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Zootaxa 1176: 41–52 (2006) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 1176 Copyright © 2006 Magnolia Press ISSN 1175-5334 (online edition)

Urolophus kapalensis sp. nov., a new stingaree (: ) off eastern Australia

GORDON K. YEARSLEY & PETER R. LAST CSIRO Marine and Atmospheric Research, GPO Box 1538, Hobart, Tasmania 7001, Australia; gordon.years- [email protected]

Abstract

A new stingaree, kapalensis sp. nov., is described from material collected off eastern Australia. It differs from the partially sympatric U. paucimaculatus, the only other known Urolo- phus with a bell-shaped internasal flap, in having a (absent in U. paucimaculatus), in col- oration (e.g. with a V-shaped band across the interorbit, which is absent in U. paucimaculatus), and in a number of morphometric and meristic characters including: a narrower disc (disc width 4.6–5.0 times distance between first gill slits vs 5.1 in U. paucimaculatus), a longer stinging spine (11.8– 14.9 vs 9.3–11.5% TL), a shorter spiracle (0.8–1.0 vs 1.1–1.2 times orbit length), and more pre- spine vertebrae (86–95 vs 79–88). The two species are also distinguished by the cytochrome oxi- dase subunit 1 (CO1) gene with a divergence of 9%. The new Urolophus is medium-sized, and occurs from Cape Moreton (Qld) south to Disaster Bay (N.S.W.) in depths of 9–79 m.

Key words: Australia, new species, southwestern Pacific Ocean, stingaree, Tasman Sea, Urol- ophidae, Urolophus

Introduction

The batoid family Urolophidae, otherwise known as stingarees, is represented in Australia by two genera Müller & Henle, 1841 and Urolophus Müller & Henle, 1837 (Last & Stevens 1994; Last & Compagno 1999). Members of Trygonoptera are readily identified by the presence of broad, flattened fleshy lobes on the mid-lateral margin of each nostril (lobes absent in Urolophus) (Last & Stevens 1994). Phylogenetically signifi- cant skeletal differences also distinguish the two genera and these are the subject of a forthcoming paper by Yearsley, Last & Gomon (in prep.). Urolophus contains 22 valid nominal species, with 15 recorded from Australian seas (Last & Stevens 1994, Séret & Last 2003). Two new Australian Urolophus species were

Accepted by M. de Carvalho: 23 Feb. 2006; published: 21 Apr. 2006 41 ZOOTAXA identified by Last & Stevens (1994): U. sp. B was recently described as U. piperatus Séret 1176 & Last, 2003; U. sp. A is described herein from specimens collected on fish surveys off eastern Australia. The new species and the sympatric U. paucimaculatus Dixon, 1969 are the only Urolophus species with a bell-shaped internasal flap but these species differ from one another in dorsal coloration, and in a number of morphological and meristic charac- ters.

Methods

Counts and measurements follow Séret & Last (2003). Morphometrics of the holotype and 5 paratypes (CSIRO H 74-05, CSIRO H 74-09, CSIRO H 999-09, CSIRO H 999-11, CSIRO H 999-13) were compared with 4 specimens (CSIRO C 4757, CSIRO H 3-01, CSIRO H 6-01, CSIRO H 337-01) of U. paucimaculatus from the southeastern coast of Australia. Measurements are expressed as a proportion of total length (TL) in Table 1. Meristics were taken from radiographs of the holotype and 7 paratypes (CSIRO H 74-03, CSIRO H 74-05, CSIRO H 74-09, CSIRO H 999-09, CSIRO H 999-11, CSIRO H 999-13, CSIRO H 1002-05), as well as from 5 specimens (CSIRO C 4757, CSIRO H 3-01, CSIRO H 6-01, CSIRO H 722-05, CSIRO H 870-10) of U. paucimaculatus. Disc width is abbrevi- ated to ‘DW’. The mouths of two paratypes, a male (CSIRO H 999-11) and a female (CSIRO H 999-09), were partially dissected to access oral characters, including their oral papillae and teeth. The morphology of the holotype is described below with variation (if any) among paratypes presented in parentheses. The holotype and 7 paratypes were deposited at the Australian National Fish Collec- tion at the Commonwealth Scientific and Industrial Research Organisation’s Marine and Atmospheric Research laboratories in Hobart, Tasmania (CSIRO). Two other paratypes were deposited at the Museum Victoria, Melbourne (NMV).

