Squatina albipunctata, Last, Peter R. & White, William T., 2008

Last, Peter R. & White, William T., 2008, Three new angel sharks (Chondrichthyes: Squatinidae) from the Indo-Australian region, Zootaxa 1734, pp. 1-26 : 5-12

publication ID	https://doi.org/ 10.5281/zenodo.181433
DOI	https://doi.org/10.5281/zenodo.5689525
persistent identifier	https://treatment.plazi.org/id/03F887AA-9C13-FFD3-FF3E-FB165A9CFB8A
treatment provided by	Plazi
scientific name	Squatina albipunctata
status	sp. nov.

Treatment

Squatina albipunctata View in CoL new species

Eastern Angel Shark

Figs 2 View FIGURE 2 , 3 View FIGURE 3 , 8 View FIGURE 8 a, Table 2.

Squatina View in CoL sp. A: Last and Stevens, 1994, Sharks and Rays of Australia, pp 277, 278, figs 31.1, pl. 38. Compagno & Niem, 1998, Squatinidae View in CoL , angelsharks, sand devils. In: Carpenter & Niem (eds), 1998, FAO species identification guide for fisheries purposes. The marine living resources of the Western Central Pacific, pp 1237, figs.

Holotype. CSIRO H 6561–01, adult male 985 mm TL, south of Lakes Entrance, Victoria, 38°42' S, 148°18' E, 84–92 m, 27 July 1993.

Paratypes. 12 specimens. C 4896, immature male 616 mm TL, southeast of Ulladulla, New South Wales, 35°27' S, 150°51' E, 15 Sep. 1976; CA 2842, immature male 407 mm TL, northeast of Sydney, New South Wales, 33°40' S, 151°50' E, 204–210 m, 25 Jan. 1982; CA 3077, immature male 296 mm TL, east of Sydney, New South Wales, 33°44' S, 151°49' E, 348–350 m, 25 Jan. 1982; CSIRO H 594–01, immature male 580 mm TL, northeast of Townsville, Queensland Plateau, 18°10' S, 147°13' E, 240–248 m, 8 Dec. 1985; CSIRO H 713–02, immature male 722 mm TL, south of Saumarez Reef, Queensland, 22°35' S, 153°40' E, 314–319 m, 16 Nov. 1985; CSIRO H 864–01, female 420 mm TL, CSIRO H 864–02, immature male 321 mm TL, east of Batemans Bay, New South Wales, 35°40' S, 150°26' E, 115–117 m, 10 Sep. 1986; CSIRO H 877–07, immature male 596 mm TL, east of Broken Bay, New South Wales, 33°34' S, 151°41' E, 131–133 m, 10 Feb. 1986; CSIRO H 1111–01, female 276 mm TL, Great Barrier Reef off Cairns, Queensland, 17°45' S, 146°53' E, 302– 306 m, 30 Nov. 1985; CSIRO H 1112–01, female 319 mm TL, west of Flinders Reef, Queensland, 17°58' S, 147°01' E, 212 m, 9 Dec. 1985; CSIRO H 1113–01, female 301 mm TL, south of Saumarez Reef, Queensland, 22°06' S, 153°18' E, 246–254 m, 19 Nov. 1985; CSIRO H 1114–01, female 566 mm TL, east of Townsville, Queensland, 19°01' S, 149°28' E, 397–415 m, 26 Nov. 1985.

Other material. 6 specimens. CSIRO H 3538–05, female 340 mm TL, east of Bermagui, New South Wales, 36°24' S, 150°11' E, 77–85 m, 14 Aug. 1993; CSIRO H 3578–02, female 722 mm TL, CSIRO H 3578–03, female 726 mm TL, south of Bermagui, New South Wales, 36°39' S, 150°02' E, 37–60 m, 27 Nov. 1993; CSIRO H 3683–01, immature male 461 mm TL, south east of Disaster Bay, New South Wales, 37°21' S, 150°11' E, 108–110 m, 23 Feb. 1994; CSIRO H 3706–01, female 476 mm TL, east of Wollongong, New South Wales, 34°24' S, 151°11' E, 135–139 m, 20 Apr. 1994; CSIRO H 6560–01, female 1177 mm TL, east of Bermagui, New South Wales, 36°26' S, 150°18' E, 229–236 m, 14 Aug. 1993.

