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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.

Aetobatus flagellum (Bloch & Schneider, 1801) (Figures 1���6, Table 1) Raja flagellum Bloch & Schneider, 1801: 361, pl. 73 (Coromandel Coast, India). Aetobatis narinari��� Day, 1878 (in part): 743, pl. 194, fig. 4 (misidentification, India). A��tobatis narinari��� Day, 1889: 59���60, fig. 24; Blegvad, 1944: 55���56, fig. 23 (brief description, illustration after Day, 1878) (misidentification, Persian Gulf). A��tobatis flagellum��� Annandale, 1909: 54���58, fig. 10 a, pl. (fig. 5) (off Orissa Coast and Chilka Lake) Aetobatis flagellum��� Fowler, 1930: 507 (Hawaii; Indian Ocean) Aetobatus narinari (in part) ��� Fowler, 1941: 471; Misra, 1947: 40; Dor, 1984: 20. Syntypes. ZMB 7845, male, coast of Coramandel, India, collected by M.E. Bloch; ZMB 31560, male, Tharangambadi (formerly Tranquebar), coast of Coramandel, India, collected by M.E. Bloch. Other material examined. (20 specimens): BMNH 89.2.1.4205-8, 5 specimens (female 243 mm DW [653 mm TL], immature male 233 mm DW [548 mm TL], female 369 mm DW [1082 mm TL], female 289 mm DW [796 mm TL], female 290 mm DW), Madras (possibly), India; CSIRO H 4426 ��� 14, subadult male 446 mm DW, Muara Angke fish landing site, Jakarta, Indonesia, 17 Oct. 1995, collected by P. Last; CSIRO H 5485 ���02, immature male 350 mm DW (1017 mm TL), Kuching fish market, Sarawak, Malaysia, 0 2 May 1999, collected by P. Last & M. Manjaji; CSIRO H 6134 ���01, subadult male 431 mm DW (1260 mm TL), Muara Angke fish landing site, Jakarta, Indonesia, 20 May 2002, collected by W. White & Dharmadi; CSIRO H 6662 ���03, immature male 346 mm DW (1156 mm TL), CSIRO H 6662 ���04, immature male 306 mm DW (956 mm TL), CSIRO H 6662 ���05, female 326 mm DW, CSIRO H 6662 ���06, immature male 305 mm DW (1027 mm TL), Muara Baru fish landing site, Jakarta, Indonesia, 19 Apr. 2004, collected by W. White & Dharmadi; CSIRO H 7252 ���01, female 388 mm DW (1029 mm TL), Persian (Arabian) Gulf, Sharq fish market, Kuwait City, Kuwait, 29 �� 23 ��� N, 47 �� 58 ��� E, probably caught off Kuwait in Specimens examined but not retained. female 746 mm DW, Persian (Arabian) Gulf, Sharq fish market, Kuwait City, Kuwait, 29 �� 23 ��� N, 47 �� 58 ��� E, probably caught off Kuwait in Specimens not examined but with images verified. USNM 206131, Caraioor fish market, near Jaffna Fort, Sri Lanka, 17 Mar. 1970; USNM 222684, fish market at Kalupittiya, Sri Lanka, 25 Jan. 1970, collected by C.C. Koenig; USNM 222690, St John���s fish market, Colombo, Sri Lanka, 23 May 1970, collected by T. Iwamoto. Diagnosis. A small Aetobatus (attaining about 900 mm DW) with the following combination of characters: dorsal surfaces uniformly brownish, without pale spots; tail very long (1.22���2.81 times DW); stinging spine(s) relatively long (6.2���16.2 % DW); head long; rostral lobe long to very long (longest in adult males) with a narrowly pointed apex; teeth plates in a single row, those in lower jaw chevron-shaped; width of lower tooth plate about two thirds mouth width; pectoral-fin radials 89���96 (excluding propterygial radials anterior of eyes); total vertebral centra (including synarcual) 85���91; males mature by about 500 mm DW and females by about 746 mm DW. n= 20 Min. Max. Mean Disc width (mm) 233 578 353.25 Total length 178.1 336.7 274.3 Pre-dorsal length 55.7 68.5 61.5 Disc, length 55.3 70.0 62.1 Snout to pectoral-fin insertion 49.4 63.0 54.9 Disc thickness 9.5 12.8 11.2 Snout to pectoral-fin origin 13.8 20.4 17.7 Posterior orbit to pectoral-fin insertion 39.5 45.8 42.2 Snout to maximum width (horiz.) 38.2 45.7 42.8 Pectoral-fin anterior margin 48.8 52.7 50.5 Pectoral-fin posterior margin 45.1 50.4 47.6 Pectoral-fin base length 38.2 43.8 40.7 Pectoral-fin inner margin 5.8 8.1 7.1 ......continued on the next page n= 20 Description. Disc diamond-shaped, broad but relatively short, width about 1.33���1.81 times disc length; anterior projection 3.15���4.22 in disc length; axis of greatest width of disc well posterior to scapular region, over abdominal cavity, its horizontal distance from snout tip 1.18���1.40 times in distance from tip of snout to pectoral-fin insertion; moderately deep, greatest thickness above scapular region and posterior head, thickness 7.83���10.57 in disc width; without denticles, or thorns; with a short, bony ridge on midline above scapular region. Pectoral fins very large, winglike, narrowly triangular, weakly falcate; anterior margin concave basally, nearly straight for first two thirds, slightly to moderately convex distally; apex narrowly rounded to subangular, pectoral angle 54���61 ��; posterior margin moderately concave near apex, almost straight posteriorly; free rear tip broadly rounded; inner margin convex distally, becoming nearly straight basally; length of anterior margin 48.8���52.7 % DW, 1.17���1.31 times its base length, inner margin 4.87���6.91 in its base; origin over anterior edge of spiracles; apex located just posteriorly to pectoral mid-base; insertion just posterior to pelvic-fin