Pseudophycis palmata ( Klunzinger, 1872 )

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 : 62-72

Pseudophycis palmata ( Klunzinger, 1872)

Proposed name: Australian Red Cod (Australian Standard)

Figures 1 , 2 , 3A , 4 , 5 ; Tables 1–4

Physiculus palmatus Klunzinger, 1872: 38 . Type locality: Hobsons Bay , Victoria, Australia. Types: SMNS 1589 .

Pseudophysis barbatus (nec Günther, 1862). McCoy, 1878: 29, pl. 20 (in part; description).

Pseudophycis bacchus (nec Forster in Bloch and Schneider, 1801). Günther,1880a:28 (Twofold Bay,NSW); Waite, 1899:119 (distribution).

Pseudophycis bachus (nec Forster in Bloch and Schneider, 1801). Ogilby, 1886: 48 (list, in part); Edgar et al., 1982:32, fig.17 (description, in part); Last et al., 1983: 234, fig. 21.7 (description, in part); Paulin, 1983: 93 (distribution, in part); Hutchins and Swainston, 1986: 34, 124, fig. 113 (description); May and Maxwell, 1986: 194 (description); Paxton and Hanley in Paxton et al., 1989: 302 (list, in part); Grant, 1991: 75, fig. 164 (list, in part); Gomon in Gomon et al., 1994: 333, fig. 297 (description, in part); Yearsley et al., 1999: 87, fig. (description); Paxton et al., 2006: 616 (taxonomy, in part); Gomon in Gomon et al., 2008: 313, fig. (description, in part).

Physiculus bachus (nec Forster in Bloch and Schneider, 1801). Waite, 1904: 24; Stead, 1906: 86 (description, in part); Waite, 1907: 18, fig. (South Australia); Stead, 1908: 48, pl. 16; Waite, 1921: 67; McCulloch, 1921: 42; McCulloch, 1922: XVII, 32; Waite, 1923: 92, fig. (description, in part); Lord and Scott, 1924: 8, 43 (description, in part); Lord, 1927: 13; McCulloch, 1927: 32, pl xi, fig. 112a (list); Waite, 1928: 6 (listed); McCulloch, 1929: 129; McCulloch, 1930: 129 (in part, listed); Norman, 1935: 3; Norman, 1937: 54, 55 (in part, listed); Munro, 1961: 62, fig. 441, (description; in part); Scott, 1962: 84, fig. (description, in part); Whitley, 1962: 58; Whitley, 1964: 40 (list, in part); Walker, 1972: 2; Suda, 1973: 2150–2152 (distribution); Scott et al., 1974: 95, 96, fig. (description, in part).

Pseudophycis barbata (nec Günther, 1862). Kuiter, 1993: 59, fig. (description, in part).

Diagnosis. First dorsal fin with 10–12 rays, second dorsal fin with 47–56 rays; anal fin with 50–57 rays; total vertebrae 47– 50; nostrils located about two-thirds to three-quarters of the way from snout tip to eye; gill rakers of outer arch of moderate length, almost as long near angle as opposing gill filaments; chin barbel short, 6–17% HL; scales above lateral line in oblique series from base of first dorsal fin ray 11–15; oblique rows of scales intersecting with lateral line 96–116; pyloric caeca 8–10; 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 not extending onto side above fleshy pectoral fin base. A large species reaching at least 598 mm TL or approximately 532 mm SL ( Kemp, 2010: 26).

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 11 (10–12, rarely 10, first ray usually minute); second dorsal fin 56 (47–56, rarely 47 or 54–55); anal fin 55 (50–57, rarely less than 51); pectoral fin 22 (22–27); pelvic fin 5 (5 or 6, rarely 6); caudal fin 37 (35–41) rays; gill rakers 4 + 10 (3–4 + 9–11 = 12–15); lateral line pores not associated with individual scales; oblique scale rows intersecting with lateral line unknown (96–116), scales in oblique series above lateral line 15 (11–15), scales in oblique series below lateral line unknown (28–40); vertebrae 16 + 34 (15–18 + 31–34) = 50 (47–50); pyloric caeca 10 (8–10).