Diagnosis. A moderately large Squatina with the following characters: dorsal coloration complex, yellowish brown with small, white flecks and spots, and pale yellowish blotches; no subdorsal saddles; anterior ventral surface of pectoral fin not black; no median row of scute-like denticles on trunk; anterior nasal flap with strongly fringed barbels; upper lip forming a relatively broad, moderately convex border with snout (lip arch width 3.3–4.5% TL, height 1.0–1.8% TL); interorbital region moderately concave; pectoral–pelvic space 11.9–13.7% TL; pectoral-fin apex broadly rounded to weakly angular, slightly less than 120°; pelvic-fin free rear tips usually extending just beyond first dorsal-fin origin; tail relatively short, pelvic-fin origin to ventral caudal lobe 3.0–3.5 times pectoral-fin base; pectoral fins long, length 34.1–37.1% TL; dorsal fins not lobelike; first dorsal-fin base longer than second dorsal-fin base; caudal fin of adult with narrowly rounded apices; monospondylous centra 48–51; total vertebral centra 134–141.

Description. Body robust, strongly depressed anteriorly, firm; trunk similarly depressed, almost ray-like, deepest over abdomen; not tapering abruptly at pelvic-fin insertion, tail strongly depressed, even at origin of caudal fin. Abdomen moderately elongate, pectoral–pelvic space 12.9 (11.9–13.7)% TL, 0.86 (0.70–0.90) of pre-branchial length; pelvic–caudal space 1.23 (1.03–1.17) times pelvic-fin length; tail short, height at pelvic insertions 1.88 (1.90–2.56) in its width; caudal peduncle strongly depressed, short; lateral keels very welldeveloped, greatly expanded anteriorly, originating slightly forward of insertion of second dorsal fin (below in most paratypes), well below lateral midline, falling short of middle of caudal fin; well-developed dorsal precaudal pit; pit absent ventrally.

Head broad, extended laterally, strongly depressed; oval in cross-section; subcircular, with truncate anterior margin when viewed from above (somewhat rounded in small paratypes); well-developed dermal folds present along ventrolateral margin of head, no lobes present near mouth corners; head width 18.7 (19.1– 21.5)% TL, 3.34 (2.76–3.41) times head height. Snout very short, bluntly rounded in lateral view, horizontal preorbital length 3.2 (3.0–3.9)% TL; slightly bulbous between nostril and eye; no obvious preorbital pit present; apex not indented anteriorly (sometimes slightly indented in paratypes). Eye dorsolateral on head, small, slightly elevated, suboval, length 1.9 (2.2–3.0)% TL, 7.98 (5.59–7.17) in pre-branchial length; interorbit and midline of preorbital snout moderately concave; no subocular pouches; widely separated, interorbital space 4.32 (2.99–3.62) times eye length, 2.55 (2.27–2.83) times horizontal preorbital length, 0.97 (0.88–0.96) times direct prespiracular length, 4.78 (3.63–4.57) times spiracle length. Spiracle small, crescentic to elongate oval, slightly oblique, well separated from eye, subequal to eye; anterior margin strongly concave, barely elevated above flattened posterior margin; posterior margin almost flat to forming a slight depression leading into spiracle; greatest diameter 4.29 (3.49–4.33) in interspiracular space. Gill slits large, pleated, lateral on head, subequal in length, located well forward of midbase of pectoral fin, intergill distance 7.5 (7.4–8.2)% TL, 2.48 (2.51–2.69) in head width; posterior lateral head fold partly overlapping anterior tip of pectoral fin (in some paratypes not overlapping when head is twisted).

