Halaelurus maculosus, William T. White, Peter R. Last & John D. Stevens, 2007

William T. White, Peter R. Last & John D. Stevens, 2007, Halaelurus maculosus n. sp. and H. sellus n. sp., two new species of catshark (Carcharhiniformes: Scyliorhinidae) from the Indo-West Pacific., Zootaxa 1639, pp. 1-21 : 2-10

Halaelurus maculosus new species

Indonesian Speckled Catshark

Figs 1-4; Tables 1 and 2.

Halaelurus cf buergeri : White et al., 2006: 178, 179.

Holotype. MZB 15499 , adult male 436 mm TL, Kedonganan fish market , Bali, Indonesia, 08°45' S, 115° 10' E, July 2002.

Paratypes. 7 specimens: CSIRO H 5889-09 , adult male 457 mm TL, CSIRO H 5889-16 , adult male 439 mm TL, CSIRO H 5889-17 , adult male 444 mm TL, CSIRO H 5889-18 , adult male 430 mm TL, CSIRO H 5890-01 , immature male 315 mm TL, MZB 15550 , adult male 421 mm TL, collected at same locality as holotype ; CSIRO H 5858-01 , female 426 mm TL, Cilacap landing site , Central Java, Indonesia, 07°40' S, 109°00' E, 2001 ; CSIRO H 6497-02 , 17 egg cases, Tanjung Luar landing site , Lombok, Indonesia, 08°45'S, 116°35'E, 2006.

Other material. 14 specimens: CSIRO H 5889-14 , adult male 434 mm TL, CSIRO H 5889-25 , immature male 344 mm TL, CSIRO H 5890-02 , adolescent male, 360 mm TL, MZB 15446 , adult male 426 mm TL, MZB 15448 , adult male 411 mm TL, MZB 15449 , adolescent male 413 mm TL, MZB 15459 , adult male 430 mm TL, MZB 15462 , adult male 433 mm TL, MZB 15464 , adult male 444 mm TL, MZB 15465 , adult male 400 mm TL, MZB 15466 , female 453 mm TL, MZB 15467 , adult male 412 mm TL, MZB 15490 , adult male 441 mm TL, collected at same locality as holotype ; JPAG 114 , female 507 mm TL, Philippines , 1999 .

Diagnosis. Member of the genus Halaelurus with the following combination of characters: a relatively short, broadly rounded snout, preoral length 3.9-4.9% TL; predorsal length 41.6-42.8% TL, 3.8-4.7 times anal -caudal space; anal -caudal space 9.0-10.9% TL; mouth moderately long, length 2.8-3.6% TL; upper labial furrows very short (8.1-14.9 in eye length), sometimes inconspicuous; eyes moderately large, eye length 3.1-4.0% TL; pectoral fin with broadly rounded apex, anterior margin 10.8-11.7% TL, 1.9-2.3 times inner margin; anal fin relatively large, length 11.0-12.2% TL, anterior margin 8.0-9.1% TL; adult clasper elongate and relatively narrow, outer length 6.4-7.5% TL, 3.6-5.0 times clasper base length, extending past pelvic-fin free rear tip by 0.25-0.36 times pelvic-fin inner margin length, clasper tip narrowly rounded; total vertebral centra 127-132, precaudal centra 81-90; dorsal surface with broad, dark brownish saddles, densely covered with numerous, dark brown spots; spots present on interspaces between saddles. Small adult size, attaining about 530 mm TL.

Description. Proportions as percentages of total length for the holotype and paratypes (7) are presented in Table 1.

