering entire body and fins; most of surface uniformly finely granular, but denticles becoming enlarged along dorsal midline and infraorbit, enlarged slightly (barely evident without magnification) on midline of tail between dorsal fins, and on precaudal midline, snout tip, lateral margin of disc anteriorly, and lateral fold; enlarged denticles often irregular in shape, globular with flattened crowns and/or crenulate anterior margins; a dense covering of small denticles over nasal lamellae. Prebranchial sensory pore patch concealed, extending to just behind level of first gill slit. Postscapular sensory canals embedded, weakly undulated anteriorly, terminating just forward of pectoral-fin insertions; sensory pores minute; sensory canal not forming a shallow groove. Rostral cartilage in paratype SUML MMLM 0 12 (sex, size) broad, its shaft barely increasing in width in a posterior direction; rostral node very broadly expanded and elongate, rounded apically, not angular, axis at widest part of node 27.1 % of length of rostral cartilage from snout tip; precerebral cavity broad and uniformly convex posteriorly, converging to a point anteriorly at rostral node, dorsolateral edges of cartilage surrounding cavity (rostral ridges on surface of snout) slightly separated posteriorly, not constricted medially; rostral cartilage about 65 % of length of neurocranium, ventral edges of rostral cartilage united; nasal capsules large, their transverse axes anterolaterally directed; maximum width across capsules 1.25 times nasobasal length of cranium (base of rostrum to occipital condyles); length of nasal capsule about equal to its width; basal plate narrow, its minimum width 4.78 times in nasobasal length; cranial roof with small oval fenestra, located well behind precerebral cavity (separated by 1.4 times its length); anterior cartilage broadly triangular, posterior apex narrowly rounded, without an anterior lobe extending beyond nasal capsules; preorbital processes indistinct; postorbital processes well developed, strongly bifurcate, posterior process most pronounced; greatest width across processes 1.49 times in nasobasal length. Nasal lamellae 51 (50���53). Pectoral skeleton with 29 (30���32, n = 5) propterygial, 7���8 (6���8) mesopterygial, 1���2 (1���2) neopterygial, 25 (26���29) metapterygial and 63 (66���70) total radials; anterior radials of propterygium (in paratype SUML F 1) extending forward of nasal capsules by about 12.7 % of rostral length. Total pelvic radials 25 (24���26 in males, n = 3) excluding clasper; 25���26 in females (n = 2). Vertebral column with 192 (185���191, n = 5 paratypes) total segments (synarcual and free), 178 (173���179) post-synarcual centra; 146 (138���147) total precaudal centra (including synarcual); 14 (11���14) synarcual segments; 25 (23���27) monospondylous precaudal centra, all posterior centra of holotype with ribs; 107 (101���107) diplospondylous precaudal centra; 46 (43���48) diplospondylous caudal centra. Colour. Fresh (based on image of holotype, PNM 15189). Dorsal surface medium greyish brown, covered with a complex (but subtle) mottled pattern of poorly defined white spots and large diffuse dusky and orange blotches; white spots scattered, small, similar in size to spiracular folds, most obvious on central disc and anterior tail; snout apex slightly darker than rest of snout; dusky blotches similar in size to eye diameter, most pronounced near outer part of disc; narrow white margins along pelvic fins and anterior disc; orange blotches most evident on central disc and near margins of tail; orbital membrane dusky. Ventral surface mostly white; pinkish to translucent beside rostral cartilage; broad dusky margins on both posterior disc and pelvic fins. First dorsal fin pale brownish and weakly blotched anteriorly, its posterior half darker, dusky to blackish; second dorsal and caudal fins dark brownish near fin origin, becoming pale yellowish medially, then semi translucent with dusky blotches posteriorly; an obvious subcircular blotch over inner margins of both dorsal fins; margin of ventral lobe of caudal fin white. In preservative. Dark greyish brown dorsally, orange blotches more obvious (each about eye size or slightly smaller, becoming increasingly more evident as skin dries, see Fig. 5 A); blotches most strongly contrasted with pale area of snout lateral to rostral cartilage; white spots barely detectable in most types; melanophores large, densely packed and prominent over most of surface (see Fig. 8 B���D for paratype CSIRO H 7542 ���04); dorsal fins pale anteriorly with darker areas posteriorly, large dark spot evident above inner margins of holotype and most paratypes; caudal fin blotched, dusky, darkest near fin apex. Ventral surface creamy white; posterior disc and pelvic fins with white-edged dark margin still evident in holotype, less so in some paratypes. Size. Available material consists of 12 males and 10 females; adult males ranged from 556���720 mm TL, and the holotype was adolescent at 593 mm TL; adult females probably larger than males, several exceeded 720 mm TL with the largest being 844 mm TL; birth size and