Pseudophycis barbata Günther 1862

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 : 77-79

Pseudophycis barbata Günther 1862

Common name: Bearded Rock Cod (Australian Standard); southern bastard cod ( New Zealand)

Figures 1 , 3B , 4 , 5 , 7 ; Tables 1–4

Pseudophycis barbatus Günther, 1863: 116 . Type locality: Victoria, South Australia. Holotype: BMNH 1863.1 .15.38 (skin in alcohol).

Pseudophycis barbatus . McCoy, 1878: 29, pl. 20 (in part, description); Johnston, 1883: 126 (description); Ayling and Cox, 1982: 144, fig. (description).

Lotella grandis Ramsay, 1881: 462 . Type locality: Wollongong, New South Wales. Holotype: AMS I.696 (decision by Paulin, 1983: 94).

Lotella grandis . Steindachner, 1901: 509 (description).

Physiculus barbatus . Zietz 1909: 266; Waite, 1923: 91, fig. (description); Lord and Scott, 1924: 8, 43 (description); McCulloch, 1927: 32 (listed); Waite, 1928: 6 (listed); McCulloch, 1929: 128 (list); Norman, 1937: 55 (listed); Munro, 1938: 62, fig. 440 (description); Whitley, 1955: 119 (taxonomy); Scott, 1962: 84, fig. (description); Whitley, 1964: 40 (list); Svetovidov, 1967: 1686; Scott et al., 1974: 95, fig. (description).

? Pseudophycis breviusculus (nec Richardson, 1846) Graham, 1939: 405 (after Paulin, 1983: 93).

Pseudophycis breviusculus (nec Richardson 1846). Habib, 1975: 32–57.

Physiculus (Pseudophycis) breviculus (nec Richardson, 1846). Graham, 1956: 173 (in part, locality implies multiple species; misspelling).

Physiculus (Pseudophycis) breviusculus (nec Richardson, 1846) Whitely, 1956: 403 (in part?).

Pseudophycis barbata . Paulin, 1983: 94, fig. 10 (description, new record for NZ); Paulin and Stewart, 1985: 22; Hutchins and Swainston, 1986: 34 124, fig. 114 (description); May and Maxwell, 1986: 195 (description); Paul, 1986: 58, fig; Francis, 1988: 21, pl. 19; Paulin, 1988: 453; Paxton and Hanley in Paxton et al., 1989: 302 (list); Paulin et al., 1989: 119 (key), 255 (listed); Cohen in Cohen et al., 1990: 374 (taxonomy); Paulin in Amaoka et al., 1990: 156 (description); Kuiter, 1993: 59 (description); Gomon in Gomon et al., 1994: 334 (description); Francis, 1996:21, pl. 19; Horn,1996:157; Kuiter,1997:50 (description); Paulin, 1998: 67, fig. (description); Paul, 2000: 58; Francis, 2001: 25, pl. 18; Hutchins, 2001: 23 (description); Paxton et al., 2006: 616 (list); Hirt-Chabbert, 2006: 40; Gomon in Gomon et al., 2008: 314 (description); Roberts et al. in Gordon et al., 2009: 532 (listed); McMillan et al., 2011a: 161, 162; McMillan et al., 2011b: 94; Francis, 2012: 52; Roberts et al., 2014: 18 (list); Struthers et al. in Roberts et al., 2015: 863, fig. 107.21 (description); Roberts et al., 2015: S164; Roberts et al., 2017: 81 (list), Roberts et al., 2019: 90 (listed).

Diagnosis. First dorsal fin with 10–11 rays, second dorsal fin with 54–59 rays; anal fin with 54–63 rays; total vertebrae 48– 51; scales above lateral line in oblique series from base of first dorsal fin ray 16–22; oblique rows of scales intersecting with lateral line 123–162; pyloric caeca 14–20; caudal peduncle short 13.5–18.6% HL; chin barbel of moderate length, 16.5– 35.1% HL; caudal fin rounded without angular corners, the middle rays equal to or longer than rays above and below; no distinct, dark blotch basally on pectoral fin although fin base sometimes overall dark; underside of head and body tan to brown; anal fin tan to brown with distinct black distal margin. A large species reaching at least 620 mm SL.

