Whitley): Whitley, 1940 : 212 Himantura uarnak Paxton et al ., 1989 : 42 Last & Stevens, 1994 : 406 Last & Stevens, 2009 : 449 Himantura uarnak Naylor et al ., 2012 : 70 Himantura australis Himantura H. uarnak H. undulata ( Bleeker, 1852 ) H. undulata H. undulata H. uarnak Three new stingrays (Myliobatiformes: Dasyatidae) from the Indo – West Pacific Last, Peter R. White, William T. Naylor, Gavin Zootaxa 2016 4147 4 377 402 3LY2Y [151,397,1652,1678] Elasmobranchii Dasyatidae Himantura Animalia Myliobatiformes 1 378 Chordata species australis sp. nov.     Himantura toshi(not  Whitley): Whitley, 1940: 212(in part), brief description (misidentification).     Himantura uarnak(not Gmelin):  Paxton et al., 1989: 42(listed);  Last & Stevens, 1994: 406-07, description, illustration;  Last & Stevens, 2009: 449-50, description, illustration (misidentifications).    Himantura uarnak2:  Naylor et al., 2012: 70, 255 (molecular data).    Himanturasp. 4: Last et al., 2016: figs. 3, 5 (molecular data).     Holotype.  CSIROH 7798-04 (tissue accession GN15798), juvenile male 415 mmDW, west of Oriomo River, Daru,  WesternProvince, Papua New Guinea, 9°04.43’S, 143°08.53’E,  25 Oct 2014.   FIGURE 1.Holotype of  Himantura australis  sp. nov., juvenile male 420 mm DW ( CSIROH 7798-04), from Papua New Guinea: (A) Dorsal view (fresh); (B) Ventral view (preserved).   FIGURE 2.Oronasal region of the holotype of  Himantura australis  sp. nov., juvenile male 420 mm DW ( CSIROH 7798-04), from Papua New Guinea (preserved).   FIGURE 3.Denticles of the mid-disc of the holotype of  Himantura australis  sp. nov., juvenile male 420 mm DW ( CSIROH 7798-04), from Papua New Guinea (preserved).   FIGURE 4.Post-caudal sting tail of the holotype of  Himantura australis  sp. nov., juvenile male 420 mm DW ( CSIROH 7798- 04), from Papua New Guinea (preserved). A. Dorsal view; B. Lateral view; C. Ventral view.    Paratypes. 13 specimens:  CSIROH 1134-1, late embryo male 292 mmDW, north of Port Hedland,  WesternAustralia, 19°35.6’S, 118°42.8’E,  32–34 mdepth,  21 Sep 1987;   CSIROH 1463-3, juvenile male 283 mmDW, north of Cape Lambert,  WesternAustralia, 20°06.1’S, 117°21.4’ E,  41 mdepth,  20 Sep 1988;   CSIROH 1920-01 (tail only), mother of  CSIROH 1463-3, North-West Shelf,  WesternAustralia,  20 Sep 1988;   CSIROH 4016-01, neonatal female 309 mmDW, north of Cape Preston,  WesternAustralia, 20°21.1’S, 116°07.3’E,  41–42 mdepth,  25 Aug 1995;   CSIROH 4422-01, juvenile male 314 mmDW, near Proserpine, Repulse Bay, Queensland, Australia, 20°38’S, 148°41.75’E,  11 Nov 1993;   CSIROH 4542-06, juvenile male 310 mmDW, Kamora Riverestuary, West Papua, Indonesia, 4°49.36’S, 136°38.17’E,  5–10 mdepth,  30 May 1996;   CSIROH 7839-01 (tissue accession GN15784), juvenile male 333 mmDW, Darufish market,  WesternProvince, Papua New Guinea, 9°03.91’S, 143°12.59’E,  21 Oct 2014;   CSIROH 7840-01 (tissue accession GN15789), juvenile male 241 mmDW, fishing camp near Daru, 9°02.26’S, 143°11.49’ E,  24 Oct 2014;  NTM S 11144-001, juvenile male 285 mmDW, King Creek, Shoal Bay, Darwin Harbour,  NorthernTerritory, Australia, 12°21.48’S, 131°1.02’E,  15 Jan 1983;  NTM S 11507-006, juvenile male 343 mmDW, Ludmilla Creek, Darwin Harbour,  NorthernTerritory, Australia, 12°24.78’S, 130°50.22’E,  19 Dec 1984;  KFRSunregistered (field accession  220349; tissue accession GN15785), juvenile female 350 mmDW, Katatai,  WesternProvince, Papua New Guinea, 9°01.25’S, 143°20.51’E,  23 Oct 2014;  KFRSunregistered (field accession  220420; tissue accession GN15790), juvenile female 286 mmDW, fishing camp near Daru,  WesternProvince, Papua New Guinea, 9°02.26’S, 143°11.49’ E,  24 Oct 2014;  KFRSunregistered (field accession  230247; tissue accession GN16607), late-term embryo 300 mmDW (from female 1400 mmDW), Gulfof Papua, Papua New Guinea, 7°55’S, 145°00’ E,  1 Dec 2014.    