Pseudophycis bachus (Forster, 1801)

Gomon, Martin, Struthers, Carl & Kemp, Jodie, 2021, A review of the Australasian genus Pseudophycis (Gadiformes: Moridae), redescribing its four species and resurrecting the name Physiculus palmatus Klunzinger, 1872, for the Australian Red Cod, Memoirs of Museum Victoria 80, pp. 59-99 : 73-76

Pseudophycis bachus (Forster in Bloch and Schneider, 1801)

Common name: New Zealand Red Cod (new Australian name); red cod, hoka ( New Zealand)

Figures 1 , 3A , 5 , 6 ; Tables 1–4

Enchelyopus bachus Forster in Bloch and Schneider, 1801: 53. Type locality: Queen Charlotte Sound, New Zealand. No types known. Based on manuscript description of Gadus bacchus by Forster.

Gadus bacchus . Cuvier, 1817: 486; Forster in Lichtenstein, 1844: 120, 420; Whitehead, 1978: 40; Hoare, 1982: 269.

Lota bacchus Cuvier, 1829: 334 ; Richardson, 1846: 61.

Brosmius venustus Richardson and Gray, 1843: 222 (generic referral of Parkinson’s unpublished manuscript name Blennius venustus , written on the painting used by Forster for his description of Gadus bacchus ). Richardson, 1843: 27; Taylor, 1855: 413.

Enchelyopus bacchus . Lichtenstein, 1844: 419 (index only).

Lotella bacchus . Günther, 1862; 347; Hutton and Hector, 1872: 46, 115; Hector, 1875: 239; Hector, 1884: 55; Hector, 1886: 28; Hutton, 1875: 134; Thomson, 1877: 485; Thomson, 1878: 326; Thomson, 1879: 382; Dambeck, 1879: 536, 547, 555; Parker, 1882: 263; Parker, 1883: 234, 235, pl. 33; Sherrin, 1886: 16, 17, 93, 304; Thomson, 1890: 370, pl. 28; Beattie, 1891: 71, 81, 82, pl. 12, pl. 13, pl. 14, pl. 15; Thomson, 1892: 212; Ayson, 1900: 14; Mair, 1903: 319; Ayson, 1907: 22; Johnson, 1921: 473; Carter and Malcolm, 1926: 647; Malcolm, 1926: 658; Svetovidov, 1937: 1285; Svetovidov, 1948: 17, 60 (anatomy).

Pseudophycis bacchus . Günther, 1880a: 26 (Port Hardy, D’Urville Island); Murray, 1895: 599 Gill, 1893: 94, 95 100, 120 (list); Karrer, 1971: 153, 179, 180, 185, 195; Habib, 1975: 1; Ayling and Cox, 1982: 142, pl 9. (description).

Pseudophycis bachus . Günther, 1880b: 542–543, fig. 248; Ogilby, 1886: 48 (list, in part); Hutton, 1890: 282; Fitch, 1972: 570, 573; Marshall and Cohen, 1973: 490; Edgar et al., 1982: 32, fig. 17 (description, in part); Paulin, 1983: 91 (description, taxonomy, in part); Paulin and Stewart, 1985: 22; Paul, 1986: 57, fig. (in part); Francis 1988: 21, pl. 18; Paulin, 1988: 450, 451, 453; Paxton and Hanley in Paxton et al., 1989: 302 (list, in part); Paulin et al., 1989: 119 (keyed), 255 (list); Cohen in Cohen et al., 1990: 373 (taxonomy, in part); Paulin in Amaoka et al., 1990: 155, fig. 103 (description, in part); Paulin and Roberts, 1992: 130, fig. 62a (description, in part); Kuiter, 1993: 59 (description, in part); Gomon in Gomon et al., 1994: 333 (description, in part); Francis, 1996: 20, pl. 18; Horn, 1996: 151, 158; Kuiter, 1997: 50 (description, in part); Paulin, 1998: 52, fig. (description); Paul, 2000: 57 (description); Francis, 2001:25, pl. 19; Beentjes and Renwick, 2001: 315, 316; Paxton et al., 2006: 616 (taxonomy, in part); Hirt-Chabbert, 2006: 39; Gomon in Gomon et al., 2008: 313 (description, in part); Roberts et al., in Gordon et al., 2009: 532 (listed); McMillan et al., 2011a: 161, 162; McMillan et al., 2011b: 94; Francis, 2012: 5; Horn et al., 2012: 624, 625, 627–629, 631–633; Roberts et al., 2014: 18 (listed); Struthers et al., in Roberts et al., 2015: 864, fig. 107.20 (description); Roberts et al., 2015: S164 (listed); Roberts et al., 2017: 81 (listed), Roberts et al., 2019: 90 (listed).

