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BULLETIN OF MARINE SCIENCE. 87(3):501–512. 2011 NEW TAXA PAPER http://dx.doi.org/10.5343/bms.2010.1108

Pristiophorus NANCyAE, A New Species of (: Pristiophoridae) from southern Africa

David A Ebert and Gregor M Cailliet

ABSTRACT A new species of sawshark, nancyae sp. nov., is described from eight specimens collected off Mozambique. The new species is the second member of the family Pristiophoridae and first member of the Pristiophorus described from the western Indian Ocean. The new Pristiophorus species can be distinguished from the sympatric occurring six-gilled sawshark, Pliotrema warreni Regan, 1906, most notably by having five paired gill openings as opposed to six. The new species is distinguished from all other Pristiophorus species by several distinctive characteristics. Most notably, the new species differs by having a very distinctive double row of four to five conspicuous large pits anterior to the nasal barbels on the underside of its snout. Other distinguishing characteristics include a broad, triangular first with a rear tip that extends behind the pelvic midbases, barbels much closer to mouth than snout tip, two rows of enlarged pits on the underside of the pre- , ridges on the base of its large lateral rostral teeth, mostly tricuspidate, flat, imbricated lateral trunk denticles, and plain color pattern. The new species is compared to the five other known Pristiophorus species and a revised key to the genus is presented.

The P( ristiophoriformes: Pristiophoridae) are a small elasmobranch group with a single family, two genera, and five or more species C( ompagno et al. 2005). This is an unmistakable order, easily distinguishable by its elongated and flattened rostrum edged with slender sharp lateral teeth and a long pair of rostral barbels in front of the nostrils. These have two dorsal fins, no anal fin, a short transverse mouth, and small cuspidate teeth in both jaws. Most species occur in the Indo-West Pacific, except for Pristiophorus schroederi Springer and Bullis, 1960, found in the western North Atlantic. Furthermore, except for Pliotrema warreni Regan, 1906, no other described pristiophorids are known from the western Indian Ocean. Compagno et al. (2005) cite an undescribed species, Pristiophorus sp. D, from the western Indian Ocean and give its range as Mozambique, and possibly Somalia to the Arabian Sea off Pakistan. In June 1994, South Africa’s Marine and Coastal Management R/V Algoa con- ducted an extensive bottom trawl survey for horse mackerel (Trachurus spp.) be- tween 37 and 517 m along the Mozambique coast. Two species of sawsharks were collected, P. warreni and a dwarf Pristiophorus (= Pristiophorus sp. D of Compagno et al. 2005), on the upper continental slope at depths of 286–517 m. Eight specimens of the new Pristiophorus species and three P. warreni were saved and deposited in the collections of the South African Museum. These Mozambique Pristiophorus speci- mens possess several very distinctive morphological characteristics that separate it from the five other known species of this genus. Here we describe this new western Indian Ocean Pristiophorus species.

Bulletin of Marine Science 501 © 2011 Rosenstiel School of Marine and Atmospheric Science of the University of Miami 502 BULLETIN OF MARINE SCIENCE. VOL 87, NO 3. 2011

Methods

Morphometric measurements follow a modification of Compagno (2001) and Yearsley et al. (2008). Meristics, including tooth, spiral valve, pectoral fin radial, and vertebral counts, were taken for all specimens. Morphometric values are presented as ratios for the holotype followed by a range of paratype values in parentheses. Comparative material was obtained from collections at the California Academy of Sciences (CAS), Australian National Fish Collection, Hobart, Australia (CSIRO), LJV Compagno field number (LJVC), Museum Comparative Zoology (MCZ), Stanford University (SU: now housed at the CAS), South African Museum (SAM), United States National Museum (USNM), and the British Museum Natural History (BMNH). Institutional abbreviations follow Sabaj Pérez (2010) and are listed at http://www.asih.org/node/204.

Pristiophorus nancyae new species [New English name: African dwarf sawshark] (Figs. 1–3, Tables 1, 2)

Pristiophorus sp. D: Compagno et al. 2005: 135, ill., pl. 16.