Othermaterial. 15 specimens:  CSIROH 1134-2, juvenile female 297 mmDW, north of Port Hedland,  WesternAustralia, 19°35.6’S, 118°42.8’E,  32–34 mdepth,  21 Sep 1987;   CSIROH 1479-03, juvenile female 259 mmDW,  CSIROH 1479-04, juvenile male 255 mmDW,  CSIROH 1479-05, juvenile male 262 mmDW,  CSIROH 1479-06, juvenile female 273 mmDW, north of Dampier Archipelago,  WesternAustralia, 20°09.5’S, 116°47.7’E,  43 mdepth,  24 Sep 1988;   CSIROH 2371-02, juvenile female 290 mmDW,  CSIROH 2371-03, juvenile female 278 mmDW,  CSIROH 2371-04, juvenile male 283 mmDW,  CSIROH 2371-05, juvenile male 293 mmDW, north of Cape Lambert,  WesternAustralia, 20°06.1’S, 117°21.4’E,  41 mdepth,  20 Sep 1988;   CSIROH 4786-01 (tissue accession GN5082), juvenile male 310 mmDW,  CSIROH 4786-02 (tissue accession GN1596), juvenile male 322 mmDW, near mouth of Buffalo Creek, Lee Point,  NorthernTerritory, Australia, 12°20.25’S, 130°54.48’E,  7 Aug 1997; CSIROH 7629-02; CSIROH 7807-02; PNG-232047; PNG- 230349.   Diagnosis.A species of  Himanturadistinguished by a combination of the following features: disc weakly rhomboidal; preorbital snout moderately short (length 19–22% DW), rather broad, angle 117–127°, with a distinct apical lobe; lateral apices narrowly rounded; orbits moderately large, often strongly protruding (particularly in young); 1–2, mostly heart-shaped suprascapular denticles (not preceded before and after by a row of smaller primary denticles); secondary denticle band developed before birth; dorsal surface of juveniles (smaller than 370 mmDW) dark spotted or with spots and weak reticulations, subadults and adults (exceeding 390 mmDW) more strongly reticulated; dorsal tail of juveniles with 3 rows of spots before caudal sting, faint dark saddles beyond sting (tail lacking alternating black and white bands); tail uniformly dark ventrally; pectoral-fin radials 1 46–152; vertebral centra (excluding synarcual) 123; including synarcual 124.   Description.Disc rhomboidal, width 1.05 in holotype( 1.01–1.06 in paratypes, all early juveniles and neonates < 350 mmDW) times length; anterior angle 110° (103–118°), pectoral angle 96° (92–94°); most robust on cranial region of head, raised slightly on mid-scapular region, maximum thickness 7.55 (6.37–9.09)% disc width (DW). Snout with a distinct apical lobe, angle 117° (117–127°); anterior margin of disc almost straight (not noticeably double convex), lateral apices narrowly rounded; posterior margin broadly convex, free rear tip narrowly rounded. Pelvic fins rather small, length 18.9 (18.3–19.5)% DW; width across base 13.9 (12.2–15.4)% DW; not protruding far beyond disc. Claspers of adult male unavailable for examination. Tail very long and slender, tapering gently from base toward caudal sting then becoming whip-like; total length 3.8–4.2 times DW when undamaged (3.0 in holotypebut tip missing), tail length 3.5–4.0 times precloacal length when intact; base narrow, slightly depressed and oval, width 1.58 (1.26–1.56) times height. No obvious skin folds on dorsal or ventral surfaces of tail, but midventral surface of tail in neonates with a long and narrow, longitudinal fleshy ridge (presumably a rudimentary