Centrophorus lesliei, 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 lesliei n. sp.

African Gulper Shark

( Figs 1 , 4–9 ; Table 1)

Centrophorus lusitanicus— Günther, 1870: 421 (Portugal—erroneously); Regan, 1908: 53 (Portugal—erroneously); Garman, 1913: 199 (Portugal—erroneously); Bigelow & Schroeder, 1957: 84 (in part); Cadenat, 1959: 743, Fig. 1A (West Africa); Cadenat, 1960: 1428, Figs 1–3 (West Africa); Blache et al., 1970: 36, Fig. 73 (Eastern Atlantic); Hureau & Monod, 1973: 39 ( Senegal; Portugal—erroneously); Cadenat & Blache; 1981: 56, Figs 33b, 38, 39 ( Senegal); Munoz-Chapuli & Ramos, 1989: 65, Figs 1b , 3b , 4d , 5c , 6b , 7b (West Africa).

Centrophorus cf. lusitanicus — Naylor et al., 2012: 59, Fig. 43 ( Mozambique Channel, Madagascar).

Holotype. SAMC-F 041921 , pregnant female (embryo removed) 863 mm TL, off Pebane District , Zambezia Province, Mozambique, 17°33’2” S, 38°27’1” E, 433 m depth, 15 Jun. 2015 .

Paratypes. (10 specimens) BMNH 1867.7 .23.2, juvenile male 742 mm TL, locality unknown [possibly collected from Angola or Mozambique by Anchieta] ; MNHN AB-248 (dried jaw, first dorsal fin and skin patch), female 950 mm TL, MNHN AB-249 (dried jaw and dorsal fins), female 930 mm TL, Kayar , Senegal, 14°52’58.8” N, 17°10’1.2” W, 25 Mar. 1958 ; MNHN AB-250 (dried jaw and skin patch), male 735 mm TL, off Côte d’Ivoire , Feb. 1960 ; MNHN 1969-0225 , juvenile male 408 mm TL, Togo , 5°57’ N, 1°34’ E, 340 m depth, 2 Oct. 1964 ; MNHN 1969-0276 , female 524 mm TL, west of Bioko Island , Equatorial Guinea, 3°45’ N, 8°22’ E, 425 m depth, 2 Nov. 1963 ; SAMC 33320 (1 of 2), late-term embryo 317 mm TL, taken from holotype; SAMC 33320 (2 of 2), female 907 mm TL, collected with holotype ; SAMC 33321 , female 942 mm TL, off Angoche Island , Nampula Province, Mozambique, 16°19’59” S, 40°7’59” E, 500 m depth, 16 Jun. 2015 ; ZMB 6455 , female 644 mm TL, locality unknown [possibly collected from Angola or Mozambique by Anchieta], donated by Bocage (as C. granulosus ).

Genetic material (specimens not retained). (5 samples) Field code GA8 (tissue accession GN 2011), female 920 mm TL, St. Augustin, Madagascar, Mozambique Channel, 16 Sep. 1999; Field code GA11 (tissue accession GN 2012), male 750 mm TL, St. Augustin, Madagascar, Mozambique Channel, 16 Sep. 1999; Field code GA12 (tissue accession GN 2013), male 740 mm TL, St. Augustin, Madagascar, Mozambique Channel, 16 Sep. 1999; Field code MG2 (tissue accession GN 1218), male 770 mm TL, south of Toliara, Madagascar, Mozambique Channel, 17 Jan. 1997; Field code MG5 (tissue accession GN 1221), female 900 mm TL, south of Toliara, Madagascar, Mozambique Channel, 17 Jan. 1997.

Diagnosis. A medium sized (<1 m maximum total length) species of Centrophorus with the following combination of characters: body relatively slender; head moderately long (21.4–26.4% TL); snout relatively short (horizontal preorbital length 6.5–8.2% TL) and rounded in dorsal view; first dorsal very long based (base length 16.8–19.7% TL, soft fin length 16.0–17.4% TL) and high (height 6.5–7.8% TL), inner margin relatively short (5.7– 7.4% TL, 2.3–2.9 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 11.9–13.4% TL), free rear tip elongate in larger individuals (2.4–4.8% 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 119–122; teeth 33–42/29–31.

Description. Body fusiform, relatively slender, nape slightly humped; deepest at mid first dorsal-fin base, trunk height 1.18 (1.06–1.30 in paratypes <525 mm TL; 1.16–1.27 in paratypes> 640 mm TL) times width, 0.97 (0.84–0.96; 0.97–1.06) times abdomen height; no lateral ridges; pre-first dorsal length 3.84 (3.45–3.87; 3.59–3.86) in TL; interdorsal space 1.19 (1.24–1.68; 1.11–1.27) in prepectoral length, 1.46 (1.53–1.93; 1.40–1.56) in pre-first dorsal length; pelvic–caudal space 3.19 (2.22–2.42; 2.72–3.19) in pectoral–pelvic space, 1.77 (1.79–2.03; 1.74– 1.81) in prepectoral length; dorsal–caudal space 2.35 (1.89–2.00; 2.15–2.57) in interdorsal space. Caudal peduncle moderately short and deep, moderately compressed, its length 12.1 (12.2–12.7; 11.7–12.7)% TL, its height 1.53 (1.67–2.38; 1.49–1.94) times its width; tapering slightly towards caudal fin; no lateral keels; precaudal pits absent.

