Squalus acanthias Linnaeus, 1758

De, Sarah T., De, Marcelo R. & Gomes, Ulisses L., 2016, Taxonomy and morphology of species of the genus Squalus Linnaeus, 1758 from the Southwestern Atlantic Ocean (Chondrichthyes: Squaliformes: Squalidae), Zootaxa 4133 (1), pp. 1-89: 8-29

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Squalus acanthias Linnaeus, 1758
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Squalus acanthias Linnaeus, 1758 

( Figs. 2–15View FIGURE 2View FIGURE 3View FIGURE 4View FIGURE 5View FIGURE 6View FIGURE 7View FIGURE 8View FIGURE 9View FIGURE 10View FIGURE 11View FIGURE 12View FIGURE 13View FIGURE 14View FIGURE 15, Tables 1 –4)

Spotted spiny dogfish; Piked dogfish; Cação-bagre-espinhoso (Portuguese)

Squalus acanthias Linnaeus, 1758: 233  (original description, “Oceano Europaeo”); Rafinesque, 1810: 45 (listed; Sicily); Gill, 1862: 405 (cited; global); Poey, 1868: 213, 454 (cited; Cuba); Berg, 1895: 5, 6 (listed; Argentina, Uruguay); Jordan & Evermann, 1896: 54 (listed; North and Central America); Schreiner & Ribeiro, 1903: 79 (listed; Brazil); Regan, 1908: 45, 46 (identification key, listed; global); Garman, 1913: 192, plates 14 (figs. 1–4), 43 (figs. 9, 10), 59 (figs. 1, 2) (description; global); Fowler, 1936: 69 –71, figs. 19, 20 (revision; Eastern Atlantic); Bigelow & Schroeder, 1948: 455 –473, figs. 87 (A– D), 88 (revision; Northwestern Atlantic); Bigelow & Schroeder, 1953: 47–51, fig. 17 (description, cited; Western North Atlantic); Bigelow, Schroeder & Springer, 1953: 221 (cited; Western Atlantic); Bigelow & Schroeder, 1957: 30, fig. 3 D (description; global); Garrick, 1960: 520, figs. 1 (A–C), 3 (G–M), 5 (revision; New Zealand); Bass et al., 1976: 13, 14, figs. 8 (F–G), 9 (revision; Eastern South Africa); Cadenat & Blache, 1981: 46 –48; fig. 28 (A–E) (revision; Mediterranean Sea); Lucena & Lucena, 1981: 2, fig. 3 (listed; Brazil); Compagno, 1984: 109 -113 (revision; global); Kondyurin & Myagkov, 1984: 118 –120, fig. 1 A (revision; Western Atlantic); Menni et al., 1984: 62, 83, 84 (listed; Argentina, Uruguay); Myagkov & Kondyurin, 1986: 1 –20, fig. 1 (A, E, F, H) (revision; Atlantic); López et al., 1996: 7, 8 (listed; Argentina); Cousseau & Perrotta, 1998: 34–35 (description; Argentina); Lessa et al., 1999: 26, 61, 150 (cited, listed; Brazil); Mazzoleni & Schwingel, 1999: 114 (listed; Itajaí, Brazil); Gadig, 2001: 29, 36, 54–57, fig. 27 (description; Brazil); Soto, 2001: 94, 95 (listed; Brazil); Compagno, 2002: 380, 381, 383 (revision; South Atlantic); Nion et al., 2002: 4 (listed; Uruguay); Haimovici et al., 2003: 38, 39 (cited; Brazil); Meneses & Paesch, 2003: 7, 25, 45 (cited; Argentina); Smith & Heemstra, 2003: 61, 62, fig. 5.24 (identification key; description; South Africa); Heemstra & Heemstra, 2004 (cited, Southern Africa): 54; Soto & Mincarone, 2004: 73, 74 (listed; Brazil); Lamilla & Bustamante, 2005: 9, 26 (cited; Chile); Nelson, 2006: 66 (listed; global); Menni & Lucifora, 2007: 3 (cited; Argentina, Uruguay); Pon & Gandini, 2007 (cited; Argentina); Carrier et al., 2010: 44, 127, 139 (cited); Gomes et al., 2010: 44, 45 (cited; Brazil); Menni et al., 2010 (cited; Southwestern Atlantic); Saéz et al., 2010: 623 (identification key; Chile); Viana, 2011: 28 –56 (description; Southwestern Atlantic); Menezes, 2011: 4 (listed; Southern Brazil); Rosa & Gadig, 2014: 92 (listed; Brazil).

Squalus fernandinus Molina, 1782: 188  , 189, 285 (original description, not illustrated; Chile); Regan, 1908: 45, 46 (cited; Chile); Bigelow, Schroeder & Springer, 1953: 220–222 (cited; Western Atlantic).

Acanthorhinus acanthias: Bigelow & Schroeder, 1948: 452  (in synonymy of S. acanthias  ).

Spinax acanthias: Cuvier, 1817: 130  (cited); Cuvier, 1863: 320 (listed); Bigelow & Schroeder, 1948: 452 (in synonymy of S. acanthias  ).

Acanthias  vulgaris Risso, 1826: 13 (original description, not illustrated; Mediterranean, Sea); Müller & Henle, 1841: 83 (description); Duméril, 1865: 437 (description); Günther, 1866: 384, 396 (listed; Central America); Macleay, 1881: 366 (description; Australia); Vaillant, 1888: 5 (listed; Argentina).

Acanthias  americanus Storer, 1846: 506 (original description; United States of America); Gill, 1861: 60 (listed; North America).

Spinax acantheus: Cuvier, 1863: 320  (cited).

Acanthias  lebruni Vaillant, 1888: 5  , 13, 14, plate 1, fig. 2 (original description; Chile).

Squalus lebruni: Berg, 1895: 6  (description; Argentina, Uruguay); Menni et al., 1984: 62, 84 (listed; Argentina, Uruguay).

Squalus blainvillei: Schreiner & Ribeiro, 1903: 79  (listed; Brazil); Miranda Ribeiro, 1907 (in part): 168 (description; Brazil).

Squalus barbouri Howell-Rivero, 1936: 47  , 48 (original description, not illustrated; Cuba).

Squalus tasmaniensis Howell-Rivero, 1936  (original description, not illustrated; Tasmania); Last et al., 2007: 109–113, figs. 1 B, 2 B (revision; Tasmania).

Flakeus tasmaniensis: Whitley, 1940: 139  , fig. 150 (listed; Australia).

Koinga whitleyi: Whitley, 1940: 139  , figs. 151, 152 (listed; Australia).

Koinga kirki: Whitley, 1940: 140  , fig. 153 (listed; Australia).

Squalus kirki Phillipps, 1931  (original description, not illustrated; New Zealand).

Squalus  sp. of acanthias  group: Figueiredo, 1981: 17 (listed; Brazil); Gomes et al., 1997: 93 (description; Brazil); Marques, 1999 (cited; Brazil).

Syntype. NRMAbout NRM 85, juvenile female, 177 mm TL, unknown locality. Donation by King Gustav IV Adolf of Sweden. Other possible syntypes in Uppsala (refuted by Wheeler, 1991): UUZMAbout UUZM 159, juvenile male, 346 mm TL; UUZMAbout UUZM 160, neonate female, 380 mm TL. Type locality: “Oceano Europaeo”. Possibly from off Sweden, Finland or Norway.

Material examined (59 SWAO specimens). AMNHAbout AMNH 4099, neonate male, 230 mm TL, between the mouth of the rivers Coyle and Gallegos, Santa Cruz, Argentina; BMNH 1936.8. 26.17, adult male, 635 mm TL, near Strait of Magellan, Argentina, Southwestern Atlantic Ocean, 52.18 °S, 68 °W; BMNH 1999.5. 4.4, juvenile male, 550 mm TL, Falkland Islands, Southwest Atlantic Ocean; BMNH 1999.5. 4.13, juvenile female, 448 mm TL, Falkland Islands; BMNH 1999.5. 4.15, adult female, 535 mm TL, Falkland Islands; BMNH 1999.5. 4.16, juvenile male, 496 mm TL, Falkland Islands; HUMZAbout HUMZ 30173, adult male, unknown TL, off Patagonia, Argentina, 47 °S, 6516 ’W (dissected); HUMZAbout HUMZ 30178, adult male, 693 mm TL, off Patagonia, Argentina, 47 °S, 6516 ’W; HUMZAbout HUMZ 30200, adult male, 655 mm TL, off Patagonia, Argentina, 47 °S, 6516 ’W; HUMZAbout HUMZ 30285, adult male, 720 mm TL, off Patagonia, 47 °S, 6516 ’W; HUMZAbout HUMZ 30291, adult male, 595 mm TL, off Patagonia, Argentina, 47 °S, 6516 ’W; HUMZAbout HUMZ 30303, juvenile female, 487 mm TL, off Patagonia, 47 °S, 6516 ’W; HUMZAbout HUMZ 30310, juvenile female, 520 mm TL, off Patagonia, Argentina, 47 °S, 6516 ’W; HUMZAbout HUMZ 30324, adult male, 760 mm TL, off Patagonia, 47 °S, 6516 ’W; HUMZAbout HUMZ 107285, juvenile female, 340 mm TL, off Argentina, 4659.5 ’S, 6516 ’W (dissected); MCTAbout MCT 7439, adult female, 587 mm TL, Tramandaí, Rio Grande do Sul, Brazil; MCZAbout MCZ 1435 -S, neonate male, 175 mm TL; neonate male, 205 mm TL, Suriname; MNRJAbout MNRJ 509, juvenile male, 403 mm TL, Rasa Island, Rio de Janeiro, Brazil; MNRJAbout MNRJ 513, juvenile female, 440 mm TL, Rasa Island, Rio de Janeiro, Brazil; NMWAbout NMW 83924View Materials, neonate male, 215 mm TL, unspecified locality, Brazil; ZMHAbout ZMH 104416, adult male, 725 mm TL; adult female, 755 mm TL, near Peninsula del Valdes, Argentina, 5948 'W, 438 'S; ZMHAbout ZMH 104461, two juvenile females, 510, 557 mm TL; one adult female, 742 mm TL, near Peninsula del Valdes, Argentina, 608 'W, 433 'S; ZMHAbout ZMH 104519, two adult females, 670, 745 mm TL, near Cabo Blanco, Argentina, 6112 'W, 478 'S; ZMHAbout ZMH 104951, adult male, 715 mm TL, near Santa Cruz, Argentina, 6340 'W, 490 'S; ZMHAbout ZMH 104955, seven neonates males, 187–192 mm TL; five neonate females, 185–200 mm TL, near Santa Cruz, Argentina, 6546 'W, 4750 'S; ZMHAbout ZMH 104968, six juvenile male, 226–233 mm TL; four juvenile female, 225–235 mm TL, near Cabo Blanco, Argentina, 6518 'W, 470 'S; ZMHAbout ZMH 107911, four juvenile males, 213– 225 mm TL; six juvenile females, 215–223 mm TL, near Santa Cruz, Argentina, 640 'W, 460 'S; ZMHAbout ZMH 108038, adult male, 670 mm TL, near Cabo Blanco, Argentina, 650 'W, 470 'S; ZMHAbout ZMH 115469, juvenile male, 322 mm TL, near Santa Cruz, Argentina, 6324 'W, 4551 'S.