Mouth large, terminal (or very slightly inferior), horizontally expanded, broadly arched, terminating at about mid-eye, width 11.8 (12.1–13.8)% TL, 1.58 (1.46–1.60) in head width, 1.46 (1.49–1.61) times interorbital space; upper lip rather broad, forming a moderately convex border with snout; upper labial furrows very large, forming deep preoral groove from symphysis to posterior of jaw angle; lower labial furrows elongate, deep, separated from each other by much less than their length, lip forming a prominent lateral fold; shorter postoral groove very deep, extending parallel to lower labial furrow from jaw angle, length about half lower labial furrow length. Teeth upright or directed orally, widely spaced, unicuspid, dagger-like with a broad base; upper jaw with 3 small rows on each side of symphysis, midlateral teeth slightly larger (about twice size of symphysial teeth in smallest paratypes) and more widely spaced than those adjacent; lower jaw edentate at symphysis, lateral teeth evenly spaced, slightly larger than those of upper jaw; no smaller, unpaired teeth at symphysis of upper jaw; teeth not visible when mouth closed (2 or 3 pairs of teeth protruding in most paratypes); teeth by row, in 8 paratypes, 18 in upper jaw, 14–18 in lower jaw.

Nostrils large, terminal on head, widely separated, internarial space 6.0 (6.1–7.1)% TL, 3.10 (2.79–3.37) in head width; adjacent upper lip of mouth. Anterior nasal flap very large, overlapping mouth, with two very well-developed, strongly fringed barbels; very large, basally flattened, posterolaterally directed barbel with a very strongly fringed tip; distal fringes much larger than incurrent aperture of nostril; broad, strongly fringed membranous basal mesial lobe (partly concealed on inner basal edge of barbel and not distinct from posterolateral extension); central margin of flap with an enlarged, moveable, filamentous barbel (thallate and very strongly fringed in some paratypes), directed ventrally or mesially, originating from a prominent, shallow pocket near border of flap; posteroventral margin of nostril with a variably developed, mostly strongly fringed, membranous flap; no nasoral or circumnarial grooves; no dermal lobes.

Dermal denticles densely covering entire dorsal surface of body (less dense, absent on posterior borders of fins in smaller paratypes, anterior apex of pectoral fins usually naked). Trunk denticles small, widely spaced in smaller paratypes, broadly subconical, raked posteriorly; anterior bases with clearly defined anterior ridges; cusps bluntly pointed. No row of enlarged scute-like denticles on midline. Pre- and postorbit with patches of enlarged, conical denticles, more widely spaced than those above midorbit; a few smaller enlarged denticles on postnarial snout. Adult male holotype with non-embedded, pungent, widely spaced, elevated to slightly posteriorly recurved denticles; much larger than adjacent trunk denticles; denticle patch extending well around pectoral-fin apex; pelvic fin with similar denticles along the outer anterior margin, extending posteriorly along posterior margin for about two thirds its length (absent in immature male paratypes); edge of pectoral and pelvic fin anterior margins with dense coverage of minute aspinous denticles (similar to those of ventral surface). Smallest paratype (CSIRO H 864–02) with less well-developed trunk squamation, denticles in ill-defined linear rows, more or less evenly spaced; enlarged denticles on head well developed, spinose, prominent, two postnasal denticles on each side of head, two on preorbit (posterior larger), two on postorbit (anterior larger); enlarged denticles becoming less distinct in larger paratypes. Ventral surface of body and fins largely naked; broad longitudinal band extending along almost entire length of anterior margin of pectoral fin, sharply demarcated from rest of fin, its mesial border regular, greatest width less than twice orbit diameter; similar but less extensive band around apex of pelvic fin, extending almost from fin origin to its free rear tip, broadest at fin apex (more restricted in smallest paratype); post-pelvic tail mostly covered with denticles, commencing at midline, well anterior of free rear tip of pelvic fins and extending to caudal fin (less welldeveloped in smallest paratypes); ventrolateral surfaces of tail with distinctly larger denticles than on rest of tail; denticles minute, densely packed, aspinous with smooth circular crowns with mostly entire margins, slightly more widely spaced near mesial border. Claspers of adult male holotype naked dorsally; posterior two thirds with a dense patch of denticles resembling those on paired fin margins (immature male paratypes with a weak development of denticles on ventral surface of clasper). Anterior half of dorsal fins with similar coverage of denticles to basal portions of pelvic and pectoral fins; similar denticles on most of caudal fin (smallest paratype with less well-developed denticle patches on dorsal and caudal fins).