origin; free rear tip partly overlapping pelvic-fin anterior margin. Head pronounced, deep, short and relatively narrow; projecting well anterior to pectoral-fin origins; subquadrangular in cross-section at pectoral-fin origin; cranial region of head broadly rounded in dorsoventral view; chondrocranium pronounced above eyes and spiracles; snout abruptly convex anterior of eyes, becoming deeply concave at origin of rostral lobe; slightly convex ventrally; ventral head length 26.0��� 33.5 % DW, 1.46���2.23 times width at pectoral-fin origins, 2.96���6.36 times preorbital length (horizontal), 2.39���3.39 times interorbital width; preoral snout length 0.99���2.30 times mouth width, 1.73���2.99 times internarial width, 0.50���0.99 times distance between first gill slits; head width at pectoral-fin origin 14.8���17.8 % DW, 1.58���1.93 times its height. Rostral lobe fleshy, long (very long in adult males); narrowly parabolic in dorsoventral view with a narrowly pointed apex; narrowly pointed in lateral view; its length 3.6���9.4 % DW, 3.57���7.28 in head length, its width 1.38��� 1.98 in head width at pectoral-fin origin. Interorbital space moderately broad, convex but with a broad medial depression, without ridges, denticles or thorns; interorbital width 9.2 ���11.0% DW, 1.72���2.37 times orbit length, 0.63���0.81 times head width at mid-eye. Eyes small, subcircular, very slightly ventrolateral on head; orbit level or only slightly elevated above dorsal head profile, diameter 2.12���3.08 in spiracle length, 5.84���8.78 in head width at pectoral-fin origin. Spiracles large, suboval to elliptical, situated dorsolaterally posterior to orbit and above pectoral-fin origin, more visible dorsally than laterally; margins without any protuberances or folds; length 4.8���6.8 % DW, 2.08���3.75 times width. Nostril narrowly suboval, immediately preceded by a broad, shallow, fleshy depression bordering anterolateral margin of the nasal curtain; anterior nasal fold thin, membranous, internal; deep oronasal groove present; internarial space 1.26���2.25 in prenasal length, 1.18���2.28 times nostril length. Nasal curtain large, elongate, lobate, width 1.32���1.94 times length; lateral margin concave, smooth edged; posterior margin divided by deep medial notch, bordered by a long, curtain-like fringe, not following contour of lower jaw; posterior margin of each lobe convex with apices narrowly rounded; most of surface finely papillate, covered with minute pores centrally; apex and posterolateral margin recessible within oronasal groove. Mouth moderate-sized, transverse, located ventrally, width 6.5���8.9 % DW, 0.44���1.01 times preoral length, 1.92���2.63 in head width at pectoral-fin origin; not protrusible, anterior teeth of lower jaw visible when mouth closed; buccal region intricately papillate; skin on chin and at margin of lower jaw fleshy, strongly furrowed, papillate, indented slightly at symphysis. Teeth in a single row in each jaw, coalesced to form plates; about 6 narrow, almost straight teeth in upper jaw, tooth plate well inside palate, its length about half its width (based on CSIRO H 4426 ��� 14); about 13 narrow, chevron-shaped teeth in lower jaw, tooth plate protruding distally, its length more than twice its width, its width about two thirds mouth width (based on CSIRO H 4426 ��� 14); roof of mouth with 2 rows of oral papillae, those in outer row slightly larger than those of inner row; floor of mouth near lingual margin of lower tooth plate with lunate fringe of about 16 variably shaped (usually pointed), irregular oral papillae. Gill openings small, elongated S-shaped, forming a weakly fringed lobe laterally; length of first gill slit 0.95���1.70 times length of fifth gill slit, 2.36���4.61 in mouth width; distance between first gill slits 2.86���3.95 times internarial space, 0.45���0.64 times ventral head length; distance between fifth gill slits 1.78���2.45 times internarial distance, 0.45���0.64 times ventral head length. Pelvic fins moderately large, slender, subquadrangular, anterior margin slightly concave to almost straight, apex moderately angular, posterior margin moderately convex, free rear tip broadly rounded, inner margin slightly convex; extending well beyond pectoral-fin free tips; pelvic-fin length 14.3���18.7 % DW, 1.12���1.74 times width across fin bases, inner margin 9.0��� 13.3 % DW. Claspers of adult male (MNHN A 7949) relatively short, outer length 6.4 % DW. Dorsal fin small, strongly raked, its origin posterior to pelvic-fin insertions by about half to two-thirds its fin base; anterior margin almost straight; apex broadly rounded, posterior to insertion of fin; posterior margin convex to nearly straight; free rear tip subangular, inner margin very short, nearly straight; predorsal length 1.46���1.80 in disc width, fin length 4.7���7.8 % DW, height 0.35���0.67 times its length, inner margin 2.77���10.4 in fin length. Tail very long, slender, whip-like, its length (from cloaca origin) 1.22���2.81 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 0.73���1.20 times height; almost