Diagnosis. Squalus acanthias  from the Southwestern Atlantic Ocean is distinguished from all local congeners by the following unique characters: rounded white spots dorsolaterally on body; anterior margin of nostrils unilobate; origin of first dorsal-fin spine behind the free rear tips of the pectoral fins; first dorsal-fin spine length much lower than fin height (first dorsal-fin spine length 1.5 %– 2.8 % TL vs. first dorsal-fin height 6.0%– 8.1 % TL); second dorsal-fin spine very elongated (1.9 %– 4.5 % TL), taller than second dorsal-fin apex; interdorsal distance 1.8 (1.5–2.2) times greater than dorsal-caudal distance; pectoral-pelvic distance 1.0 (0.6–1.3) times pelvic-caudal distance; dermal denticles unicuspid; coracoid bar with segmented processes anterolaterally.

Description. External morphology. Measurements and counts are summarized in Tables 1 –2 (in the description of S. acanthias  below, single values represent the mean followed by the range for non-type specimens from which data were taken). Body fusiform and slender, slightly more arched anteriorly as of posterior margin of spiracle to level of pectoral fin insertion; head height 0.8–1.1 times trunk height, and 0.8–1.5 times abdomen height. Head flattened, small (its length 18.4 %– 23.3 % TL) and broad, its width 1.2 (1.0– 1.5) times trunk width and 1.4 (1.1–2.2) times abdomen width. Snout somewhat elongate (5.8 %– 8.1 % TL), slightly obtuse in ventral view. Nasal apertures located laterally, small and slightly oblique; anterior margin of nostril unilobate or slightly bilobate in young specimens ( Fig. 3View FIGURE 3); distance from snout tip to nostrils 1.0 (0.9–1.2) times distance from nostrils to upper labial furrow; prenarial length corresponding to half of preoral length; internarial distance 0.9 (0.6–1.4) times eye length. Eyes oval with anterior margin rounded and posterior margin angular, not notched; length 2.0 (1.4–3.5) times eye height; eyes closer to snout tip than to first branchial arch. Prespiracular length 1.7 (1.5 –2.0) times preorbital length and 0.6 (0.5–0.6) times pre-pectoral length. Spiracles crescent-shaped, located posterodorsally to eye, oblique in relation to longitudinal axis of body; diameter at least one-quarter eye length. Prebranchial length 1.5 (1.3–1.7) times prespiracular length. Gill slits concave, located in front of pectoral fins, tall with fifth gill slit 1.2 (0.7–1.7) greater in height than first gill slit.

Measurements Holotypes Squalus S. barbouri  S. tasmaniensis  MCZAbout MCZ 1463 -S MCZAbout MCZ 146 -S N Range x SD TL (mm) 267.0 245.0 21 200.0–880.0 487.7 212.3 PCL 77.2 77.6 21 31.2–81.1 76.3 10.4 PD2 59.2 57.1 21 53.4–62.2 58.8 2.1 PD1 34.5 30.6 21 29.1–33.7 32.4 0.9 SVL 52.4 49.0 21 44.2–52.8 49.7 2.3 PP2 54.3 44.9 21 41.9–50.3 46.8 2.1 PP1 21.8 19.6 21 17.8–22.8 20.4 1.2 HDL 22.4 19.9 21 18.4–23.3 20.9 1.3 PG1 18.5 16.4 21 15.5–20.5 17.4 1.2 PSP 12.6 9.8 21 9.9–14.1 11.3 1.2 POB 7.0 5.4 21 5.8–8.1 6.9 0.6 PRN 5.1 3.3 21 3.8–5.1 4.5 0.4 POR 9.9 7.6 21 7.8–10.9 9.1 1.0 INLF 4.9 3.6 21 3.7–5.6 4.4 0.5 MOW 6.6 5.6 21 6.5–8.1 7.2 0.4 ULA 2.5 2.2 21 2.0–2.9 2.3 0.2 INW 2.8 3.2 21 2.8–4.8 3.4 0.4 INO 7.2 6.2 21 6.5–9.7 7.4 1.0 EYL 3.5 3.1 21 2.7–5.7 3.7 0.9 EYH 0.9 1.6 21 1.4–3.6 1.9 0.5 SPL 1.3 1.8 21 0.9–2.1 1.4 0.4 GS1 1.7 1.0 21 1.3–2.3 1.7 0.3 GS5 1.8 1.6 21 1.2–2.9 2.0 0.3 IDS 18.9 19.9 21 17.1–22.7 20.2 1.7 DCS 10.8 11.9 21 9.6–11.9 11.1 0.6 PPS 24.9 22.2 21 16.5–26.8 22.8 2.5 PCA 22.2 22.4 21 20.3–28.1 22.8 1.9 D1L 12.1 13.2 21 10.5–13.1 12.1 0.7 D1A 11.5 11.2 21 8.5–11.6 9.5 0.8 D1B 7.0 7.9 21 6.4–7.5 7.0 0.3 D1H 7.8 6.9 21 6.0–8.1 6.7 0.4 D1I 5.7 5.8 21 4.4–6.0 5.2 0.4

Mouth arched and evidently broad, its width 1.6 (1.4–1.9) times prenarial length and 2.1 (1.6–2.5) times internarial space; upper labial furrow slender and large, its length corresponding to 2.0%– 2.9 % TL; lower labial furrow inconspicuous. Teeth unicuspid, similar in both jaws, flattened labial-lingually and laterally overlapped; teeth broad, although short at crown; upper teeth smaller than lower teeth; cusp thick and elongate, oblique and directed laterally (cusp strongly vertical in most adult males); convex mesial cutting edge; both distal and mesial heels rounded; apron well elongated in both jaws; one intermediate tooth present at least in upper jaw, usually smaller than the following teeth, with cusp completely vertical, short apron, and both distal and mesial heels markedly rounded; two series of functional teeth in upper and lower jaws; teeth rows varying from 14 – 1–14 in upper jaw and 11 – 1–12 in lower jaw ( Fig. 4View FIGURE 4).Origin of first dorsal fin posterior to free rear tips of pectoral fin; in young specimens, origin over free rear pectoral tips (pre-first dorsal length 29.1 %– 33.7 % TL in neonates and juveniles); horizontal distance between origins of pectoral fin and first dorsal fin 1.8 (1.2–2.3) times preorbital length. First dorsal fin vertical and markedly wide at its fin web, relatively large (10.5 %– 13.1 % TL) and low, its length 1.8 (1.5–2.1) times its height, its height and base length 1.0 (0.9–1.2) times preorbital length; anterior margin convex, posterior margin moderately concave near free rear tip; apex rounded, and free rear tips pointed; inner margin length 0.8 (0.6–0.9) times fin height. Origin of first dorsal-fin spine slightly posterior to free rear tips of pectoral fins. First dorsal-fin spine conspicuously slender and low, its length not greater than half dorsal-fin height, never reaching fin apex. Interdorsal distance 1.0 (0.8–1.3) times prepectoral length and 1.8 (1.5–2.2) times dorsal-caudal distance. Pre-second dorsal length 4.3 (3.7–5.2) times the anterior margin length of pectoral fin and 2.9 (2.6–3.2) times dorsal caudal lobe length. Second dorsal fin rather oblique and broad at its fin web, relatively large, its length 1.0 (0.9–1.2) times first dorsal-fin length, and 2.5 (1.9 –3.0) times second dorsal-fin height; anterior margin convex and posterior margin falcate; apex rounded, and free rear tip pointed; inner margin elongate, its length 1.0 (0.8–1.1) times fin height. Second dorsal-fin spine slightly inclined, elongated (1.9 %– 4.5 % TL) and reaching the fin apex (in some adult specimens, second spine exceeds the fin apex); slender and low, its length 0.7 (0.4–0.9) times second dorsal-fin height and 1.8 (1.3–2.5) times first dorsal-fin spine length; second dorsal-fin spine broader than first dorsal-fin spine, its base length 1.3 (1.0– 1.5) times first dorsal-fin spine base length.