Holotype.—SAM 34013, 616 mm TL, mature male, R/V Algoa, Mozambique Scad Survey, bottom trawl, Station C00840 014 037 3074, 22°07´S, 35°45´E, 500 m, 19 June, 1994. Paratypes.—7 specimens: SAM 33477, 440 mm TL, maturing male taken with ho- lotype, R/V Algoa, Mozambique Scad Survey, bottom trawl, Station C00840 014 037 3074, 22°07´S, 35°45´E, 500 m, 19 June, 1994; SAM 33502, 573 mm TL, maturing female, R/V Algoa, Mozambique Scad Survey, bottom trawl, Station C00848 014 045 3179, 25°21´S, 34°30´E, 286 m, 21 June, 1994; SAM 33511 (five specimens), 314 and 358 mm TL, both immature males, 391 mm TL immature female, 522 and 550 mm TL, both mature males, R/V Algoa, Mozambique Scad Survey, bottom trawl, Station C00841 014 038 3118, 23°32´S, 35°51´E, 490 m, 20 June, 1994. Diagnosis.—A small five-gilled sawshark distinct from its congeners by the pres- ence of two rows of enlarged pre-barbel pits on the rostral ventral surface; rostral barbels closer to mouth than snout tip; base of large rostral teeth with ridges. Eyes very large, oval-shaped, emerald green in life. First dorsal fin broadly triangular, with rear tip extending behind pelvic fin midbases. Lateral trunk denticles imbricated. Coloration in life brownish gray laterally, becoming lighter ventrally; fin margins lighter along posterior edges; dorsal rostrum surface with two distinct longitudinal dark stripes; lateral rostral teeth dark-edged. Description.—Body slender, snout flattened, forming very elongated, blade-like rostrum, pre-barbel length 17.0% (16.6%–17.5%) total length (TL). Head very nar- row, subtriangular, strongly depressed over eyes, height greatest at gill opening, head width at mouth 6.9% (6.2%–7.9%) TL; head width at nostril 4.8 (4.1–5.0) times in pre- orbital length. Trunk narrow, tapering toward caudal origin; subcircular in cross- section; pectoral-pelvic space 12.9% (12.9%–15.0%) TL. Head length 0.5 (0.5) times trunk length; pectoral-pelvic length short 12.9% (12.9%–15.0%) TL; dorsal-caudal ebert and cailliet: new from southern africa 503

Figure 1. Pristiophorus nancyae sp. nov., holotype, SAM 34013, 616 mm TL, mature male. space 8.4% (7.8%–9.2%) TL; caudal peduncle height 3.5 (3.4–4.1) into dorsal caudal space, and width 0.9 (0.8–1.0) in height. Lateral keels present; no precaudal pits. Rostrum is very long, narrow, and tapering; pre-oral length 28.4% (27.7%–29.6%) TL; sides of rostrum slightly convex from tip to barbels and nearly straight between barbels and nostrils. Ratio of pre-orbital length to rostrum width at nostrils 4.8 (4.1– 5.0). Two parallel rows of 4 (4–5) enlarged pits present on underside of pre-barbel snout. Barbels closer to mouth than rostral tip, pre-barbel length 59.8% (58.4%– 60.7%) of pre-oral length; space from barbel to nostril 0.8 (0.7–0.9) times space from nostril to first gill opening; barbels ending 0.4 (0.2–0.5) eye length anterior to nos- trils in adults, posterior to them by 0.3 to 0.7 of eye length in juveniles. Large lateral rostral saw-teeth along sides of snout and head total 25 (21–32), including 16 (15–22) teeth in front of barbels and 9 (6–10) teeth behind barbels. Post-narial lateral rostral teeth extending 0.4 (0.1–0.5) times eye length behind eye. Adults (> 400 mm) have up to 4 smaller rostral teeth between large lateral rostral saw-teeth; juveniles < 400 mm with 1–3. Prominent longitudinal ridges on bases of lateral and ventral rostral teeth. Ventral rostral teeth anterior to barbels number 15 (13–18), between barbels and nostril 6 (5–7). A ventral rostral tooth in front of nostril but this not enlarged. Eyes oval, very large, lateral on head, length 3.5% (3.5%–4.6%) TL; distance to last rostral tooth 1.5 (0.3–1.9); inter-orbital concave width 4.4% (4.2%–5.0%) TL; sub-ocular groove present. Spiracles small, length 3.7 (2.3–5.4) times eye length, crescent-shaped, slightly oblique; located just posterior to eye. Gill openings small, lateral on head, closer to ventral surface than dorsal; each similar in length, evenly spaced except for fourth and fifth openings slightly closer to each other; fifth gill opening arches slightly around pectoral fin origin. Nostrils small, anterior nasal flap lobe-like and well-developed; excurrent aper- tures transversely oval, nearly circular, width 0.8% (0.7%–1.0%) TL, 3.9 (3.2–4.0) in internarial width. Distance from nostrils to mouth 1.2 (1.1–1.3) times internarial space. Distance from nostrils to barbels 1.3 (1.1–1.4) times distance between nostrils and mouth corners. Mouth broadly arched, extending forward to below posterior third or fourth of eye, width 4.3% (4.0%–4.3%) TL. Labial furrows short, upper and lower about equal 504 BULLETIN OF MARINE SCIENCE. VOL 87, NO 3. 2011