Body moderately slender, compressed laterally (fig. 2), greatest depth at anal fin origin 23.6 (18.7–26.5)% SL, tapering uniformly from second dorsal fin origin to shallow caudal peduncle; caudal peduncle moderately short, 8.9 (6.7– 10.8)% SL, strongly compressed, depth subequal to orbital diameter. Distance between middle of anus and base of anal fin slightly less than half suborbital depth. Body cavity extending to above origin of anal fin.

Head acute, of moderate size, length 26.1 (25.9–30.0)% SL, width 14.5 (15.3–21.6)% SL and depth 12.8 (10.2–18.4)% SL; snout of moderate length 7.6 (6.7–9.1)% 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 less than (equal to or less than) diameter of combined nostrils; nostrils positioned above horizontal through centre of eye; posterior opening larger than (about half size to larger than) anterior opening, separated from it by raised skin flap; most of margin (most or just posterior opening) encircled by tubular flap. Interorbital broad, slightly convex. Eye of moderate size, orbital diameter 28.0 (18.9–31.5)% HL, 1.33 (0.66–1.87) times in interorbital space, 2.41 (1.65–7.06) 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, 0.91 (0.85–1.01) times length of upper jaw. Mouth large, terminal, upper jaw terminating posterior to (at or posterior to) 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, about 0.8 to 1.3 times length of opposing gill filaments, 8.4–12.2 times in head, their inner margin denticulate. Chin barbel short, subconical 11.3 (6.0–17.4)% HL.

Small cycloid scales covering all of head, including gular region and isthmus, 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, 10th to 12th 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, narrower posteriorly. Profile of anal fin like second dorsal fin with comparable unbranched and branched rays. Anal fin enclosed in broad fleshy sheath like 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 slight 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 reaching (not quite to or to) 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, 15.1 (11.7–17.3)% SL, nearly twice length of subsequent ray, reaching vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 2A, B.) Medium brown above extending ventrally to about ventral portion 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, sometimes tinged with pink. Dorsal, caudal and distal half of anal fins medium brown; proximal half of anal fin white, especially anteriorly, with pink hue; dorsal and anal fins with fine black edge; posterior edge of caudal fin with broad black margin. Pectoral fin medium brown with semi-circular black basal spot covering dorsal 80% of proximal edge of fin, extending little if at all onto side dorsal to fin base. Pelvic fin rays white with pink hue.

Preserved colour. Mostly pale (upper half of head and body pale dusky to dusky, lower half very pale, frequently pearly white. Dorsal, caudal and distal half of anal fins pale dusky; distal edges of dorsal and anal fins with fine dark margin; posterior edge of caudal fin broadly dark; pelvic fin, pectoral fin and basal half of anal fin very pale; pectoral fin with prominent dark spot (faded in type) covering dorsal half or more of basal edge, not extending onto side of body dorsally.

Etymology. The specific epithet palmata appears to be Latin for “embroidered with palm branches”, although the reason for the name is unknown.

Distribution. Endemic to coastal temperate waters of southeast Australia at least from Port Lincoln, Spencer Gulf, South Australia (34° 44' S, 135° 52' E, SAMA F2766), to Port Stephens, New South Wales (32° 49' S, 152° 05' E, AMS I.25865-003), including all of Tasmania (fig. 3a). An unverified record at Coles Point, South Australia, 34° 22' 06” S, 135° 21' 09” E (SAMA F11864), may extend the distribution slightly farther to the west (fig. 3A). Occurs on soft bottom habitat at 2– 115 m.