fold) extending posteriorly from about level of caudal sting for a distance equivalent to tail length before sting; no evidence of ventral ridge in large individuals. Snout rather short, angular, strongly depressed; preoral snout length 2.04 (2.42–2.55) times mouth width, 2.26 (2.34–2.73) times internarial distance, 20.6 (21.2–21.8)% DW; direct preorbital snout length 1.46 (1.34–1.66) times interorbital length; snout to maximum disc width 40.3 (38.9–43.3)% DW, interorbital space almost flat; eye moderately large, length 2.08 (1.80–2.64) in spiracle length; orbits protruding well beyond disc in young, exceedingly so in neonates and less so in large individuals; diameter 1.21 (1.08–1.58) in spiracle length, interorbital distance 2.54 (2.04–2.56) times orbit. Spiracles large, subrectangular, situated laterally or dorsolaterally. Nostrils moderately large, narrowly elongate, oblique, posterior half recurved in posterolateral direction; lateral margin with weak double concavity, length 1.94 (1.85–2.28) in internasal distance; internasal distance 1.75 (1.78–2.03) of prenasal length. Nasal curtain subrectangular, skirt-shaped, relatively broad, width 1.85 (1.77–2.11) times length; lateral margin almost straight, smooth edged; posterolateral apex depressible into shallow groove; posterior margin weakly fringed, weakly concave. Mouth moderately arched; prominent knob at symphysis of upper jaw, retractable mesially into deep notch at symphysis of lower jaw; oronasal groove shallow, extending posteriorly from posterolateral edge of mouth to chin; skin on ventral surface of lower jaw moderately papillose, not confined to narrow strip around lips; no circumoral grooves. Jaws of typesnot dissected to reveal details of mouth, but images of oral region of discarded material indicate a mouth floor with mainly 4 well-developed papillae (medial pair occasionally separated by a smaller papilla); medial pair simple, broad, flattened, rounded distally, subequal in size (slightly larger than outer pair), located near to each other; single outer papilla located near each corner of mouth, well separated from inner pair. Teeth in a juvenile paratype( CSIROH 4542-06) small, subequal in size in upper and lower jaws; narrowly rhombic with 1–2 low, transverse ridges on crown, ridges separated by prominent groove; ~59 vertical rows in upper jaw. Gill openings S-shaped, strongly arched posteriorly, margins smooth; length of first gill slit 1.64 (1.11–1.50) times length of fifth, 2.78 (2.59–3.37) in mouth width; distance between first gill slits 2.39 (2.18–2.54) times internasal distance, 0.48 (0.44–0.48) of ventral head length; distance between fifth gill slits 1.46 (1.45–1.66) times internasal distance, 0.29 (0.29) in ventral head length.  Squamation.Ontogenetic stages 2 and 4 present in available material; stages 0, 1, 3, 5 and 6 not applicable (data on large individuals inadequate). Denticle development relatively rapid; late-term embryos display welldeveloped suprascapular denticles and a loose band of primary denticles along median disc.  Stage 2: Suprascapular and narrow secondary denticle band present in late embryos. Secondary denticles extending from interorbital region, along median disc, almost to pectoral-fin insertions at birth ( 240–350 mmDW); 1–2 (usually 2), well-developed, heart-shaped (occasionally pearl-shaped) suprascapular denticles; first suprascapular denticle largest, with convex crown (length 7.2–9.5 mmin morphometric types); second with flatter crown than first; secondary denticles heart-shaped, similar in size to each other and none enlarged beside suprascapular denticles. Denticles absent on tail of neonates.  