rhomboidal in cross section near origin of stinging spine, width 0.59���1.36 times height at first spine origin; no skin folds present; a weak naked groove on dorsal surface of tail immediately posterior to base of stingingspine(s), almost fully housing spines. Stinging spines 0���2, very elongate, slender, moderately broad-based, strongly tapered, almost fully serrated laterally; distance from sting base to pectoral-fin insertion 10.4���14.8 % DW; longest stinging spine 9.4���16.2 % DW, 1.86���3.44 times dorsal-fin length. Vertebral centra total (including synarcual) 85���91 (n= 7); total (excluding synarcual) 80���87 (n= 7); monospondylous (including synarcual) 33���42 (n= 8); monospondylous (excluding synarcual) 29���38 (n= 8); predorsal diplospondylous 13���29 (n= 4); post-dorsal diplospondylous 27���34 (n= 3). Total pectoral-fin radials (excluding propterygial radials anterior of eyes) 89���96 (n= 7); propterygium (anterior of eyes) 13 *��� 16 *, propterygium (posterior of eyes) 10���14, mesopterygium 27���33, metapterygium 48���54. Pelvic-fin radials: 1 (4���6 fused elements) + 14���16 (n= 7). Colour (when fresh). Dorsal surface uniformly brownish (sometimes greenish brown), without distinct markings; eye bluish black; dark (dorsal) and pale (ventral) surfaces well demarcated (waterline) at pectoral-fin origin at junction with head; waterline extending anteriorly to mid eye and onto forehead; dark dorsal surface on rostral lobe similar, contrasted with its paler ventral surface and posteriorly with pale mid-snout; tail uniform greyish brown. Ventral surface mostly whitish; broad brownish margin along most of disc, junction between brown margin and whitish ventral colour strongly mottled, broadest on posterior margin, narrowest anteriorly; distal third of pelvic fins brownish; rostral lobe mostly whitish, anteriormost margin narrowly brownish. Size. The male and female specimens of A. flagellum measured in this study ranged in size from 233���543 and 243���578 mm DW, respectively. Two male specimens of 431 and 446 mm DW were adolescent, and one specimen of 543 mm DW was mature. Moore et al. (2012) reported 36 individuals from the Persian Gulf with males and females ranging from 277���580 and 330���746 mm DW, respectively; males mature by ~ 500 mm DW. Birth size unknown; smallest free-swimming individual examined was 233 mm DW. Specimens of up to 900 mm DW have been recorded from northern Kuwait (J. Bishop, Kuwait Institute for Scientific Research, unpubl. data). Sujatha (2002) recorded two specimens off Visakhapatnam in northeastern India, which were 790 and 830 mm DW, but no sex was given. A single female of 746 mm DW (not retained), was mature and had functional, but empty, uteri. Distribution. Patchily distributed in the Indo ���West Pacific; known from the Western Indian Ocean, from Kuwait in the Persian Gulf to Pakistan and India; and the Eastern Indian Ocean, from India and Sri Lanka to Indonesia (Kalimantan) and Malaysia (Sarawak). Not recorded from the east coast of South Africa (S. Dudley, KwaZulu-Natal Sharks Board, pers. comm. July 2009), Madagascar (A.J. Cooke, Blue Ventures, pers. comm.; Robinson & Sauer, 2013), Oman (Randall, 1995; Henderson & Reeve, 2011) nor the southern Persian Gulf (Moore et al., 2012; A. Moore unpubl. data). Reported presence in the Red Sea (Bonfil & Abdallah, 2004) was not based on records (R. Bonfil, pers. comm.) and requires confirmation, as it has not been reported previously from this region (Gohar & Mazhar, 1964). The Red Sea records are possibly due to its inclusion (mistakenly) as a synonym of A. narinari in species lists for the Red Sea (e.g. Fowler, 1956; Dor, 1984). Compagno & Last (1999) mentioned that records of A. flagellum from Hawaii and the Eastern Atlantic require confirmation. During this study, no specimens or accurate records of this species from these two regions were found and experts on the chondrichthyans faunas of Hawaii (J. Randall, BPBM, pers. comm. July 2009) and West Africa (B. Ser��t, pers. comm. March 2009) had no records. Records of this species from southern China require validation., Published as part of White, William T. & Moore, Alec B. M., 2013, Redescription of Aetobatus flagellum (Bloch & Schneider, 1801), an endangered eagle ray (Myliobatoidea: Myliobatidae) from the Indo ��� West Pacific, pp. 199-213 in Zootaxa 3752 (1) on pages 201-207, DOI: 10.11646/zootaxa.3752.1.12, http://zenodo.org/record/218575

Lamnidae (Mackerel sharks) No confirmed Gulf records. A Kuwait record (Khalaf 1987) of the great white shark Carcharodon carcharias (Linnaeus, 1758) was a misidentification of Carcharias taurus (Moore et al. 2007). Tourenq et al. (2008) note an unsubstantiated Carcharodon carcharias record from the United Arab Emirates and Musandam area, and a skin specimen registered as C. carcharias from nearby Oman (BMNH 1891.2.9.41) has apparently been lost (Patrick Campbell, BMNH, pers. comm.). Randall (1986) noted the shortfin mako Isurus oxyrinchus Rafinesque, 1810 as not yet reported from the Gulf and our data support this.