Remarks. Pseudophycis palmata is closely related to the New Zealand P.bachus , with which it was confused ( Günther, 1880a: 28; McCulloch, 1921: 42; Ogilby, 1886: 48; Waite, 1904: 24). Both differ from the remaining two congeners by the possession of a black spot or blotch at the base of the pectoral fin and a truncated caudal fin margin in adults. The two also differ from the others in having the lower half of the body and the basal half or more of the anal fin mostly white, rather than tan to brown, although these areas are occasionally suffused with orange to pink in specimens of all four species. Morphometrically, the pair appear to have a deeper caudal peduncle (3.8–5.9, mean 4.7% SL vs. 3.3–5.2, mean 4.4% SL), shorter barbel (1.7–4.9, mean 3.5% SL vs. 4.7–9.6, mean 6.5% SL) and shorter or longer head (25.9–30.0, mean 27.4% SL vs. 23.3–31.6, mean 28.8 in P. barbata and 23.4–26.0, mean 24.8% SL in P. breviuscula ), as well as other relative lengths of features that correspond with the general body form. All four species in the genus are separable by vertical fin ray, scale and vertebral counts, as identified in the above key and in Table 2 . Both P. palmata and P. bachus attain a large size, as does Pseudophycis barbata (well over 600 mm SL), while P. breviuscula is the smallest species, reaching only about 150 mm SL.

The two P. bachus -like species are very similar to each other proportionally, although the eye of the Australian species is slightly larger than that of P. bachus , the orbital diameter 5.3–9.7, mean 7.4% SL vs. 4.6–8.1, mean 6.2% SL, the paired fins proportionally shorter, pectoral fin 13.5–17.7, mean 15.8% SL vs. 14.6–19.4, mean 17.1% SL and pelvic fin 11.7–17.3, mean 14.3% SL vs. 11.5–21.1, mean 16.7% SL, and caudal fin similarly shorter 9.9–15.3, mean 12.2% SL vs. 11.2–15.6, 13.7% SL. Pseudophycis palmata is readily separable from P. bachus by the smaller pectoral fin blotch that fails to extend onto the body above the pectoral fin base. It also differs subtly from P. bachus in having the distal half of the anal fin brownish with a fine black margin rather than whitish, like the basal half, and the black margin confined to the posterior lobe of the fin, if it is present at all. The Australian cognate has fewer transverse scale rows (96–116 vs. 102–136 in P. bachus ) but more pyloric caeca (8 – 10 vs. 6), anal fin rays (50–57 vs. 42–48) and second dorsal fin rays (47–56 vs. 40–45).

In the course of exploring the taxonomic identity of the Australian Red Cod, the species to which the name Physiculus palmata Klunzinger, 1872 , is referable became less and less clear. The name was regarded as a junior synonym of P. barbata for more than 80 years ( McCulloch, 1929: 129). Klunzinger’s (1872) original description is inadequate for separating the three species of Pseudophycis occurring at or in the vicinity of the type locality. A specimen (SMNS 1589) in the Stuttgart Museum, the repository of Klunzinger’s type material, registered the year the name was published, apparently collected at the type locality and clearly identified by Klunzinger as that species, was regarded as a syntype by Fricke (1992: 13; 2005: 48). It is much smaller (173 mm SL) than the 50 cm length given in the description, implying Klunzinger was aware of or had other material. The dark spot on the base of the pectoral fin that is diagnostic for the species, as well as dark pigment on the distal edge of the caudal fin, has completely faded. The only other specimen of Pseudophycis (SMNS 2242) dating from that approximate time currently in the Stuttgart collection was identified by Klunzinger in 1877 and is unlikely to have been available when the description was published. We therefore follow Fricke in regarding SMNS 1589 as the only known type.

