Centrophorus longipinnis, White & Ebert & Naylor, 2017

White, William T., Ebert, David A. & Naylor, Gavin J. P., 2017, Revision of the genus Centrophorus (Squaliformes: Centrophoridae): Part 2 - Description of two new species of Centrophorus and clarification of the status of Centrophorus lusitanicus Barbosa du Bocage & de Brito Capello, 1864, Zootaxa 4344 (1) : -

Centrophorus longipinnis n. sp.

Longfin Gulper Shark

( Figs 11–15 ; Table 2)

Centrophorus lusitanicus — Teng, 1958: 25, fig. 16 (Yilan and Kaohsiung, Taiwan) ; Teng, 1962: 155, fig. 39 (Tashi); Chen, 1963: 92 ( Taiwan) ; Yang, 1979: 205, Fig. 2 ( Taiwan) ; Chen & Cheng, 1982: 143, Fig. 1 ( Taiwan) ; Chen & Yu, 1986: 112 ( Taiwan) ; Compagno et al., 2005: 52 ( Philippines); Hsu & Joung, 2004: 184 ( Taiwan) ; Shen & Wu, 2011: 82, Fig. ( Taiwan) .

Centrophorus niaukang — Chen & Joung 1993: pl. 2 (fig. 1) (Taiwan).

Centrophorus cf. lusitanicus— White et al., 2006: 50, Fig. (Indonesia); White & Dharmadi, 2010: 1364, Fig. 6a, b (Indonesia); Ebert et al., 2013: 289 (Taiwan).

Centrophorus sp. 2— Naylor et al., 2012: 59, Fig. 43 (Taiwan).

Centrophorus sp. 3— Naylor et al., 2012: 59, Fig. 43 ( Philippines).

Holotype. NMMB-P 15756 (tissue accession GN10189), adult male 720 mm TL, Cheng-gong , Taiwan, 30 Jul. 2011.

Paratypes. (15 specimens) CSIRO H 5788-02 (tissue accession GN11178), female 899 mm TL, Tanjung Luar fish market, Lombok, Indonesia, 11 Apr. 2001; CSIRO H 8104-01 (tissue accession GN11174), female 855 mm TL, CSIRO H 8104-02 (tissue accession GN11175), adult male 679 mm TL, CSIRO H 8104-04 (tissue accession GN11177), female 872 mm TL, Tanjung Luar fish market, Lombok, Indonesia, 19 Aug. 2005; CSIRO H 7990-02, female 825 mm TL, Pelabuhanratu fish market, West Java, Indonesia, 10 Mar. 2009; CSIRO H 8103-01, pregnant female 890 mm TL, Huon Gulf, off Lae, Papua New Guinea, 6°45.147’ S, 147°2.783’ E, 460 m depth, 4 May 2017; CSIRO H 8103-02, late-term embryo 346 mm TL, taken from CSIRO H 8103-01; CSIRO H 8171-01, adult male 761 mm TL, Da-xi, Taiwan, 14 Apr. 2012; FRIP 0 3628 (1 of 2), adult male 719 mm TL, FRIP 0 3628 (2 of 2), juvenile male 637 mm TL, Taiwan, 11 Mar. 1958; NMMB-P 25361, 736 mm TL, adult male, Da-xi, Taiwan, 20 Mar. 2013; NMMB-P 14051, juvenile female 408 mm TL, Cheng-gong, Taiwan, 2 Oct. 2011; NMMB-P 15813, female 905 mm TL, Cheng-gong, Taiwan, 2 Oct. 2011; NMMB-P 15814 (tissue accession GN10190), adult male 745 mm TL, Cheng-gong, Taiwan, 18 Jul. 2011; NMMB-P 15859, adult male 761 mm TL, Cheng-gong, Taiwan, 29 Mar. 2011; BMNH 2017.8.29.1 (to be donated to BMNH), adult male 776 mm TL, Da-xi, Taiwan, 14 Apr. 2012.

Other specimens. (1 specimen) CSIRO H 8104-03 (tissue accession GN11176), adult male 764 mm TL, Tanjung Luar fish market, Lombok, Indonesia, 19 Aug. 2005.