Table 1. Pristiophorus nancyae total length proportions (TL%) of the holotype (SAM 34013) and seven paratypes.

Measurement Holotype Paratypes Total length (mm) 616.0 314–573 Pre-caudal length 80.0 78.3–80.7 Pre-barbel length 17.0 16.6–17.5 Pre-narial length 24.2 23.6–25.4 Pre-oral length 28.4 27.7–29.6 Pre-orbital length 25.8 25.1–26.6 Pre-spiracle length 29.4 28.8–30.9 Pre-gill length 33.4 33.2–35.0 Head length 37.0 36.4–38.5 Pre-pectoral length 37.0 36.4–38.5 Pre-pelvic length 52.9 51.6–54.1 Snout–vent length 55.7 54.1–56.4 Pre-first dorsal fin length 45.3 44.2–47.1 Pre-second dorsal fin length 64.8 62.4–65.6 Inter-dorsal fin length 12.0 9.8–11.5 Dorsal–caudal length 8.4 7.8–9.2 Pectoral–pelvic length 12.9 12.9–15.0 Pelvic–caudal length 23.1 22.0–24.8 Pectoral origin–1st dorsal origin distance 0.0 0.0–1.7 Pelvic insertion–2nd dorsal origin distance 7.2 2.6–7.1 Eye length 3.5 3.5–4.6 Eye height 1.4 1.1–1.9 Eye to last rostral tooth length 1.5 0.3–1.9 Inter-orbital length 4.2 4.2–5.0 Inter-barbel width 2.9 3.0–3.9 Barbel length 6.0 6.2–9.8 Barbel–nostril length 7.0 6.2–7.7 Barbel tip–nostril 1.3 −3.1–1.8 Nostril width 0.8 0.7–1.0 Internarial length 3.1 3.0–3.6 Anterior nasal flap length 0.6 0.6–1.1 Nostril–mouth symphysis length 3.7 3.2–4.1 Nostril–mouth rictuses (corners) length 5.5 5.0–5.6 Nostril–1st gill opening length 9.1 8.0–9.7 Spiracle length 1.0 0.7–1.6 Spiracle height 0.9 0.6–1.0 Inter-spiracular width 3.8 3.6–4.6 Eye–spiracle length 0.3 0.3–0.5 Mouth length 2.0 1.5–2.1 Mouth width 4.3 4.0–4.3 Upper labial furrow 0.2 0.2–0.4 Lower labial furrow 0.2 0.2–0.4 1st gill height 1.0 1.0–1.5 2nd gill height 1.2 1.0–1.3 3rd gill height 1.2 1.0–1.4 4th gill height 1.0 1.0–1.3 5th gill height 1.0 1.2–1.4 ebert and cailliet: new shark from southern africa 505

Table 1. Continued.