Sphyrnidae (Hammerhead sharks) The winghead shark Eusphyra blochii (Cuvier, 1816) has been reported from the Gulf but without evidence (Randall 1986; Carpenter et al. 1997 a), although its occurrence in the Gulf of Oman near Jask is proven (Blegvad 1944, as Zygaena blochii). Reports of the smooth hammerhead shark Sphyrna zygaena (Linnaeus, 1758) in the Gulf were doubted by Randall (1996), and our data support this, although a number of specimens of this species have recently been confirmed from Oman (Henderson & Reeve 2011)., Published as part of Moore, Alec B. M., Ward, Robert D. & Peirce, Richard, 2012, Sharks of the Persian (Arabian) Gulf: a first annotated checklist (Chondrichthyes: Elasmobranchii), pp. 1-16 in Zootaxa 3167 on page 13, DOI: 10.5281/zenodo.279779, {"references":["Cuvier, G. (1816) Le Regne Animal distribue d'apres son organisation pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparee. Les reptiles, les poissons, les mollusques et les annelides. Deterville, Paris.","Randall, J. E. (1986) Sharks of Arabia. Immel, London.","Carpenter, K. E., Krupp, F., Jones, D. A. & Zajonz, U. (1997 a) The living marine resources of Kuwait, Eastern Saudi Arabia, Bahrain, Qatar, and the United Arab Emirates. FAO, Rome.","Blegvad, H. (1944) Danish Scientific Investigations in Iran. Part III. Fishes of the Iranian Gulf. Einar Munksgaard, Copenhagen.","Linnaeus, C. (1758) Systema Naturae, Ed. X. (Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Holmiae (Stockholm).","Henderson, A. C. & Reeve, A. J. (2011) Noteworthy elasmobranch records from Oman. African Journal of Marine Science, 33, 171 - 175."]}

Himantura randalli sp. nov. Figures 1���6, Table 1 Trygon gerrardi (not Gray): Blegvad, 1944 (in part), 50���51, Plate II figs 2, 2a (brief description, illustrations of 880 mm TL individual and denticle band) (misidentification). Himantura gerrardi (not Gray): Randall, 1995, 45, fig. 62 (brief description, figure) (misidentification). Himantura sp. B: Manjaji 2004, 5 (136���144), fig. 5.2. 27 (description, key 5 (10). Dasyatis bennettii (as bennetti) (not M��ller & Henle): Assadi & Dehghani, 1997, 212 (diagnosis, illustration of 270 mm DW female) (misidentification). Holotype. CSIRO H 7254 ���01, adult male 412 mm DW, Persian Gulf, Sharq fish market, Kuwait City, Kuwait, 29 �� 23 N, 47 �� 58 ��� E, probably caught off Kuwait in less than 40 m, collected 0 5 Apr 2011. Paratypes. 5 specimens. BPBM 33201 (2 specimens), female 151 mm DW, female 245 mm DW, Persian Gulf, Kuwait Bay, Kuwait, R/V Bahith, trawl 8���10 m, collected 21 Aug 1985; BPBM 29480, early adolescent male 325 mm DW, Persian Gulf, Bahrain fish market, collected 0 7 Nov 1983; MTUF 20642, female 414 mm DW, Persian Gulf (as Arabian Gulf); CSIRO H 7296 ���01, female 251 mm DW, Persian Gulf, Sharq fish market, Kuwait City, Kuwait, 29 �� 23 ��� N, 47 �� 58 ��� E, probably caught off Kuwait in less than 40 m, collected 13 Apr 2011. Other material. CSIRO H 6889 ���01, BW���A 6090 (dried, salted tail section, including sting), mature male ca 500 mm DW, Persian Gulf, Fahaheel fish market, Kuwait, 13 Apr 2008; BW���A 6091 (tissued, not retained), male ca 490 mm DW, Kuwait, 15 Apr 2008; BW��� 2113 (tissued, not retained), R/V Flinders, Kuwait, 2 Sep 2002; BW���A 6089 (tissued, not retained), female ca 550 mm DW, Kuwait, 16 Apr 2008; BW���A 6303 (tissued, not retained), male ca 500 mm DW, Qatar, 12 Apr 2009; BW���A 6308 (tissued, not retained), male ca 240 mm DW, Qatar, 13 Apr 2009. Diagnosis. A medium-sized species of Himantura (to at least 62 cm DW) distinguished by the following combination of features: disc weakly rhomboidal; preorbital snout moderately elongate with weak apical lobe, snout angle 113���119 ��; pectoral-fin apices rounded, angle 96���98 ��; orbits small, protrusible; mouth relatively broad, width 0.9���1.2 in internasal width; distance between first gill slits 2.5���2.6 times internasal distance; distance between fifth gill slits times 1.5���1.7 times internasal distance, 27���29 % of ventral head length; pelvic-fin base broad, 13���17 % DW; tail behind sting of juveniles subcircular with deep longitudinal ventral groove and prominent mid-lateral ridge, weakly depressed in adults; 1���2 (usually 1) small, broadly heart-shaped to seed shaped suprascapular denticles, primary denticle band and thorns absent; secondary denticle band irregularly suboval, relatively narrow (its maximum width across scapulocoracoid barely exceeding its width at spiracles), with well-defined lateral margins, narrowly tapering near tail base; band fully developed and covering entire dorsal surface of tail by 33 cm DW; dorsal surface mainly uniformly coloured (occasionally with dark flecks in specimens smaller than 25 cm DW), disc margin sometimes paler dorsally; ventral disc uniformly whitish, not black edged; in adults, darker dorsal surface of tail sharply demarcated from paler ventral surface; in neonates and juveniles, tail dark with conspicuous white saddles, its distal portion usually almost uniformly dark; pectoral-fin radials 124���129; total vertebral count (excluding 1 st synarcual centra) 108���111, monospondylous centra 43���44, pre-sting diplospondylous centra 65���68. Description. Disc rhomboidal, width 1.02 times length in adult male holotype (1.02���1.08 in the four largest paratypes,> 245 mm disc width, DW); robust, raised at mid-scapular region (more pronounced in largest specimens), maximum thickness 14 % (12���17 %) of DW; preorbital snout moderately long, with small medial lobe at the snout tip, angle 97 �� (96���98 ��) (no obvious ontogenetic or sexual variability); anterior margins of disc almost straight (sometimes weakly concave), apices rounded rather than angular, posterior margins broadly and evenly convex, free rear tip narrowly rounded. Pelvic fins subtriangular, short, length 19.4 % (18.5���21.2 %) DW; lateral margin almost straight, apex narrowly rounded; free rear tip broadly