Genetic material (specimens not retained). (3 samples) Tissue accession GN973, Da-xi, Taiwan; Tissue accession GN974, Da-xi, Taiwan; Tissue accession GN1007, Da-xi, Taiwan.

Diagnosis. A medium sized (<1 m maximum total length) species of Centrophorus with the following combination of characters: body relatively slender; head moderately long (20.5–25.1% TL); snout relatively short (horizontal preorbital length 6.4–8.4% TL) and rounded in dorsal view; first dorsal extremely long based (base length 20.8–23.3% TL, soft fin length 16.9–20.2% TL) and relatively high (height 5.7–7.2% TL), inner margin relatively short (5.3–6.8% TL, 2.6–3.2 in soft fin length); second dorsal fin much smaller in area to first, similar in height to first dorsal fin (height 1.0– 1.3 in first dorsal-fin height); pectoral fins large (anterior margin length 11.0– 13.0% TL), free rear tip elongate in larger individuals (2.2–4.0% TL); lateral trunk denticles of larger individuals sessile (not raised on pedicels), block-like, not elevated; upper teeth of larger individuals with erect to slightly oblique cusps; lower teeth of all sized specimens much larger than upper teeth, strongly oblique, blade-like; total vertebral centra 112–122; teeth 38–43/29–31.

Description. Body fusiform, relatively slender, nape only slightly humped; deepest near first dorsal-fin spine, trunk height 1.30 (0.91–1.34 in paratypes> 700 mm TL) times width, 0.96 (0.82–1.05 in paratypes> 700 mm TL) times abdomen height; no lateral ridges; interdorsal ridge absent; pre-first dorsal length 4.23 (4.12 in 408 mm TL paratype; 3.62–4.26 in paratypes> 700 mm TL) in TL; interdorsal space 1.28 (1.61; 1.03–1.38) in prepectoral length, 1.45 (1.69; 1.28–1.62) in pre-first dorsal length; pelvic–caudal space 3.28 (2.60; 2.70–3.50) in pectoral– pelvic space, 1.82 (1.87; 1.62–2.00) in prepectoral length; dorsal–caudal space 2.05 (1.81; 2.09–2.65) in interdorsal space. Caudal peduncle moderately short and deep, moderately compressed, its length 11.5 (12.4; 10.7–12.8)% TL, its height 1.47 (2.11; 1.61–2.15) times its width; tapering slightly towards caudal fin; ventral midline with a groove; dorsal midline with a weak ridge anteriorly becoming a weak groove posteriorly; no lateral keels; precaudal pits absent.

Head moderately long, moderately broad, width 1.72 (1.44; 1.04–1.73) times trunk width, 1.75 (1.85; 1.18– 1.81) times abdomen width, length 21.1 (24.3; 20.5–22.4)% TL, 2.96 (2.47; 2.76–3.09) in pre-vent length, height 0.67 (0.75; 0.65–0.96) times width; slightly depressed forward of spiracles, somewhat broadly pear-shaped in cross-section at pectoral-fin origin. Band of transverse dermal folds on ventral surface of head broadly rounded with apex about three quarters of horizontal prenasal length behind symphysis of lower jaw, extending from below lower edges of first four gill slits on either side; up to 19 folds present.

Snout moderately long, narrowly triangular in lateral view, apex bluntly pointed; lateral prenarial margin rounded; rounded in dorsal view; horizontal length 1.24 (1.28; 1.07–1.29) times eye length, 0.78 (0.96; 0.68–0.84) times interorbital space; horizontal prenarial length 1.44 (1.43; 1.40–1.57) times in preoral length. Nostrils small, slightly oblique; anterior nasal flap with a large, narrowly triangular lobe, with a very small, sometimes pale lobe at inner corner of large lobe; internarial space 2.80 (2.67; 2.43–2.95) in preoral length, 1.77 (1.85; 1.53–2.05) times nostril length. Eye moderately large, elongate, length 4.04 (3.98; 3.96–4.66) in head, 2.46 (3.30; 1.94–3.81) times height; notched anteriorly; strongly notched posteriorly, notch not extending towards spiracle. Spiracle moderately large, semicircular; located dorsolaterally on head, entirely visible in dorsal view; lower margin about level with upper eye, slightly less than its diameter away from eye; no lobe-like fold on posterior margin; greatest diameter 3.05 (3.19; 2.57–3.83) in eye length. Gill slits directed anteroventrally from top to bottom, fifth angled more than first; first shortest then becoming progressively longer to fifth; fifth longest, its height 2.6 (3.2; 2.4–3.5)% TL.