reproductive information unknown. Distribution. Collected from fish markets in the central and southern Philippines at Danao and Cebu Cities (Cebu), Baliangao and Dipolog City (Mindanao), and Bacolod, Cadiz and Dumaguete Cities (Negros). Probably more widely distributed in seas of the Philippine Archipelago but exact information available from fishermen. Etymology. Epithet in recognition of the considerable efforts of colleague Dr William White who has contributed so much to the taxonomic and biological knowledge of sharks and rays of the Western Central Pacific over the past decade. The vernacular name, Philippine guitarfish, follows Compagno et al. (2005) with reference to its known distribution in the Philippines, and where it was first collected as part of a WWF���funded project to investigate the biodiversity of chondrichthyans of the region. Molecular analysis. The analysis of the NADH 2 data suggests that Rhinobatos whitei represents a monophyletic lineage that is distinct from, but closely related to, R. jimbarensis and that these two species, in turn, are sister to a clade containing R. punctifer from the Arabian Sea and R. penggali from Indonesia. We caution however that this inference is based on a single mitochondrial marker. Inclusion of multiple nuclear markers could affect the presented inference. A closely related form from nearby Borneo differs slightly from R. whitei and may be another undescribed species. Comparisons. Several guitarfishes in the Western Central Pacific have been confused with Rhinobatos schlegelii M��ller & Henle. Herre (1953) did not report a ray referable to R. whitei in his checklist of Philippine fishes. The Philippine species, subsequently referred to by Compagno et al. (2005) as Rhinobatos cf. schlegelii, is distinct from R. schlegelii (occurring off Japan and Taiwan) in morphology (i.e. body form, squamation, and colour pattern) and its NADH 2 gene sequence. The types of R. schlegelii were examined recently by Bernard Seret and the first author as part of a broader investigation of Rhinobatos of the western North Pacific. Rhinobatos schlegelii differs from R. whitei in some meristic and morphological details. Rhinobatos whitei has a much lower free precaudal centra count (126���133 vs. 142���148 centra in R. schlegelii) but typically more caudal centra (43���48 vs. 31���43 centra), fewer pelvic rays (24���26 vs. 27���30 elements in males, excluding the clasper), a broader pectoral disc (width 30.9���34.6 % vs. 27.9���30.8 % TL) and mouth (width 5.7���6.5 % vs. 5.5���5.6 % TL), and larger spiracles (width 2.4���2.8 % vs. 1.9���2.2 % TL). Rhinobatos schlegelii has a more bottlenose shaped snout with a black teardrop marking on its ventral surface (most obvious in young and absent in R. whitei). Rhinobatos whitei belongs to a group of plain or weakly patterned (rather than distinctly white spotted) guitarfishes of the genus that includes two other valid nominal species in the Western Central Pacific, R. jimbaranensis and R. sainsburyi. Another nominal species considered valid by several authors (see Fricke & Eschmeyer, 2014), R. formosensis, is thought to be a synonym of R. schlegelii (Seret et al., in prep). Rhinobatos whitei differs from R. jimbaranensis from nearby Indonesia in its NADH 2 sequence (see Fig. XX). Both species are covered with rusty brown blotches but unlike R. jimbaranensis, R. whitei is also covered with faint white spots and has bicoloured dorsal fins. Rhinobatos whitei has fewer pelvic radials (females 25���26 vs. 27���29 in R. jimbaranensis), slightly fewer monospondylous precaudal centra (23���27 vs. 26���2, Published as part of Last, Peter R., Corrigan, Shannon & Naylor, Gavin, 2014, Rhinobatos whitei, a new shovelnose ray (Batoidea: Rhinobatidae) from the Philippine Archipelago, pp. 31-47 in Zootaxa 3872 (1) on pages 33-45, DOI: 10.11646/zootaxa.3872.1.3, http://zenodo.org/record/287210, {"references":["Fowler, H. W. (1941) Contributions to the biology of the Philippine archipelago and adjacent regions. The fishes of the groups Elasmobranchii, Holocephali, Isospondyli, and Ostariophysi obtained by the United States ... \" Albatross \" in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. Bulletin of the United States National Museum 100 (13), 1 - 879.","Compagno, L. J. V. & Last, P. R. (1999) Rhinobatidae. In: Carpenter, K. E. & Niem, V. H. (Eds.), FAO Species Identification Guide for Fishery Purposes. The living marine resources of the Western Central Pacific. Vol. 3. Batoid fishes, chimaeras and bony fishes, part 1 (Elopidae to Linophyrnidae) FAO, Rome, pp. 1423 - 1430.","Compagno, L. J. V., Last, P. R., Stevens J. D. & Alava, M. N. R. (2005) Checklist of Philippine Chondrichthyes. CSIRO Marine Laboratories Report 243, 103 pp.","Herre A. W. C. T. (1953) Check list of Philippine fishes. United States Fish and Wildlife Service Research Report 20, 977 pp.","Fricke, R. & Eschmeyer, W. N. (2014) Catalog of Fishes - Guide to fish collections. Electronic version. Available from: http: // research. calacademy. org / research / ichthyology / catalog / collections. asp (accessed 15 August 2013)"]}