Pre-pectoral length 0.6 (0.6–0.7) times pre-first dorsal length, 0.4 (0.4–0.5) times pre-vent length. Pectoral fins with both anterior and inner margins convex, and posterior margin straight; anterior margin length 1.6 (1.4–1.6) times posterior margin length, but its tip reaches same level of its apex; apex and free rear tips evidently rounded, but not lobe-like; pectoral fin base length varying from 3.2 %– 5.2 % TL.

Pelvic fins narrow and elongate, their length 8.6 %– 12.4 % TL anterior margin; slightly convex and posterior margin straight; apex and free rear tips rounded, the latter lobe-like; origin of pelvic fins 1.8 (1.5 –2.0) times distance between origins of the two dorsal fins, nearest to second dorsal fin (in young specimens, nearest to first dorsal fin); pectoral-pelvic distance 1.0 (0.6–1.3) times pelvic-caudal distance, usually equal in adults. Clasper cylindrical, compressed dorsoventrally throughout its extension, greatly extended beyond free rear tips of pelvic fin, its outer length 1.1 %– 5.8 % TL; siphon large, located medioventrally from the anteriormost end of puboischiadic bar level of intermediate cartilage; clasper groove longitudinal and dorsomedially, sinuous and deep; apopyle broad, placed more anteriorly in clasper groove; hypopyle narrow, located posteriorly to rhipidion; rhipidion elongate, blade-like, located medially at distal end of clasper ( Fig. 5View FIGURE 5).

Caudal peduncle with inconspicuous lateral keels, originating behind insertion of second dorsal fin; upper and lower precaudal pits marked. Caudal fin conspicuously rectangular on dorsal lobe with dorsal-caudal margin straight and upper postventral margin strongly convex; dorsal-caudal margin length 1.0 (0.9–1.1) times head length and 1.8 (1.7 –2.0) times preventral margin length; caudal fork markedly concave and broad, its width corresponding to 6.0%– 7.4 % TL; lower postventral margin also convex; preventral caudal margin convex and large, its length 2.4 (1.7–3.5) times pelvic fin inner margin length.

Dermal denticles ( Fig. 7View FIGURE 7). Denticles unicuspid, their length and width equivalent, relatively sparsely positioned, not imbricated; median ridge pronounced, elongate and narrow, projecting anteriorly beyond crown base; median cusp pointed posteriorly. In neonates, denticles smaller, markedly sparse and underdeveloped throughout body.

Coloration ( Figs. 2View FIGURE 2, 6View FIGURE 6). Body dark gray dorsal and laterally, and light gray more posteriorly from the lower lateral half of the body just in front of the pectoral fin origin to the caudal fin; white ventrally; large and few paired white spots dorsolaterally, usually anterior to second dorsal fin. Both dorsal fins dark gray with apex slightly darker at tip than rest of fin, white at fin base and on free rear tips; dorsal-fin spines dark gray to brownish anteriorly and whitish at tips. Pectoral fins also dark gray dorsoventrally with both inner and posterior margins broadly whitish. Pelvic fins gray dorsally and ventrally, lighter ventrally, with anterior and posterior margins evidently white. Caudal fin gray, darker at posterior tip, slightly white at dorsal caudal margin, forming a white caudal bar; black caudal stripe strong and wide, dorsal to vertebral column on caudal fin; postventral caudal margins discreetly white; preventral caudal margin white, including at ventral tip ( Fig. 2View FIGURE 2). In young specimens (between 200–330 mm TL), body usually light gray with numerous white spots, often fused ( Fig. 6View FIGURE 6); dorsal fins markedly darker at tip; caudal fin conspicuously dark gray with broad white caudal bar at dorsal caudal margin; black upper caudal blotch near the terminal region from vertebral column to posterior caudal tip; preventral and postventral margins largely white.

Skeletal morphology. Measurements and counts are summarized in Tables 2–3.

Neurocranium ( Fig. 8View FIGURE 8). Neurocranium greatest width at level of postorbital processes (50.0%– 53.4 % CL), narrower at interorbital region (28.5 %– 31.8 % CL) and between the opisthotic processes in the otic region (32.6 %– 35.5 % CL). Rostrum elongate (its length 36.0%– 42.3 % CL), slender proximal and distally, and wide medially, its width 11.5 %– 16.9 % CL; lateral rostral cartilages cylindrical, somewhat depressed anteriorly; ventrally, two lateral rostral appendices thick and hook-like; median rostral prominence small; rostral keel conspicuous and well elongate, extending anterior to nasal capsules (its length 21.9 %– 26.1 % CL). Prefrontal fontanelle rounded, located at base of rostrum and anterior to cerebrum. Nasal capsules oval and strongly oblique, very narrow, width across nasal capsules 44.0%– 48.7 % CL; small fissures evident ventrally on nasal capsules (not shown); anterior nasal margin with a unique cartilaginous lobe; subnasal fossa oval and large, located ventrally on each side of rostral keel.

Cranial roof strongly concave medially at interorbital region with a prominent lateral supraorbital crest; longitudinal sulcus deeper anteriorly, carrying the profundus canal for the ophtalmicus profundus and a series of foramina (8–9 foramina) of the superficial ophthalmic branches of trigeminal (V) and facial (VII) nerves; preorbital canal rounded and markedly large, placed anteriorly to a series of foramina; canal for the ophtalmicus profundus rounded, located anterior to the preorbital canal and beside the ethmoidal canal. Preorbital processes small, neurocranium broad between them (width 47.9 %– 50.2 % CL). Postorbital processes prominent and triangular, not elongate (length 7.8 %– 9.7 % CL). Distance between orbital processes 28.4 %– 28.7 % CL.

Supraethmoidal processes in ethmoidal region prominent and triangular not elongate, and placed at dorsal base of prefrontal fontanelle; epiphyseal pit rounded and broad, posterior to supraethmoidal processes; ethmoidal canal rounded and narrow, placed at the base of each nasal capsule; ethmoidal chamber flattened, somewhat constricted ventrally with a triangular and prominent ectethmoid process on each side; conspicuous subethmoidal ridge, posterior to rostral keel, extending to subethmoidal region; subethmoidal region elongate and very narrow, its width 10.5 %– 13.6 % CL.

Otic region deep anterodorsally between the otic capsules; otic capsules narrow and heptagonal on each side; dorsally, two conspicuous anterior and two posterior semicircular canals, the former placed almost in parallel to neurocranial longitudinal axis; endolymphatic fossa oval and large, with two anterior endolymphatic foramina, slightly oblique, and two posterior and vertical perilymphatic foramina; strong otic crest located posteriorly to endolymphatic fossa; prominent sphenopterotic ridge at sides of otic capsules; opisthotic process rather discreet at distal portion of sphenopterotic ridge; laterally, otic wall delimited by a prominent lateral semicircular canal below the sphenopterotic ridge, a shallow ventrally situated hyomandibular facet, and anteriorly by a deep lateral auditory groove; width across hyomandibular facets between 36.8 %– 44.1 % CL.

Orbital region very narrow with concave preorbital wall, with orbitonasal canal at base; optic foramen (II) large, placed midventrally in interorbital wall; trochlear foramen (IV) small dorsoanterior to optic foramen (II); eye-stalk located more posteriorly in this region between the oculomotor foramen (III) and abducens foramen (VI), positioned, respectively, dorsally and ventrally to eye-stalk; a broad foramen prooticum for trigeminal (V) and facial (VII) nerves positioned in the posterior edge of the interorbital wall, just anterior to the postorbital process; the foramen prooticum also opens posteriorly for the hyomandibular branch of the facial nerve (VII) at the base of hyomandibular facet; transbasal canal small located ventroposterior to the anterior opening of the foramen prooticum.

Basal plate flattened and large (length 36.9 %– 42.2 % CL), narrower anteriorly at basitrabecular process, broader posteriorly with its width between 17.8 %– 20.7 % CL; basitrabecular processes conspicuous and beanshaped, perpendicular to basal plate axis; basal angle width 16.8 %–19.0% CL; two sinuous lateral prominences on each side at basitrabecular processes and the first cartilaginous process; a single cylindrical cartilaginous process on each side of the basal plate, width across processes 28.1 %– 32.8 % CL; single foramen for carotid artery anteromedially located in basal plate; foramina for orbital artery with its ventral opening in the anterior base of the cartilaginous process and its lateral opening in the lateral otic wall.

Occipital region with two broad occipital condyles distally, and a wide foramen magnum between them, its width 7.2 %– 10.7 % CL; vagus foramen (X) rounded, lateral to occipital condyles; thick and subtriangular glossopharyngeal base, although not very prominent, located more laterally in the occipital region, with an oval and broad foramen for glossopharyngeal nerve (IX).

Pectoral fin and girdle ( Figs. 9View FIGURE 9, 10View FIGURE 10). Coracoid bar unpaired, strongly concave dorsally, convex ventrally, flattened and slender at midline, and thick and cylindrical at dorsal tips; two conspicuous facets anteriorly, an anterior facet and the m. depressor pectoralis facet, located more laterally and bearing the diazonal foramen; a pair of longitudinal segmented processes attached ventrally to the coracoid bar, comprised by 3–4 units of small barrelshaped cartilages, evident laterally and dorsally to the m. depressor pectoralis facet; posteriorly, a pair of medioventral facets support the origin of the m. parietalis pars hypaxonica; a conspicuous caudal process present on each side of the hindmost part of this facet that, with the base of the articular process, supports the origin of m. pterygii ventralis muscle. Scapulae dorsal to, and continuous with, the coracoid bar, forming a U-shaped scapulocoracoid cartilage; articular process of scapulocoracoid with a single rounded condyle articulating with mesopterygium of pectoral fin; the m. levator pectoralis facet, located more distally to the articular process of pectoral girdle, supports the origin of m. pterygii dorsalis. Suprascapulae paired and triangular, cylindrical distally and somewhat flattened proximally, attaching dorsally to each side of scapulae, directed upward toward the vertebral column.