Physiculus bacchus . Günther, 1887: 87; Goode and Bean, 1895: 365, 549; Hutton, 1896: 316; Hutton, 1904: 48 (listed); Thomson, 1906: 551; Thomson and Anderton, 1921: 74; Thomson and Thomson, 1923: 111; Frost, 1924: 609; Frost, 1926: 488, 490; Young, 1925: 370; Archey in Speight, Wall and Laing, 1927: 203; Anonymous, 1931: 32; Frost, 1933: 140; Benham, 1934: 31; Benham, 1935: 22; Benham, 1938: 56; Graham, 1938: 405; Graham, 1939: 364; Doogue and Moreland, 1960: 197, 288; Doogue and Moreland, 1961: 208, 316; Moreland, 1963: 20; Webb, 1966: 52, fig. 2.8, 70, 128, table 3.4, 164, 209, 230, 231, 234, 238–240, 257, 263, 266, 280, 294b, 294c; Webb, 1972b: 43; Webb, 1973: 307–309; Heath and Moreland, 1967: 37, 56; Whitehead, 1969: pl. 11; Anonymous, 1971: 17; Anonymous, 1972c: 47; Watkinson and Smith, 1972: 31; Knox and Kilner, 1973: 354; Vooren, 1974: 43, 44.

Physiculus bachus . Stead, 1906: 86 (in part); Waite, 1907: 18 (listed); Zietz, 1909: 266; Waite, 1909: 51, 52, 57, 134; Waite, 1911: 162, 183, 259, 265, 270, pl. 31; Thomson, 1913: 233; Phillipps, 1918: 271; Phillipps, 1921: 121, 125; Phillipps and Hodgkinson, 1922: 95; Waite, 1923: 92 (in part); Ayson, 1924: 7; Lord and Scott, 1924: 8, 43; Phillipps, 1926: 528; Lord, 1927: 13; Phillipps, 1927a: 128; Phillipps, 1927b:23, 60; Phillipps, 1927c:12; Waite, 1928: 6 (listed); McCulloch, 1929: 129 (in part); Young, 1929: 141; Anonymous, 1930: 28; Finlay, 1930: 47; Anonymous, 1934: 43; Anonymous, 1935: 34; Norman, 1935: 3; Benham, 1936: 26; Hefford, 1936: 71, 74; Cunningham, 1937: 898, Norman, 1937: 54, 55 (in part, listed); Shorland, 1937: 223; Wilson, 1937: 31; Johnston, 1938: 47; Munro, 1938: 62; Graham, 1939 b: 364; Fowler, 1940: 758; Rapson, 1940: 35; Phillipps, 1947: 42; Phillipps, 1948: 129; Shorland, 1948: 109; Laird, 1949: 14, 19, 36, 37, 39, 53, 56, 60, 61, 137, 145, 146; Phillipps, 1949: 24, 59; Shorland, 1950: 35; Laird, 1951: 287, 298, 306, 308; Laird, 1952: 589, 590, 595, 596, 600; Graham, 1953: 166, 173, 399; Manter, 1954. 498, 545, 547, 549, 559; Robinson, 1955: 10, 71, 105, pls 9 and 14; Anonymous, 1957: 69; Kaberry, 1957: 90; Moreland, 1957: 34, 36; Parrott, 1957: 47, 175; Anonymous, 1958: 73; Parrott, 1958: 117; Anonymous, 1959: 70; Robinson, 1959: 152; Anonymous, 1960: 70; Parrott, 1960: 67, 164; Meglitsch, 1960: 321–323; Scott, 1962: 84 (in part); Gorman, 1963: 29; Graham, 1963: 167; Doogue and Moreland, 1964: 205, 311; Street, 1964: 18; Anonymous, 1965: 15, 22; Churchman, 1965: 56; Doogue and Moreland, 1966: 205, 311; Elder, 1966: 96, 97; Howell, 1966: 33; Paul, 1966: 372, 373; Svetovidov, 1967: 1685, 1686, 1689, 1690; Sorensen, 1968: 148; Tong and Elder, 1968: 64; Doogue and Moreland, 1969: 205, 311; Russell, 1969: 108; Cowper, 1970: 45; Godfriaux, 1970: 257; Iwai et al., 1970: 21; Shuntov, 1970: 373, 374, 376; Sorensen, 1970: 4, 17; Coakley, 1971: 24; Russell, 1971a: 9, 19, 21, fig. 3, 38, 41, 94, 173, table 10, fig. 13, 191, 197; Russell, 1971b: 83; Anonymous, 1972a: 229; Anonymous, 1972b: 187–189, 278; Doak, 1972: 19, 101, 131; Hewitt and Hine, 1972: 92; Iwai et al., 1972: 29, 36; Shuntov, 1972: 339; Walker, 1972: 2; Webb, 1972a: 16; Suda, 1973: 2150–2152; Waugh in Williams, 1973: 257, 274; Williams, 1973: 427; Godfriaux, 1974: 502; Ryan, 1974: 133, 135; Scott et al., 1974: 95, 96 (in part).