Description. (Values for non-type specimens when different from type in parentheses; see Tables 2–4 for summary of selected meristic and comparative morphometric values.) First dorsal fin 10 (10–11), first ray small to minute; second dorsal fin 59 (54–59, rarely 59); anal fin 59 (54–63); pectoral fin 25 (24– 27, rarely 24); pelvic fin 5; caudal fin 33 (32–35 rays); gill rakers unknown (3–4 + 8–11 = 12–14); lateral line pores not associated with individual scales; oblique scale rows intersecting with lateral line unknown (123–162); scales in oblique series above lateral line unknown (16–22), scales in oblique series below lateral line unknown (38–56, rarely less than 45); vertebrae unknown (14–16 + 33–37, rarely 37 = 48–51); pyloric caeca unknown (14–20).

Body of moderate depth, moderately compressed laterally (fig. 7b), greatest depth at anal fin origin unknown (19.7– 29.3)% SL, tapering uniformly from second dorsal fin origin to shallow caudal peduncle; caudal peduncle short, 5.4 (5.2– 9.2)% SL, strongly compressed, depth subequal to orbital diameter. Distance between middle of anus and base of anal fin slightly less than suborbital depth. Body cavity extending to above origin of anal fin.

Head acute, of moderate size, length 23.3 (23.3–31.6)% SL, width unknown (14.9–19.8)% SL and depth unknown (11.0– 17.9)% SL; snout of moderate length 7.5 (6.5–9.6)% 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 about equal to or greater than diameter of combined nostrils; nostrils positioned above horizontal through centre of eye, both with low tubular rim; posterior opening about half size of anterior, separated from it by raised skin flap. Interorbital of moderate width, very slightly convex. Eye of moderate size, orbital diameter 20.8 (16.8–28.5)% HL, unknown (0.91–1.68) times in interorbital space, unknown (1.46–4.71) times suborbital distance, circular, upper edge of eye adjacent to dorsal margin of head in lateral view. Postorbital moderately long, 1.00 (0.52–1.21) 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, moderately short, unknown (about 0.6–0.8) times length of opposing gill filaments near angle, unknown (19–22) times in head length, slightly denticulate. Chin barbel of moderate length, subconical 21.1 (16.5–25.3)% HL.

Very small cycloid scales covering all of head 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 distinctly behind vertical through 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, last ten or so rays progressively shorter; rays of both dorsal fins mostly unbranched, only last 13 to 16 rays, 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 slightly 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 less deep 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 distinctly rounded without obvious dorsoposterior and ventroposterior corners in adults; all but anterior-most rays of similar length; base of fin covered by indistinct sheath with much smaller scales (sheath sharply demarcated from scales of caudal peduncle). Pectoral fin tip variably reaching not quite to, or past vertical through anal fin origin, sixth or seventh ray longest. Pelvic fin inserted anterior to vertical through posterior edge of preopercle; outer two rays longer than inner rays; second ray longest, 13.3 (13.3– 24.9)% SL, nearly twice length of subsequent ray, reaching vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 7A.) Medium to dark reddish brown above, extending ventrally to level of ventral portion of pectoral fin base, white below in smaller individuals, large individuals becoming brown ventrally and often much darker; lateral line not distinctively coloured. Underside of head, jaws and barbel white, tinged with pink to orange, becoming more orange to brown in larger individuals; ventral margin of suborbital dusky to dark. Dorsal and caudal fins of similar colour to brown of sides, anal fin pink or orange in smaller individuals, becoming brown like other median fins in large individuals, dorsal, caudal and anal fins with distinct broad to narrow black margin distally, margin on caudal fin broadest. Pectoral fin orange to brown with faintly darker semi-circular brown blotch basally, covering dorsal 80% of edge of fin proximally. Pelvic fin rays white with pink to orange hue, becoming brownish in large individuals.

Preserved colour. Head and body dusky to dark dusky; underside of head, belly and side adjacent to anal fin pale, especially in small specimens, extent of pale underside less in large individuals. Dorsal, caudal and anal fins dusky to dark dusky, anal fin often less dark than dorsal and caudal; distal edges of dorsal, caudal and anal fins with distinctly dark margins, dark margin on caudal fin broad; pectoral fin broadly dusky basally, becoming pale toward outer margin. Pelvic fin pale dusky.