Head moderately long, moderately broad, width 1.14 (1.17–1.46; 1.18–1.37) times trunk width, 1.16 (1.14– 1.79; 1.28–1.85) times abdomen width, length 22.0 (22.7–26.4; 21.4–23.0)% TL, 2.83 (2.21–2.58; 2.68–2.93) in pre-vent length, height 0.89 (0.69–0.93; 0.78–0.90) 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 about 20 folds present.

Snout moderately long, narrowly triangular in lateral view, apex bluntly pointed; lateral prenarial margin rounded; rounded in dorsal view; horizontal length 0.94 (1.02–1.06; 1.05–1.25) times eye length, 0.67 (0.66–0.95; 0.84–0.88) times interorbital space; horizontal prenarial length 1.70 (1.63–2.13; 1.82–2.23) times in preoral length. Nostrils small, slightly oblique; anterior nasal flap with a large, narrowly triangular lobe, with a very small lobe at inner corner of large lobe; internarial space 2.48 (2.59–3.31; 2.48–2.81) in preoral length, 2.37 (1.63–1.74; 1.39– 1.50) times nostril length. Eye moderately large, elongate, length 3.84 (3.70–3.84; 3.61–4.36) in head, 3.18 (3.14– 3.87; 3.02–3.86) 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 above level of upper eye, slightly more than its diameter away from eye; no lobe-like fold on posterior margin; greatest diameter 3.48 (2.84–4.28; 2.80–3.81) in eye length. Gill slits directed slightly anteroventrally from top to bottom; relatively equal in size, becoming progressively longer from first to fifth; fifth longest, its height 3.1 (2.5–3.0; 2.6–3.0)% TL.

Mouth almost transverse, upper jaw slightly concave, width 1.15 (1.32–1.50; 1.06–1.21) in preoral length; lower labial furrows slightly shorter than upper furrows; prominent postoral groove, usually 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 MNHN AB-248, AB- 249 and AB-250) moderately large, with erect cusps, becoming slightly oblique posteriorly towards mouth corners, bases slightly overlapping ( Fig. 8a ). Lower teeth much larger than uppers, cusps very strongly oblique, blade-like, overlapping, edges with fine serrations ( Fig. 8b ).

Dermal denticles on flank below first dorsal fin varying greatly in shape between juveniles and adults; absent from insertions of fins. Denticles of a near-term embryo (317 mm TL) small, upright, slender, unicuspid, slightly overlapping. Denticles of larger specimens (based on a 742 mm TL specimen) block-like, sessile (not raised on pedicels), close set but rarely overlapping; anterior edges of crowns shallowly scalloped, posterior edge moderately or bluntly pointed ( Fig. 9 ). Denticles of adult holotype (863 mm TL) similar to 742 mm TL specimen but with less pointed and more rhomboidal crowns.

First dorsal fin very long, relatively high; length 3.81 (3.21–3.76; 3.11–3.86) times its height, 1.69 (1.55–1.65; 1.59–1.74) times second dorsal-fin length; soft-fin length 2.43 (2.13–2.43; 2.24–2.49) times its height; height 1.23 (1.05–1.24; 1.12–1.28) times second dorsal-fin height; anterior margin moderately convex; apex moderately rounded; posterior margin slightly concave, slanting well posteroventrally from top to bottom; free rear tip moderately long, relatively thick basally; inner margin nearly straight, 2.27 (2.49–2.85; 2.37–2.57) in soft-fin length, 1.07 (0.81–0.97; 0.91–0.97) 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 17.0 (13.0–15.2; 15.4–17.6)% 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, strongly tapering distally, anterior margin slightly convex; exposed portion of spine sloping posterodorsally from base (of exposed portion) to apex, shorter in length to exposed portion of second dorsal-fin spine, exposed first dorsal spine length 0.28 (0.24– 0.32; 0.28–0.38) times height of fin.

Second dorsal fin moderately large, much smaller in area compared to first dorsal fin, relatively short; second dorsal-fin length 2.76 (2.35–2.84; 2.06–2.84) times its height; anterior margin slightly to moderately convex; apex narrowly rounded; posterior margin weakly concave, sloping strongly posteroventrally from apex; free rear tip moderately long, thick basally, inner margin length 1.96 (1.78–2.02; 2.00–2.84) in soft-fin length, 0.85 (0.67–0.88; 0.64–0.81) times fin height; spine length 0.30 (0.38–0.53; 0.35–0.47) in height of fin; base of exposed fin spine level with anterior part of pelvic-fin inner margin, exposed just above level of junction with spine and soft portion of fin; exposed fin spine relatively long, robust, broad based, strongly tapering distally, recurved.