Genus Rhinobatos Linck Rhinobatos Linck 1790, Mag. Neueste Phys. Naturg. Gotha 6, 32. Type not designated in original description, but Raja rhinobatos Linnaeus, 1758 is possibly the type by absolute tautonymy., Published as part of Last, Peter R., Corrigan, Shannon & Naylor, Gavin, 2014, Rhinobatos whitei, a new shovelnose ray (Batoidea: Rhinobatidae) from the Philippine Archipelago, pp. 31-47 in Zootaxa 3872 (1) on page 32, DOI: 10.11646/zootaxa.3872.1.3, http://zenodo.org/record/287210, {"references":["Linck, H. F. (1790) Versuch einer Eintheilung der Fische nach den Zahnen. Magazin fur das Neueste aus der Physik und Naturgeschichte, Gotha, 6 (3), 28 - 38.","Linnaeus, C. (1758) Systema naturae. Vol. 1. 10 th Edition. Laurentii Salvii, Holmiae, 824 pp. http: // dx. doi. org / 10.5962 / bhl. title. 542"]}

Order Chimaeriformes

Order Squatiniformes

Family Triakidae

Apristurus macrostomus Chu, Meng & Li, 1985 Type locality: off Zhujiang, South China Sea. The 13 Taiwanese samples of A. macrostomus formed a single cluster that includes other previously collected samples from Taiwan (Supplementary Material 1). Sample GN 1013 is labeled as A. macrorhynchus in Naylor et al. (2012 a, 2012 b). Based on results of this study, the sample may have to be transferred to A. macrostomus. This clade is closest to, but very distinct from, an A. exsanguis clade from New Zealand., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 268, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Urolophus aurantiacus M��ller & Henle, 1841 Type locality: Gotto Island, Nagasaki, Japan (the additional type locations of Westernport Victoria, and Tasmania in Australia are incorrect and refer to Urolophus cruciatus). The two samples of U. aurantiacus from Taiwan formed a single cluster, sister to U. paucimaculatus (GN 2554) from southern Australia (Supplementary Material 1). This represents the Taiwan range of this species, which was described from southern Japan., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 275, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Etmopterus brachyurus Smith & Radcliffe, 1912 Type locality: off Jolo Island, Philippines, 6 ��02'00" N, 120 �� 44 ' 40 " E. The two Taiwanese samples of E. brachyurus formed a single cluster together with a sample from Japan (GN 7423), and were sister to the E. cf. molleri clade from Taiwan (Figure 2, Supplementary Material 1)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 260, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Order Orectolobiformes