Body tapering, anterior trunk wide and somewhat depressed; length of trunk from fifth gill openings to vent 1.06 (1.02-1.30) times head length; tail long and slender, almost circular in cross-section at second dorsal-fin insertion; no lateral keels; height 1.21 (0.91-1.10) in width at second dorsal-fin insertion, 3.28 (2.62- 3.08) in dorsal -caudal space; no postnatal ridge between anal fin base and lower caudal-fin origin; no longitudinal ridges on dorsal surface, no obvious lateral ridges. Head short, length 1.31 (1.04-1.33) to pectoral -pelvic space; wide and somewhat depressed, more so between eyes and ventrally; narrowly pointed, not upturned or knob-like in lateral view (slightly upturned in some paratypes); in dorsoventral view anterior to gill openings broadly parabolic; subocular ridge evident, variably angular. Preoral snout short, 0.68 (0.49-0.76) times mouth width; broadly rounded in dorsoventral view, no indentations adjacent to anterior of orbits. Eyes small, spindle-shaped; dorsolateral on head, with lower edges well medial to lateral head margin in dorsal view; subocular ridges weak, not well defined, subequal in length to eye, almost a spiracle length from eye margin; no supraorbital crest; length 4.95 (4.80-6.00) in head length, 6.90 (4.42-6.14) times eye height.

External eye openings with or without weak anterior and posterior notches. Nictitating lower eyelids of rudimentary type. Spiracles small, variable, subcircular, cresentic to suboval, length 3.65 (3.17-5.05) in eye length, 0.17 (0.16-0.26) eye length behind and below posterior eye notch. Gill openings straight to slightly concave; slightly elevated on dorsolateral surface of head, upper margins slightly below lower edge of eye; anteriormost openings slightly larger than those posterior (heights variable, first four subequal or with anteriormost taller in paratypes); fifth opening much shorter than anterior four; fourth opening over origin of pectoral fin; height of fifth 0.57 (0.54-0.61) of third; height of third 14.32 (11.09-14.91) in head, 0.35 (0.35-0.51) of eye length. Gill filaments not visible from outside. Nostrils separated from each other and not reaching mouth; very small incurrent apertures; width 0.94 (1.02-1.29) in internarial space, 1.42 (1.40-1.66) in eye length, 0.49 (0.50-0.77) in third gill-slit height. Anterior nasal flaps large, forming a right angle at apex (sometimes with a weak lobe in paratypes); posterior margin nearly straight or weakly corrugated, innermost edge short of upper lip by more than a tooth length; covering excurrent apertures; posterior nasal flaps present but inconspicuous, directed internally, visible when upper flap is lifted (but exposed in paratype CSIRO H 5889-17); skirt-shaped fold around excurrent aperture (nasal curtain and fold overlapping).

Mouth moderately large, very broadly arched; upper jaw horizontal at symphysis, forming a sharp angle just lateral to insertion of nasal curtain, posterior half of jaw almost straight, directed posterolaterally; central part of upper jaw overhanging lower jaw so that upper teeth are exposed in ventral view, edge of lower jaw similar in shape to edge of upper jaw; width 2.64 (2.27-2.99) in head length; length 2.19 (1.98-2.84) in width. Tongue large, flat, rounded anteriorly, filling most of floor of mouth (from paratype MZB 15550). Labial furrows very short but usually conspicuous (labial furrows on left side inconspicuous in some paratypes), confined to jaw angle; upper furrows shorter than lower furrows, 0.58 (0.32-0.69) of lower furrows. Teeth in 56/ 53 rows; 5/4 series functional; usually with long central cusp flanked by either one or two short cusplets on each side; lateral teeth with progressively smaller cusps; no toothless spaces at symphysis; not strongly differentiated in upper and lower jaws; medials, anterolaterals and posteriors in both jaws weakly defined; central upper anterolateral teeth largest. Tooth formula (from paratype MZB 15550) is: 27-2-27 / 25-4-24

Sexual heterodonty present; upper anterolateral teeth in an adult male paratype (MZB 15550) with enlarged central cusp compared to those in the female paratype (CSIRO H 5858-01), teeth height ~1.3 vs. 0.6-0.7 mm.

Lateral trunk denticles below first dorsal fin small (0.5-0.6 mm long), dense, strongly imbricate. Crowns mostly tricuspid, with long pungent central cusp flanked by short, sharp lateral cusps; pair of strong, converging medial ridges extending entire length of crown; pair of low but prominent ridges on lateral cusps; crown length 1.5 to 2 times width, medial cusp about half or less total length of denticle.