Urolophus kapalensis sp. nov. Kapala Stingaree (Figs. 1–5)

Urolophus sp. A: Last & Stevens 1994, 428, fig. Trygonoptera testacea (non Müller & Henle): Kuiter 1996, p. 20, fig. Urolophus sp.: Edgar 1997, p. 400, fig.

Holotype. CSIRO H 999-02, 345 mm TL, mature male, Jervis Bay, N.S.W. (35°02’S, 150°45’E), Tasman Sea, southwestern Pacific Ocean, 18–22 m depth, coll. by FRV Kapala, 14 Nov. 1984.

42 © 2006 Magnolia Press YEARSLEY & LAST ZOOTAXA 1176

FIGURE 1. Urolophus kapalensis sp. nov., holotype, CSIRO H 999-02, mature male, 345 mm TL, dorsal view.

Paratypes. 9 specimens. All from the Tasman Sea, southwestern Pacific Ocean. CSIRO H 74-03, 361 mm TL, female, CSIRO H 74-05, 403 mm TL, female, CSIRO H 74- 09, 440 mm TL, female, Jervis Bay, N.S.W. (35°05’S, 150°44’E), 20–30 m depth, coll. by FRV Kapala, 4 Apr. 1984; CSIRO H 999-09, 429 mm TL, female, CSIRO H 999-11, 343 mm TL, mature male, CSIRO H 999-13, 365 mm TL, mature male, CSIRO H 1002-05, 276 mm TL, female, same data as holotype; NMV A 25097-001, 220 mm TL, male, Disas- ter Bay, N.S.W. (37°16’S, 149°58’E), 22–31 m depth, coll. by FRV Southern Surveyor, 3 Aug. 1993; NMV A 25098-001, 250 mm TL, female, east of Lake Macquarie, N.S.W. (33°05’S, 151°42’E), 27–33 m depth, coll. by FRV Kapala, 30 Mar. 1993. Other material. All from the southwestern Pacific Ocean. CSIRO C 2850, 144 mm TL, immature male, Botany Bay, N.S.W. (approximately 33°59’S, 151°11’E), depth unknown, coll. by FV Jaybee, 27 May 1953; CSIRO H 74-04, 370 mm TL, female, Jervis Bay, N.S.W. (35°05’S, 150°44’E), 20–30 m depth, coll. by FRV Kapala, 4 Apr. 1984; CSIRO H 874-01, 327 mm TL, mature male, east of Yamba, N.S.W. (29°28’S, 153°30’E), 50–54 m depth, coll. by FRV Kapala, 26 May 1986; CSIRO H 875-07, 387 mm TL, mature male, east of Crowdy Head, N.S.W. (31°50’S, 152°51’E), 76 m depth, coll. by FRV Kapala, 23 Jul. 1986; CSIRO H 908-01, 360 mm TL, female, CSIRO H 908-04, 399 mm TL, mature male, east of Sydney, N.S.W. (33°58’S, 151°17’E), 47–64 m depth, coll. by FRV Kapala, 25 Mar. 1987; CSIRO H 916-05, 357 mm TL, mature male, east of Port