Lotella bachus . Ayson, 1908: 28; Rendahl, 1926: 2.

Physiculus backus . Fowler, 1940: 758 (near Loba, New Zealand).

Physiculus (Pseudophycis) bachus . Graham, 1956: 166; Whitley, 1956: 403; Whitley, 1968: 40 (list).

Pscudophycis bachus . Doogue and Moreland, 1982: 218, fig. (misspelling).

Diagnosis. First dorsal fin 11–13 rays, second dorsal fin 40–45 rays; anal fin 42–48 rays; total vertebrae 44–46; scales above lateral line in oblique series from base of first dorsal fin ray 10–12; oblique rows of scales intersecting with lateral line 102–136; pyloric caeca 6; caudal peduncle moderately short 28.0–38.9% HL; barbel short, 8.6–16.9% HL; caudal fin truncate with angular corners in specimens larger than about 150 mm SL, middle rays shorter than rays extending to corners; dark blotch basally on pectoral fin extending well onto side above fleshy pectoral fin base; underside of head and body whitish; anal fin mostly whitish with faint narrow black distal margin only posteriorly, if at all. A large species reaching at least 650 mm SL.

Description. (Values for neotype with those for non-type specimens when different in parentheses; see Tables 2–4 for summary of selected meristic and comparative morphometric values.) First dorsal fin 11 (11–13, rarely 13, first ray usually minute); second dorsal fin 42 (40–45, usually 42 or 43); anal fin 45 (42–48, usually 45–47); pectoral fin 24 (22–25, rarely 22); pelvic fin 5; caudal fin 36 (33–39); gill rakers 4 + 10 (4–5 + 10–12 = 14–16); lateral line pores not associated with individual scales; oblique scale rows intersecting with lateral line 112 (102–136); scales in oblique series above lateral line 11 (10– 12), scales in oblique series below lateral line 30 (26–34, rarely 33 or 34); vertebrae 16 + 29 (16–17 + 27–30 = 44–46); pyloric caeca 6.

Body moderately slender, compressed laterally (fig. 6), greatest depth at anal fin origin 19.5 (18.4–23.7)% SL, tapering uniformly from second dorsal fin origin to shallow caudal peduncle; caudal peduncle short, 8.5 (7.7–10.5)% SL, strongly compressed, depth less than orbital diameter. Distance between middle of anus and base of anal fin less than suborbital depth. Body cavity extending posteriorly to above origin of anal fin.

Head acute, of moderate size, length 26.5 (25.9–29.6)% SL, its width 14.4 (13.4–17.7)% SL and depth 11.7 (10.2– 15.6)% SL; snout of moderate length 7.5 (7.1–9.0)% SL, not projecting in advance of upper jaw, rounded in dorsal view. Nostrils small, located about two-thirds to three-quarters of the way from snout tip to eye, distance from nostril to orbital margin equal to or greater than diameter of combined nostrils; nostrils positioned above horizontal through centre of eye; posterior opening larger than anterior, separated from it by raised skin flap; posterior opening with tubular flap encircling margin. Interorbital of moderate width, very slightly convex. Eye of moderate size, orbital diameter 21.1 (17.0–30.4)% HL, 1.58 (0.98–2.00) times in interorbital space, 2.20 (1.50–4.25) times suborbital distance, circular, upper edge of eye adjacent to dorsal margin of head in lateral view, transparent skin covering eye faintly pigmented near orbital dorsal margin. Postorbital moderately long, 1.01 (0.89–1.22) times length of upper jaw. Mouth large, terminal, upper jaw terminating at vertical through posterior margin of eye. Jaw teeth caniniform, slightly curved, depressible, band of up to four or five irregular rows in upper jaw with distinct hiatus at symphysis; teeth in outer row more regularly arranged than in inner rows; band tapering near rear of jaw. Teeth of lower jaw similar in form to those of upper jaw; band of two or three rows with broader patch on either side of symphysis, tapering to single, widely spaced row posteriorly; rows almost contiguous across symphysis. Vomerine teeth absent. Opercular bones strong; upper extremity of gill opening at horizontal through middle of eye; gill membranes continuous across isthmus. Gill rakers on outer arch slender, of moderate length, almost as long near angle as opposing gill filaments, 12.8 (9.5–12.8) times in head, their inner margin denticulate. Chin barbel short, subconical 12.0 (8.6–16.9)% HL.