Pelvic fin and girdle ( Fig. 11View FIGURE 11). Pelvic girdle with puboichiadic bar rectangular, short and slightly slender, with anterior margin somewhat straight and posterior margin convex; two conspicuous expansions posteriorly articulate puboischiadic bar to the basipterygium and to the anterior pelvic fin element; a single pelvic nerve foramen present at each side of pubosichiadic bar. Anterior pelvic fin basal element subrectangular and wide with three series of irregular and small radials; basipterygium elongate markedly slender, cylindrical and sinuous; pelvic radials thin and large, cylindrical, and segmented into proximal and distal elements, the former much larger; 14 total pelvic radials.

Claspers ( Fig. 12View FIGURE 12). Intermediate element barrel-shaped, attaching pelvic fin basipterygium to axial cartilage; beta cartilage single, slender and cylindrical, located laterodorsally between intermediate element and axial cartilage; axial cartilage small, slender, conspicuously sinuous medioproximally; end-style elongate and thin, mediodistal to axial cartilage; dorsal marginal cartilage slim, located dorsolaterally to axial cartilage; dorsal terminal cartilage conspicuously elongate, not reaching distal end of clasper, hook-like, connected proximally to dorsal marginal cartilage and axial cartilage, and medially to end-style; dorsal terminal 2 cartilage leaf-like and flattened, also elongate, with concave lateral margin, attached medially to dorsal terminal cartilage, and proximally to dorsal marginal cartilage, externally supporting the rhipidion; ventral marginal cartilage slightly thick and broad, concave and grooved laterally at its inner side, emerging as a folded plate at insertion of accessory terminal cartilage; ventral terminal cartilage also large, slender and spatula-like with rounded distal tip somewhat sinuous laterally, located at distal end of clasper, and attached proximally to ventral marginal cartilage and medially to endstyle; accessory terminal 3 cartilage (or spur) markedly slim and elongate with an evident dorsal groove, distally pointed, with a rounded proximal edge, partially attached to ventral margin and ventral terminal cartilages.

Vertebral counts ( Table 2). Monospondylous vertebrae with mode value 44 (43–47); diplospondylous vertebrae 61 (57–63); precaudal vertebrae 75 (75–81); total vertebrae 101 (101–109).

Geographical distribution ( Fig. 13View FIGURE 13). In the SWAO, S. acanthias  is more commonly found from southern Brazil off Tramandai, Rio Grande do Sul state to the southern coast of Argentina in Santa Cruz, Patagonia, as well as in the Falkland Islands. Few occurrences are observed in warmer equatorial waters from Suriname and southeastern Brazil (e.g. Rasa Island, Rio de Janeiro state). For its distribution elsewhere, see Ebert et al. (2013).

Etymology. The epithet acanthias  , from the Greek αγκάθι, means spine.

Remarks. In the 10 th edition of the Systema Naturae, Linnaeus (1758) defined this species through a combination of superficial characters that may be applied to any species of shark of the order Squaliformes  , among them: absence of anal fin; presence of dorsal-fin spines, and slender body. The type locality is regarded as “Europaeo Ocean without further specifications. Type specimens were not designated in the original description. The lack of information in fish collections contributed to mystery of the whereabouts of the types, and there have been speculations regarding possible syntypes in the fish collection of the Natural History Museum in Stockholm based on pre-Linnean and Linnean data in the literature ( Fernholm & Wheeler, 1983; Wheeler, 1991). This situation may have contributed to confuse the identity of S. acanthias  , as seen by the numerous misidentifications in the literature, and compounded by the fact that the species was not well characterized.

This species also has a worldwide distribution and exhibits varying patterns of its biological functions such as reproduction and growth, distribution and abundance, and populational structure ( Gallucci et al., 2009; Ebert et al., 2013). Thus, it is commonly considered a polytypic species or a complex of cryptic species or populations, supported by morphological and molecular data, which have allowed for the recognition of new species ( Myagkov & Kondyurin, 1986; Veríssimo et al., 2010; White & Last, 2012).

The taxonomy of S. acanthias  is not yet completely clarified due to the morphological and molecular variation between specimens from different localities worldwide (e.g. Bigelow & Schroeder, 1948, 1957; Springer & Garrick, 1964; Bass et al., 1976; Ward et al., 2007). Our results corroborate these studies from a morphological standpoint, despite that external measurements mostly overlap between specimens of S. acanthias  from SWAO and other localities, indicating that they appear to be conspecific ( Table 4). Specimens of S. acanthias  share the following characteristics ( Table 2): anterior margin of the nostrils with a simple lobe, not bifurcated when adults; 14 – 1–14 teeth in the upper jaw and 12 – 1–12 in the lower jaw; first dorsal-fin spine located posterior to the free rear tips of the pectoral fins; unicuspid dermal denticles without lateral expansions; presence of white spots on dorsum.

Significant differences are noticed compared to specimens from the Southeastern Pacific Ocean ( SEPO): smaller direct length for SWAO specimens such as precaudal length (31.2 %– 81.1 % TL vs. 81.8 %– 82.1 % TL), prefirst dorsal fin length (29.1 %– 33.7 % TL vs. 34.0%– 34.3 % TL), pre-second dorsal fin length (53.4 %– 62.2 % TL vs. 63.4 %– 64.8 % TL) and interdorsal space (17.1 %– 22.7 % TL vs. 24.5 %– 24.6 % TL); greater first dorsal fin-spine length in SWAO than in SEPO (1.5 %– 2.8 % TL vs. 0.8 %– 1.3 % TL); and pre-second dorsal fin length 2.6–3.2 times dorsal caudal margin length (vs. 3.3 –4.0 times in SEPO). Two nominal species are often applied in the Southeastern Pacific Ocean, S. fernandinus Molina, 1782  from Juan Fernandez Island, Chile, and S. lebruni (Vaillant, 1888)  from Tierra del Fuego, Argentina, which are both recognized as junior synonym of S. acanthias  (e.g. Garrick, 1960; White et al., 2007). The former species, however, is also frequently treated as a junior synonym of S. blainvillei  (e.g. Fowler, 1936; Compagno, 1984) due to the size of its dorsal spines (approximately 63.5 mm according to Molina, 1782). The presence of white spots refers undoubtedly to S. acanthias  but the dorsal spine length represents a morphological regional peculiarity ( Garrick, 1960). Few authors support S. lebruni  as a valid species distinct from S. acanthias  (e.g. Berg, 1895; Menni et al., 1984) based on the presence of upper central teeth with a vertical cusp and lateral expansions that provide a tricuspid shape, although this was not noticed in the present study. Specimens from the Southeastern Pacific Ocean are also distinct in external morphometrics, color pattern, and vertebral counts from specimens from the Northwestern Atlantic Ocean ( NWAO), Northeastern Atlantic Ocean ( NEAO), Northeastern Pacific Ocean ( NEPO), and Mediterranean Sea, calling for further studies regarding the taxonomy of S. acanthias  in the SEPO.

We noticed slight but possibly significant variations in external measurements among specimens of S. acanthias  worldwide ( Table 4): mouth width (7.4 %– 7.7 % TL in NEAO; 7.3 %– 9.4 % TL in NEPO; 7.0%– 7.1 % TL in SEPO); first dorsal fin-spine length (1.6 %– 3.1 % TL in NWAO; 2.0%– 4.6 % TL in the Mediterranean Sea; 1.5 %– 2.2 % TL in NEAO; 1.5 %– 2.5 % TL in NEPO; 0.8 %– 1.3 % TL in SEPO); pectoral-fin posterior margin length (10.5 %– 13.7 % TL in Mediterranean Sea; 9.4 %– 10.3 % TL in NEAO; 5.5 %– 10.3 % TL in NEPO); and dorsal caudal margin length (16.0%– 19.5 % TL in SEPO; 19.8 %– 22.2 % TL in Mediterranean Sea; 19.9 %– 21.9 % TL in NEAO; 20.4 %– 24.6 % TL in NEPO). Specimens from NEPO and SEPO have pelvic fins located almost at midspace between origins of the dorsal fins, while in the remaining specimens these fins are placed closer to the second dorsal fin. Small juvenile specimens have pelvic fins usually nearest the first dorsal fin, which is in agreement with the ontogenetic observations of Garrick (1960).

Variations in monospondylous, diplospondylous, precaudal, and total vertebrae ( Table 2) are also present among specimens of S. acanthias  from the Northwestern Atlantic, Southeastern Atlantic, Northwestern Pacific, Northeastern Pacific, and Southeastern Pacific oceans, in agreement with the previous results of Springer & Garrick (1964). These results, however, do not demonstrate consistent morphological variation for a North Pacific population separate from a second population that includes South Pacific and Atlantic specimens, as supported in a molecular analysis ( Veríssimo et al., 2010) and in terms of data on general biology (e.g. Galucci et al., 2009).

Squalus suckleyi (Girard, 1854)  from the Northwest Pacific Ocean has also been considered a junior synonym of S. acanthias  (e.g. Bigelow & Schroeder, 1948, 1957; Compagno, 1984), but which has been recently resurrected as valid ( Ebert et al., 2010). Molecular analysis (e.g. Ward et al., 2007; Veríssimo et al., 2010) strongly support its separation, despite the fact that mostly overlapping morphological characters were used to distinguish this species from S. acanthias  (e.g. Jordan & Evermann, 1896; Regan, 1908; Garman, 1913; Ebert et al., 2010), such as a less obtuse snout, the position of the first dorsal-fin spine located right after the free rear tip of the pectoral fin, pelvic fins located slightly closer to the second dorsal fin than to the first dorsal fin, and a smaller number of total vertebrae. These differences, however, were not observed by us with exception of the position of the first dorsal spine in relation to the pectoral fin. Other morphological characters are still needed to better distinguish S. suckleyi  from S. acanthias  .