ONE NEW UROLOPHUS © 2006 Magnolia Press 43 ZOOTAXA Stephens, N.S.W. (32°41’S, 152°16’E), 45–63 m depth, coll. by FRV Kapala, 11 Apr. 1176 1985; CSIRO H 930-04, 326 mm TL, mature male, north of North Solitary Island, N.S.W. (29°49’S, 153°24’E), 36–54 m depth, coll. by FRV Kapala, 24 Mar. 1985; CSIRO H 1002-04, 309 mm TL, mature male, same data as holotype; CSIRO H 3540-02, 433 mm TL, mature male, CSIRO H 3540-03, 521 mm TL, female, east of Bermagui, N.S.W. (36°21’S, 150°07’E), 27–28 m depth, coll. by FRV Southern Surveyor, 14 Aug. 1993; CSIRO H 6153-01, 341 mm TL, mature male, CSIRO H 6153-02, 212 mm TL, immature male, CSIRO H 6153-03, 213 mm TL, female, off Cape Moreton, Qld (about 27°02’S, 153°29’E), 79 m depth, coll. by FV Billy Joe, 7 Feb. 2004; NMV A 28034-001 346 mm TL, mature male, NMV A 28034-002, 250 mm TL, female, NMV A 28034-003, 167 mm TL, immature male, Sydney Harbour, N.S.W. (33°49’S, 152°17’E), 9–15 m depth, coll. by FRV Kapala, 9 Sep. 1981. Diagnosis. A medium-sized Urolophus, a that lacks broad lobes on posterolat- eral border of nostrils, with the following combination of characters: subcircular to weakly rhomboidal disc, width less than 62% TL; bell-shaped internasal flap; tail with well-devel- oped lateral cutaneous folds; stinging spine long, 12–15% TL; dorsal fin low but promi- nent, free-rear tip over spine origin; total vertebrae 156–170; pre-spine vertebrae 86–95; dorsal surface of disc greenish, paler laterally; usually with dark suborbital blotch and V- shaped interorbital bar. Description. Disc subcircular (subcircular to weakly rhomboidal in paratypes), not especially broad, wider than long; width 1.1 times length, 5.0 (4.6–5.0) times distance between first gill slits; broadest about 1.5 (about 1–2) eye diameters behind level of spira- cles; anterior profile obtuse; anterior disc margins almost straight (very weakly concave to weakly convex); pectoral apices broadly rounded; posterior disc margins convex (Fig. 1). Snout fleshy, tip angular, barely extended; preoral snout length 3.1 (2.7–3.2) times interna- rial distance, 1.1 (0.9–1.0) times distance between first gill slits; direct preorbital snout length 3.2 (2.6–3.8) times interorbital distance; snout to maximum disc width 2.5 (2.7–2.8) times in DW; interorbital space broad, almost flat (sometimes slightly concave); orbital region barely distinguishable from head (sometimes elevated well above interorbit, eleva- tion also evident from underwater photographs of non-types), orbit diameter 0.8 (0.8–1.0) in spiracle length; eye of moderate size (small to moderate), lateral, 24% (22–29%) preoc- ular snout length, 12% (10–13%) ventral head length, diameter 1.1 (1.1–1.4) in spiracle length; lower half of eye separated from spiracle by thick, fleshy curtain consisting of sev- eral irregular longitudinal skin folds (when eyes raised, curtain wholly below eye); curtain originating just in advance of mid-eye, inserted near posterior margin of orbit; inter-eye distance 2.5 (2.5–3.1) times eye diameter. Spiracles narrowly tear-shaped and recurved medially, greatly enlarged, opening dorsally (dorsolaterally in paratypes with raised eyes), slightly smaller than orbit, posterior margin bluntly rounded (bluntly rounded to slightly angular), interspiracular distance 1.8 (1.8–2.2) times interorbit distance. Mouth small, slightly convex (slightly convex, occasionally almost straight), width 2.5 (2.1–2.7) in