Pectoral fins very large, expanded, base very fleshy, length 34.1 (35.1–37.1)% TL, 1.73 (1.83–2.03) times width; anterior margin weakly convex, 1.57 (1.49–1.69) times inner margin; apex broadly rounded to weakly angular, angle slightly less than 120°; posterior margin slightly concave; inner margin very strongly convex, free rear tip broadly rounded. Pelvic fins large, elongate, length 24.1 (22.8–25.8)% TL, 1.42 (1.44–1.57) in pectoral-fin length; anterior margin strongly convex, apex broadly rounded; posterior margin strongly convex (weakly convex in some paratypes); inner margin almost straight; free rear tip bluntly rounded (narrowly pointed in smaller paratypes), not quite reaching origin of first dorsal fin (extending beyond fin origin in most paratypes); pelvic-fin origin anterior to pectoral-fin free rear tip. Claspers of adult male slender, tapering, elongate, somewhat pointed and depressed distally, apex membranous, cartilage at glans opening not sharpedged, outer margin 7.6% TL, 2.20 times base width. Dorsal fins similar in size and shape, somewhat upright (strongly raked in smaller paratypes), close together, height of first 1.06 (0.99–1.12) times height of second; anterior margins straight to slightly convex; apices narrowly rounded (somewhat angular in some paratypes), well posterior to fin insertions; posterior margins slightly convex (almost straight), almost upright; inner margin weakly convex, free rear tip obtuse, broadly rounded; posteriorly located, interdorsal space 1.65 (1.37– 1.81) times first dorsal-fin base, 1.00 (0.96–1.09) times dorsal–caudal space. Caudal fin hypocercal, short, dorsal caudal margin 11.9 (9.9–12.1)% TL, 2.50 (2.18–2.71) in pelvic–caudal space, height 15.7 (13.3– 15.8)% TL; lower lobe much larger than upper lobe, preventral caudal margin 1.62 (1.39–1.75) times dorsal caudal margin; apices narrowly rounded; dorsal caudal margin almost weakly convex or straight; preventral caudal margin strongly convex; postventral margin strongly notched, deepest slightly above its midlength.

Vertebral counts: monospondylous centra 50 (48–51); precaudal centra 108 (102–106); caudal centra 33 (31–34); total centra 141 (134–139).

Colour. In preservative (when mucous removed): Dorsal surface coloration complex; dark yellowish brown, punctuated with dense coverage of small, sharp-edged, white flecks and spots (paratype CSIRO C 4896 appears to be bleached through preservation but still evidence of spots on anterior head) and paler yellowish blotches; edges of lateral keel pale yellowish; no subdorsal saddles; no blotch on caudal fin near its base; short, broad white stripe present on anterior basal half of caudal fin; spiracles brownish; anterior nasal flap brownish and filaments yellowish or brownish. White markings not densest on head, irregularly scattered over dorsal surface; variable in size and shape (more consistent in juveniles), largest slightly less than eye diameter; prominent median post-spiracular spot present; narrow white bar at insertion of pectoral fin; some markings with slightly darker borders than ground colour; scattered paler, diffuse-edged, yellowish blotches over S. albipunctata n. sp. S. australis S. tergocellata