Pectoral fins large; anterior margin weakly convex, its length 13.0 (12.1–13.1; 11.9–13.4)% TL; base very short, 2.65 (2.24–2.84; 2.44–3.09) in anterior margin length; apex moderately rounded, not falcate; posterior margin almost straight to weakly convex from apex angle of free rear tip then broadly concave; inner margin weakly convex anteriorly and weakly concave posteriorly; free rear tip moderately elongate in adults (less produced in juveniles and embryos), free rear tip 1.13 (0.79–1.24; 1.10–1.23) in inner margin, extending to 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 11.7 (10.4–10.9; 10.7–12.2)% TL, 1.24 (1.15–1.31; 1.21–1.35) times second dorsal-fin soft length; anterior margin nearly straight; apex moderately rounded; posterior margin weakly concave; free rear tip acutely pointed. Claspers of adult males not available for examination.

Caudal fin relatively long, deep, broad; dorsal margin almost straight to very slightly concave, 1.23 (1.07–1.24; 1.12–1.29) in head length, 1.33 (1.43–1.54; 1.35–1.49) times preventral margin; preventral margin slightly convex, apex moderately rounded to subangular; upper postventral margin nearly straight, lower postventral margin slightly convex, angle between postventral margins moderately concave; terminal lobe moderately large but short, lobe length 2.15 (2.20–2.64; 2.08–2.31) in dorsal caudal margin, terminal margin slightly undulating; apex of upper lobe narrowly rounded.

Meristic data. Total vertebral centra 119–122 (based on 3 paratypes), monospondylous precaudal centra 59–60 (n = 3), diplospondylous precaudal centra 28–29 (n = 2), total precaudal centra 88 (n = 2) and diplospondylous caudal centra ~31–32 (n = 2). Tooth count (n = 6): 17–22 + 16–20 / 14–16 + 14–16; total 33–42/29–31.

Holotype Paratypes Paratypes

(<524 mm TL) (> 644 mm TL)

Min. Max. Min. Max. Holotype Paratypes Paratypes

(<524 mm TL) (> 644 mm TL) ......continued on the next page Holotype Paratypes Paratypes

(<524 mm TL) (> 644 mm TL) Colour. In preservative: Dorsal and lateral surfaces medium brownish to greyish brown; ventral surfaces slightly paler; waterline between dorsal and ventral colour shades very diffuse on lower sides. Fins without distinct markings in preserved specimens examined (no fresh material observed); dorsal fins and ventral caudal lobe of near-term embryo blackish distally, with upper caudal lobe with a broad whitish margin with a dusky marking anteriorly.

Size. Type specimens ranged in size from 317 to 950 mm TL. The smallest free-swimming individual was 408 mm TL; a late-term embryo of 317 mm TL still possessed a small external yolk sac; Cadenat (1960) recorded an embryo of 395 mm TL from a pregnant female. A female of 863 mm TL was mature and pregnant. Cadenat (1960) reported adult males between 715 and 765 mm TL and adult females between 890 and 990 mm TL.

Distribution. Type specimens were from the Mozambique Channel ( Madagascar and Mozambique), and off Senegal, Côte d’Ivoire, Equatorial Guinea, and Togo at depths of 340– 500 m. The BMNH and ZMB specimens were most likely collected by Anchieta from Angola or Mozambique in the 1800’s. Additional specimens examined by Munoz-Chapuli & Ramos (1989) were recorded as being collected from Morocco, the Canary Islands, and Ghana at depths of 370– 610 m.

Etymology. Named after Dr Robin Leslie (Fisheries Branch, Department of Agriculture, Forestry and Fisheries in South Africa) who has contributed greatly to our knowledge of southern African chondrichthyans and provided numerous important specimens and tissue samples for various projects.

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 3. Excerpt from the specimen register showing the batch of three sharks, including the possible syntype of Centrophorus lusitanicus (as C granulosus) BMNH 1867.7. 23.2, donated by Bocage. Note the lack of ditto marks beneath Portugal for Centrophorus granulosus and no mention of it being a type.

FIGURE 4. Lateral view of ZMB 6455 (female 644 mm TL), a second specimen donated from the Lisbon Museum by Bocage as Centrophorus granulosus (now a paratype of Centrophorus lesliei n. sp.).

FIGURE 5. Lateral view of Centrophorus lesliei n. sp.: (A) pregnant female holotype (SAMC-F 041921, 863 mm TL); (B) lateterm embryo paratype (SAMC 33320, 317 mm TL, taken from holotype).

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

FIGURE 7. Ventral view of pectoral fin of Centrophorus lesliei n. sp.: paratype BMNH 1867.7. 23.2, juvenile male 742 mm TL.

FIGURE 8. Teeth of adult female paratype of Centrophorus lesliei n. sp. (MNHN AB- 249, 930 mm TL): (A) upper; (B) lower.

FIGURE 9. Lateral trunk denticles (from below first dorsal fin) of Centrophorus lesliei n. sp.: paratype BMNH 1867.7. 23.2, juvenile male 742 mm TL.