44 © 2006 Magnolia Press YEARSLEY & LAST snout tip to lower jaw; in dissected paratypes CSIRO H 999-09 and CSIRO H 999-11, 7 ZOOTAXA oral papillae on floor, not arranged in a single series, scattered on a ’W-shaped’ line; single 1176 mesial papilla bifid; 3 lateral pairs located distally to mesial papilla, more-or-less equidis- tant, first closer to mesial papilla than to second papilla; those near angle of jaw about half height of other papillae; external patch of papillae below symphysis of lower jaw well- developed, appearing as ridges or short longitudinal folds. Internasal flap small, distinctly bell-shaped, distinctly broader distally than anteriorly (Fig. 2A), width 1.9 (1.7–2.0) times length, 1.6 (1.3–1.7) times internasal distance; anterolateral margin convex, elevated slightly, forming a low cutaneous ridge; posterior lateral lobe apex extended, narrow, pro- nounced, depressible into a deep oronasal groove; distal fringe well developed, its margin irregular, separated from anterior portion of flap by weak groove, connected directly to posterior lateral lobe. Nostril slightly oblique, slightly shorter than internasal flap, with approximately 38 nasal lamellae in holotype; posterolateral margin entire, concave, form- ing a weak cutaneous knob just in advance of its posterior margin; no broad, flattened fleshy lobe. Jaws strongly asymmetric; upper jaw subrectangular, labial margin almost straight, directed posterolaterally near angle, without exposed tooth rows (Fig. 3A); lower jaw subtriangular, labial margin near symphysis straight, consisting of 9 (6–8 in dissected CSIRO H 999-09 and CSIRO H 999-11) outer exposed tooth rows (Fig. 3B). Teeth quin- cuncial; in female (CSIRO H 999-09), rhomboidal labially in lower jaw; labial margin of cusp less angular than lingual margin; lingual teeth larger than labial teeth, not abutting those adjacent; cusps of teeth towards middle of jaw more pronounced than those distally; teeth in upper jaw similar in size and shape to labial teeth of lower jaw; in male (CSIRO H 999-11), labial upper-jaw teeth weakly rhomboidal, labial margins blunt or weakly con- cave, much less angular than lingual margin, 5 inner series of lower jaw with prominent posteriorly directed cusps, most pronounced towards lingual margin of tooth band; in upper jaw, teeth near labial margin similar to those of lower jaw, series 4 and further lin- gually with prominent cusps.

A B FIGURE 2. Oronasal region of: A) Urolophus kapalensis sp. nov., paratype, CSIRO H 74-09; B) U. viridis, CSIRO H 877-03.

Gill openings moderately S-shaped, non-fringed, margin membraneous; length of first 1.6 (1.4–2.1) times length of fifth gill slit, 3.2 (2.5–2.9) times in mouth width; dis-

ONE NEW UROLOPHUS © 2006 Magnolia Press 45 ZOOTAXA tance between first gill slits 2.8 (2.7–3.2) times internasal distance, 0.4 (0.4–0.5) times 1176 ventral head length; distance between fifth gill slits 1.9 (1.8–2.3) times internasal distance, 0.3 times ventral head length.

AB FIGURE 3. Oral region of Urolophus kapalensis sp. nov., paratype, CSIRO H 999-09: A) upper jaw; B) lower jaw and oral papillae.

Body entirely smooth, sensory pores indistinct dorsally; subcutaneous canal system evident ventrally. Tail rather elongate, postcloacal length 87% (82–90%) disc length; very depressed anteriorly, tapering strongly near pelvic-fin insertions, less strongly tapered near caudal fin; subcircular above anterior part of hypocaudal lobe of caudal fin, compressed below epicaudal lobe; flattened beneath stinging spine; lateral cutaneous tail folds well devel- oped, terminating abruptly near posterior base of stinging spine (Fig. 4). Dorsal fin low, base relatively long, height slightly less than half eye diameter, apex broadly rounded, free rear tip short, over spine origin. Pelvic fins semicircular, moderate (small to moderate), length 1.5 (1.4–1.7) in greatest width across both fins, outer margin very broadly rounded, anterior and posterior margins strongly convex, free rear tip weakly angular. Caudal fin usually narrowly lanceolate (broader in some paratypes), epicaudal-lobe length 3.2 (2.8– 3.5) times fin height; hypocaudal lobe 5.3 (3.7–6.0) times fin height. Clasper robust, short, subconical, slightly depressed, weakly tapering, bluntly pointed distally.

FIGURE 4. Lateral view of posterior tail of Urolophus kapalensis sp. nov., holotype, CSIRO H 999-02.

Stinging spine very elongate, length 0.9 (0.9–1.1) times in epicaudal lobe length, ser- rated for about posterior half, point pungent; holotype with 16 recurved serrations on left- hand side of spine.