Small cycloid scales covering all of head, including isthmus and gular region, and body except for branchiostegal membranes, surface of maxilla and premaxilla, lower lip and distal parts of fins; thick mucus covering obscuring scales and pores in freshly preserved material. Most head pores tiny, following main cephalic sensory canals; row of slightly raised pores from nostrils to tip of snout and then posteriorly just above lower edge of suborbital; row of enlarged mandibular pores on underside of lower jaw. Lateral line comprising widely spaced pores on short tubes arising from narrow scale-less gap, anterior end curved upwards slightly, then gradually descending to lateral midline below posterior 25% of second dorsal fin, remaining on lateral midline posteriorly.

Membranes of first and second dorsal fins continuous at base; first dorsal originating vertically above pectoral fin base; anterior two-thirds of second dorsal of uniform height, 7th to 9th ray from posterior end of fin longest, its height about 1.4 times length of 2nd ray at anterior end of fin, subsequent rays progressively shorter; rays of both dorsal fins mostly unbranched, only last 13 to 16 rays of second dorsal, apart from last one to three, branched. Fleshy, fine scale-covered basal sheath on third or more of first dorsal and anterior portion of second dorsal fins; fin rays interconnected by membranes to tips, sheath gradually decreasing in coverage and thickness posteriorly, encompassing about half of second dorsal fin near its insertion; sheath extending onto body for anterior third of combined dorsal fin base, broadest anteriorly, narrowing posteriorly. Profile of anal fin like that of second dorsal fin, although shallower posteriorly, with comparable unbranched and branched rays. Likewise, anal fin enclosed in broad fleshy sheath that is like that of dorsal fins; sheath also extending onto body for anterior third or less of fin. Caudal fin truncate with distinct dorsoposterior and ventroposterior corners in adults, posterior margin with little if any convex curve; longest rays to corners, middle rays about 85–90% length of longest rays; fin more rounded in juveniles; base of fin covered by indistinct sheath sharply demarcated from scales of caudal peduncle. Pectoral fin tip variably reaching almost to, to or past vertical through anal fin origin, sixth or seventh ray longest. Pelvic fin inserted anterior to vertical through posterior edge of preopercle (more posteriorly in some specimens); outer two rays longer than inner rays; second ray longest, 16.0 (11.5–21.1)% SL, nearly twice length of subsequent ray, reaching past a vertical through first dorsal fin origin.

Fresh colour. (Based on images of neotype; fig. 6A, B.) Medium brown above extending ventrally to level of ventral edge of pectoral fin base, white below, suffused with pinkish hue, especially above anal fin base; lateral line slightly paler at least anteriorly. Underside of head, jaws and barbel white, tinged with pink. Dorsal and caudal fins medium brown, caudal paler with reddish tinge ventrally; anal fin white with pink hue; dorsal fins with fine black distal margin; caudal fin margin with broader, more diffuse black margin; anal fin with faint black distal margin, darkest posteriorly. Pectoral fin medium brown with semi-circular black basal spot covering dorsal half of fin edge proximally, extending well onto side of body above fin base; ventroposterior margin of fin pale to translucent. Pelvic fin rays white with pink hue.

Preserved colour. Upper half of head and body pale dusky to dusky, lower half very pale, frequently pearly white. Dorsal and caudal fins pale dusky; distal edge of dorsal fin with fine dark margin; posterior edge of caudal fin broadly dark; anal fin very pale, distal edge of fin posteriorly with fine dark margin in some; pelvic fin, pectoral fin and basal half of anal fin very pale; pectoral fin with prominent dark spot covering dorsal half or more of basal edge that extends onto side above pectoral fin base.

Etymology. The name bachus is an altered spelling of bacchus , the Greek god of wine, adopted for the wine-red colouration assumed shortly after capture but lost soon after death.