Many attempts (e.g. Rafinesque, 1810; Müller & Henle, 1841; Duméril, 1865) have been made to provide additional characters for S. acanthias  . Garman (1913), however, was the first author to provide detailed distinctive characters and important information about its synonymy. He based his conclusions on a specimen from the Mediterranean Sea whose characteristics are similar to the specimens of S. acanthias  examined by us from the SWAO. However, few variations exist between specimens of these two regions: shorter interdorsal space in specimens from SWAO (17.1 %– 22.7 % TL vs. 22.9 %– 24.8 % TL), and claspers more elongate in specimens from the Mediterranean Sea (clasper inner length 8.8 %– 9.9 % TL vs. 2.2 %– 5.7 % TL for SWAO).

Later, Bigelow & Schroeder (1948, 1957) provided some diagnostic characters of S. acanthias  from the North Atlantic Ocean, which overlap with those of specimens from the Southwestern Atlantic Ocean. These authors further emphasized that sexual dimorphism may be present in this species, as the first dorsal-fin spine length is greater in males than in females, and the second dorsal-fin spine reaches the dorsal-fin apex only in males. Our results, however, do not support this dimorphism once all specimens have the first dorsal-fin spine always below half of the first dorsal-fin height, and the second dorsal-fin spine is always taller than the second dorsal-fin in both males and females.

The dorsolateral white spots in S. acanthias  usually are smaller and in greater quantity in neonates and young juveniles than in adults as they reduce with maturity, as supported by previous authors (e.g. Jordan & Evermann, 1896; Bigelow & Schroeder, 1948; Myagkov & Kondyurin, 1986). Bass et al. (1976) and Cadenat & Blache (1981) also demonstrated variation in dermal denticles with growth, although this was not observed by us once; even the largest specimens also had unicuspid dermal denticles without any lateral expansions.

Myagkov & Kondyurin (1986) recognized four subspecies of S. acanthias  due its great apparent morphological variation: S. acanthias acanthias  (sensu Lindberg & Legeza, 1956) from the North Atlantic, S. acanthias  ssp. from the North Pacific, S. acanthias ponticus Myagkov & Kondyurin, 1986  from the Black Sea, and S. acanthias africana Myagkov & Kondyurin, 1986  from the Southeastern Atlantic Ocean. The first two subspecies had been differentiated anteriorly by Lindberg & Legeza (1956, 1959) by pectoral-fin length. According to these authors, pectoral length is greater in S. acanthias suckleyi  than in S. acanthias acanthias  , as was also reported by Regan (1908). Myagkov & Kondyurin (1986) present additional characters to separate them, concerning tooth formula and the number of total vertebrae, in which the latter is greater in specimens from the North Atlantic Ocean. However, these characters were based on neonates and used an unclear methodology that leaves doubt regarding its reliability. Myagkov & Kondyurin (1986) also provided the following features for S. acanthias acanthias  : base length of pelvic fins 1.5 times greater than base length of the second dorsal fin; distance from snout tip to nostrils much greater than internarial distance; teeth varying from 14 – 14 on the upper jaw and 11–12 on the lower jaw; number of total vertebrae greater than 106. These characters, however, overlap with other species of Squalus  described in their work.

We analyzed the holotypes of Squalus tasmaniensis Howell-Rivero, 1936  from Tasmania and Squalus barbouri Howell-Rivero, 1936  from Cuba. The characters used by Howell-Rivero (1936 a) to distinguish these two species from congeners, such as small white spots on the dorsum and the position of the first dorsal-fin spine in relation to the pectoral fins, are problematic as they vary ontogenetically. Both holotypes correspond to juveniles (245 mm and 267 mm, respectively) very similar to typical young specimens of S. acanthias  regarding external morphology and dermal denticles ( Figs. 14View FIGURE 14, 15View FIGURE 15). Some body measurements ( Table 1), however, are quite distinct, such as pre-first dorsal fin length, prepelvic length, second dorsal-fin height, and mouth width, although many of these variations are not relevant due to the lack of accuracy for the measurements of the holotypes that are in poor condition.

Squalus acanthias  can be easily identified by the presence of white spots on the dorsum once no other congener has this particular color pattern (other than S. suckleyi  ), associated with the more posterior position of the first dorsal-fin spine in relation to the free rear tips of the pectoral fin, and the presence of a single simple lobe on the anterior margin of the nostrils, as corroborated by the present study and earlier studies, such as Meneses & Paesch (2003) for Mar del Plata, Argentina, Gomes et al. (1997) for Brazil, and Lamilla & Bustamante (2005) for Chile.

Despite the morphological particularities observed in specimens of different geographical regions, it is still difficult to subdivide S. acanthias  into different species based on morphology (including S. suckleyi  ); recognizing a complex of cryptic species with the tenuous data presently in hand may disrupt nomenclatural stability. Revising S. acanthias  taxonomically on a global scale through a detailed morphological analysis (external and skeletal) and taking into account all of its regional variations is ongoing (Viana & Carvalho, in prep.).