46 © 2006 Magnolia Press YEARSLEY & LAST TABLE 1. Selected morphometrics of Urolophus kapalensis (holotype, CSIRO H 999-02; and ZOOTAXA range for five paratypes, CSIRO H 74-05, CSIRO H 74-09, CSIRO H 999-09, CSIRO H 999-11, 1176 CSIRO H 999-13) and U. paucimaculatus (range for 4 specimens, CSIRO C 4757, CSIRO H 3-01, CSIRO H 6-01, CSIRO H 337-01). Data are expressed as percentages of total length.

U. kapalensis U. paucimaculatus Holotype Paratypes Min Max Min Max Total length (mm) 344 342 440 342 430 Disc width 61.9 58.4 62.0 60.9 66.7 Disc length (direct) 55.1 53.8 57.1 54.0 59.0 Disc length (horizontal) 53.8 53.1 56.4 53.5 58.4 Snout to maximum width 25.0 20.6 23.4 22.0 24.2 Snout length—preorbital (direct) 15.1 12.5 13.8 13.1 13.5 Snout length—preorbital (horizontal) 12.8 11.6 12.7 11.3 12.7 Snout to spiracle—ant. (horizontal) 14.1 13.4 14.6 12.6 14.4 Snout to spiracle—post. (direct) 19.6 18.0 19.1 18.1 19.8 Orbit diameter 4.8 4.1 4.9 4.3 4.6 Eye diameter (cornea) 3.6 2.7 3.6 3.2 3.6 Orbit and spiracle length 6.0 5.5 6.1 5.9 6.4 Spiracle length 4.0 3.6 4.3 4.9 5.1 Distance between eyes 9.0 7.7 9.2 8.1 8.9 Distance between orbits 4.7 3.4 5.2 4.8 5.3 Distance between spiracles 8.8 7.6 9.3 8.2 8.5 Distance-snout to post. cloaca 51.9 51.8 54.2 50.9 56.7 Distance-cloaca to caudal fin tip 48.1 45.8 48.2 43.3 49.1 Spine orig to tail tip 25.0 22.8 24.6 20.4 23.1 Spine length—longest 13.7 11.8 14.9 9.3 11.5 Epi-caudal lobe length 14.6 12.5 13.6 11.3 15.1 Hypo-caudal lobe length 21.6 17.7 21.2 18.0 22.6 Max. caudal height 4.1 3.5 4.9 3.8 4.7 Dorsal fin total length 4.0 3.6 5.3 – – Dorsal fin max. height 1.2 0.9 1.4 – – Snout tip to lower jaw 13.8 11.5 12.9 11.7 12.7 Prenasal length (direct) 11.2 8.6 10.0 9.6 10.0 Head length to fifth gill (direct) 29.5 26.2 28.6 26.6 27.8 Mouth width (to corners) 5.4 4.8 6.0 5.3 5.5 Distance between nostrils 4.4 3.8 4.8 3.7 4.7 ...... continued

ONE NEW UROLOPHUS © 2006 Magnolia Press 47 ZOOTAXA TABLE 1 (continued) 1176 U. kapalensis U. paucimaculatus Holotype Paratypes Min Max Min Max Nasal curtain-length 3.9 3.3 4.1 3.6 3.9 Nostril length 2.8 2.1 2.8 2.1 2.5 Nasal curtain (maximum width) 7.3 6.1 7.2 6.7 7.6 Width of first gill opening 1.7 1.7 2.1 1.8 2.1 Width of third gill opening 2.0 1.9 2.1 1.6 2.2 Width of fifth gill opening 1.1 1.0 1.3 1.0 1.3 Distance between first gill openings 12.4 11.8 13.5 11.9 13.0 Distance between fifth gill openings 8.3 8.2 9.2 7.6 8.7 Clasper-post cloacal length 10.9 10.1 10.6 10.5 10.9 Length of (max.) 14.1 13.5 15.4 13.7 17.6 Width across pelvic base 15.8 15.3 18.6 14.3 17.4 Width across pelvics (max.) 21.6 21.4 24.4 19.6 25.5 Tail width (pelvic axil) 5.0 5.5 5.8 5.4 6.6 Tail depth (pelvic axil) 2.6 2.6 3.2 2.8 3.2 Tail width (spine origin) 2.0 2.1 2.6 2.3 3.1 Tail depth (spine origin) 2.0 1.6 1.8 1.6 2.0