Etymology. The specific epithet barbatus is Latin for “bearded” in reference to the chin barbel, which is characteristic of this species, its congeners and many other members of the family.

Distribution. Endemic to coastal waters of Australia and New Zealand from Rottnest Island, Western Australia (32° 01' S, 115° 30' E), to Port Stephens, New South Wales (32° 30' S, 152° 35' E), including all of Tasmania and around both the North and South Islands of New Zealand, reaching the Chatham Islands in the east (fig. 3B). A demersal species on rocky bottom at 1 to at least 520 m. Although found at considerable depths in New Zealand waters, this is a shallow reef-dwelling species in southern Australia, where it shelters in caves and beneath overhangs during daylight hours, venturing out at night to feed.

Remarks. Pseudophycis barbata has an overall darker colouration than its congeners, often with a reddish hue in fresh material, and distinct black margins to the dorsal, caudal and anal fins. In particularly dark individuals, the basal portion of the pectoral fin is similarly dark but lacks the distinct black spot present in P. bachus and P. palmata . Pseudophycis barbata is easily separable from P. breviuscula by the more numerous scales (16–22 vs. 7–9) in a transverse row between the dorsal fin origin and lateral line.

As mentioned above under Remarks for P. palmata , McCoy’s (1878: 29) treatment of P. barbata , the first published after Günther’s original description, was largely based on specimens of P. palmata , but the accompanying illustration is of a large specimen of P. barbata . Although Ramsay’s (1881) description of Lotella grandis is inadequate for a conclusive identification of the species, the lone type specimen (AMS I.696) is clearly P. barbata . Early taxonomic confusion resulted in Steindachner (1901: 509) reporting P. barbata from New Zealand as L. grandis, Norman (1937: 55) speculating that P. barbata (as Physiculus barbatus ) may be identical with P. bachus , and Graham (1938: 399) and Habib (1975) treating it in New Zealand as P. breviusculus . The last stems from our assumption that P. breviusculus does not occur in the waters of the South Island based on specimens examined and reliable identification records.

Material examined. Types. Pseudophycis barbatus BMNH 1863.1.15.38 (394, holotype) Victoria, South Australia (fig. 7a); Lotella grandis AMS I. 696 (503, holotype) Wollongong , New South Wales .