Mouth almost transverse, upper jaw slightly concave, width 1.22 (1.29; 1.06–1.22) in preoral length; lower labial furrows slightly longer than upper furrows; prominent postoral groove, more than twice length of upper labial furrows, extending posterolaterally from angle of jaws. Teeth strongly differentiated in upper and lower jaws, with upper teeth much smaller than lower teeth. Upper teeth of adults (based on CSIRO H 7990-02) moderately large, with erect cusps, becoming slightly oblique posteriorly towards mouth corners, bases slightly overlapping ( Fig. 13a ). Lower teeth much larger than uppers, cusps very strongly oblique, blade-like, overlapping, edges with fine serrations ( Fig. 13b ).

Dermal denticles on flank below first dorsal fin varying greatly in shape between juveniles and adults; absent from insertions of fins and most of the dorsal surface of claspers. Denticles of a juvenile (408 mm TL) small, upright, slender, unicuspid, closely spaced (slightly overlapping), and posteriorly curved; about 0.4 mm long ( Fig. 14a ). Denticles of adults (based on a 855 mm TL specimen) block-like, sessile (not raised on pedicels), rhomboidal, close set but not overlapping; anterior edges of crowns shallowly scalloped, posterior edge bluntly pointed ( Fig. 14b ).

First dorsal fin extremely long, moderately high; length 4.18 (4.18; 3.37–5.03) times its height, 1.76 (1.77; 1.76–1.98) times second dorsal-fin length; soft-fin length 2.86 (2.55; 2.48–3.47) times its height; height 1.14 (1.13; 0.99–1.25) times second dorsal-fin height; anterior margin slightly convex; apex narrowly rounded; posterior margin moderately concave, slanting well posteroventrally from top to bottom; free rear tip moderately long, relatively thick basally; inner margin nearly straight, 2.88 (2.94; 2.60–3.23) in soft-fin length, 1.00 (0.87; 0.82– 1.20) times its height; insertion level with mid pectoral–pelvic space, extremely well forward of pelvic-fin origin, pelvic-fin midpoint to first dorsal-fin insertion 15.1 (13.1; 13.2–17.6)% TL; base of exposed fin spine level with pectoral-fin free rear tip; spine base moderately broad, exposed anteriorly just above junction of spine and soft portion of fin; exposed fin spine relatively short, robust, tapering distally, anterior margin slightly convex; exposed portion of spine sloping strongly posterodorsally from base (of exposed portion) to apex, shorter in length to exposed portion of second dorsal-fin spine, exposed first dorsal spine length 0.39 (0.30; 0.23–0.39) times height of fin.

Second dorsal fin moderately large, much smaller in area compared to first dorsal fin, relatively short; second dorsal-fin length 2.69 (2.66; 2.25–2.84) times its height; anterior margin slightly convex; apex narrowly rounded; posterior margin weakly concave, sloping strongly posteroventrally from apex; free rear tip relatively short, thick basally, inner margin length 2.09 (2.05; 1.83–2.40) in soft-fin length, 0.78 (0.81; 0.63–0.83) times fin height; spine length 0.42 (0.32–0.50 in paratypes> 700 mm TL) in height of fin; base of exposed fin spine level with mid pelvicfin inner margin, exposed just above level of junction with spine and soft portion of fin; exposed fin spine relatively long, robust, broad based, tapering distally.

Pectoral fins large; anterior margin weakly convex to nearly straight, its length 12.1 (12.3; 12.0–13.0)% TL; base very short, 2.68 (2.77; 2.26–2.99) in anterior margin length; apex somewhat angular, not falcate; posterior margin nearly straight from apex angle of free rear tip then broadly concave; inner margin slightly concave; free rear tip elongate in adults (less so in juveniles), free rear tip 1.18 (1.04; 1.01–1.14) in inner margin, extending past level of exposed first dorsal-fin spine; origin situated at level of mid-fifth gill slit, partially obscured by gill membrane.