Stage4: Secondary denticle band developing more widely over central disc and on head. In juvenile male holotype( CSIROH 7798-04, 415 mmDW), band moderately dense, covering entire interorbit, width at scapular region ~24% DW; small flattened denticles scattered over median dorsal surface of pre-sting tail; remaining disc smooth. In CMO3-65 ( 560 mmDW) denticles minute (not tightly spaced), extending well forward of the eyes. In adult ( CSIROH 1920-01) denticles present over nearly all of tail (absent near ventral base); flattened denticles interspersed with slightly larger and more widely spaced, upright, stellate-based tubercular denticles.  Meristics.Total pectoral-fin radials (non-types) 146–152 (n=3); propterygium 60–64, mesopterygium 17–21, metapterygium 68–70. Pelvic-fin radials difficult to count, possibly 25–27 (n=4). Vertebral centra (excluding synarcual) 123 (n=1), (including synarcual) 124 (n=1); monospondylous (including 2nd synarcual) 50–52 (n=2), pre-sting diplospondylous 66–73 (n=4); and post-sting diplospondylous 0 (n=4).  Colour. When fresh( holotype): Dorsal disc entirely covered with dark brown, coarsely reticulate colour pattern; reticulate markings differing in length, formed from clusters of sequentially coalesced spots; width of reticulations about half of pupil diameter; reticulations separated from each other by narrower and paler yellow wavy lines (mostly much narrower than dark reticulations); dark spots not fused around outer disc and pelvic-fin margins; tail before caudal sting with 3 irregular rows of dark spots (medially and dorsolaterally), beyond sting more uniformly greyish (blotched but not with alternating light and dark bands), darkest distally. Ventral surface of disc largely white; narrow outer margin of disc and pelvic fins dusky with some small darker markings (margins of paratypesoften densely covered with black spots); tail white forward of caudal sting, dark to black posterior to sting, similar to dorsal surface and not banded.   FIGURE 6.Map showing locations of the type material of  Himantura australis  sp. nov.(red),  Taeniura lessoni  sp. nov.(yellow), and  Telatrygon biasa  sp. nov.(blue). Stars denote primary types (holotypes) and solid circles denote secondary types (paratypes). (Map data ©2016 NASA, TerraMetrics, Google Earth).   FIGURE 7.Late embryos of  Himantura leoparda(left, CSIROH 635-02, 200 mm DW) and  H. australis(right, CSIROH 1134-01, 292 mm DW). Other material: Displays two primary developmental colour morphs (based on all available images, both retained and non-retained material): a dark spotted or spotted/weakly reticulated juvenile form (largest observed 370 mmDW) and subadult and adult forms which are more strongly reticulate (smallest observed 390 mmDW). The smallest individuals (e.g. CSIROH 7840-01, 241 mmDW, Fig. 5a; CSIROH 1463-03, 283 mmDW, Fig. 5b) have a honeycomb pattern consisting of irregularly shaped brownish black spots (of more or less similar size and similar to pupil diameter) on central disc separated by narrow yellowish lines; some spots coalesced to form short wavy lines; spots on head and around disc margin typically smaller; dorsal tail before caudal sting with 3 irregular rows of similar dark blotches, not obviously banded beyond sting. Some individuals (e.g. CMO 3-57, 290 mmDW) of this morph had the bulk of their