Tooth rows (in paratypes CSIRO H 999-09 and CSIRO H 999-11) 25 in upper jaw, 31–32 in lower jaw. Pectoral-fin radials 93–94 (93–105) in total, 47–48 (43–50) proptery- gial, 10–11 (9–13) mesopterygial, 35–36 (36–47) metapterygial. Total pelvic-fin radials 20 (22–23 in males; 22–26 in females). Postsynarcual vertebral centra 170 (156–169) in total, 86 (86–95) in pre-spine total, 30 (30–33) monospondylous, 56 (56–63) pre-spine diplo- spondylous, 84 (68–83) post-spine tail vertebrae. Color when fresh (based on all material available). Dorsal surface of central disc and tail greenish (live coloration shown in Fig. 5); outer margin of disc posterior to eye paler yellowish or brownish; pelvic fins similar to posterior disc margin; whitish where skin removed. Variable dark pattern usually present on upper surface; dark subtriangular blotch adjacent eye, its length slightly exceeding combined orbit and spiracle length (blotch usually pronounced but sometimes faint); thin, dark V-shaped bar extending across interorbit, its apex directed posteriorly, variable in intensity and width; sometimes with dark mid-pectoral blotch and stripe from interorbit to tail base, absent in holotype; dark, irregular blotch at base of pelvic fin, indistinct in holotype, often prominent and extending anteriorly onto pectoral-fin base; clasper base pale, dark mid-dorsally. Ventral surface whitish with darker, broad marginal band on pectoral-fin; marginal band yellowish or

48 © 2006 Magnolia Press YEARSLEY & LAST dusky, extending from anterior disc forward of mouth to pectoral-fin insertion, maximum ZOOTAXA width (near pectoral-fin apex) subequal to prenasal length; similar marginal band on pelvic 1176 fin. Tail mainly whitish dorsally, with darkish, variable, mid-dorsal stripe; mainly whitish on and below lateral fold; ventral tail darker posterior to fold, sometimes with scattered darker blotches, usually dusky or brownish beside hypocaudal lobe. Dorsal fin greenish brown; caudal fin dark greyish brown, darker in juveniles. Clasper pale. Stinging spine yellowish or whitish.

FIGURE 5. Underwater color photograph of Urolophus kapalensis, specimen not collected, from off Bermagui, N.S.W. (about 36°24’S, 150°05’E), depth about 20 m (photograph by R. Kuiter).

Color of preserved specimens. Upper surface uniformly brownish, without evidence of green coloration; dark patches on suborbit persisting and, to a lesser extent on interorbit and pelvic-fin bases (dark patches more or less evident in paratypes). Ventral surface gen- erally pale, yellowish; marginal band distinct, often faint; oronasal region, teeth and buccal cavity uniformly pale; dark markings on tail persistent. Size. Largest specimen examined was a female of 521 mm TL and 312 mm DW (CSIRO H 3540-03). The largest male examined was 433 mm TL (CSIRO H 3540-02). The smallest mature male examined was 309 mm TL (CSIRO H 1002-04). The smallest specimen examined, lacking an umbilical scar and presumably postnatal, was a 144 mm TL juvenile male (CSIRO C 2850). Distribution and . Known only from the off eastern Austra- lia, mostly in the northern Tasman Sea. Based on material listed above, and additional material held at CSIRO, the species ranges from Cape Moreton, Qld (about 27°02’S, 153°29’E), south to Disaster Bay, N.S.W. (about 37°15’S, 149°58’E), in depths of 9–79 m.