rounded, combined with inner and posterior margins to form strongly convex edge; width across base 15.4 % (13.3���16.8 %) DW. Claspers of adult male moderately elongate, stout basally, tapering strongly, depressed slightly; lining of pseudopera smooth; hypopyle short, about 1 / 3 rd of length of clasper outer margin; without prominent anterior notch; distance from cloaca origin to sting 0.38 (0.37���0.43) of disc length. Tail slender, whip-like; tapering gradually and evenly toward sting, then with very weak taper beyond sting to tail tip; its length 1.8 (2.2���2.4 in 3 smallest paratypes, all undamaged) times DW; base relatively narrow, weakly depressed, typically suboval in cross-section, its width 1.47 (1.07���1.45) times its height at base (usually slightly more depressed in juveniles); below sting base in holotype and adult female, subcircular, in juveniles (less than 250 mm DW) somewhat quadrangular; deep dorsal groove housing stinging spine persistent well beyond stinging spine tip, best developed in juveniles; in adults posterior tail slightly depressed, its dorsal surface strongly convex, ventral surface almost flat, lateral edge subangular; juveniles similar, but with a deep longitudinal groove on mid-ventral surface (variable in length, originating beneath or slightly posterior to stinging spine), flanked by low ventral ridges; juveniles also with prominent ridge along mid-lateral edge of tail, originating near stinging spine, extending almost to tip of tail. Snout moderately elongate, depressed; preoral snout length 2.72 (2.57���3.22) times mouth width, 2.79 (2.69���2.78) times internarial distance, 22.6 % (21.9 % in all 4 paratypes) DW; direct preorbital snout length 1.75 (1.47���1.63) times interorbital length; snout to maximum disc width 40 % (39���43 %) DW; interorbital space almost flat; eye small, diameter 1.98 (1.78���2.64) in spiracle length; orbits capable of being protruded, diameter 1.17 (1.11���1.47) in spiracle length, interorbital distance 2.46 (2.31���3.22) times orbit, intereye distance 3.36 (3.21���4.43) times orbit. Spiracles large, subrectangular to oval, situated dorso-laterally. Nostrils narrow, slightly oblique, outer margin with a weak double concavity; internasal distance 2.14 (2.02���2.12) in prenasal length, 2.19 (1.82���2.42) times nostril length. Nasal curtain relatively narrow, width 1.61 (1.65���1.78) times length; lateral margin almost straight, directed posterolaterally, smooth edged; apices narrowly rounded, lying within broad groove; posterior margin smooth or very finely fringed, weakly concave to weakly double concave (sometimes expanded slightly medially). Holotype Paratypes Neonate Range Mouth large, width 0.98 (0.92���1.20) in internasal width; profile arched strongly but variably, not obviously more so in adult male holotype than in large females; upper jaw strongly double concave, dorsal to lower jaw; lower jaw concave near symphysis, slotting into an expanded symphysial knob of upper jaw; oronasal groove prominent, deep; skin along margin of lower jaw moderately well-corrugated, confined to narrow strip around lips. Mouth floor with 2 well-developed papillae, situated close together near centre of mouth; papillae simple, slender, elongate. Teeth small, subequal in size in upper and lower jaws; cone-shaped with blunt apices. Tooth rows (in paratype BPBM 29480) 34 in upper jaw, 40 in lower jaw; holotype not dissected. Gill opening margins moderately S-shaped, smooth-edged; length of first gill slit 1.41 (1.20���1.28) times length of fifth, 2.99 (2.21���2.65) in mouth width; distance between first gill slits 2.46 (2.49���2.62) times internasal distance, 0.43 (0.45���0.47) of ventral head length; distance between fifth gill slits 1.58 (1.59���1.67) times internasal distance, 0.27 (0.29 in all types) in ventral head length. Squamation. Ontogenetic stages of squamation (definitions following Manjaji, 2004) 0, 2, and 4, evident from type series; no evidence of stages 1, 3, 5 and 6 in this species. Main denticle band well developed in adults; apart from suprascapular denticles, no enlarged thorns-like denticles on disc or tail; denticle development rapid with all stages occurring across a narrow size range; tail with scattered post-sting denticles even in young (> 250 mm DW), with main denticle band extending onto tail at about 320 mm DW; posterolateral series of denticle band on disc converging posteriorly rather than broad and truncate posteriorly. Development of secondary denticle band coincident with raising of central disc. Stage 0: early juvenile (ca. 150 mm DW) ������ Disc entirely smooth, apart from 2 small, broadly seed-shaped to heart-shape suprascapular denticles (anteriormost longest, ca 2 mm long). Primary median denticle band absent (i.e. Stage 1 absent); existence of suprascapular denticles before birth unknown. Stage 2: (ca. 250 mm DW) ������ Initial stages of development of secondary band not available from the type series, presumably rapid. A 245 mm DW female (BPBM 33201 [1 of 2]) and CSIRO H 7296 ���01 are at late stage 2 with a well-developed denticle band on disc; band commencing immediately forward of orbit and terminating just forward of pectoral-fin insertion, tapering posteriorly, indented immediately behind spiracles. Stage 4: (> 300 mm DW) ������ Secondary denticle band well-developed, narrow, with well-defined margin; extending as a continuous, longitudinal band along trunk from immediately forward of orbit, terminating immediately forward of pectoral-fin insertion or onto anterior part of tail (ca 310 mm DW) but well short of sting; margin sharply defined but slightly irregular, slightly indented over gills, expanded slightly over scapulocoracoid, converging over posterior disc toward tail. A 325 mm adolescent male paratype (BPBM 29480) has a fully formed denticle band extending over full width of dorsal tail to sting; widely spaced; denticles absent from ventral pre-sting tail; post-sting