markings coalesced to form a distinct reticulate pattern; dark markings only slightly broader than pale lines separating them. Tail of smallest individuals prominently marked; on pre-sting tail upper surface with single median row of dark spots, dorsalateral surfaces with row of similar spots, ventral surface white; lateral spots persist slight beyond caudal sting; anterior tail beyond sting not strongly banded, but with vague light and dark dorsal saddles, sides of tail pale and ventral surface uniformly dark; posterior most part of tail beyond sting entirely black. Smallest fully reticulate form (CMO 3-13, 390 mmDW) with very dark, coarse reticulate markings covering entire disc; pale lines separating them much less than half their width; tail markings before caudal sting similar, tail dark greyish or black beyond sting; pattern persisting until about 55 cmDW (CMO 3-10, 550 mmDW). Latter stages becoming more finely reticulate (CMO 3-65, 560 mmDW, Fig. 5d; PNGnot retained 100043, 830 mmDW, Fig. 5e; PNGnot retained 130028, 1120 mm DW; PNGnot retained 130022, 1400 mmDW, Fig. 5f) or reticulated and partly ocellated ( PNGnot retained 100096, 1140 mm DW).   TABLE 1.Morphometric data for the holotype ( CSIROH 7798-04) and five paratypes of  Himantura australis  sp. nov., with ratios expressed as percentages of disc width. Holotype Paratypes Range ......continued on the next pageHolotype Paratypes    Disc width (mm) 415 241 350  Total length damaged 306.1 419.1  Disc length 95.2 94.0 98.5  Snout to pectoral-fin insertion 84.3 83.6 87.4  Orbit to pectoral-fin insertion 59.9 58.7 61.6  Snout to maximum disc width 40.3 38.9 43.3  Snout to origin of cloaca 79.0 77.2 83.3  Cloaca origin to tail tip damaged 228.3 335.8  Cloaca origin to caudal sting 39.0 42.0 47.9  Pectoral-fin insertion to caudal sting (horiz) 35.1 34.7 45.6  Disc thickness 13.3 11.0 15.7  Snout (preorbital) length 20.0 19.0 22.4  Snout (preorbital horiz.) length 16.7 16.4 18.9  Orbit diameter 5.4 5.6 6.6  Eye diameter 3.1 3.3 4.0  Spiracle length 6.5 7.1 9.3  Orbit and spiracle length 10.1 9.8 11.3  Interorbital width 13.7 12.0 14.6  Inter-eye width 22.2 21.6 25.2  Distance between spiracles 20.3 19.5 21.6  Head length 45.3 43.5 47.2  Preoral length (to lower jaw) 20.6 21.2 21.8  Prenasal length 16.0 16.0 17.4   TABLE 1.(Continued) Range Size.One paratype( CSIROH 1463-3), a late-term embryo recovered from an adult female ( CSIROH 1920- 01), was 283 mmDW. Two neonates with strong evidence of umbilical scars were 292 and 309 mmDW. A smaller immature male has a healed scar at 241 mmDW. Smallest confirmed adult male 1120 mmDW; largest specimen a 1400 mmDW pregnant female containing 2 embryos 300 mmDW (White, unpublished).   Distribution.Once considered to be conspecific with  Himantura uarnak( Gmelin, 1789)and widespread in the Indo–West Pacific. Now appears to be confined to the Australasian Plate; known from off  PapuaNew Guineaand northern Australia, from Shark Bay (off  Western Australia) to Brisbane (off Queensland); typematerial displayed in Fig. 6. Depth distribution not well documented, but primarily in shallow-water from near the shore to at least 45 mdepth.   Etymology.Noun in apposition referring to the tropical SouthernHemisphere distribution of this  Himantura. Vernacular name: Australian Whipray.  Comparisons.  