Comparative material. Squalus acanthias  : Northwestern Atlantic Ocean (151 specimens): AMNHAbout AMNH 1, neonate female, 165 mm TL, neonate male, 153 mm TL, no data; AMNHAbout AMNH 652, neonate female, 209 mm TL, New York, U.S.A.; AMNHAbout AMNH 656, neonate female, 158 mm TL, New Jersey, U.S.A.; AMNHAbout AMNH 3518, neonate female, 186 mm TL; three neonate males, 177–187 mm TL, Canada; AMNHAbout AMNH 7175, juvenile female, 305 mm TL, New Jersey, U.S.A.; AMNHAbout AMNH 8955, nine neonate females, 155–205 mm TL, three neonate males, 169–190 mm TL, Massachusetts, U.S.A.; AMNHAbout AMNH 36980, neonate female, 207 mm TL, neonate male, 198 mm TL, Monhegan Island, Maine, U.S.A.; AMNHAbout AMNH 40205, three neonate females, 242–254 mm TL, two neonate males, 252–253 mm TL, Rhode Island, U.S.A.; AMNHAbout AMNH 40802, two neonate females, 190–207 mm TL, two neonate males, 189–190 mm TL, Woods Hole, Massachusetts, U.S.A.; AMNHAbout AMNH 44117, adult male, 755 mm TL, Rhode Island, U.S.A.; AMNHAbout AMNH 56194, neonate male, 226 mm TL, New York, U.S.A.; AMNHAbout AMNH 65177, juvenile female, 445 mm TL, North Carolina, U.S.A.; AMNHAbout AMNH 73637, neonate female, 163 mm TL, North Carolina, U.S.A.; AMNHAbout AMNH 76046, two neonate females, 188– 198 mm TL, South Carolina, U.S.A.; AMNHAbout AMNH 216291, two neonate females, 93–118 mm TL, U.S.A.; AMNHAbout AMNH 221591, four neonate male, 250–287 mm TL, New York, U.S.A.; AMNHAbout AMNH 3517 (skeleton, no data), Weymouth, St. Mary Bay, Nova Scotia, Canada; AMNHAbout AMNH 53052 (skeleton, no data), U.S.A.; AMNHAbout AMNH 97553 (skeleton, no data), U.S.A.; AMNHAbout AMNH 221706 (skeleton), adult female, 860 mm TL, Worcester County, Maryland, U.S.A.; AMNHAbout AMNH 221711, neonate female, 139 mm TL, Worcester County, Maryland, U.S.A.; AMNHAbout AMNH 225783 (skeleton, no data), North Carolina, U.S.A.; CASAbout CAS 11226View Materials, neonate male, 200 mm TL, Massachusetts, U.S.A.; CASAbout CAS 11227View Materials, neonate female, 166 mm TL, Massachusetts, U.S.A.; CASAbout CAS 11228View Materials, neonate male, 185 mm TL, Massachusetts, U.S.A.; SU 3990, neonate male, 167 mm TL, Massachusetts, U.S.A.; MCZAbout MCZ 168 -S, adult female, 910 mm TL, Grand Manan Island, New Brunswick, Canada; MCZAbout MCZ 172 -S, 18 neonate females, 155–205 mm TL, Massachusetts Bay, U.S.A.; MCZAbout MCZ 454 -S, adult male, 720 mm TL, Massachusetts Bay, U.S.A., 42.41 °N, 70.88 °W; MCZAbout MCZ 458 -S, adult female, 820 mm TL, Waquoit, Massachusetts, U.S.A., 41.58 °N, 70.52 °W; MCZAbout MCZ 520 -S, five neonate females, 180–215 mm TL, seven neonate males, 160–210 mm TL, Massachusetts Bay, U.S.A., 42.42 °N, 70.90 °W; MCZAbout MCZ 840 -S, adult female, 675 mm TL, New Hampshire, U.S.A.; MCZAbout MCZ 851 -S, adult female, 625 mm TL, Massachusetts Bay, U.S.A., 42.37 °N, 70.74 °W; MCZAbout MCZ 872 -S, adult female, 870 mm TL, Gloucester, Massachusetts, U.S.A., 42.61 °N, 70.66 °W; MCZAbout MCZ 1426 -S, two neonate females, 190 mm TL, nine neonate males, 163–210 mm TL, Cape Ann, Massachusetts, U.S.A., 42.64 °N, 70.64 °W; MCZAbout MCZ 34406View Materials, neonate female, 280 mm TL, Rhode Island, U.S.A., 41 ° 4 ´N, 70 ° 46 ´W; MCZAbout MCZ 35862View Materials, adult female, 805 mm TL, Buzzard Bay, Massachusetts, U.S.A., 41.56 °N, 70.74 °W; MCZAbout MCZ 35863View Materials, adult female, 790 mm TL, Massachusetts, U.S.A., 41.56 °N, 70.74 °W; MCZAbout MCZ 35864View Materials, adult male, 695 mm TL, Buzzard Bay, Massachusetts, U.S.A., 41.56 °N, 70.74 °W; MCZAbout MCZ 39679View Materials, adult female, 720 mm TL, no data; MCZAbout MCZ 58675View Materials, 10 neonate females, 188–210 mm TL, seven neonate males, 180–210 mm TL, Buzzard Bay, Massachusetts, U.S.A., 41.70 °N, 70.75 °W; MCZAbout MCZ 99596View Materials, four neonate females, 178–210 mm TL, five neonate males, 180–215 mm TL, Massachusetts, U.S.A., 43 ° 39 ´N, 69 ° 58 ´W; MCZAbout MCZ 167209, juvenile female, 540 mm TL, adult female, 660 mm TL, New England, U.S.A.; MCZAbout MCZ 502 -S, juvenile male, 470 mm TL, unknown locality (probably Europe); MCZAbout MCZ 861 -S, adult female, 860 mm TL, unknown locality; MCZAbout MCZ 51312View Materials -S, neonate female, 265 mm TL, unknown locality; NRMAbout NRM 36067View Materials, adult female, 935 mm TL, Gulf of Maine, Nahant, Massachusetts, U.S.A., 4242 ’N, 709 ’W; UFPBAbout UFPB 1480, neonate female, 222 mm TL, Virginia, U.S.A.; USNMAbout USNM 31965, adult male, 605 mm TL, British Columbia, Canada; USNMAbout USNM 72285, adult female, 670 mm TL, New River Rocks, North Carolina, U.S.A.; USNMAbout USNM 201930, adult female, 755 mm TL, two adult males, 683–690 mm TL, juvenile male, 565 mm TL, Cape Hatteras, North Carolina, U.S.A., 36 ° 90 ’N, 74 ° 65 ’W; USNMAbout USNM 201943, adult male, 670 mm TL, Cape Hatteras, Virginia, U.S.A., 36 ° 90 ’N, 74 ° 65 ’W; USNMAbout USNM 205051, adult female, 680 mm TL, adult male, 665 mm TL, Silver Bay, North Carolina, U.S.A., 35 ° 50 ’N, 75 ° 45 ’W; USNMAbout USNM 205052, adult male, 615 mm TL, Virginia, U.S.A.; USNMAbout USNM 205056, adult male, 650 mm TL, Silver Bay, South Carolina, U.S.A., 32 ° 80 ’N, 79 ° 60 ’W; USNMAbout USNM 386056, juvenile female, 548 mm TL, subadult juvenile male, 600 mm TL, Massachusetts, U.S.A., 42 ° 50 ’N, 68 ° 56 ’W; ZMBAbout ZMB 10237View Materials, adult female, 720 mm TL, Woods Hole, Massachusetts, U.S.A.; ZMHAbout ZMH 101004, juvenile male, 555 mm TL, Newfoundland, Canada, 5924 'W, 4740 'N. Western Central Atlantic (17 specimens): MCZAbout MCZ 201 -S, adult female, 880 mm TL, Minas Bay, Cuba, 21 ° 30 ´N, 77 ° 38 ´W; MCZAbout MCZ 1463 -S (holotype of Squalus barbouri  ), neonate female, 267 mm TL, Jaimanitas Beach, Cuba; USNMAbout USNM 116902, three neonate females, 143–225 mm TL, four neonate males, 143–223 mm TL, Tortugas, Florida, U.S.A.; USNMAbout USNM 201917, adult male, 685 mm TL, Silver Bay, Florida, U.S.A., 29 ° 93 ’N, 81 ° 23 ’W; USNMAbout USNM 205049, juvenile male, 452 mm TL, Silver Bay, Florida, U.S.A., 29 ° 90 ’N, 80 ° 18 ’W; USNMAbout USNM 205050, two juvenile females, 455–465 mm TL, Silver Bay, Florida, U.S.A., 29 ° 97 ’N, 80 ° 12 ’W; USNMAbout USNM 205053, juvenile male, 520 mm TL, Silver Bay, Florida, U.S.A., 29 ° 97 ’N, 80 ° 12 ’W; USNMAbout USNM 205054, adult female, 725 mm TL, Silver Bay, Georgia, U.S.A., 31 ° 73 ’N, 79 ° 63 ’W; USNMAbout USNM 205055, adult male, 615 mm TL, Silver Bay, Florida, U.S.A., 29 ° 90 ’N, 80 ° 18 ’W; USNMAbout USNM 205057, juvenile female, 470 mm TL, Silver Bay, Florida, U.S.A., 29 ° 97 ’N, 80 ° 12 ’W. Northeastern Atlantic Ocean (12 specimens): MCZAbout MCZ 85 -S, adult male, 650 mm TL, Le Havre, English Channel, France; MCZAbout MCZ 408 -S, juvenile male, 650 mm TL, Trieste, Italy; MCZAbout MCZ 466 -S, juvenile male, 520 mm TL, Maritime Alps, Nice, France; MCZAbout MCZ 467 -S, juvenile female, 590 mm TL, Maritime Alps, Nice, France; MCZAbout MCZ 468 -S, adult female, 640 mm TL; two neonate males, 140–145 mm TL; neonate male, 145 mm TL, Le Havre, English Channel, France; MCZAbout MCZ 478 -S, adult female, 720 mm TL, North Sea; MCZAbout MCZ 905 -S, adult female, 780 mm TL, Bohustan, Sweden; MCZAbout MCZ 906 -S, juvenile female, 375 mm TL, Venice, Italy; UERJ 182, adult female, 1000 mm TL, Gulf of Lion, Mediterranean Sea; UERJ 185, adult male, 710 mm TL, Gulf of Lion, Mediterranean Sea. Southeastern Atlantic Ocean (13 specimens): MCZAbout MCZ 346 -S, adult male, 1010 mm TL, South Africa; SAIABAbout SAIAB 21873, adult male, 675 mm TL, Cape Columbine, South Africa, 3259 ’S, 1736 ’E; SAIABAbout SAIAB 25918, adult male, 700 mm TL, West coast of South Africa, 3148 ’S, 1727 ’E; SAIABAbout SAIAB 26301, adult female, 670 mm TL, West coast of South Africa, 3270 ’S, 1720 ’E; SAMAbout SAM 32584View Materials, adult male, 705 mm TL, West coast of South Africa, 3108 ’S, 1651 ’E; SAMAbout SAM 33184View Materials juvenile female, 527 mm TL, West coast of South Africa, 2991 ’S, 1617 ’E; UF 46768, juvenile male, 570 mm TL, southwestern region of Cape Town, South Africa; USNMAbout USNM 197692, adult female, 733 mm TL, west coast of Cape Town, South Africa; USNMAbout USNM 199655, adult female, 640 mm TL; adult male, 700 mm TL, Gabon; ZMBAbout ZMB 21982View Materials, juvenile male, 555 mm TL, Southeastern Atlantic Ocean (no specific locality); ZMBAbout ZMB 22989View Materials, juvenile male, 562 mm TL, Southeastern Atlantic Ocean (no specific locality); ZMHAbout ZMH 151302, juvenile male, 520 mm TL, off Namibia. Northwestern Pacific Ocean (7 specimens): HUMZAbout HUMZ 87733, juvenile male, 495 mm TL, off Shiretoko, Hokkaido, Japan (dissected); HUMZAbout HUMZ 68927, adult female, 952 mm TL, Yamasedomari Fish Market, Hakodate, Hokkaido, Japan; HUMZAbout HUMZ 107865, juvenile female, 465 mm TL, off Sekinai, Kumaishi, Hokkaido, Japan; HUMZAbout HUMZ 123859, adult male, 815 mm TL, north Japan, 4400.1 ’N, 15500.1 ’E; MCZAbout MCZ 158057 -S, juvenile male, 490 mm TL, Korea; NSMT-P 92640, adult female, 740 mm TL, Northern Japan, Japan; SU 23469, neonate female, 280 mm TL, Japan. Northeastern Pacific Ocean (58 specimens): CASAbout CAS 11229View Materials, neonate male, 210 mm TL, Steinhart Aquarium, San Francisco, California, U.S.A.; CASAbout CAS 13038View Materials, neonate male, no data, Northeast California, U.S.A.; CASAbout CAS 13127View Materials, neonate male, 380 mm TL, Roberts reef, Puget Sound, Washington, U.S.A.; CASAbout CAS 19149View Materials, neonate male, 210 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21292View Materials, neonate male, 225 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21424View Materials, five neonate females, 235–290 mm TL, neonate male, 235 mm TL, two juvenile males, 300– 375 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21444View Materials, neonate female, 180 mm TL, neonate male, 195 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21468View Materials, neonate female, 290 mm TL, neonate male, 185 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21767View Materials, juvenile female, 410 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21898View Materials, neonate female, 250 mm TL, neonate male, 260 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 21971View Materials, seven neonate females, 250–285 mm TL, two juvenile females, 310–315 mm TL, four neonate males, 232–305 mm TL, three juvenile males, 310–330 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 40863View Materials, juvenile male, 340 mm TL, California, U.S.A.; CASAbout CAS 40865View Materials, neonate male, 220 mm TL, San Francisco, California, U.S.A.; CASAbout CAS 40866View Materials, neonate female, 212 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 40872View Materials, two neonate males, 245 mm TL, San Francisco Bay, California, U.S.A.; CASAbout CAS 40873View Materials, two juvenile females, 340–355 mm TL, five juvenile males, 320–390 mm TL, Roberts reef, Puget Sound, Washington, U.S.A.; CASAbout CAS 56093View Materials, four neonate females, 270 – 265 mm TL, neonate male, 260 mm TL, San Francisco Bay, California, U.S.A.; SU 13023, adult male, 710 mm TL, San Diego Bay, California, U.S.A.; SU 58376, juvenile male, 480 mm TL, Monterey Bay, California, U.S.A.; MCZAbout MCZ 167 -S, two adult males, 770–810 mm TL, San Francisco, California, U.S.A.; MCZAbout MCZ 36466View Materials -S, juvenile male, 670 mm TL, La Jolla, California, U.S. A.. Southwestern Pacific (3 specimens): MCZAbout MCZ 146 -S (holotype of Squalus tasmaniensis  ), neonate female, 245 mm TL, Hobart, Tasmania, Australia; USNMAbout USNM 176796, adult female, 780 mm TL, New Zealand; USNMAbout USNM 176800, adult male, 740 mm TL, New Zealand. Southeastern Pacific (3 specimens): MZUSPAbout MZUSP 37366, neonate female, 260 mm TL, Valdivia, Chile; SU 13381, adult male, 670 mm TL, Gregory Bay, Strait of Magallanes, Chile; USNMAbout USNM 208074, adult female, 795 mm TL, Chile, 39 ° 70 ’S, 73 ° 45 ’W.