denticles granular, present on all surfaces of tail but much less dense on posteroventral surface of tail near its apex. Holotype (412 mm DW) at Stage 4, secondary denticle band irregularly suboval, with well-defined lateral margins, band extending from preorbit and onto tail, its maximum width (just posterior to spiracle) barely exceeding its width at spiracles; band not extending onto orbit nor present on suborbit; band narrow (not broadly truncate) posteriorly on disc, converging onto tail; scapular denticle single, small (length ca 4 mm); claspers, pelvic fins and ventral surface of disc naked; stinging spine elongate, ca 22 % DW, narrow-based, very slender; ventral-most stinging spine in paratype MTUF 20642 better developed than dorsal sting. In largest whole paratype (female, 414 mm DW), secondary denticle band extending from just forward of orbits to cover central disc and entire dorsal tail; ventral tail naked in region from its base to level of sting base; naked snout ratio in type series 72 % (64���69 %) in direct preorbital snout length; band margin convex anteriorly; broadest just behind spiracles, but similar to width across scapulocoracoid; no denticles lateral to spiracle or orbit; margin irregular, narrowing very gradually posteriorly from mid-disc, before continuing to tail; denticle band similar in largest specimens (480���620 mm DW) based on photographs. Denticles of cranial and scapular regions with flat crowns, varying from ovate to heart-shape; closely-spaced; minute, much smaller than suprascapular denticles; compact, closely-set, uniform in size in young; small- and large-size denticles interspersed (not imbricated) with age; largest along mid-trunk and around tail base, decreasing in size towards tail tip and toward lateral margins of main band; covering entire post-sting tail, minute and densely granular. Disc lateral to denticle band smooth, without obvious denticles. Meristics. Pectoral-fin counts (n= 5): total radials 124���125 (125���128, n= 5), propterygial radials 50���51 (49���52), mesopterygial radials 16 (17���20), metapterygial radials 58 (57���59). Pelvic-fin radials: in males 1, 22 (1, 22, n= 1), in females 1, 24��� 25 (n= 4). Vertebral centra: total (excluding 1 st synarcual) 110 (108���111, n= 5), monospondylous 43 (43���44), pre-sting diplospondylous 67 (65���68), and post-sting diplospondylous 0 (0). Coloration. When fresh (based on photograph of holotype CSIRO H 7254 ���01). Dorsal surface of disc and pelvic fins greenish grey with a broad pale pinkish outer margin (infused with blood and probably whitish naturally); marginally paler on denticle band; anterior orbit pale; tail similar to central disc before sting, darker greyish to blackish beyond sting, without evidence of paler bands; clasper pale pinkish basally, blackish distally; stinging spine greyish white. Ventral surface white (covered in pinkish areas where bleeding has occurred); tail basal half white, posterior half dusky but distinctly paler than and sharply demarcated from its dorsal surface. Fresh juvenile male specimen from Qatar (ca 240 mm DW, image only) with uniformly pale yellowish brown disc, pale greyish brown over main denticle band, broadly translucent along the outer disc margin; tail before stinging spine uniformly pale greyish brown dorsally, white ventrally, no white spots along its lateral margin; tail beyond sting much darker, upper half greyish with about 39 white saddles, ventral half uniformly dusky, not penetrated by white saddles. Tail of a foetus (ca 170 mm DW, fresh image only) fully banded with about 42 bands; mother (female ca 590 mm DW, fresh image only) and largest specimen (female ca 620 mm DW, image only) plain coloured on disc without bands or saddles on tail. Adult males (ca 440���490 mm DW, fresh image only) with a plain yellowish of brownish dorsal surface but usually with some scattered white spots on sides of tail after sting; largest males (ca 500 mm DW, fresh image only) without white markings on tail. In preservative. Holotype largely similar, more brownish dorsally and with narrow dark margin around disc ventrally; two smallest types (BPBM 33201, 151 ��� 245 mm DW) with faded dark brownish specks over entire dorsal disc surface, more so along the trunk and adjacent areas, but absent from disc margins. Skeletal morphology. The types were not dissected but based on radiographic information, the shape of the neurocranium is that of a ���typical��� Himantura (Manjaji, 2004). Size. Birth size around 150���170 mm DW, based on a presumably free swimming individual of 151 mm DW (BPBM 33201, smallest paratype) and an aborted foetus with unpigmented translucent disc, of 170 mm DW (AM, pers. obs.). Based on field data, males mature by approximately 400 mm DW. A 325 mm DW male specimen (BPBM 29480) was determined as being an adolescent (early maturity stage 3), while the 412 mm DW male holotype (CSIRO H 7254 ���01) was mature (maturity stage 4). Maximum recorded size of females (620 mm DW) larger than that of males (540 mm DW) (AM, pers. obs.); an uncatalogued female specimen in the South African Museum collection (ca 700 mm DW) referred to by Manjaji (2004) is probably not this species. Distribution. Known from the Persian Gulf (off Kuwait, Bahrain, Qatar and Iran) from where it is possibly endemic. The Persian Gulf, which is mostly shallower than 40 m and rarely exceeds 60 m depth, has soft substrates dominated by sand and mud. Considered to occur in the Gulf of Oman off Oman and off southern Africa (e.g. Randall, 1995; Manjaji, 2004) but these rays appear to be other species (see discussion below). Etymology. Named after J.E. Randall of the Bishop Museum whose work on the taxonomy of Indo ���Pacific fishes is legendary, and who was amongst the first authors to publish a photographic image of this species (as H. gerrardi) in his guide to the fishes of Oman (Randall, 1995). Vernacular name: Arabian banded whipray (based