Measurement Holotype Paratypes Head height at eye 2.9 2.6–3.2 Head height at mouth 4.8 4.5–5.5 Head width at eye 6.6 6.3–7.7 Head width at mouth 6.9 6.2–7.9 Head width at nostril 5.3 5.1–6.3 Trunk height 5.1 4.8–6.0 Trunk width 5.7 5.2–6.4 Caudal peduncle height at 2nd dorsal 2.4 2.2–2.4 Caudal peduncle height at caudal origin 1.6 1.5–2.0 Caudal peduncle width at 2nd dorsal 2.7 2.4–3.0 Caudal peduncle width at caudal origin 1.8 1.6–2.0 Girth 18.8 18.7–20.7 Pectoral fin length 8.5 8.7–9.9 Pectoral fin anterior margin length 10.1 8.9–10.4 Pectoral fin base length 2.6 2.9–3.8 Pectoral fin height 7.4 6.7–8.9 Pectoral fin inner margin length 5.6 6.1–6.7 Pectoral fin posterior margin length 7.2 5.8–8.7 Pelvic fin length 8.1 7.1–8.7 Pelvic fin anterior margin length 5.2 5.0–6.2 Pelvic fin base length 3.9 3.1–4.6 Pelvic fin height 3.0 2.4–3.4 Pelvic fin inner margin length 4.4 2.8–4.9 Pelvic fin posterior margin length 4.5 3.7–4.4 1st dorsal fin length 11.5 11.2–12.5 1st dorsal fin anterior margin length 8.3 7.7–9.9 1st dorsal fin base length 7.7 7.1–8.5 1st dorsal fin height 5.5 5.3–6.2 1st dorsal fin inner margin length 4.1 4.2–4.5 1st dorsal fin posterior margin length 7.6 7.0–8.7 2nd dorsal fin length 9.4 9.5–10.7 2nd dorsal fin anterior margin length 9.0 9.7–11.1 2nd dorsal fin base length 6.6 6.3–7.8 2nd dorsal fin height 6.0 5.6–6.8 2nd dorsal fin inner margin length 2.8 2.9–3.5 2nd dorsal fin posterior margin length 7.1 5.8–6.8 Caudal fin dorsal margin 19.0 18.6–20.0 Caudal fin pre-ventral margin 6.3 5.1–8.6 Caudal fin lower–upper margin 8.0 6.1–8.6 Caudal fin subterminal margin 2.5 2.5–3.4 Caudal fin terminal margin 3.5 2.9–4.2 Caudal fin terminal lobe 4.3 2.9–5.3 1st dorsal fin midpoint–pectoral fin insertion 8.9 7.2–9.9 1st dorsal fin midpoint–pelvic fin origin 4.0 2.5–4.8 Pelvic fin midpoint–1st dorsal fin insertion 3.0 1.5–3.4 Pelvic fin midpoint–2nd dorsal fin origin 9.3 8.3–10.0 506 BULLETIN OF MARINE SCIENCE. VOL 87, NO 3. 2011 in length. Teeth with a single prominent acute cusp, in well defined rows, bases flat- tened and rounded. Tooth row counts 36 (31–35) on upper jaw and 34 (29–34) on lower jaw; series of functional teeth about 3–4. Lateral trunk dermal denticles closely imbricated, with flat mostly tricuspidate crowns; outer margins of dorsal and pectoral fins are densely covered with fine den- ticles in subadults and adults. Pectoral fins large, anterior margin weakly convex, anterior margin length 10.1% (8.9%–10.4%) TL, 32%–38% of pre-oral length; posterior margin weakly concave, apex broadly rounded; inner margin nearly straight; height about equal to posterior margin length; free rear tip terminates anterior to origin of first dorsal fin. Pecto- ral fin skeletal radials (SAM 33511) divided into 2–3 segments; propterygium with one radial, mesopterygium with 7 radials, metapterygium with 13 radials, and total radial count 21. Pelvic fins large, anterior margin 5.2% (5.0%–6.2%) TL, length 1.4 times into first dorsal fin length, 1.2 times in second dorsal fin length; anterior -mar gin slightly convex, posterior margin nearly straight, broadly rounded at apex; inner margin nearly straight, length about equal to posterior margin length, free rear tip narrowly acute. First dorsal fin broad, triangular, anterior margin slightly convex, with broadly rounded apex anterior to insertion, posterior margin slanting posteroventrally; in- ner margin straight, free rear tip narrowly acute; origin about opposite pectoral free rear tips or in front of them by 0.5 times eye length or less; insertion over pelvic fin origins or slightly anterior to them by < 0.25 times fin base, free rear tip extending far posterior and about opposite or slightly in front or behind pelvic insertions and behind pelvic midbases. Second dorsal fin subequal to first, with base shorter but height as great or greater; origin behind pelvic insertions by distance about 0.4–1.0 times first dorsal base. Caudal fin long, dorsal margin slightly convex, length 19.0% (18.6%–20.0%) TL, lower post-ventral lobe absent, upper post-ventral margin broadly rounded; terminal lobe well developed, caudal terminal margin slightly concave, api- ces angular. Total vertebral count 136 (132–139), monospondylous precaudal count 42 (42–45), precaudal count 91 (90–93). Intestinal valve turns 7 (6–7). Coloration.—In life, large, oval-shaped eyes emerald green; lateral trunk brownish gray dorsally, slightly darker above, becoming lighter below lateral line; ventral sur- face whitish with some darker mottling; dorsal, pectoral, and pelvic fins with light to white posterior fin edges; caudal fin terminal margin and upper post ventral margins white; anterior margin and apex of dorsal fins and dorsal caudal margin darker, more conspicuous in juveniles than adults. Two distinct dark longitudinal stripes extend length of lighter colored dorsal rostral surface; lateral rostral teeth dark-edged, ven- tral rostral teeth dark and conspicuous against light underside of snout. After pres- ervation, these sharks are a uniform medium brown above, white below. Size.—A very small sawshark, with a maximum total length for adult males of about 62 cm. Birth size is unknown. The smallest specimen examined, a 31 cm im- mature male, lacked an umbilical scar. Males immature at 31–36 cm, adolescent at 45 cm, and adult at 52–62 cm; female immature at 39 cm and mature or nearly so at 57 cm, with large eggs in its ovaries. ebert and cailliet: new shark from southern africa 507