Order Heterodontiformes

Pseudotriakis microdon de Brito Capello, 1868 Type locality: Setubal, Portugal, Northeastern Atlantic. The five Taiwanese samples of P. microdon included in this analysis formed a single cluster, distinct from, but sister to, the P. microdon samples from the southern Madagascar Ridge and the Mid-Atlantic Ridge (Figure 7). These results challenge the currently accepted view that this species is circumglobal in distribution (e.g. Compagno et al., 2005) and suggest that the western North Pacific population is distinct from the Northern Atlantic/ southwestern Indian populations. A taxonomic revision of this genus is required to determine whether these two clades represent distinct species and if so, determine whether Pseudotriakis acrales Jordan & Snyder 1904 described from Suruga Bay in Japan should be resurrected as a valid species. Gollum attenuatus New Zealand Pacific Ocean GN 1470 100 0.03, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on pages 269-270, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Apristurus herklotsi (Fowler, 1934) Type locality: Cagayan Island, Jolo Sea, Philippines The single Taiwanese sample of A. herklotsi included in this analysis formed a distinct lineage which is close, and sister to, the morphologically similar A. australis (GN 4877) from Western Australia (Supplementary Material 1). Mitsukurina owstoni U.S.A. Pacific Ocean GN 1798 Isurus oxyrinchus Taiwan Pacific Ocean GN 939 100 Isurus oxyrinchus Mexico Pacific Ocean GN 1126 93 Isurus oxyrinchus Vietnam Pacific Ocean GN 7069 100 Isurus oxyrinchus U.S.A. Atlantic Ocean GN 16 98 Isurus oxyrinchus U.S.A. Atlantic Ocean GN 241 65 Isurus oxyrinchus U.S.A. Pacific Ocean GN 1036 76 IsurusIsurus oxyrinchusoxyrinchus TaiwanTaiwan KeelungKeelung FishmarketFishmarket GN 10186 100 Pseudocarcharias kamoharai U.S.A. Pacific Ocean GN 2000 100 PseudocarchariasPseudocarcharias kamoharaikamoharai GN 10263 HO 20110526 tsHO 20110526 ts 100 Pseudocarcharias kamoharai Taiwan Pacific Ocean GN 1034 90 Pseudocarcharias kamoharai U.S.A. Pacific Ocean GN 2634 85 Pseudocarcharias kamoharai Taiwan Pacific Ocean GN 1033 92 Alopias pelagicus Taiwan Pacific Ocean GN 1074 80 Alopias pelagicus Mexico Pacific Ocean GN 5402 98 Alopias pelagicus Mexico Atlantic Ocean GN 3130 100 Alopias pelagicus Taiwan Pacific Ocean GN 924 88 Alopias pelagicus Taiwan Pacific Ocean GN 927 97 Alopias pelagicus India Indian Ocean GN 1642 65 AlopiasAlopias pelagicuspelagicus GN 10065 10065 HOHO 178178 0.05, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on pages 267-268, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Centrophorus granulosus (Bloch & Schneider, 1801) Type locality: unknown. The 13 Taiwanese samples of C. granulosus included in this analysis formed a single cluster that are nearly identical to a sample from Japan (GN 7425) and one from Portugal (GN 6615) (Supplementary Material 1, Figure 1). These samples were previously identified as either C. niaukang or C. acus but a revision of this species revealed these names are both junior synonyms of C. granulosus (White et al., 2013). The cluster of samples for this species was sister to the C. squamosus cluster (see below)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 259, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Family Parascylliidae