TABLE 2. Meristic data fοr Squa ̸ us acanthias. N ∶ number οf specimens.

S0uthwestern Character N Range Atiantic Mοde N0rthwestern N Range Atiantic Mοde Mediterranean Sea N Range Mοde S0utheastern N Range Atiantic Mοde N0rthwestern N Range Pacific Mοde S0utheastern N Range Pacific Mοde
precauda1 vertebrae 7 75-81 75 3 80-85 - 3 80-81 80 3 75-78 - 2 71-72 - 2 77-77 77
cauda1 vertebrae 7 26-30 27 3 29-31 31 3 28-29 28 3 27-30 - 2 27-30 - 2 28-33 -
tοta1 vertebrae 7 101-109 101 3 111-116 111 3 108-110 108 3 104-107 - 2 99-101 - 2 105-110 -
mοnοspοndy1οus vertebrae 7 43-47 44 3 46-47 46 3 40-41 40 3 43-45 - 2 40-41 - 2 44-44 44
dip1οspοndy1οus vertebrae 7 57-63 61 3 64-70 - 3 67-70 - 3 60-63 - 2 58-61 - 2 61-66 -
upper teeth rοws (right) 16 13-15 14 6 12-14 13 4 10-14 - 8 9-14 13 4 12-14 14 2 12-14 -
upper teeth rοws (1eft) 16 13-15 13 6 13-14 14 4 10-14 14 8 9-14 13 4 12-14 12 2 12-13 -
1οwer teeth rοws (right) 16 10-12 12 6 11-14 11 4 10-13 - 8 7-12 11 4 10-11 10 2 13-13 13
1οwer teeth rοws (1eft) 16 11-12 12 6 11-14 12 4 11-13 12 8 9-12 12 4 10-12 10 2 12-13 -
upper teeth series 16 1-3 2 6 2-3 2 4 2-2 2 8 2-2 2 4 1-3 3 2 2-2 2
1οwer teeth series 16 2-3 2 6 2-2 2 4 2-2 2 8 2-2 2 4 2-2 2 2 2-2 2
prοpterygium radia1s 5 1-1 1 3 1-2 1 3 1-1 1 2 1-1 1 - - - 1 1 -
mesοpterygium radia1s 5 8-10 9 3 7-9 - 3 8-9 9 2 9-11 - - - - 1 9 -
metapterygium radia1s 1 8 - 3 6-10 6 3 7-8 7 2 6-10 - - - - 1 6 -
tοta1 pectοra1 radia1s 1 19 - 3 14-21 - 3 17-17 17 2 16-22 - - - - 1 16 -
tοta1 pe1vic radia1s 5 13-16 15 1 13 - 1 14 - 1 14 - - - - 2 13-14 -

TABLE 4. E terna 1 measurements e pressed as percentage οf tοta 1 1 engt $ (% ’ () οf frοm diferent 1 οca 1 ities. N ∶ num, er οf specimens­ x ∶ mean­ SD ∶ standard de. iatiοn.

