Memoirs of Museum Victoria 80: 59–99 (2021) Published 2021 1447-2554 (On-line) https://museumsvictoria.com.au/collections-research/journals/memoirs-of-museum-victoria/ DOI https://doi.org/10.24199/j.mmv.2021.80.04 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 MARTIN F. GOMON1, 4, CARL D. STRUTHERS2 and JODIE KEMP3 1 Museums Victoria, Melbourne Museum, 11 Nicholson Street, Carlton, Victoria, 3053, Australia. Email: [email protected] 2 Museum of New Zealand Te Papa Tongarewa, 467 Cable Street, Wellington, New Zealand. Email: [email protected] 3 25 Grigg Court, Wallington, Victoria, 3222, Australia. Email: [email protected] 4 Corresponding author Abstract Gomon, M.F., Struthers, C.D. and Kemp, J. 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: 59–99. The commercially fished Red Cod Pseudophycis bachus Forster in Bloch and Schneider, 1801, is shown on the basis of both morphological and genetic evidence to be confined to New Zealand waters. The separation of the closely related but distinct Australian cognate brings the number of species in the Australasian genus Pseudophycis to four. The name Physiculus palmatus Klunzinger, 1872, which was long thought to be a junior synonym of Pseudophycis barbata Günther, 1862, is demonstrated to have been originally proposed for the Australian endemic. Pseudophycis palmata (new combination) differs from P. bachus in having a prominent black blotch or spot basally on the pectoral fin that does not extend dorsally onto the body, in contrast to distinctly extending onto body, the second dorsal fin with 47–56 rather than 40–45 rays, the anal fin with 50–57 rather than 42–48 rays and the vertebral column with 47–50 rather than 44–46 total vertebrae. Diagnoses, colour descriptions, images and distributions for all four species and a key to members of the genus are provided. Keywords Teleostei, Actinopterygii, morphology, re-description, comparison, distribution, CO1 Introduction Yearsley et al., 1999) dealing with temperate Australian fish biodiversity followed Paulin (1983) and Cohen et al. (1990) in In an annotated catalogue of gadiform fishes of the world, treating the Australian Red Cod as conspecific with the New Cohen et al. (1990) treated the morid genus Pseudophycis as Zealand species. comprising three species: Pseudophycis bachus Forster in Bloch and Schneider, 1801, P. barbata Günther, 1863, and P. Smith et al. (2008) detected genetic differences between breviuscula Richardson, 1846, all confined to the temperate Australasian samples of Pseudophycis using the CO1 locus. waters of New Zealand and Australia. The taxonomy followed Their work included eight samples of P. bachus, four each Paulin’s (1983) review of New Zealand species of Moridae from Australia and New Zealand, six samples of P. barbata that concluded that only three are present in New Zealand and from Australia, but no samples of the third species, P. Australian waters, despite the historical confusion over which breviuscula. They found differences between the P. bachus names apply to the various Pseudophycis species. Paulin did, samples from the two countries that were equivalent to however, observe meristic variations in fin ray and scale differences between P. barbata and each of the two P. counts between New Zealand specimens and the limited bachus morphs, suggesting species-level differences number of Australian specimens of P. bachus at his disposal, between all three. These genetic discrepancies were saying “examination of a larger sample of Australian supported by work of Kemp (2010), who observed that there specimens might show some differences but, in view of the are a number of differences between individuals from variability within species of Moridae, there would probably Australian and New Zealand waters and that morphometric be little justification for separating the populations at the and meristic characteristics of Pseudophycis species in species level” (Paulin, 1983, p. 92). Subsequent authors (e.g. Australian waters have been poorly documented. She Gomon in Gomon et al., 1994; Kuiter, 1993; Last et al., 1983; compared 50 Australian specimens with 160 from New Paxton and Hanley in Paxton et al., 1989; Paxton et al., 2006; Zealand, both series identified as P. bachus, and found 60 M.F. Gomon, C.D. Struthers & J. Kemp significant differences in mean values for the second dorsal Australian National Fish Collection (CSIRO), Museums (50.2 vs. 42.8, respectively) and anal (52.1 vs. 45.0) fin ray Victoria (NMV), Museum of New Zealand Te Papa counts, as well as pyloric caeca (9.6 vs. 6.0). These Tongarewa (NMNZ), South Australian