Other material. (38 specimens examined for meristic or morphometric values, 111–531 mm SL; see Appendix 2 for additional material in Australasian collections.) Australia, Victoria: CSIRO H 3791-01 † (340) south of Gabo Island , 37° 43.30' S, 149° 55.20' E – 37° 41.10' S, 149° 57.10' E, 116– 107 m, A. Graham, 17 September 1994; CSIRO H 4500-01 † (410) south of Cape Everard , 38° 07.81' S, 149° 30.87' E, 156 m, A. Williams and M. Lewis, 13 January 1997; NMV A2248 (5, 134–185) Portland Harbour , halfway along lee breakwater, adjacent to oil wharf, 38° 20.916' S, 141° 37.398' E, 7–12 m, R. Kuiter , R. Wilson and I. Head, 22 October 1981; NMV A2261-001 (4, 149–180) Portland Harbour , end of lee breakwater, 38° 21' S, 141° 36' E, 12 m, R. Kuiter , R. Wilson and I. Head, 21 October 1981; NMV A13046 (203) eastern Bass Strait , 110 km E of Paradise Beach, 38° 18.7' S, 148° 50.7' E, 201–208 m, M. Gomon, 7 August 1993 . Tasmania: AMS I.20079- 009 (90.1) south side of Rocky Cape, in National Park , 40° 51' S, 145° 31' E, 2–8 m, B.C. Russell, 8 December 1977; CSIRO H 7539-03 † (185) south-east of Orford , rock point W of Triabunna woodchip mill, 42° 32.92' S, 147° 54.80' E, 3 m, P.L Last, W. White and J. Pogonoski, 01 April 2014 . South Australia: CSIRO CA 3526 (410 mm SL) Great Australian Bight , 32° 11.8' S, 131° 22.4' E – 32° 13.4' S, 131° 22.2' E, 60 m, 7 December 1981; CSIRO H 7949-13 † (2, 47–70) Great Australian Bight , 33° 20.20' S, 130° 15.42'E – 33° 20.20' S, 130° 16.26' E, 191– 188 m, A. Graham, J. Pogonoski, M.F. Gomon and D.J. Bray, 15 December 2015 . Western Australia: WAM P.26621.003 (135) Porpoise Bay , 32°00' S, 115°30' E, 1 m. New Zealand, North Island : NMNZ P.01930 (514) Wellington, Ohau Point, Makara Coast, 41 14.00' S, 174 39.00' E, 18 m, J. Moreland, 10 June 1956; NMNZ P.014921 (2, 207–240) Three Kings Islands, west end of Great Island , 34° 8.5' S, 172° 9.1' E, 5–7 m, G.S. Hardy and A.L. Stewart, 28 November 1983; NMNZ P.037109 † (518) Mahina Knoll, NW of White Island , Bay of Plenty, 37° 20.46' S, 177° 5.43'E, 303 m, NMNZ /MARS White Island field team, 22 March 2009; NMNZ P.037124 † (450) east of north-east point; White Island , Bay of Plenty, 37° 30.96' S, 177° 13.54' E, 296 m, NMNZ /MARS White Island field team, 19 March 2009; NMNZ P.044103 † (250) southeast of Club Rock; White Island , Bay of Plenty, 37° 32.35' S, 177° 11.92' E, 99 m, NMNZ /MARS White Island field team, 16 March 2009; NMNZ P.044272 (376) and NMNZ P.044273 (361) Three Kings Islands, northern edge of Three Kings Shelf; north off Princess Islands , 114 m, NMNZ /MARS Three Kings field team, 10 March 2010; NMNZ P.045835 † (197) east off Parengarenga Harbour entrance, 34° 30.93' S, 173° 18.05' E, 176 m, RV Kaharoa , 4 August 2009; NMNZ P.046563 (339) Three Kings Islands, at anchor, Northwest Bay; Three Kings Island , 34 9.16' S, 172 8.06' E, 27 m, C.D Struthers, 6 March 2010; NMNZ P.052585 (339) Northland, off North Cape , 34° 16.47' S, 173° 0.80' E, 130–132 m, RV Tangaroa , 25 March 2011; NMNZ P.052745 † (443) eastern side of Motkokako Island, Cape Brett , 35° 9.90' S, 174° 20.27'E, S. Tindale, 7 September 2011; NMNZ P.053551 † (531) Kingfish Reef, Bay of Islands, 35° 11.20' S, 174° 15.40' E, 35 m, S. Tindale, 8 April 2012. South Island : NMNZ P.035978 (2, 234–266) West Coast, Open Bay Island, south end Popotai Islet , 43° 51.82' S, 168° 52.48' E, 14–19 m, NMNZ Jackson-Haast field team, 14 February 1999; NMNZ P.036535 (3, 186–256) West Coast, Open Bay Island, south end Popotai Islet , 43° 51.82' S, 168° 52.48' E, 14–19 m, NMNZ Jackson-Haast field team, 14 February 1999; NMNZ P.037525 (4, 111–148) West Coast, North of Murphy Beach, Whakapohai Rocks , 43° 42.27' S, 169° 14.33' E, 12–14 m, NMNZ Haast-Buller field team, 18 February 2000. NMNZ P.053753 (281) off Snares Islands , 47° 59.99' S, 166° 36.25' E, 160 m, NMNZ /MARS Auckland Island field team, 21 February 2012; NMNZ P.053754 (390) Stewart Island, North Arm, Port Pegasus, 47° 10.73' S, 167° 40.95' E, 38 m, NMNZ /MARS Auckland Island field team, 20 February 2012 .

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 4. Plots of proportional measurements (% SL) against standard lengths for non-type specimens of Pseudophycis palmata, P. barbata and holotype of P. palmata. A, caudal peduncle length; B, interorbital width; C, barbel length; D, pelvic fin length.

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 7.Pseudophycis barbata. A, NMNZ P.037124, 450 mm SL, Bay of Plenty, east of north-east point, North Island, New Zealand (photograph C. Struthers, NMNZ); B, BMNH 1863.1.15.38, holotype of Pseudophycis barbata, 394 mm SL, Victoria, South Australia (photograph © The Trustees of the Natural History Museum, London).