ONE NEW UROLOPHUS © 2006 Magnolia Press 49 ZOOTAXA In N.S.W. waters, it is common in the southern part of its range in depths less than 50 m, 1176 and is relatively rarely caught north of Clarence River (about 29°29’S 152°40’E) (K. Gra- ham pers. comm.). However, in a series of south-east Qld surveys, in which the new spe- cies was the subject of a biological study, it was locally common, and the majority of specimens (86%) were found at depths greater than 62 m (P. Kyne, pers. comm.). Etymology. This new species was discovered among specimens collected by the FRV Kapala (formerly of the N.S.W. Research Institute, Australia). The species is named kapalensis to honour the extremely valuable fish collections made by the vessel over almost three decades. Remarks. In addition to dorsal coloration, a combination of the presence of lateral cutaneous tail folds, a dorsal fin, and a bell-shaped internasal flap distinguish Urolophus kapalensis from all other known Urolophus species. The presence or absence of lateral cutaneous tail folds in U. javanicus (Martens) and U. kaianus Günther is unknown, but these non-Australian species differ from U. kapalensis in having a skirt-shaped internasal flap; U. kaianus also lacks a dorsal fin. Of the species where the condition of the lateral cutaneous tail folds is known, U. aurantiacus Müller & Henle, U. circularis McKay, U. cruciatus (Lacepède), U. deforgesi Séret & Last, U. gigas Scott, U. orarius Last & Gomon and U. sufflavus Whitley all lack tail folds, whereas U. kapalensis has well-developed folds. The other remaining species have variably developed tail folds (sometimes intraspe- cifically present or absent) but, of these, U. armatus Müller & Henle, U. expansus McCul- loch, U. lobatus McKay, U. mitosis Last & Gomon, U. neocaledoniensis Séret & Last, U. paucimaculatus and U. viridis McCulloch lack a dorsal fin, which is present in U. kapalen- sis. In U. westraliensis Last & Gomon, the dorsal fin is variably present but, along with U. bucculentus Macleay, U. flavomosaicus Last & Gomon, U. papilio Séret & Last and U. piperatus, the internasal flap is skirt shaped (see example in Fig. 2B), compared with bell shaped (Fig. 2A) in U. kapalensis. In fact, U. kapalensis is one of only two known Urolo- phus species with a bell-shaped internasal flap. The other species, U. paucimaculatus, occurs sympatrically with U. kapalensis off central and southern N.S.W., but its distribu- tion extends further south, around Tasmania and west across the Great Australian Bight to about Perth, Western Australia. Urolophus kapalensis is distinguished from U. paucimacu- latus by the presence of a dorsal fin (lacking in U. paucimaculatus), a narrower disc (disc width 4.6–5.0 vs about 5.1 times distance between first gill slits; see Table 1), a longer stinging spine (11.8–14.9 vs 9.3–11.5% TL), a shorter spiracle (0.8–1.0 vs 1.1–1.2 times orbit length), more pre-spine vertebrae (86–95 vs 79–88), and by a dark, V-shaped band across the interorbit, usually accompanied by dark blotches adjacent to each eye (absent) and dark blotches at the base of the pelvic fins (absent). The dorsal colour pattern of this species is considerably variable within a discrete range of tones. In some material, the dark interorbital bar was absent, the mid-interorbital region sometimes being paler than adjacent surfaces. There was also evidence of a weak, dusky marking between the spiracles. The blotch beside the eye was often weak or absent,