Pelvic fins large, length 11.6 (11.4; 11.0–12.4)% TL, 1.20 (1.14; 1.18–1.44) times second dorsal-fin soft length; anterior margin slightly convex; apex narrowly rounded; posterior margin slightly concave; free rear tip acutely pointed, inner margin very slightly convex. Claspers of adult males relatively short, slender; tapering to a fleshy, narrowly rounded tip ( Fig. 15 ); outer length 2.9 (2.5–3.3)% TL, 3.16 (2.98–3.81) times its base length (n = 8); clasper glans about 0.3 in clasper inner length; apopyle and hypopyle connected by long clasper groove; rhipidion moderately large, laterally expanded, extended from hypopyle to anterior of clasper tip; lateral edge with a slender, straight spine; dermal denticles mostly absent from dorsal surface.

Caudal fin relatively long, deep, broad; dorsal margin almost straight to slightly concave, 1.17 (1.18; 1.11– 1.22) in head length, 1.42 (1.70; 1.32–1.51) times preventral margin; preventral margin slightly convex (more so distally), apex narrowly rounded; upper postventral margin slightly convex, lower postventral margin nearly straight to slightly convex, angle between postventral margins moderately concave; terminal lobe moderately large but short, lobe length 2.31 (2.50; 2.05–2.63) in dorsal caudal margin, terminal margin slightly to moderately convex and slightly undulating; apex of upper lobe narrowly rounded.

Meristic data. Total vertebral centra 118 (112–122), monospondylous precaudal centra 59 (56–62), diplospondylous precaudal centra 28 (25–29), total precaudal centra 87 (83–91) and diplospondylous caudal centra 31 (26–34). Tooth count (n = 5): 20 (19–22) + (21) 19–21 / 15 (15) + 14 (15–16); total 41 (38–43)/29 (30–31).

Colour. When fresh: Dorsal and lateral surfaces brownish, sometimes with a reddish hue (grey in some specimens, particularly smaller specimens); ventral surfaces paler (only slightly paler in paratype CSIRO H 7990- 02); waterline between dorsal and ventral colour shades diffuse and poorly defined on body, more distinct on head and caudal peduncle. Fins without markings in larger specimens; tip of free tip of first dorsal fin often paler; nearterm embryo with blackish dorsal and caudal fins and black anterior margins to paired fins, with narrow white posterior margin to dorsal and paired fins. Similar colour in preservation; fin markings of embryo less distinct.

Size. Postnatal type specimens ranged from 408 to 905 mm TL; a 637 mm TL male was immature; males adult between 719 and 776 mm TL; an 890 mm TL female was pregnant with a 346 mm TL late-term embryo (both paratypes). White & Dharmadi (2010) recorded: females up to 930 mm TL; an 873 mm TL pregnant female with a single 45 mm TL early-term embryo; males adult between 679 and 775 mm TL, while a 657 mm TL male was immature. Compagno et al. (2005) reported on a 737 mm TL immature male.

Distribution. Type material from off Taiwan (Cheng-gong and Da-xi fish landing sites—local fishing grounds), Indonesia (off southwest Java and eastern Lombok), and Papua New Guinea (Huon Gulf) ( Fig. 10 ). Compagno et al. (2005) reported this species off Puerto Princesa City in the Philippines (JPAG 226, tissue accession GN4348); differed slightly in ND2 sequence and specimen not examined in this study. Limited depth information available as most specimens collected from fish landing sites; caught from depths of 330–460 m in Papua New Guinea (P. Neira, pers. comm.).

Etymology. Specific name a combination of the Latin longus (long) and pinna (fin) in allusion to the very distinctive long-based first dorsal fin this species possesses.

Holotype Paratypes Paratypes

(<524 mm TL) (> 644 mm TL) ......continued on the next page ......continued on the next page The Maximum Likelihood tree generated from the ND2 sequences obtained provide further support for the separation of C. lesliei and C. longipinnis as valid species ( Fig. 16 ). Centrophorus lesliei samples form a distinct group nested within, but separate from, the three long-snout species groups (i.e. isodon-tesselatus-westraliensis, harrissoni , and isodon-tesselatus). The long-snout species group requires further attention and will be dealt with in a subsequent part of this revision series for this genus. It should be noted that none of the sequenced specimens from Madagascar are type species as whole specimens were not retained, however, they were obtained from the Mozambique Channel, close to the locality of the holotype and three paratypes of C. lesliei . Centrophorus longipinnis forms a distinct group well separated from the long-snout and C. lesliei groups. Within the C. longipinnis group, two samples (GN 11175 from Indonesia and GN 4348 from the Philippines) show some differences to the main group ( Fig. 16 ). But morphologically, the Indonesian specimen is identical to the other C. longipinnis specimens and is considered to be this species.