Distribution. Endemic to New Zealand, occurring around both the North and South Islands, and reaching the Chatham Islands in the east (fig. 3A). A demersal species on soft bottom at 2–570 m depth.

Remarks. Pseudophycis bachus was originally described as Enchelyopus bachus Forster in Bloch and Schneider, 1801, based on Forster’s manuscript description of “ Gadus bachus ”, subsequently published by Lichenstein (1844: 120). Forster’s drawing No. 180, on the back of which is pencilled “ex Queen Charlotte Sound”, is evidently an illustration of the fish on which the description was based. Although both Bloch and Schneider (1801: 53) and Lichtenstein gave the origin of the specimen simply as “in the seas around New Zealand ”, Whitehead (1969: pl. 11), Paulin (1983: 91) and Fricke et al. (2019) regarded Queen Charlotte Sound, the presumed collection locality of the specimen implied in the note on Forster’s illustration, as the type locality. However, Hoare’s (1982: 269) reproduction of Forster’s Resolution Journal has a footnote stating “ Gadus bacchus of Descr. Anim., p. 120 and Forster drawing 180 (undated). The description is from a specimen of 22 inches (559 mm), dated 13 April 1773 ”. That date puts the type locality as Dusky Sound from a larger fish. Paulin (1983: 92) provided a detailed discussion of the origin and various iterations of Bloch and Schneider’s name.

An unfinished drawing by Parkinson (vol. 2, no. 5, ex Totarra ‘nue, 404 mm tot. l.) reproduced by Whitehead (1969: pl. 11; here as fig. 6C), has “ 18, Blennius venustus ” pencilled on the reverse side and is the basis for Richardson and Gray’s (1843: 222) Brosmius venustus . The fish figured is clearly an example of P. bachus , despite the incomplete nature of the illustration. Totaranui is the Maori name for Queen Charlotte Sound. The scientific name is regarded as unavailable because no distinguishing features were provided by the authors ( Fricke et al., 2019). The illustrated characters agree with those of specimens from New Zealand identified as P. bachus .

Because no type specimen of P. bachus is known ( Fricke et al., 2019), the designation of a neotype for P. bachus is considered justified. A specimen (NMNZ P.054828, 351 mm SL; fig. 6a) collected from 2.5 km off the coast from Whanganui River bar, Whanganui, New Zealand, is proposed as the neotype for P. bachus . Accordingly, Whanganui, New Zealand, becomes the type locality of P. bachus under Article 76.3 (ICZN, 1999), replacing the original type locality Queen Charlotte Sound, New Zealand.

Material examined. Neotype. NMNZ P.054828 † (351) 2.5 km off the coast from Whanganui River bar, Whanganui, New Zealand, 39° 53' S, 174° 49' E, 18 m, C. Papple, 10 October 2012.