Paratypes (n = 12) n = 5 n = 7

Holotype Min. Max. Min. Max. Min. Max. TL – Total length 985 276 722 251 599 291 655 PCL – Precaudal length 85.3 84.3 85.9 83.1 84.8 84.7 86.4 PD1 – Predorsal length 64.0 63.6 65.5 63.3 65.0 62.5 64.8 PP2 – Prepelvic length 42.8 42.2 45.7 39.0 41.7 41.5 43.4 PP1 – Prepectoral length 18.5 19.4 22.0 19.9 21.1 19.5 20.5 PG1 – Prebranchial length 15.0 15.2 17.2 16.0 16.6 15.5 16.7 PSP – Prespiracular length 8.4 8.4 9.8 8.9 9.6 8.7 9.3 PEY – Pre-eye length 5.6 5.7 6.7 5.7 6.3 5.9 6.4 POB – Preorbital length 3.2 3.0 3.9 3.7 4.5 3.2 4.0 HDW – Head width 18.7 19.1 21.5 19.5 20.7 19.4 20.7 MOW – Mouth width 11.8 12.1 13.8 12.5 14.0 12.4 13.5 HDH – Head height 5.6 6.1 7.5 7.1 8.5 5.9 7.5 INO – Interorbital space 8.1 8.0 9.1 7.5 8.4 8.2 8.8 EYL – Eye length 1.9 2.2 3.0 1.9 2.6 2.5 3.1 INW – Internarial space 6.0 6.1 7.1 6.9 7.4 6.1 6.8 INS – Interspiracular space 7.3 7.5 8.8 8.9 10.1 7.6 8.4 SPL – Spiracle length 1.7 1.8 2.3 1.8 2.0 1.8 2.1 ING1 – Intergill width 7.5 7.4 8.2 8.2 8.9 7.3 8.1 ING – Intergill length 2.2 2.0 2.8 2.2 2.8 2.1 3.0 IDS – Interdorsal space 6.9 5.8 7.1 6.3 7.2 6.0 6.9 DCS – Dorsal–caudal space 6.9 6.2 7.5 5.6 6.4 6.3 7.4 PPS – Pectoral–pelvic space 12.9 11.9 13.7 7.4 8.6 10.5 12.6 POCS – Pelvic (origin)–caudal space 42.6 38.8 42.1 42.2 44.7 41.0 43.8 PCA – Pelvic–caudal space 29.7 25.4 28.1 27.0 28.9 26.6 29.5 WP1 – Width at pectoral origins 14.5 14.5 16.2 16.6 18.2 14.3 15.2 TRW – Trunk width 15.7 15.4 18.0 18.6 20.4 16.1 18.7 TAW – Tail width 9.5 9.0 11.9 11.7 12.1 9.8 11.9 TAH – Tail height 5.0 4.1 5.1 5.0 5.6 4.2 4.9 P1L – Pectoral-fin length 34.1 35.1 37.1 31.3 33.8 34.2 36.9 P1A – Pectoral-fin anterior margin 27.5 27.3 29.9 27.4 29.5 28.6 30.5 P1B – Pectoral-fin base length 12.3 11.9 13.4 13.0 13.9 12.3 12.9 P1W – Pectoral-fin width 19.7 17.5 19.4 17.4 19.3 17.5 19.4 P 1I – Pectoral-fin inner margin 17.5 17.6 19.0 13.5 14.7 16.4 18.9 P2L – Pelvic-fin length 24.1 22.8 25.8 22.6 24.2 24.6 26.4 P2W – Pelvic-fin width 14.7 12.6 14.4 11.5 14.1 12.3 14.1 P 2I – Pelvic-fin inner margin 11.0 9.1 11.8 8.0 8.5 9.1 11.6 D1B – First dorsal-fin base length 4.2 3.8 4.4 3.8 4.3 4.1 4.6 to be continued... surface, obscure on holotype but pronounced on smaller individuals. Dorsal and caudal fins similar to body, white markings less evident; posterior margins almost translucent, distinctly paler than rest of fin; pectoral and pelvic-fin margins whitish or yellowish.

Paratypes CSIRO H 1112–01 and CSIRO H 1114–01 have retained mucous; dorsal coloration greyish pink, white markings less evident; yellowish blotches appearing as dark greyish ocelli and blotches (these markings presumably embedded in the mucous coating and are removed with abrasion); prominent ocelli on pectoral fin adjacent its origin and on mid-portion of the rear lobe. Ventral surface and sides of abdomen usually uniformly whitish, holotype with some darker areas (somewhat translucent along posterior margins of pectoral and pelvic fins in smaller paratypes); denticulated areas usually more yellowish than rest of ventral surface; mouth, lateral head fold and lips whitish. Clasper of adult male holotype almost uniformly pale, distal dorsal portion of glans dusky.

Size. Specimens examined ranged from 276 mm TL (paratype) to 1177 mm TL (female non-type); no information available on birth size. The adult male holotype was 985 mm TL whereas a 722 mm TL paratype was immature.

Distribution. Occurs off eastern Australia between Cairns, Queensland (17°45′ S) and Lakes Entrance, Victoria (38°42′ S) in depths of 37– 415 m.