Figure 2. Pre-branchial head of Pristiophorus nancyae sp. nov., holotype, SAM 34013, 616 mm TL. (A) Dorsal view; (B) ventral view.

Distribution.—Western Indian Ocean from central and southern Mozambique. Records from off Somalia and the Arabian Sea off Pakistan appear to be a different, possibly undescribed species. General Biology.—This sawshark is a poorly known inhabitant of the upper conti- nental slope in tropical waters found at depths of 286–500 m. Of the eight individuals examined (type series) for stomach contents, two were empty, one had unidentified organic material, and five had more or less digested shrimplike , and in one case a small decapod shrimp. No fish or cephalopod remains were apparent in these samples. Larger individuals had small puncture marks and elongated parallel cuts on their backs and tails suggestive of combat injuries from the rostra of other sawsharks. Litter size unknown. It may be caught and discarded as bycatch by off- shore shrimp trawlers and other deepwater fisheries operations off Mozambique. Etymology.—The species name nancyae is in honor of Nancy Packard Burnett for her gracious support of chondrichthyan research at the Pacific Shark Research Center at Moss Landing Marine Laboratories. The proposed common name “African dwarf sawshark” is an allusion to its known occurrence and it attaining the smallest maximum total length for a member of this genus. Comparison.—Pristiophorus nancyae is the only described member of this genus known to occur in the western Indian Ocean. The only other described member of this family has six paired gill openings and is in a separate genus, Pliotrema, which is endemic to Southern Africa. Both the new species and P. warreni were captured in the same trawl off Mozambique and are easily separable by the number of gill open- 508 BULLETIN OF MARINE SCIENCE. VOL 87, NO 3. 2011

Table 2. Maximum total length (TL) and vertebral counts, including monospondylous (MP), total precaudal (PC), and total vertebrae (Total) for all seven known sawshark species. Pristiophorus nancyae new species in bold.