on Manjaji, 2004). Life history and fisheries. There appears to be some variation in the birth size of H. randalli. An aborted, 170 mm DW unpigmented foetus was observed in July 2002 (AM pers. obs.), but a presumed free-swimming 151 mm DW neonate (BPBM 33201, smallest paratype) was collected in August 1985, both off Kuwait. The smallest individuals recorded in recent commercial landing surveys by one of the authors (AM) in the month of April were notably larger (ca 240 DW), possibly indicating that birth takes place well before April or that small juveniles are either not caught or not landed locally. Sex ratios significantly different from parity in favour of males were recorded in commercial landings in both Qatar in April 2009 and Kuwait in April 2011 (Moore et al., in press). Significant bias towards females was also recorded for Himantura sp. (presumably H. randalli) in research trawls in the central southern Persian Gulf off Iran, in November���December 2007 (Haseli et al., 2010 and Moore, 2011). None of the specimens in the present study was examined for stomach contents, although 32 stomachs containing food items of ��� Dasyatis gerrardi��� (probably H. randalli) from Kuwait contained predominantly shrimp (particularly a small sergestid, Acetes sp.), and to a much lesser extent stomatopods (Euzen, 1987). Records of Himantura gerrardi and Dasyatis bennetti from the Persian Gulf in the last century are either, wholly or possibly in-part, misidentifications of H. randalli. Trygon gerrardi was listed as the most abundant batoid (77.6 % of 1119 individuals) in trawl surveys along the Iranian coast in January���April 1937 and 1938 (Blegvad, 1944), and Dasyatis bennetti was found to be the most abundant batoid (36 % of 366 individuals) on the Hormuz coast of Iran, easternmost part of the Persian Gulf (Vossoughi and Vosoughi, 1999). Himantura randalli sp. nov. is an important component of landings from gillnet fisheries in Kuwait and Qatar, and was the most commonly recorded batoid in surveys of landings in Qatar in April 2009 (Moore et al., in press). It was also taken in research trawls in Kuwait���s waters in July and September 2002; despite its relatively high abundance there, it is of low commercial value and often dumped (AM pers. obs.). Comparisons. Of members of the genus Himantura occurring in the western Indian Ocean, H. randalli is most morphologically similar to forms referable to H. gerrardi. However, H. gerrardi is a species complex (White et al., 2006; Last et al., 2010) and the taxonomy and distribution of these forms in the Indo ���Pacific are under review by two us (PL & MM). Using molecu, Published as part of Last, Peter R., Manjaji-Matsumoto, Mabel & Moore, Alec B. M., 2012, Himantura randalli sp. nov., a new whipray (Myliobatoidea: Dasyatidae) from the Persian Gulf, pp. 20-32 in Zootaxa 3327 on pages 21-31, DOI: 10.5281/zenodo.212521, {"references":["Blegvad, H. (1944) Danish Scientific Investigations in Iran. Part III. Fishes of the Iranian Gulf. Einar Munksgaard, Copenhagen.","Randall, J. E. (1995) Coastal Fishes of Oman. University of Hawai'i, Honolulu.","Manjaji, B. M. (2004) Taxonomy and phylogenetic systematics of the Indo-Pacific whip-tailed stingray genus Himantura Muller & Henle 1837 (Chondrichthyes: Myliobatiformes: Dasyatidae). Unpublished PhD Thesis Dissertation, University of Tasmania. Volumes 1 & 2, i - xxii; 607 pp.","Assadi, H. R. & Dehghani, P. (1997) Atlas of the Persian Gulf and the Sea of Oman Fishes. Iranian Fisheries Research and Training Organisation, Tehran.","Haseli, M., Malek, M. & Palm, H. W. (2010) Trypanorhynch cestodes from the Persian Gulf. Zootaxa, 2492, 28 - 48.","Moore, A. B. M. (2011) Elasmobranchs of the Persian (Arabian) Gulf: ecology, human aspects and research priorities for their improved management. Reviews in Fish Biology and Fisheries, DOI: 10.1007 / s 11160 - 011 - 9222.","Euzen, O. (1987) Food habits and diet composition of some fish of Kuwait. Kuwait Bulletin of Marine Science, 9, 65 - 85.","Vossoughi, G. H. & Vosoughi, A. R. (1999) Study of batoid fishes in northern part of Hormoz Strait, with emphasis on some species new to the Persian Gulf and Sea of Oman. Indian Journal of Fisheries, 46, 301 - 306.","White, W. T., Last, P. R., Stevens, J. D., Yearsley, G. K., Fahmi & Dharmadi (2006) Economically important sharks and rays of Indonesia. Canberra: ACIAR Publishing, 329 pp.","Last, P. R., White, W. T., Caira, J. N., Dharmadi, Fahmi, Jensen, K., Lim, A. P. K., Manjaji-Matsumoto, B. M., Naylor, G. J. P., Pogonoski, J. J., Stevens, J. D., & Yearsley, G. K. (2010) Sharks and Rays of Borneo. CSIRO, Australia. 304 pp.","Bleeker, P. (1852) Bijdrage tot de kennis der Plagiostomen van den Indischen Archipel. Verhandelingen van het Bataviaasch Genootschap van Kunsten en Wettenschappen, 24 (12), 1 - 92.","Carpenter K. E, Krupp F., Jones D. A. & Zajonz, U. (1997) The living marine resources of Kuwait, Eastern Saudi Arabia, Bahrain, Qatar, and the United Arab Emirates. FAO, Rome.","Wallace, J. H. (1967) The Batoid Fishes of the East Coast of Southern Africa. Part II: Manta, Eagle, Duckbill, Cownose, Butterfly and Stingrays. South African Association for Marine Biological Research. Oceanographic Research. Oceanographic Research Institute Investigational Report, No. 16.","Compagno, L. J. V. (1986). Family Dasyatidae. In: Smith, M. M. and Heemstra, P. C. (Ed) Smiths' Sea Fishes. Macmillian, Johannesburg. pp. 135 - 142."]}

Alopiidae (Thresher sharks) No confirmed Gulf records. Goubanov & Shleib (1980) reported thresher shark Alopias vulpinus (Bonnaterre, 1788) as “very seldom seen in the Gulf, only in its southern areas”, although Randall (1996) regarded this as doubtful. Bigeye thresher A. superciliosus (Lowe, 1841) was regarded by Compagno et al. (2005) as possibly occurring in the Gulf, and both this species and the pelagic thresher A. pelagicus Nakamura, 1935 are known from Oman (Henderson et al. 2007).