Himantura australisand  H. leoparda Manjaji-Matsumoto & Last, 2008are the only members of the genus  Himantura( sensu Last et al., 2016)occurring in Australasian seas. The species are similar but differ in coloration:  Himantura australishas a more reticulated pattern on the dorsal disc in adults (adult  H. australishave an ocellated pattern, typical of  H. leoparda, but the ocelli are smaller and remain dominated by reticulations), the suprascapular denticles are few (1–2, rather than being preceded and followed by a row of slightly smaller primary denticles), and the snout is broader (rather than being produced slightly and more angular) in young and mostly in adults. Juveniles differ in the following morphometric details: preoral length 2.04–2.55 times mouth width (vs. 2.79–3.30 in  H. leoparda), and 20.6–21.8% DW (vs. 23.3–27.6%); distance between first gill slits 2.18–2.54 times internasal distance (vs. 1.98–2.18); distance between fifth gill slits 1.45–1.66 times internasal distance (vs. 1.38– 1.40), ~ 0.29 inventral head length (vs. 0.25–0.28).    Himantura australisis not sympatric with its other congeners and its relationship to these species is part of a revision of the group in progress. It exhibits strong molecular divergence from the other reticulate  Himanturaspecies,  H. uarnakand  H. undulata( Bleeker, 1852)(see Last et al., 2016, Fig. 3). Morphologically it differs from  H. undulatain having smaller reticulations, a less elongate snout, and lacks a pair of pearl-shaped suprascapular denticles characteristic of  H. undulata. Its reticulate pattern in adults is typically more pronounced than in  H. uarnak, but elucidating characters to separate them across all size groups is a work in progress. 1290990289 2014-10-25 CSIRO Papua New Guinea -9.073833 Oriomo River 13 143.14217 Daru 1 378 1 Western Province holotype 1290990293 1987-09-21 CSIRO 33 -19.593334 Port Hedland 127 118.71333 4 381 13 Western Australia paratype 1290990282 1988-09-20 CSIRO 41 -20.101667 Cape Lambert 126 117.35667 4 381 1 Western Australia paratype 1290990269 1988-09-20 CSIRO North-West Shelf 4 381 1 Western Australia paratype 1290990268 1995-08-25 CSIRO 42 -20.351667 Cape Preston 126 116.121666 4 381 1 Western Australia paratype 1290990258 1993-11-11 CSIRO Australia -20.633333 Proserpine 925 148.69583 Repulse Bay 4 381 1 Queensland paratype 1290990255 1996-05-30 CSIRO Indonesia 8 -4.8226666 Kamora River 13 136.63617 4 381 1 West Papua paratype 1290990294 2014-10-21 CSIRO Papua New Guinea -9.065166 Daru 13 143.20984 4 381 1 Western Province paratype 1290990274 2014-10-24 CSIRO Papua New Guinea -9.037666 Daru 13 143.1915 4 381 1 Western Province paratype 1290990262 1983-01-15 NTM Australia Darwin Harbour -12.358 King Creek 13 131.017 Shoal Bay 4 381 1 Northern Territory paratype 1290990250 1984-12-19 NTM Australia -12.413 Ludmilla Creek 13 130.837 Darwin Harbour 4 381 1 Northern Territory paratype 1290990251 2014-10-23 49-03-22 2014-10-23 KFRS Papua New Guinea -9.020833 Katatai 13 143.34183 4 381 1 Western Province paratype 1290990267 20-04-22 2014-10-24 20-04-22 KFRS Papua New Guinea -9.037666 Daru 13 143.1915 4 381 1 Western Province paratype 1290990249 2014-12-01 47-02-23 2014-12-01 KFRS Papua New Guinea -7.9166665 Gulf 1302 145.0 4 381 1 Gulf paratype 1290990271 1987-09-21 CSIRO 33 -19.593334 Other 127 118.71333 Port Hedland 4 381 15 Western Australia 1290990254 1988-09-24 CSIRO 43 -20.158333 Dampier Archipelago 126 116.795 4 381 1 Western Australia 1290990266 1988-09-20 CSIRO 41 -20.101667 Cape Lambert 126 117.35667 4 381 1 Western Australia 1290990281 1997-08-07 CSIRO Australia -12.3375 Buffalo Creek 13 130.908 Lee Point 4 381 1 Northern Territory