Species TL (mm) n MP PC Total Pliotrema warreni 136 15 48–53 101–108 146–157 Pristiophorus cirratus 149 1 53 101 140 Pristiophorus japonicus 153 5 51–52 99–109 147–150 Pristiophorus nancyae 62 7 42–45 90–93 132–139 Pristiophorus nudipinnis 124 3 53 98–103 143–153 Pristiophorus delicatus 85+ 12 49–55 101–105 149–156 Pristiophorus schroederi 81+ 1 – 100 155 ings. The new species is separable from all other known species of the genus Pris- tiophorus by several distinctive characteristics, but most notably it differs by having a very distinctive double row of 4–5 conspicuous large pits anterior to the nasal bar- bels on the underside of its snout (Fig. 2). These pre-barbel rostral pits are absent in all other members of the genus Pristiophorus. The large rostral teeth of P. nancyae have prominent transverse ridges on their bases, a characteristic not found on any other species in this genus. The first dorsal fin is very broadly triangular, with its apex located well anterior of the first dorsal fin insertion, and the free rear tip is located about over the pelvic insertions. The first dorsal fin on all other known Pristiophorus species is narrowly triangular to semifalcate, with the fin apex located over or poste- rior to the insertion, and the free rear tip positioned over or anterior to the first third of the pelvic fin bases. Pristiophorus nancyae has the lowest monospondylous, pre- caudal, and total number of vertebrae of any member of the Pristiophoridae (Table 2). The new species is also the smallest member of the genus with a maximum size of 62 cm TL for an adult male. The next two smallest members of the genus, Pristiophorus delicates Yearsley, Last, and White 2008 and P. schroederi, have a minimum total length of 85 and 81 cm, respectively. Remarks.—The present study describes the sixth species of the genus Pristiophorus and the first member of this genus from the western Indian Ocean. Western Indian Ocean records of five-gilled sawsharks are few. Compagno (1984) reported on the first confirmed records of a five-gilled sawshark that had been collected in deep water in the Arabian Sea off Karachi, Pakistan. These specimens were of uncertain identity, and Compagno (1984) suggested that they might represent a new species. Other records of sawsharks from the western Indian Ocean include a small five- gill sawshark collected during Cruise 17 (1988) of the Soviet R/V Vityaz in the northwestern Indian Ocean on the upper slope off Socotra Island, Somalia (M Stehmann, Ichthyological Research Laboratory, Hamburg, Germany, pers comm). Timokhin and Usachev (1993) also mention a Pristiophorus species that was independently collected from the northern Indian Ocean, apparently off Socotra Island and possibly the Gulf of Aden. It is possible that P. nancyae ranges along the continental slope of the western Indian Ocean from Southern Africa to Somalia and even Pakistan, although this requires confirmation by detailed comparison of specimens. The deep-water shark fauna is sketchily known along the continental slope of east Africa and it is possible that the new species range may extend into the northern Indian Ocean. Also, it could be expected that P. nancyae will eventually be recorded in South African waters as ebert and cailliet: new shark from southern africa 509

Figure 3. (A) Lateral and dorsal view and (B) pre-branchial ventral head view of Pristiophorus nancyae sp. nov., paratype, SAM 33511, 522 mm TL, mature male. Arrow denotes conspicuous large pits on underside of snout in front of barbels. Illustration by JS DaSilva. 510 BULLETIN OF MARINE SCIENCE. VOL 87, NO 3. 2011 it occurs only ~250 km northeast of the South African border east of Xai Xai, Mo- zambique. Comparative Material.—Pliotrema warreni.—Seven specimens: BMNH-1905.6.8.9, syntype, female, 800 mm TL, coast of Natal (kwaZulu-Natal), 73 m, received from E Warren of the Natal Government Museum; BMNH-1899.2.10.4, female, ca. 700 mm TL, False Bay, Cape of Good Hope, received from JDF Gilchrist, possible syntype although listed as “Pristiophorus cirratus” in the collection, this is actually a Pliotrema warreni; SU-31456, immature female, 345 mm TL, Cape of Good Hope, South Africa, received from C Von Bonde; four specimens collected by R/V Africana on the Agulhas Bank, South Africa: LJVC-860705, mature male, 1023 mm TL, LJVC-870909, mature male, 1030 mm TL, LJVC-870114, immature female, 870 mm TL, LJVC-920505, immature female, 857 mm TL. Pristiophorus cirratus.—Four specimens: MCZ-38611, all females, 557, 600, and 965 mm TL, New South Wales, Australia, received 1954; SU-20805, immature fe- male, 337 mm TL, Port Jackson, New South Wales, Australia, collected DG Stead. Pristiophorus japonicus.—Nine specimens: MCZ-1045, female, 865 mm TL, Suruga Bay at Yenoura, Japan, 9 March, 1903, collector A Owston; MCZ-1283 male embryo, 304 mm TL, 4 March, 1906, Sagami Sea, Japan, collector A Owston; SU-7258, immature female, 499 mm TL, Aomori, Honshu, Japan, collected by DS Jordan and JO Snyder; SU-7258, immature male, 578 mm TL, Aomori, Honshu, Japan, collected by DS Jordan and JO Snyder; USNM-99500, mature male, 1072 mm TL, Hokkaido Prefecture, Hakodate, Japan, Albatross, Station TT 3657; BMNH-1862.11.1.37, immature male, 734 mm TL, Japan; BMNH-1953.8.10.6, immature female, ca. 700 mm TL, Japan; BMNH 1867.2.20.1–2 (two specimens), females, ca. 100 and 120 mm TL, Japan. Pristiophorus nudipinnis.—Five specimens: BMNH 1863.1.15.45, syntype, mature male, 1040 mm TL; BMNH 1859.9.11.1, holotype Pristiophorus oweni, immature fe- male, 323 mm TL; BMNH uncat., female embryo, 315 mm TL; MCZ-665, female, 1000 mm TL, F, New South Wales, received in mid-1800s; SU-29452, embryo or neo- nate female, 269 mm TL, received from the Australian Museum. Pristiophorus schroederi.—Three specimens:U SNM-185946 holotype, immature female, 383 mm TL, collected 24 June, 1957, 15 mi east of Dog Rocks, Cay Sal Bank, Bahamian Archipelago, Bahamas Islands, 24°05´N, 79°46´W, Vessel Combat, Station C-449, depth 640 m. USNM-185947, paratypes (2), 645 and 805 mm TL, immature male and female, 9 June, 1958, Little Bahama Bank, 28°03´N, 78°46´W, Vessel Silver Bay, Station 445, depth 1000 m.