Carcharhinidae (Whaler or requiem sharks) The silvertip shark Carcharhinus albimarginatus (Rüppell, 1837) was recorded by Goubanov & Shleib (1980; subsequently noted in Bishop, 2003) as seldom encountered in the southern Gulf although they did not present evidence; Randall (1996) considered this record doubtful. This species is generally associated with shelf waters, offshore islands, reefs, and banks, and near drop-offs, with the nearest confirmed records from the Red Sea and the Maldives (Compagno et al. 2005); it was not recorded in Omani surveys (Henderson et al. 2007; Henderson & Reeve, 2011); it is considered unlikely to be present in the Gulf. A single specimen of a shark morphologically similar to, but clearly distinct from, C. dussumieri, was photographed by the lead author in Kuwait in April 2008 but not retained. Whole specimens are needed to resolve the identity of this species (W. White, CSIRO Marine Research, pers. comm.). The silky shark Carcharhinus falciformis (Müller & Henle, 1839) is abundant in Omani waters (Henderson et al. 2007) and considered to be possibly present in the Gulf (Compagno et al. 2005), although no records are known to the current authors; this species may be more likely to occur in the deeper waters of the northeastern Gulf. Isolated records of the spadenose shark Scoliodon laticaudus (Müller & Henle, 1838) from fish community studies in Kuwait Bay (Wright et al. 1990, as S. laticaudatus; Wright et al. 1996) require confirmation. The circumglobal sandbar shark Carcharhinus plumbeus (Nardo, 1827) has been reported (e.g. Goubanov & Shleib, 1980; Herdson 1981; Krupp & Almarri 1996) and photographs of a specimen seen by the authors are more than likely to have originated from the Gulf, although unequivocal evidence is required. Compagno (1984 b) raised the possibility that the critically endangered Ganges shark Glyphis gangeticus (Müller & Henle, 1839) may occur in the Gulf, and although this author did not pursue the idea (e.g. Compagno 1984 a; Compagno et al. 2005) it was considered to be plausible given environmental changes over recent geological history (Moore 2011). Mahdi (1962) noted two separate species of freshwater shark occurring in the Tigris River as far upstream as Baghdad, although these may simply represent misidentification of C. leucas. Nonetheless the undersampled (and highly threatened) Tigris-Euphrates-Karun system remains intriguing. Randall (1996) noted that the whitetip reef shark Triaenodon obesus (Rüppell, 1837) has not been recorded from the Gulf and our data support this.

Triakidae (Houndsharks) A record of the blacktip tope Hypogaleus hyugaensis (Miyosi, 1939) in the Gulf was first reported in Compagno (1984), apparently inserted without this author���s knowledge and on an uncertain basis (LJV Compagno, pers. comm.). Randall (1986, Fig. 39) presented a photograph of the then undescribed Paragaleus randalli from Bahrain as H. hyugaensis. Currently H. hyugaensis is known from southern and eastern Africa, Australia, Taiwan and Japan, and seems to be a deeper-water species than P. randalli (LJV Compagno, pers. comm.). As yet there is no evidence for this species in the Gulf., Published as part of Moore, Alec B. M., Ward, Robert D. & Peirce, Richard, 2012, Sharks of the Persian (Arabian) Gulf: a first annotated checklist (Chondrichthyes: Elasmobranchii), pp. 1-16 in Zootaxa 3167 on page 12, DOI: 10.5281/zenodo.279779, {"references":["Miyosi, Y. (1939) Description of three new species of elasmobranchiate fishes collected at Hyuga Nada, Japan. Bulletin of the Biogeographical Society of Japan, 9, 91 - 97.","Randall, J. E. (1986) Sharks of Arabia. Immel, London."]}

[[Carcharias taurus Rafinesque]] Records of the great white shark Carcharodon carcharias (Linnaeus) (Lamniformes: Lamnidae) in the Persian/Arabian Gulf (hereafter referred to as ���The Gulf���) are limited to a single report from Kuwait, in the northwest Gulf, which has subsequently been noted in key references (e.g. Compagno, 2001). Khalaf (1987) recorded C. carcharias based on an observation of a mounted specimen in Kuwait Science and Natural History Museum on a visit there in March 1987. The specimen is reported as being a female C. carcharias of 3m in length, caught off Kuwait ���s coast and supplied to the museum on the 14th of April 1984. A black and white photograph of the anterior portion (from just posterior of the pelvic fins) was included in Khalaf���s account (Fig. 1 here), but no measurements or morphological data were presented. The present authors examined the photograph and identified the specimen as Carcharias taurus Rafinesque (Lamniformes: Odontaspididae) based on the first dorsal fin origin being well behind the inner margins of the pectoral fins (cf. over the pectoral inner margins in C. carcharias), the first dorsal fin insertion being about over the pelvic fin origin (cf. being well ahead of pelvic fin origin), a short flattened snout (cf. moderately long conical snout), moderately long gill openings not extending onto the dorsal surface (cf. long gill openings), and characteristic protruding teeth. The photograph presented did not allow for examination of the caudal region or for effective examination of colouration. Carcharodon carcharias has been recorded from a wide range of habitats and with an extensive distribution ranging from the equatorial tropics to the sub-Arctic, and from the intertidal down to the continental slope (Compagno, 2001). Cliff et al. (2000), Compagno (2001), and Zuffa et al. (2002) report white sharks from the tropical Western Indian Ocean off Mozambique and Madagascar north to Tanzania and Kenya and including the Seychelles, R��union and Mauritius. There is therefore no theoretical reason why this species should not occur in the Persian Gulf or the wider northwestern Indian Ocean, although the nearest confirmed record is from Sri Lanka and possibly the Red Sea (Compagno, 2001). In addition, there are unconfirmed, anecdotal records of white sharks from the Gulf of Aden coast of Yemen, Djibouti and northern Somalia (Conan Doyle, 1963). Gubanov & Schleib (1980) reported C. taurus as sometimes being encountered in Kuwait waters, although Krupp et al., (2000) noted that there was no material provided to support this. Krupp et al., (2000) reported the first record of C. taurus from The Gulf supported by reference material based on a 2.98m specimen caught off Abu Halifa, Kuwait, in 1997. The C. taurus reported here predates this by thirteen years., Published as part of Alec B. M. Moore, Leonard J. V. Compagno & Ian K. Fergusson, 2007, The Persian / Arabian Gulf's sole great white shark Carcharodon carcharias (Lamniformes: Lamnidae) record from Kuwait: misidentification of a sandtiger shark Carcharias taurus (Lamniformes: Odontaspididae)., pp. 67-68 in Zootaxa 1591 on pages 67-68