Hemitriakis japanica (M��ller & Henle, 1839) Type locality: Japan. The five Taiwanese samples of H. japanica included in this analysis formed a single cluster that included previously collected samples of this species from Taiwan, Japan and Vietnam (Supplementary Material 1; Naylor et al., 2012 a). This cluster is closely related and sister to the H. leucoperiptera clade consisting of samples from the Philippines (Supplementary Material 1)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 270, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Family Alopiidae

Family Etmopteridae, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 260, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Zameus ichiharai (Yano & Tanaka, 1984) Type locality: Suruga Bay off Okitsu, 35 ��02.9'N, 138 �� 34.6 'E, Japan The four samples formed a single cluster closely related to Proscymnodon plunketi (GN 5607) (Supplementary Material 1). Analysis of additional sequences of P. plunketi from New Zealand (the type locality) and the southern Madagascar Ridge further support that Z. ichiharai samples from Taiwan are more closely related to Proscymnodon than Zameus squamulosus (Figure 3). Taxonomic investigation of this species is currently underway to clarify the generic relationships of the genera Zameus and Proscymnodon. This species had not been previously recorded from Taiwanese waters. Somniosus pacificus U.S.A. Pacific Ocean GN 1528 Zameus squamulosus Australia Pacific Ocean GN 4986 80 Centroselachus crepidater Portugal Atlantic Ocean GN 6617 100 Centroscymnus coelolepis Portugal Atlantic Ocean GN 6626 Centroscymnus owstonii Portugal Atlantic Ocean GN 6600 74 Scymnodon ringens Ireland Atlantic Ocean GN 1703 ZameusZameus ichiharaiichiharai GN GN 10228 HO HO 20111002 ck005 98.67 100 ZameusZameus ichiharaiichiharai GN 10231 HO 20111002 ck012 100 ZameusZameus ichiharaiichiharaii GN 10224 HO 20110730 ck002 98 96 ZameusZameus ichiharaiichiharaii GN 10227 HO 20111002 ck004 Proscymnodon plunketi New Zealand Pacific Ocean GN 5607 100 Proscymnodon plunketi New Zealand Pacific Ocean GN 5621 60.67 Proscymnodon plunketi Madagascar Ridge Indian Ocean GN 11707 60 Proscymnodon plunketi Madagascar Ridge Indian Ocean GN 11781 Proscymnodon plunketi Madagascar Ridge Indian Ocean GN 11669 65.33 Proscymnodon plunketi Madagascar Ridge Indian Ocean GN 11783 0.04, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on pages 262-263, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Family Myliobatidae, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 275, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Rhinobatos hynnicephalus Richardson, 1846 Type locality: probably China Seas, Canton, China (no types known). The three Taiwanese samples included in this study formed a single cluster, sister to R. schlegelii from Taiwan (Supplementary Material 1). The samples were preliminary identified as R. schlegelii but later assigned to R. hynnicephalus (B. S��ret, pers. comm.)., Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 272, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Family Urolophidae, Published as part of Straube, Nicolas, White, William T., Ho, Hsuan-Ching, Rochel, Elisabeth, Corrigan, Shannon, Li, Chenhong & Naylor, Gavin J. P., 2013, A DNA sequence-based identification checklist for Taiwanese chondrichthyans, pp. 256-278 in Zootaxa 3752 (1) on page 275, DOI: 10.11646/zootaxa.3752.1.16, http://zenodo.org/record/285357

Rhynchobatus immaculatus Last, Ho & Chen, 2013 [this issue] Type locality: Keelung, Taiwan The two samples of Rhynchobatus immaculatus n. sp. included in our analysis represent a distinct lineage, sister to a clade comprising several Rhynchobatus species from Malaysia and Australia, i.e. R. australiae (GN 2996), R. cf. laevis (GN 2065), R. laevis (GN 3004) and R. palpebratus (GN 2044) (Supplementary Material 1). This species was determined as a new and previously undescribed species during this study based on both, the molecular results and morphological information.