Key to the Genus Pristiophorus

1a. A double row of 4–5 conspicuous large pits on underside of snout in front of barbels. Larger rostral teeth with prominent transverse ridges on their bases. First dorsal fin very broadly triangular, with apex well in front of insertion and free rear tip about over pelvic insertions...... Pristiophorus nancyae new species 1b. No enlarged pits on pre-barbel snout. Larger rostral teeth apparently lacking basal ridges. First dorsal fin narrower and triangular to semifalcate, with apex over or behind insertion and free rear tip over or anterior to first third of pelvic bases...... 2 ebert and cailliet: new shark from southern africa 511

2a. Rostrum short, broad, and strongly tapering, pre-oral length 22%–24% of total length. Snout width at nostrils 2.7–4.3 times in pre-orbital length. Nostrils about equidistant be- tween barbels and mouth corners. Nostrils diagonally oval and elongated...... Pristiophorus nudipinnis Günther, 1870 2b. Rostrum more elongated, narrower, and less strongly tapering, pre-oral length 25%–32% of total length. Snout width at nostrils 4.4–6.1 times in pre-orbital length. Nostrils consider- ably closer to mouth corners than barbels. Nostrils transversely oval, nearly circular...... 3 3a. Body and fins with a color pattern of dark blotches and spots (occasionally faint). Pre-barbel length from snout tip to barbels only 40%–43% of pre-oral length...... Pristiophorus cirratus (Latham, 1794) 3b. Body and fins uniform in color. Pre-barbel length from snout tip to barbels 44%–55% of pre-oral length...... 4 4a. Snout greatly elongated, pre-oral length 31%–32% of total length. Lateral trunk denticles largely tricuspidate...... Pristiophorus schroederi Springer and Bullis, 1960 4b. Snout shorter, pre-oral length 25%–31% of total length. Lateral trunk denticles unicuspi- date...... 5 5a. Mouth extending forward to below posterior third of eye. 15–26 large lateral rostral saw- teeth in front of barbels. Tooth rows 35–58 in upper jaw, increasing with growth. Lateral trunk denticles with erect crowns that are not closely imbricated...... Pristiophorus japonicus Günther, 1870 5b. Mouth extending forward to below rear margin of eye. 11–15 large lateral rostral saw-teeth in front of barbels. Tooth rows 31–42 in upper jaw. Lateral trunk denticles with flat crowns that are imbricated...... Pristiophorus delicates Yearsley, Last and White, 2008

Acknowledgments

We thank the following individuals for assistance on various aspects of this study: R Leslie (Marine and Coastal Management, Cape Town, South Africa) for collecting the specimens, J Schaaf DaSilva (California Department Fish and Game) for the illustrations of the new spe- cies, D Catania (California Academy Sciences), M Bougaardt (South African Museum), WT White (CSIRO Marine and Atmospheric Research), and J Kemper (Pacific Shark Research Center/Moss Landing Marine Laboratories). We especially thank LJV Compagno (South Af- rican Museum) for providing comparative material and general assistance on various aspects of this project. Support for this project was provided by NOAA/NMFS to the National Shark Research Consortium and the David and Lucile Packard Foundation to the Pacific Shark Re- search Center.

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Addresses: (dAe, Gmc) 1: Pacifi c shark research Center, Moss Landing Marine Laboratories, Moss Landing, California 95039. (dAe) 2: research Associate, south African institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown, 6140, south Africa. 3: Department of ichthyology, California Academy of sciences, 55 Music Concourse Drive, san Francisco, California 94118. corresPonding Author: (dAe) E-mail: .