50 © 2006 Magnolia Press YEARSLEY & LAST and no other obvious blotches or bands were evident. Two other specimens also displayed ZOOTAXA unusual coloration. CSIRO H 74-04 differed from 8 typical forms of the species collected 1176 at the same station in having a more rounded disc, a shorter caudal fin, and a mottled colour pattern on the dorsal surface and on the ventral marginal bands. CSIRO H 6153-03 was almost black dorsally, and the ventral marginal band and the caudal fin very dark; black areas were also present ventrally on the tail. These atypical specimens, which could constitute another species, were excluded from the type series. However, a concurrent study that aims to genetically barcode Australia’s fish species (Ward et al. 2005) failed to differentiate between the ‘normal’ colour form of the types and those with the atypical barred coloration discussed above (B. Innes & R. Ward pers. comm.). In the study, cytochrome oxidase subunit 1 (CO1) sequence divergences of U. kapalensis specimens (n = 13) were calculated using Kimura’s (1980) two-parameter model (MEGA version 3; Kumar et al. 2004) and found to be just 0.2–0.3% (B. Innes & R. Ward pers. comm.), well within acceptable intraspecific variation levels. The atypical specimens need further investigation. The same genetic study distinguished the two Urolophus species possessing a bell- shaped internasal flap (U. kapalensis and U. paucimaculatus), with a divergence of 9% (B. Innes & R. Ward pers. comm.). Comparative material examined. All from southwestern Pacific Ocean. Urolophus paucimaculatus—CSIRO C 4757, 384 mm TL, mature male, south-west of Cape Otway, Vic. (39°09’S, 143°15’E), 50–67 m depth, coll. by FRV Courageous, 20 Jun. 1976; CSIRO H 3-01, 430 mm TL, female, Frederick Henry Bay, Tas. (42°55’S, 147°36’E), 20 m depth, coll. by FV Allanwood, 16 Sep. 1983; CSIRO H 6-01, 342 mm TL, mature male, same data as CSIRO H 3-01; CSIRO H 337-01, off Stanley, Tas. (exact coordinates unknown), 30 m depth, collector unkown, 19 May 1980; CSIRO H 722-05, south-east of Broken Bay, N.S.W. (33°38’S, 151°27’E), 63–76 m depth, coll. by FRV Kapala, 20 Nov. 1985; CSIRO H 870-10, east of Port Stephens, N.S.W. (32°42’S, 152°32’E), 124–133 m depth, coll. by FRV Kapala, 22 Apr. 1986. Urolophus viridis—CSIRO H 877-03, east of Broken Bay, N.S.W. (33°34’S, 151°41’E), 131–133 m depth, coll. by FRV Kapala, 10 Feb. 1986.

Acknowledgments

We are especially grateful to K. Graham (N.S.W. Fisheries) for kindly supplying the authors with numerous specimens of the new species that were collected by the FRV Kapala, and to A. Graham (CSIRO) for his helpful curatorial assistance during this study. T. Fountain and S. Riddoch (CSIRO) assisted with the collection of meristic data, and D. Gledhill (CSIRO) obtained some morphometric data. We thank M. Gomon and D. Bray (NMV) for providing comparative material on loan from the National Museum of Victoria. Photographs were taken by L. Conboy, T. Carter (CSIRO) and R. Kuiter (Aquatic Photo-

ONE NEW UROLOPHUS © 2006 Magnolia Press 51 ZOOTAXA graphics), and were digitised and etched by L. Conboy. P. Kyne (University of Queen- 1176 sland) supplied specimens from off Cape Moreton, Queensland. M. Berry (University of Tasmania) provided helpful advice on the Latinisation of Kapala. R. Ward and B Innes (CSIRO) kindly supplied CO1 sequence divergence data. M. Gomon, P. Kyne, K. Graham, and two anonymous referees, provided very useful comments on the manuscript.

References

Edgar, G.J. (1997) Australian Marine Life: the Plants and of Temperate Waters. Reed Books, Sydney, 544pp. Kimura, M. (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120. Kuiter, R.H. (1996) Guide to Sea Fishes of Australia. New Holland (Publishers) Inc., Frenchs For- est, 433pp. Kumar, S., Tamura, K. & Nei, M. (2004) MEGA3: Integrated software for molecular evolutionary genetics analysis and sequence alignment. Briefings in Bioinformatics, 5, 150–163. Last, P.R. & Compagno, L.J.V. (1999) Urolophidae. In: Carpenter, K.E. and Niem, V.H. Species identification guide for fisheries purposes. The living marine resources of the western central Pacific. Volume 3. Batoid fishes, chimeras and bony fishes part 1 (Elopidae to Linophrynidae). FAO, Rome, i–vi, pp 1397–2068. Last, P.R. & Stevens, J.D. (1994) Sharks and Rays of Australia. CSIRO Publishing, Melbourne, 513pp. Séret, B. & Last. P.R. (2003) Description of four new stingarees of the genus Urolophus (: Urolophidae) from the , south-west Pacific. Cybium, 27(4), 307–320. Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R. & Hebert, P.D.N. (2005) DNA barcoding Austra- lia’s fish species. Philosophical Transactions of the Royal Society of London B, 360, 1847– 1857.

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