First dorsal fin low, angled rearwards, not falcate; slightly convex anterior margin, apex broadly rounded; posterior margin almost straight (sometimes weakly convex), sloping slightly posteroventrally from apex (upright in some paratypes); free rear tip angular, inner margin nearly straight; insertion slightly anterior to fin apex; origin barely anterior to pelvic-fin insertions (further forward in female paratype CSIRO H 5858-01); insertion slightly posterior to pelvic-fin free rear tips (over free rear tips in female paratype), well anterior to anal-fin origin. First dorsal-fin base 1.87 (1.68-1.92) in interdorsal space, 2.68 (2.08-2.81) in dorsal caudalfin margin; height 1.59 (1.55-1.96) in base length; inner margin 2.14 (1.87-2.55) in height, 3.41 (3.09-4.51) in base length. Second dorsal fin low, subtriangular, generally lower and more elongate than first dorsal fin; height 0.88 (0.77-0.87) of first dorsal-fin height, base length 1.08 (0.87-1.13) of first dorsal-fin base length; anterior margin nearly straight or slightly convex, with bluntly angular to narrowly rounded apex; posterior margin sloping posteroventrally from apex, weakly concave; free rear tip angular, extended slightly; inner margin straight; origin slightly forward to insertion of anal-fin (over anal-fin insertion in smallest paratype); insertion well posterior to anal-fin free rear tip, about opposite second dorsal-fin apex. Second dorsal-fin base length 1.10 (0.91-1.12) in dorsal -caudal space; height 1.94 (1.82-2.34) base length; inner margin 2.00 (1.50- 2.13) in height, 3.89 (3.26-4.26) in base length.

Anal fin low, subtriangular, smaller in area to second dorsal fin; height 0.76 (0.67-0.80) in second dorsalfin height, base 1.27 (1.06-1.21) times second dorsal-fin base; base sometimes with a preanal ridge; anterior margin nearly straight or weakly convex, apex broadly rounded; posterior margin weakly concave, slanting strongly posterodorsally; free rear tip angular, extended slightly, inner margin nearly straight; origin about 1.26 (1.26-1.48) times base length behind pelvic-fin insertions; insertion well posterior to apex. Anal-fin base 0.98 (0.83-1.07) in anal-caudal space; height 3.25 (3.07-3.73) in base length; inner margin 1.43 (1.14-1.45) in height, 4.66 (3.77-4.46) in base length.

Pectoral fin small, somewhat lobate, apex almost forming a right angle; anterior margin weakly convex, 1.02 (0.97-1.07) times its length; base narrow; posterior margin almost straight; free rear tip broadly rounded, inner margin moderately to strongly convex. Pelvic fin weakly triangular; anterior margin slightly convex, 0.60 (0.58-0.64) of pectoral-fin anterior margin; apex broadly rounded; posterior margin slightly convex; free rear tip angular, slightly attenuated in adult males; inner margins straight, not fused together over claspers of adult males.

Claspers of the adult male holotype elongate, moderately slender, nearly straight and tapering on lateral edge, not undulated; extending well behind pelvic-fin free rear tips, but terminating well anterior to anal-fin origin; tip sharply pointed, not fleshy. Most of ventral and distal portion of dorsal clasper with small clasper denticles with anteriorly directed cusps; anterior dorsal surface of glans largely naked. Apopyle and hypopyle connected by long clasper groove, with its dorsal margins fused over clasper canal. Lateral fold well developed, originating along length of dorsal midline and enveloping part of inner dorsal half of clasper; terminating distally just posterior to origin of cover rhipidion. Cover rhipidion very short, narrow, formed as distally tapering wedge, mesial margin with fleshy, longitudinal fold, without mid-lateral tab; separated posteriorly from posterior end of external edge of clasper by a deep groove; partly posterior to exorhipidion, well ahead of the rear end of the rhipidion. Rhipidion present, large, extending along most of clasper glans; formed as a flat blade with posterior tip beside apex of exorhipidion. Pseudosiphon absent. Pseudopera indistinct. Exorhipidion elongate, tapering distally, not flap-like, originating near posterolateral edge of cover rhipidion; a series of about 20 small clasper hooks along outer rim of hypopyle, mostly in multiple rows; apex sharply pointed.