Museum (SAMA), observations provided the incentive to further compare the Staatliches Museum für Naturkunde, Stuttgart (SMNS) and Australian and New Zealand populations with each other Western Australian Museum (WAM). and with the other two long-recognised members of the Genetic material and sequences. A genetic analysis of genus, P. barbata and P. breviuscula. The outcome was a mitochondrial cytochrome oxidase subunit I (COI) gene was recognition of four easily distinguished species in the genus carried out on 72 sequences from muscle tissues, representing and the initiation of research on the scientific name for the 5 terminal taxa of the family Moridae. Australian Red Cod. The dataset included 61 sequences from the BOLD The Australian Red Cod is a relatively common demersal database (www.boldsystems.org) and 11 sequences newly species, distributed in the shelf waters of south-eastern generated for this study (see Appendix 1 for list of material Australia (Gomon in Gomon et al., 2008). It is an important and BOLD and GenBank accession numbers; vouchers for prey species for some high trophic level predators, including tissues yielding sequences in the material examined lists are little penguins, Eudyptula minor (Chiaradia et al., 2003; indicated by a dagger †), including 18 specimens of Cullen et al., 1992; Montague and Cullen, 1988), Australian Pseudophycis bachus from New Zealand, 23 P. barbata from fur seals, Arctocephalus pusillus doriferus (Gales and Australia and New Zealand, 12 P. breviuscula from Australia Pemberton, 1994; Hume et al., 2004; Kirkwood et al., 2008; and New Zealand, 12 P. palmata from Australia, and 7 Littnan et al., 2007), and New Zealand fur seals, Auchenoceros punctatus from New Zealand as a morid Arctocephalus forsteri (Page et al., 2005). The trophic role of outgroup. Australian Red Cod as prey, coupled with its relatively Genomic DNA was extracted from the 11 (7 P. bachus common occurrence, suggests that this species is particularly [including neotype], 2 P. breviuscula and 2 P. barbata) important in Australian waters. Humans include these fishes additional ethanol-preserved muscle tissues (Appendix 1) as part of the non-targeted bycatch consumed annually, using DNeasy blood and tissue kit (Qiagen). A fragment of although the species is more frequently discarded than the COI gene was PCR amplified and sequenced using retained (pers. comm. M. Tudman, Bycatch Reduction thermocycling conditions and primers (FishF2 and FishR1) Program Manager, Australian Fisheries Management from Ward et al., 2005). PCR products were purified by Authority, 2008). digestion with the ExoProStar (Illustra) or with the Zymo This contribution provides a name for the endemic DNA Clean and Concentrator (Zymo Research, USA), Australian Red Cod, a detailed description that distinguishes following the manufacturer’s instructions. PCR products it from the New Zealand P. bachus, descriptions for the four were sequenced in both directions on an ABI-3730 (Massey species in the genus and a dichotomous key to the species. University Genome Service, Palmerston North, New Zealand or Macrogen, Seoul, South Korea). Newly Methods generated sequences were deposited in GenBank (see Morphometric methodology mostly followed Paulin (1983), Appendix 1 for a list of voucher specimens and their Trunov (1990), and Markle and Melendez (1988); accession numbers). measurements were taken directly (point-to-point) except Phylogenetic analyses. All 72 sequences were individually that measurements between fins were horizontal distances aligned in ClustalW using the software Geneious Prime between vertical projections at the origins of each fin, v10.0.5. The COI alignment contained no insertions or caudal peduncle length was taken from the posterior end of deletions. Maximum likelihood, neighbour joining and the anal fin base to the hypural joint, caudal-fin length was maximum parsimony trees were used to visualise distance from the hypural joint to the fin tip, and lengths of gill relationships within and between species; node support was rakers and filaments were taken at or adjacent to the angle assessed with 1,000 bootstrap replications of each tree. For of the gill arch. In lists of specimens examined, numbers of the maximum likelihood analyses, the most appropriate specimens and their standard lengths in mm are placed in model of sequence evolution for each dataset was determined parentheses after registration numbers, except for lots using the Akaike information criterion in ModelTest; the containing only a single specimen where only the length is HKY + G model was selected