Although the three species occurring near the type locality are clearly separable by morphological characters, the colours and markings on the type specimen that would have been diagnostic have faded. Meristic characters including vertebral numbers, fin ray and scale counts have ranges that overlap slightly in the two most likely candidate species, P. barbata and the Australian P. bachus cognate. Unfortunately, the type specimen of P. palmata has meristic values that fall in the overlap zone for all but three characters, the number of scales above the lateral line 15 (11–15 in P. bachus -like vs. 16–22 in P. barbata ), total caudal fin rays 37 (35–41 in P. bachus -like vs. 32–35 in P. barbata ) and pyloric caeca 10 (8–10 in P. bachus -like vs. 14–20 in P. barbata ) that favour its identity as the P. bachus cognate ( Table 1 ). A comparison of relative morphometric values for the type with those of these two species, however, revealed clear support for the identity of the type as the P. bachus -like species with eight of the type’s relative measurements (head length, caudal peduncle length, caudal peduncle depth, post-orbital length, interorbital width, barbel length, pelvic fin length and caudal fin length) positioned closer to the P. bachus -like species proportional curves relative to standard length (e.g. fig. 4) and curves for the same measurements (not head length) equally close to the proportional curve relative to its head length. Only predorsal fin length has clearly different proportional curves for the P. bachus -like species and P. barbata , with the predorsal value for the type of P. palmata falling equidistant between the curves of the two species for that measurement.

The diagnostics of a CAP analysis revealed that 96.8% of 124 specimens of Pseudophycis were correctly classified as their respective species based on the 20 characters examined. By species, the percentage of individuals correctly classified by the CAP discriminant model was 96.6% for P. palmata (with one misclassified as P. bachus ), 97.3% for P. bachus (with one misclassified as P. palmata ), 94.3% for P. barbata (with one misclassified as P. palmata ) and 100% for P. breviuscula . Separation of the four species of Pseudophycis is shown in fig. 5, with some overlap of P. palmata and P. bachus . SMNS 1589 (holotype of P. palmata ) is grouped within the other specimens of P. palmata and is well separated from P. barbata . Consequently, the CAP analysis and fig. 5 support the taxonomic decision to resurrect Pseudophycis palmata as the appropriate name for the Australian endemic previously thought to be conspecific with P. bachus . Overlap of P. palmata and P. bachus is consistent with the historical confusion involving the two species.

Despite its close relationship with P. bachus , the first known detailed description of this species appears to be that of McCoy (1878) who based his description of P. “ barbatus ” almost entirely on specimens of the Australian P. bachus -like species. That species was probably the common representative of the genus in the Melbourne markets at the time. Meristic and morphometric data presented by McCoy appear to be entirely attributable to specimens of P. palmata , with only the illustration ( McCoy, 1878: pl. 20) based on a specimen of P. barbata . Although several of McCoy’s specimens were either lost or obscured by early collection practices, NMV A23366- 001 is likely to be the specimen featured in the illustration, while counts and measurements were probably taken from NMV 43104, 43105, A841 and A23366-002. Other old NMV collection specimens of the new species that lack documented provenance probably comprised the remaining three. Considering the frequent occurrence of this species at the type locality of P. palmata , together with the strong morphological support discussed above, we consider the name to be applicable to the Australian P. bachus -like cognate. Paulin (1983) failed to deal with the name in the synonymies of the three species he recognised.

This species has the most restricted range of Australian Pseudophycis species, with no records of it west of Spencer Gulf, South Australia. Its latitudinal limits approach those of P. bachus in New Zealand, as do those of the two species occurring in both Australia and New Zealand where bathymetrically feasible.

Material examined. Type. Physiculus palmatus SMNS 1589 (172, holotype) Port Phillip, Hobsons Bay (northernmost section of Port Phillip Bay immediately south of Melbourne ), Victoria.