Caudal fin narrow and asymmetrical; terminal lobe large; ventral lobe very low; dorsal margin moderately long, nearly straight with weak lateral undulations, 3.96 (3.67-4.60) in precaudal length; preventral margin nearly straight, 1.68 (1.53-1.82) in dorsal caudal-fin margin; tip of ventral caudal-fin lobe broadly rounded; postventral margin weakly concave, not well differentiated into upper and lower parts. Subterminal notch a narrow, shallow slot; margin straight, its length 0.71 (0.82-1.16) in terminal region. Terminal margin slightly convex, sometimes notched, dorsal and ventral tips broadly rounded, its length 3.66 (3.39-4.62) times dorsal caudal-fin margin.

Egg cases medium-sized, 45-50 mm in length from anterior to posterior margins (excluding horns); smooth-walls without longitudinal striations or ridges; relatively flat, its height 18.1 (16.0-19.4)% of case length; posterior width 39.2 (38.3-40.2)% of case length; lateral flanges absent. Anterior border nearly straight, broad; anterior horns small and nearly vestigial; tendrils long, wiry and tightly coiled. Posterior border narrow, concave; posterior horns short, curved strongly inwards but not touching; tendrils very long, wiry, highly convoluted, tightly coiled. Egg cases, in alcohol, are uniformly golden yellow to brown in colour; similar, but slightly darker, when fresh.

Vertebral centra 132 (127-132 in paratypes), monospondylous 33 (33-34), precaudal 90 (81-88, mostly less than 83-85) and caudal 37 (38-41). Last few MP centra before MP-DP transition hardly enlarged, not forming ‘stutter zone’ of alternating long and short centra.

Colour. In preservative: Dorsal surface pale brown, with about 10 vague, widely separated, darker brown saddles and densely peppered with brownish black spots; margins of saddles not demarcated by darker transverse lines; saddles mostly subequal in breadth to their interspaces, extending to about lateral midline on tail and slightly more ventrally on trunk; interspaces densely spotted; spots variable in size, largest (slightly larger than spiracle) mostly widely spaced on side on abdomen, smallest (about quarter of spiracle diameter) densest along dorsal midline and on head (spots consistently much smaller than spiracles in some paratypes). Head with two weakly defined saddles, across orbito -spiracular region and between gills; anterior margin (origin) of first saddle through middle of interorbit, posterior margin (insertion) between spiracles, well-developed extensions of saddle directed antero-ventrally below eyes; second saddle not obviously recurved, origin through second gill slit, insertion well posterior to last gill slit; small dark spots densely scattered over snout (~10-13 spots per cm2, denser in some paratypes). Supra-abdominal region with two similar saddles; anterior saddle originating just posterior to pectoral-fin insertion, terminating well-behind level of its free rear tip; second saddle centered over pelvic-fin origin. Tail and caudal fin with 6 saddles; one saddle centered under each dorsal-fin base; slightly broader, central interdorsal saddle; narrower saddle centered on dorsal -caudal space; two broad saddles on caudal fin, anterior saddle above lower caudal lobe, narrower than posterior saddle at tail apex. Dorsal fins densely spotted, with evidence of rusty saddles anteriorly, paler posteriorly. Paired fins evenly spotted on dorsal surface, not paler distally; ventral surface mostly uniformly yellowish. Lower lobe of caudal fin spotted. Anal fin uniform yellowish. Ventral surface of body pale yellow or creamish, dusky on tail (holotype and most paratypes with dark, embedded, sand granules); mouth white (more yellowish is some paratypes).