Moura et al. (2015) concluded that the BMNH specimen was C. uyato based on DNA sequence comparisons for CO1. This result is inconsistent with the findings we present herein. Morphological data suggest that the BMNH specimen is C. lesliei and furthermore, is very distinct from C. uyato in possessing a much longer first dorsal-fin base. Munoz-Chapuli & Ramos (1989) recorded vastly different first dorsal-fin ray counts between C. lusitanicus (= C. lesliei ) and C. granulosus (= C. uyato ), i.e. 7–9 vs. 16–19, reflecting the large difference in length of the first dorsal fin. Thus, it is not conceivable that these two species could be considered conspecific, or that the BMNH specimen is a long finned variant of C. uyato . Moura et al. (2015) state in their acknowledgements that they obtained a tissue sample from the BMNH specimen and also from ‘the Aquário Vasco da Gama Museum, and Aldina Inácio …’ and state at the end of the sentence state, in parentheses, that ‘DNA extraction was unsuccessful’. One possibility is that the BMNH specimen was the one which DNA extraction was not successful on and that a sample from the other source(s) was C. uyato . A tissue sample from the BMNH specimen was also collected as part of a broader Chondrichthyan Tree of Life project led by one of us (GN) but despite numerous attempts, no valid DNA sequence could be obtained.

Figures

FIGURE 1. Lateral view of BMNH 1867.7. 23.2 (juvenile male 742 mm TL), previously considered to be a possible syntype of Centrophorus lusitanicus (now a paratype of Centrophorus lesliei n. sp.).

FIGURE 2. Illustration of Centrophorus lusitanicus in the original description (Barbosa du Bocage & Brito Capello, 1864).

FIGURE 6. Ventral view of head of the pregnant female holotype of Centrophorus lesliei n. sp. (SAMC-F 041921, 863 mm TL).

FIGURE 10. Map showing the collection locations, where known, of specimens of Centrophorus lesliei n. sp. (yellow and green) and Centrophorus longipinnis n. sp. (cyan and pink). Stars denote the holotypes, circles denote paratypes, and triangles denote non-type specimens (Image © NASA, TerraMetrics, Google Earth).

FIGURE 11. Lateral view of Centrophorus longipinnis n. sp.: (A) adult male holotype (NMMB-P 15756, 720 mm TL); (B) female paratype (CSIRO H 8104 - 01, 855 mm TL); (C) female paratype (CSIRO H 7990 - 02, 825 mm TL); (D) late-term embryo paratype (CSIRO H 8103 - 02, 346 mm TL).

FIGURE 12. Ventral view of head and pectoral fins of Centrophorus longipinnis n. sp.: female paratype (CSIRO H 7990 - 02, 825 mm TL).

FIGURE 13. Upper and lower teeth of female paratype of Centrophorus longipinnis n. sp. (CSIRO H 7990 - 02, 825 mm TL).

FIGURE 14. Lateral trunk denticles (from below first dorsal fin) of Centrophorus longipinnis n. sp.: (A) juvenile female paratype, NMMB-P 14051, 408 mm TL; (B) female paratype, CSIRO H 8104 - 01, 855 mm TL.

FIGURE 15. Clasper (right) of Centrophorus longipinnis (paratype CSIRO H 8104 – 02, adult male 679 mm TL). A, Glans not dilated; B, Glans spread. Abbreviations: AP, apopyle; CG, clasper groove; CS, clasper spine; HP, hypopyle; P 2, pelvic fin; RH, rhipidion.

FIGURE 16. Maximum Likelihood tree estimated under the General Time Reversible model (GTR) with model terms to accommodate both Invariant site (I) and Gamma Distributed rates (G). Bootstrap support values are shown from a separate ML bootstrap analysis. Sequences used in this tree are part of the Chondrichthyan Tree of Life project (http: // sharksrays. org /).