Other material. (51 non-type specimens examined for meristic or morphometric values, 90.1–380 mm SL; see Appendix 2 for additional material in Australasian collections.) Australia, New South Wales: AMS I.34462-004 (2, 230–233) north-east of Lookout Point, Twofold Bay , 37° 4.2'S, 149° 56.1'E, 100 m, J.K. Lowry and S.J. Keable, 26–27 November 1988 ; AMS I.34567-001 (146) middle of Long Beach, Batemans Bay , 35° 42'S, 150° 13' E, 50 m, J.K. Lowry and S.J. Keable, 23–24 November 1988 ; AMS I.34569-003 (112) east of Lookout Point, Twofold Bay , 37° 46' S, 149° 56' E, 50 m, J.K. Lowry and S.J. Keable, 26–27 November 1988 ; AMS I.34570-001 (3, 117–190) east of Lookout Point, Twofold Bay , 37° 46' S, 149° 55' E, 50 m, 26–27 November 1988 . Victoria: NMV A840-001 (2, 95.5–211) old collection, no data ; NMV A840-002 (2, 198–202) old collection, no data ; NMV A3859 (187) eastern Bass Strait, 6 km west-south-west of Cape Conran , 37° 49.8' S, 148° 40' E, 26 m, BSS 208 T, M.F. Gomon and R.S. Wilson , 30 July 1983 ; NMV A8870-001 (4, 95.7–113) and NMV A8870-002 (121) Bass Strait, 1 km off Lake Tyers , 37° 51.1' S, 148° 9' E, 15 m, Marine Science Laboratories, 5 June 1984 . Tasmania: AMS I.23880-004 (192) north of George Town , 40 57.90' S, 146 46.20' E, 50 m, K. Graham and FRV Kapala, 13 July 1980 ; AMS I.34952-003 (258) mouth of Fortescue Bay , 43° 7.77' S, 147° 59.47' E, 50 m, J.K. Lowry and K. Dempsey, trawl, 9–10 April 1994 ; CSIRO H 4229-01 † (380) south of Port Arthur , 43° 17.3' S, 147° 47.1' E – 43° 16.5' S, 147° 49.6' E, 115–119 m, B. Evans, 27 May 1996 ; CSIRO H 6205-02 (136) Battery Point , CSIRO wharf, 42° 53' S, 147° 20' E, H. Motomura, 20 February 2005 ; CSIRO H 7366-01 † (265) Storm Bay, east of Variety Bay on North Bruny Island , 43° 12.17'S, 147° 27.46'E, 40–45 m, hook and line, A. Pender, 6 May 2012 ; CSIRO H 7716-01 † (350) Munro Bight , 43° 11' S, 147° 59' E, 22 m, A. Pender, January 2012 ; CSIRO H 7717-01 † (362) Hinsby Beach, Taroona , 42° 57.22' S, 147° 20.82' E, 2 m, A. Pender, February 2012 ; CSIRO T 1186-02 † (8, 110-262) Nutgrove Beach, Sandy Bay , Derwent River, P . R. Last , 19 May 1980 ; CSIRO T 1417 (113 mm SL) Derwent Estuary, 7 m, P . R. Last ; NMNZ P.024340 (4: 142–197) Nakyrare Beach, Derwent Estuary , 43° 3.000' S, 147° 22.000' E, FV Ophelia, 29 March 1988 ; NMV A 1218-001 (316) , NMV A 1218-002 (318) central Bass Strait, 20 km north-north-east of North Point , 40° 31.8' S, 145° 22.8' E, 44 m, BSS 116 T, M.F. Gomon, G.C.B. Poore and P. Forsyth, 4 November 1980 ; NMV A1275 (378) central Bass Strait, 23 km east of Cape Rochon, Three Hummock Island , 40° 22.8' S, 145° 16.998' E, 40 m, BSS 112 T, M.F. Gomon, G.C.B. Poore and P. Forsyth, 3 November 1980 ; NMV A1528-001 (6, 73.5–161) and NMV A1528-002 (3, 93.6–151) central Bass Strait, 30 km north of Wynyard , 40° 33.07' S, 145° 44.69' E, 67.7 m, 4 February 1981 ; NMV A 31132 -001 † (318) off Taroona between Taroona and Alum Cliffs , 42° 57.470' S, 147° 20.797' E, 6 m, hook and line, B. Barlow, 10 May 2014 .

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 2. Pseudophycis palmata. A, B, CSIRO H 7366-01, 265 mm SL, Storm Bay, east of Variety Bay on North Bruny Island, Tasmania, Australia, fresh specimen, lateral view of body and lateral view of head and anterior body with black pectoral blotch, respectively, (photographs C. Devine, CSIRO); C, SMNS 1589, holotype, 172 mm SL, Port Phillip, Hobsons Bay, Victoria, Australia (photograph C. Struthers, NMNZ).

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.