Size. Attains a maximum size of about 530 mm TL; males mature at ~400 mm TL; five females between 476 and 528 mm TL were pregnant, with between three and six egg cases in each of the two uteri.

Distribution. Known from the southern coast of Java, and off the islands of Bali and Lombok in eastern Indonesia, and has also been recorded from several landing sites in the Philippines (see Fig. 9).

Habitat. Caught occasionally by demersal fisheries, but no depth distribution information available.

Etymology. Derived from the Latin maculosus (speckled) in allusion to the speckling of small dark spots over the dorsal and lateral surfaces of the body.

Comparisons. Halaelurus maculosus was first recognised from the Philippines as being similar but possible distinct from H. buergeri (Compagno et al., 2005). However, it differs from H. buergeri in having a colour pattern of numerous, small to moderately large dark spots (usually much smaller than spiracle and covering the entire dorsal and lateral surfaces of the body), rather than fewer, larger dark spots (usually larger than spiracle and arranged along the saddle-marking borders but which are absent from the paler areas between saddles) (see Figs 1 and 11). Halaelurus maculosus also differs from H. buergeri in a number of morphological characters: more posteriorly positioned first dorsal fin (predorsal length 41.6-42.8 vs. 39.3-41.0% TL), dorsal fins closer together (interdorsal space 13.1-15.0 vs. 16.0-17.1% TL), slightly longer head (head length 17.8- 19.4 vs. 16.5-17.8% TL), shorter caudal peduncle (dorsal-caudal space 7.7-8.9 vs. 9.2-9.9% TL, anal-caudal space 9.0-10.9 vs. 11.3-12.8% TL), longer pectoral fin (anterior margin 10.8-11.7 vs. 9.6-10.6% TL) and longer second dorsal (length 9.7-11.3 vs. 8.8-9.5% TL) and anal (length 11.0-12.2 vs. 10.3-10.5% TL) fins.

The length of the labial furrows has been previously used to separate Indo-West Pacific species H. boesemani (long furrows, up to 5 mm in length) and H. buergeri (furrows absent or very short, <2 mm) (see e.g. Springer, 1979; Compagno 1984; Figs 10b and 11b). Halaelurus maculosus possess very short (<1.8 mm) labial furrows, similar to H. buergeri . Springer and D’Aubrey (1972) noted that the degree of development of the labial furrows in H. buergeri is variable, with some specimens lacking noticeable furrows and one specimen with furrows present on only one of the mouth corners. The labial furrows of H. maculosus are also variably developed in some paratypes and are barely visible in the holotype (Fig. 2).

The colour pattern of H. maculosus is very similar to that of H. boesemani , but differs in having a denser coverage of variable-sized dark spots, with those over the darker saddle markings often much larger (Figs 1 and 10). Halaelurus maculosus also differs from H. boesemani in a number of morphometric values and ratios: mouth length 2.8-3.6 vs. 2.2% TL, 2.0-2.8 vs. 3.7 times mouth width; lower labial furrow length 0.4- 0.9 vs. 1.6% TL, 4.0-9.6 vs. 2.0 times in eye length; interdorsal space 13.1-15.0 vs. 17.1% TL; anterior nasal flap length 1.3-1.7 vs. 2.0% TL; head width 11.1-12.2 vs. 13.6% TL,>28 vs. 7.4 times lower labial furrow length; caudal peduncle height 0.9-1.2 vs. 1.6 times its width; clasper outer length 3.6-5.0 vs. 5.7 times clasper base; claspers extending past pelvic-fin free tips by 5.8-7.1 vs. 0.9 times eye length. They also differ in the total number of vertebrae, 127-136 (usually 130-132) vs. 135-139 (usually 136-138).

Halaelurus maculosus differs from two southern African endemics, H. lineatus and H. natalensis , in having a rounded snout tip which is not upturned, compared to a knob-like, upturned snout tip in the latter two species. It differs from Halaelurus quagga from the northwestern Indian Ocean, in having a more heavily spotted colour pattern (rather than with>20 dark vertical bars without dark spots).