Other material. (38 non-type specimens examined for meristic or morphometric values, 110–539 mm SL; see Appendix 2 for additional material in Australasian collections.) New Zealand, North Island: NMNZ P.001718 (539) off Castlepoint, Wairarapa , 40° 54' S, 176° 13' E, 73–110 m, F. Abernethy, 8 July 1955 ; NMNZ P.033899 (5, 121–202) off Raglan , Waikato, 37° 48.33' S, 174° 34.00' E, RV Kaharoa , 61–63 m, 26 October 1996 . South Island: NMNZ P.025037 (2, 479–512) Blueskin Bay, Otago Peninsula , Otago, 45° 43.28' S, 170° 40.33' E, 20–22 m, C.D. Roberts and C.D. Paulin, 9 May 1990 ; NMNZ P.025129 (4, 137–192) Entrance mole, Otago Harbour , Otago, 45° 46.45' S, 170° 43.23' E, 6–9 m, C.D Roberts, 9 May 1990 ; NMNZ P.027580 (239) Harrold’s Bay, Halfmoon Bay, Stewart Island , 46° 53.75' S, 168° 9.25' E, 2–5 m , NMNZ Stewart Island field team, 4 March 1992 ; NMNZ P.032385 (2, 328–395) Mooring, head of Gold Arm, Charles Sound , Fiordland , 45° 8.55' S, 167° 8.78' E, 5–10 m , NMNZ Fiordland 1995 field team, 24 March 1995 ; NMNZ P.032393 (3, 292–309) Mooring at Toe Cove, head of Nancy Sound , Fiordland , 45° 10.55' S, 167° 8.85' E, 25 m , NMNZ Fiordland 1995 field team, 26 March 1995 ; NMNZ P.035991 (388) Flowerpot Rock, Jackson’s Bay , West Coast, 43° 58.03' S, 168° 37.30' E, 5–8 m , NMNZ Jackson-Haast 1999 field team, 7 February 1999 ; NMNZ P.044338 † (136) south of Timaru , Canterbury, 44° 45.04' S, 171° 18.40' E, 26–31 m, RV Kaharoa , 3 June 2007 ; NMNZ P.044339 † (176) south off Timaru , Canterbury, 44° 45.04' S, 171° 18.40' E, 26–31 m, RV Kaharoa , 3 June 2007 ; NMNZ P. 047709 (229) Lyttelton Port , Canterbury, 43° 36.33' S, 172° 43.10' E , NIWA Port Survey , 4 November 2004 ; NMNZ P.049392 (343) c. 7 km eastnorth-east of Kaikoura Peninsula , 42° 24.77' S, 173° 48.22' E, 93 m, C.D. Struthers, 16 November 2010 ; NMNZ P.049401 (370) c. 6 km north-east of Kaikoura Peninsula , 42° 23.60' S, 173° 47.59' E, 49 m, C.D. Struthers, 16 November 2010 ; NMNZ P.049402 (545) c. 5 km east-south-east of Kaikoura Peninsula , 42° 26.77' S, 173° 46.39' E, 94 m, C.D. Struthers, 17 November 2010 ; NMNZ P.049403 (519) c. 5 km south off Kaikoura Peninsula , 42 28.09' S, 173 43.02' E, 91 m, C.D. Struthers, 19 November 2010 ; NMNZ P.049406 (485) and NMNZ P.049407 (467) c. 3 km south off Kaikoura Peninsula , 42° 24.72' S, 173° 45.22' E, 47 m, C.D. Struthers, 17 November 2010 ; NMNZ P.049678 (358) Bay View, end of Franklin Road , Hawke's Bay, 10–15 m, C.D. Struthers and D.H. Struthers, 27 December 2010 ; NMNZ P.052508 (297) and NMNZ P.052509 (290) south of Pegasus Canyon mouth, Canterbury Bight , 43° 33.99' S, 173° 33.44' E, 86 m , RV Tangaroa , 13 May 2011 ; NMNZ P.053748 † (383), NMNZ P.053749 † (405), NMNZ P.053750 † (368), NMNZ P.053751 † (373) and NMNZ P.053752 † (337) Port Pegasus; Stewart Island , 47° 13.20' S, 167° 41.38' E, 64 m, C.D. Struthers, 20 February 2012 . Chatham Rise: NMNZ P.044348 † (273) central Chatham Rise , 43° 47.42' S, 179° 30.01' W, 327–328 m, RV Tangaroa , 31 December 2007 ; NMNZ P.044349 † (382) central Chatham Rise , 43° 47.42' S, 179° 30.01' W, 327–328 m, RV Tangaroa , 31 December 2007 ; NMNZ P.044350 † (297) central Chatham Rise , 43° 43.78' S, 179° 24.00' W, 367–375 m, RV Tangaroa , 31 December 2007.

Figures

Figure 1. Hasegawa-Kishino-Yano maximum likelihood tree of 65 COI sequences from four purported species of Pseudophycis, along with seven sequences for an outgroup species, Auchenoceros punctatus. Percentage bootstrap values followed by bootstrap values in p-distance neighbour joining and maximum parsimony trees are given for all well supported nodes (2 of 3 exceeding 80%). Specimen numbers are those of registered museum vouchers or sequence numbers in the Barcode of Life Database (BOLD, http://www.barcodinglife.org/).

Figure 3. Collection localities for specimens of Pseudophycis spp. in museums. A, P. bachus, green and P. palmata, orange; B, P. barbata, blue; C, P. breviuscula, red.

Figure 5. Scatter plot of canonical analysis of principal coordinates scores on the first and second canonical axes for 124 specimens of Pseudophycis, including types.

Figure 6. Pseudophycis bachus. A, B, NMNZ P.054828, neotype, 351 mm SL, 2.5 km off coast from Whanganui River bar, Whanganui, New Zealand, fresh specimen, lateral view of body and anterior body showing black pectoral blotch respectively (photographs C. Struthers, NMNZ); C, unpublished line drawing by S. Parkinson of Enchelyopus bachus Forster in Bloch and Schneider, 1801 (reproduced from Whitehead, 1969: pl. 31).