Measurements N N0rthwestern Atiantic Range x SD Centrai Western Atiantic N N Mediterranean Sea Range x SD N N0rtheastern Atiantic Range x SD N N0rtheastern Pacific Range x SD N S0utheastern Pacific Range x SD
ΤL 17 210.0 - 910.0 618.4 203.4 1 685.0 4 520.0 - 650.0 591.3 53.9 4 570.0 - 700.0 640.0 53.5 13 260.0 - 710.0 379.6 123.8 2 670.0 - 795.0 732.5 88.4
PCL 17 76.7 - 82.1 80.1 1.3 1 80.3 4 77.9 - 80.8 79.6 1.2 4 78.1 - 82.5 80.0 2.0 13 75.0 - 81.0 78.3 1.7 2 81.8 - 82.1 81.9 0.2
PD 2 17 59.2 - 64.9 62.4 1.6 1 62.0 4 58.7 - 62.8 60.8 1.7 4 59.3 - 63.8 62.2 2.0 13 58.8 - 65.5 61.4 1.7 2 63.4 - 64.8 64.1 1.0
PD 1 17 31.7 - 36.7 33.7 1.3 1 33.6 4 29.8 - 33.1 31.9 1.5 4 32.5 - 35.0 33.8 1.0 13 32.4 - 37.1 35.5 1.2 2 34.0 - 34.3 34.1 0.3
SvL 17 48.0 - 55.9 53.1 1.8 1 52.6 4 45.2 - 54.2 51.5 4.3 4 51.4 - 53.9 52.6 1.3 13 50.0 - 53.3 51.7 1.1 2 50.9 - 54.5 52.7 2.5
PP 2 17 45.0 - 53.1 50.1 1.8 1 49.6 4 42.3 - 51.7 48.8 4.4 4 47.1 - 51.1 49.2 1.7 13 47.4 - 50.0 48.9 0.9 2 48.4 - 50.7 49.6 1.6
PP 1 17 19.9 - 23.8 21.4 1.2 1 20.4 4 19.0 - 22.4 20.8 1.4 4 17.5 - 21.9 19.9 1.9 13 21.1 - 25.9 23.2 1.3 2 20.8 - 21.6 21.2 0.6
HDL 17 20.9 - 25.2 22.4 1.4 1 20.1 4 19.4 - 22.8 21.5 1.5 4 19.7 - 21.9 21.1 1.0 13 21.3 - 24.5 22.7 1.0 2 22.0 - 23.1 22.6 0.8
PG 1 17 16.7 - 20.0 18.3 1.0 1 16.6 4 16.2 - 17.8 17.2 0.7 4 15.9 - 18.3 17.4 1.2 13 17.4 - 20.4 18.9 0.8 2 17.9 - 18.7 18.3 0.6
PSP 17 10.3 - 13.4 11.3 0.9 1 10.8 4 10.8 - 11.4 11.1 0.2 4 10.1 - 11.4 10.7 0.5 13 11.1 - 22.7 13.1 3.0 2 10.8 - 11.2 11.0 0.3
PΟB 17 5.8 - 8.2 6.9 0.7 1 6.2 4 6.1 - 7.0 6.5 0.4 4 5.5 - 6.9 6.4 0.6 13 6.7 - 8.7 7.5 0.5 2 6.4 - 7.2 6.8 0.6
PRN 17 3.9 - 5.2 4.6 0.4 1 3.8 4 3.9 - 4.9 4.3 0.4 4 3.9 - 4.9 4.4 0.4 13 4.4 - 6.0 5.1 0.5 2 3.9 - 4.5 4.2 0.4
PΟR 17 7.4 - 11.0 9.4 1.0 1 8.4 4 8.4 - 9.1 8.8 0.4 4 8.1 - 9.2 8.5 0.5 13 8.4 - 11.6 10.0 0.9 2 8.3 - 8.7 8.5 0.3
INLF 17 3.8 - 5.3 4.5 0.4 1 4.0 4 3.9 - 4.9 4.3 0.4 4 3.8 - 4.0 3.9 0.1 13 3.2 - 5.3 4.7 0.6 2 4.0 - 4.1 4.0 0.0
MΟW 17 6.2 - 7.9 7.1 0.4 1 6.7 4 6.6 - 7.9 7.4 0.6 4 7.4 - 7.7 7.5 0.1 13 7.3 - 9.4 8.0 0.5 2 7.0 - 7.1 7.1 0.1
ULA 17 1.8 - 3.0 2.3 0.3 1 2.3 4 2.1 - 2.5 2.3 0.2 4 2.1 - 2.6 2.4 0.2 13 2.1 - 3.0 2.6 0.3 2 2.0 - 2.4 2.2 0.2
INW 17 3.0 - 3.7 3.4 0.2 1 3.1 4 2.8 - 3.6 3.3 0.4 4 3.0 - 3.8 3.4 0.4 13 3.2 - 4.6 4.0 0.4 2 3.1 - 3.5 3.3 0.3
INΟ 17 6.6 - 8.2 7.2 0.5 1 5.2 4 5.9 - 7.8 6.9 0.8 4 6.6 - 7.6 7.1 0.4 13 0.9 - 9.1 7.7 2.1 2 6.6 - 7.0 6.8 0.3
E L 17 1.9 - 3.6 2.7 0.5 1 2.7 4 2.5 - 3.5 2.9 0.4 4 2.2 - 2.6 2.4 0.2 13 2.6 - 4.3 3.3 0.5 2 2.0 - 2.5 2.3 0.3
E H 17 0.8 - 3.1 1.9 0.6 1 0.9 4 0.8 - 1.8 1.2 0.5 4 1.3 - 1.9 1.6 0.3 13 1.5 - 3.0 2.2 0.4 2 1.4 - 1.6 1.5 0.1
SPL 17 0.8 - 2.0 1.2 0.3 1 0.7 4 1.0 - 1.1 1.1 0.0 4 1.1 - 1.4 1.3 0.1 13 1.0 - 1.9 1.5 0.3 2 0.9 - 1.2 1.0 0.2
GS 1 17 1.4 - 2.3 1.6 0.2 1 1.2 4 1.1 - 2.0 1.5 0.4 4 1.3 - 2.1 1.7 0.4 13 1.3 - 1.9 1.7 0.2 2 1.4 - 1.5 1.5 0.1
GS 17 1.8 - 2.6 2.2 0.2 1 2.1 4 1.9 - 2.3 2.1 0.2 4 1.9 - 2.4 2.1 0.2 13 1.9 - 2.5 2.1 0.2 2 2.0 - 2.1 2.1 0.0
IDS 17 18.7 - 26.5 22.5 2.2 1 22.6 4 22.9 - 24.8 23.5 0.9 4 17.9 - 24.6 22.1 3.0 13 17.9 - 24.1 20.6 1.9 2 24.5 - 24.6 24.6 0.1
DCS 17 9.7 - 12.0 11.2 0.6 1 11.7 4 10.9 - 13.5 12.0 1.1 4 9.7 - 12.2 10.9 1.0 13 10.5 - 13.8 12.0 0.9 2 11.2 - 12.5 11.8 0.9
PPS 17 23.0 - 28.6 25.9 1.9 1 21.9 4 21.2 - 26.4 24.6 2.5 4 21.4 - 27.2 24.4 2.4 13 20.1 - 26.8 23.9 2.1 2 25.2 - 27.6 26.4 1.7
PCA 17 19.7 - 25.3 22.7 1.2 1 24.1 4 24.0 - 27.9 25.5 1.7 4 20.6 - 24.6 22.0 1.7 13 20.4 - 25.9 22.8 1.4 2 23.1 - 23.3 23.2 0.1
D1L 17 11.2 - 13.6 12.3 0.7 1 12.1 4 11.5 - 14.3 12.5 1.2 4 12.9 - 14.8 13.4 0.9 13 11.4 - 12.9 12.0 0.5 2 12.0 - 13.6 12.8 1.1
D1A 17 8.1 - 12.0 9.6 0.9 1 8.6 4 9.1 - 11.9 10.4 1.1 4 9.3 - 10.1 9.6 0.4 13 9.0 - 10.4 9.7 0.5 2 8.8 - 10.3 9.6 1.1
D1B 17 6.2 - 8.5 7.2 0.6 1 7.0 4 7.3 - 7.8 7.5 0.2 4 7.1 - 8.6 7.8 0.7 13 6.3 - 7.9 7.0 0.5 2 7.2 - 8.7 7.9 1.1
D1H 17 6.0 - 7.6 6.8 0.4 1 5.8 4 6.4 - 9.2 7.4 1.2 4 5.9 - 6.6 6.3 0.3 13 6.5 - 8.0 6.9 0.4 2 5.8 - 6.9 6.4 0.8
D1 I 17 5.0 - 6.5 5.6 0.4 1 5.5 4 4.6 - 6.8 5.4 1.0 4 5.2 - 6.2 5.7 0.4 13 4.9 - 5.9 5.3 0.3 2 4.9 - 5.2 5.0 0.2
D1P 17 6.0 - 8.3 7.2 0.7 1 7.5 4 5.3 - 7.5 6.8 1.0 4 7.1 - 7.4 7.3 0.1 13 5.9 - 8.0 6.7 0.6 2 6.4 - 8.4 7.4 1.4
D1ES 15 1.6 - 3.1 2.0 0.4 1 1.6 4 2.0 - 4.6 2.7 1.2 4 1.5 - 2.2 1.9 0.3 13 1.5 - 2.5 2.0 0.2 2 0.8 - 1.3 1.1 0.3
D1BS 17 0.4 - 0.8 0.6 0.1 1 0.5 4 0.5 - 0.8 0.6 0.1 4 0.5 - 0.6 0.5 0.1 13 0.4 - 0.7 0.5 0.1 2 0.4 - 0.5 0.5 0.1
D2L 17 10.0 - 13.2 11.9 0.8 1 12.2 4 11.5 - 13.4 12.2 0.9 4 10.9 - 13.5 12.1 1.4 13 9.4 - 14.3 11.3 1.2 2 10.8 - 11.0 10.9 0.2
D2A 17 7.6 - 10.1 9.0 0.7 1 8.5 4 8.6 - 11.5 9.5 1.3 4 7.7 - 9.7 8.6 1.1 13 6.8 - 10.8 8.4 1.1 2 6.8 - 8.2 7.5 1.0
D2B 17 5.7 - 8.1 7.0 0.6 1 7.2 4 7.1 - 7.7 7.5 0.3 4 6.3 - 8.3 7.1 1.0 13 4.8 - 9.5 6.6 1.1 2 5.8 - 7.1 6.4 0.9
D2H 17 4.4 - 6.9 5.1 0.7 1 4.9 4 4.6 - 6.6 5.2 1.0 4 4.5 - 4.7 4.6 0.1 13 4.1 - 5.4 4.7 0.4 2 3.7 - 4.4 4.1 0.5
D2 I 17 4.4 - 6.1 5.1 0.5 1 5.2 4 4.1 - 5.8 4.8 0.7 4 4.5 - 6.1 5.1 0.7 13 4.0 - 5.4 4.9 0.4 2 4.3 - 4.9 4.6 0.5
D2P 17 4.2 - 6.7 5.4 0.6 1 5.8 4 4.3 - 5.5 5.0 0.5 4 4.6 - 5.6 5.3 0.4 13 4.1 - 5.7 5.2 0.5 2 4.7 - 5.4 5.0 0.5
D2ES 16 2.5 - 4.1 3.4 0.5 1 3.4 4 3.1 - 6.0 4.0 1.4 3 2.5 - 3.4 3.0 0.4 13 3.1 - 4.6 3.6 0.4 2 1.5 - 2.9 2.2 1.0

C x

TABLE 1. External measurements expressed as percentages of total length (% TL) of Squalus acanthias from the Southwestern Atlantic Ocean, and for holotypes of Squalus barbouri (MCZ 1463 - S) and Squalus tasmaniensis (MCZ 146 - S). TL expressed in mm. N: number of specimens; x: mean; SD: standard deviation.


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NRM

Swedish Museum of Natural History - Zoological Collections

UUZM

Uppsala University, Zoological Museum

AMNH

American Museum of Natural History

HUMZ

Hokkaido University, Laboratory of Marine Zoology

MCT

Michigan Technological University

MCZ

Museum of Comparative Zoology

MNRJ

Museu Nacional/Universidade Federal de Rio de Janeiro

NMW

Naturhistorisches Museum, Wien

ZMH

Zoologisches Museum Hamburg

CAS

California Academy of Sciences

UFPB

Departamento de Sistematica e Ecologia

USNM

Smithsonian Institution, National Museum of Natural History

ZMB

Museum f�r Naturkunde Berlin (Zoological Collections)

SAIAB

South African Institute for Aquatic Biodiversity

SAM

South African Museum

MZUSP

Museu de Zoologia da Universidade de Sao Paulo

Kingdom

Animalia

Phylum

Chordata

Class

Elasmobranchii

Order

Squaliformes

Family

Squalidae

Genus

Squalus

Loc

Squalus acanthias Linnaeus, 1758

De, Sarah T., De, Marcelo R. & Gomes, Ulisses L. 2016

2016
Loc

Squalus lebruni:

Menni 1984: 62Berg 1895: 6

1984
Loc

Squalus

Gomes 1997: 93
Figueiredo 1981: 17

1981
Loc

Spinax acanthias:

Bigelow 1948: 452

1948
Loc

Squalus fernandinus

Regan 1908: 45Molina 1782: 188

1908
Loc

Acanthias

Gill 1861: 60

1861
Loc

Acanthias

Macleay 1881: 366
Dumeril 1865: 437
Muller 1841: 83
Risso 1826: 13

1826
Loc

Squalus acanthias

Rosa 2014: 92
Viana 2011: 28
Gomes 2010: 44
Lamilla 2005: 9
Soto 2004: 73
Meneses 2003: 7
Compagno 2002: 380
Gadig 2001: 29
Soto 2001: 94
Lessa 1999: 26
Myagkov 1986: 1
Compagno 1984: 109
Kondyurin 1984: 118
Menni 1984: 62
Cadenat 1981: 46
Lucena 1981: 2
Bass 1976: 13
Garrick 1960: 520
Bigelow 1957: 30
Bigelow 1948: 455
Fowler 1936: 69
Garman 1913: 192
Regan 1908: 45
Schreiner 1903: 79
Jordan 1896: 54
Berg 1895: 5
Gill 1862: 405
Rafinesque 1810: 45Linnaeus 1758: 233

1810
Loc

Acanthorhinus acanthias:

Bigelow 1948: 452

Loc

Squalus blainvillei:

Schreiner 1903: 79