Etymology. Combination of the Latin albi (meaning white) and punctata (meaning spots) referring to the distinctive white-spotted colour pattern.

Comparisons. Squatina albipunctata differs from its sympatric congener S. australis ( Fig. 9 View FIGURE 9 ) in coloration, morphometrics and meristics. Both species have whitish markings on the dorsal surface, however, in S. albipunctata the markings are represented as regular-sized white spots on yellowish brown skin rather than pale irregular flecks on greyish brown. These species differ markedly in morphometrics: prepelvic length of S. albipunctata 42.2–45.7% vs. 39.0–41.7% TL in S. australis ; interspiracular space 7.3–8.8% vs. 8.9–10.1% TL; intergill width 7.4–8.2% vs. 8.2–8.9% TL; pectoral–pelvic space 11.9–13.7% vs. 7.4–8.6% TL; pectoralfin length 34.1–37.1% vs. 31.3–33.8% TL; pectoral-fin inner margin 17.5–19.0% vs. 13.5–14.7% TL; pelvicfin inner margin 9.1–11.8% vs. 8.0–8.5% TL; pelvic (origin)–caudal space 38.8–42.6% vs. 42.2–44.7% TL; tail height 4.1–5.1% vs. 5.0–5.6% TL and width 9.0–11.9% vs. 11.7–12.1% TL; trunk width 15.4–18.0% vs. 18.6–20.4% TL; and width at pectoral origins 14.5–16.2% vs. 16.6–18.2% TL. They also differ in the following ratios: pectoral–pelvic space 0.7–0.9 vs. about 0.5 times prebranchial length; interorbital space 2.3–2.8 vs. 1.8–2.2 times preorbital length; interspiracular space 3.5–4.3 vs. 4.4–5.5 times spiracle length; head width 2.5–2.7 vs. 2.3–2.4 times intergill width; and pectoral-fin anterior margin 1.5–1.7 vs. 1.9–2.2 times its inner margin. Squatina albipunctata also has far more vertebrae than S. australis (total centra 134–141 vs. 123). Squatina tergocellata ( Fig. 10 View FIGURE 10 ) is very similar in shape to S. albipunctata and, apart from their coloration, they differ only subtly in the observed mean values of some characters: prepectoral length 18.5–22.0 (mean 20.6)% TL in S. albipunctata vs. 19.5–20.5 (mean 19.9)% TL in S. tergocellata ; predorsal length 63.6–65.5 (mean 64.5)% vs. 62.5–64.8 (mean 63.7)% TL; prepelvic length 42.2–45.7 (mean 43.6)% vs. 41.5–43.4 (mean 42.5)% TL; pelvic-fin length 22.8–25.8 (mean 24.3)% vs. 24.6–26.4 (mean 25.4)% TL; pelvic (origin)–caudal space 38.8–42.6 (mean 40.7)% vs. 41.0–43.8 (mean 42.7)% TL; pelvic–caudal space 25.4–29.7 (mean 26.9)% vs. 26.6–29.5 (mean 28.5)% TL; 1st to 2nd dorsal-fin origins 9.6–10.8 (mean 10.4)% vs. 10.6–11.6 (mean 11.0)% TL; and first dorsal-fin base length 3.8–4.4 (mean 4.1)% vs. 4.1–4.6 (mean 4.4)% TL. Squatina albipunctata is white spotted and lack the large, granular ocelli as in S. tergocellata . They also differ slightly in the number of caudal vertebrae, 31–34 in S. albipunctata vs. 27–31 in S. tergocellata .

Of the other Squatina species, only S. aculeata , S. oculata and S. tergocellatoides Chen have strongly fringed or bifurcated nasal barbels and moderately or strongly fringed lateral margins of the anterior nasal flaps. Squatina albipunctata lacks ocellate markings (present in S. tergocellatoides ), lacks spines along the midline of the back (present in S. aculeata ), and the pelvic-fin free rear tips usually extend beyond the first dorsal-fin origin (well forward in S. oculata ) and the anterior nasal barbel is strongly fringed (weakly fringed according to Compagno, 1984).

Squatina albipunctata also differs from all other Australian species and the undescribed Indonesian species in its genetic barcode ( Ward et al., 2008).