Published December 31, 2021 | Version v1
Taxonomic treatment Open

Pseudophycis breviuscula

Description

Pseudophycis breviuscula (Richardson 1846)

Common name: Bastard Red Cod (Australian Standard); northern bastard cod (New Zealand)

Figures 1, 3C, 5, 8; Tables 1–4

Lota breviuscula Richardson, 1846: 61, Pl. 38 (figs 1–2). Type locality: Bay of Islands, New Zealand. Holotype: BMNH 1855.9.19.1182.

Pseudophycis breviusculus. Günther, 1862: 350; Günther, 1863: 116; Hector in Hutton and Hector, 1872: 116, pl. 8; Hutton in Hutton and Hector, 1872: 47; Dambeck, 1879: 536, 547, 555; Hector, 1884: 55; Hector, 1886: 28; Sherrin, 1886: 304; Gill, 1893: 120 (list); Hutton, 1904: 48; Thomson, 1906: 551; Young, 1925: 370; Graham, 1938: 405; Graham, 1939 b: 365; Ayling and Cox, 1982: 143, fig. (description).

Austrophycis megalops Ogilby, 1897: 91. Type locality: Maroubra Bay, New South Wales, Australia. Holotype: AMS I.3655.

Austrophycis megalops. McCulloch, 1927: 32 (list); McCulloch, 1929: 129 (list); Munro, 1938: 62, fig. 443 (description); Whitley, 1964: 40 (list); Paxton and Hanley in Paxton et al., 1989: 302 (list).

? Pseudophycis breviusculus. Graham, 1939: 399.

Physiculus (Pseudophycis) breviusculus. Graham 1956: 173, 174 fig.; Whitley, 1956: 403 (list); Svetovidov, 1967: 1686, 1689; Whitley, 1968: 40 (list).

Pseudophycis breviuscula. Paulin, 1983: 93 (description, taxonomy); Francis, 1988: 21; Paxton and Hanley in Paxton et al., 1989: 302 (list); Paul, 1986: 58; Paulin, 1988: 450, 451, 453; Paulin et al., 1989: 119 (key), 255 (list); Cohen in Cohen et al., 1990: 375 (taxonomy); Paulin and Roberts, 1992: 128, fig. 68a (description); Kuiter, 1993: 60 (description); Gomon in Gomon et al., 1994: 335 (description); Francis, 1996: 21; Paul, 2000: 58; Francis, 2001: 25; Hutchins, 2001: 23 (distribution); Sazonov, 2001: 293 (taxonomy of A. megalops); Paxton et al., 2006: 616 (list); Gomon in Gomon et al., 2008: 314 (description); Roberts et al. in Gordon et al., 2009: 532 (list); McMillan et al., 2011a: 161, 162; McMillan et al., 2011b: 94; Francis, 2012: 52; Struthers et al., in Roberts et al., 2016: 864, fig. 107.22 (description); Roberts et al., 2015: S164; Roberts et al., 2017: 81, Roberts et al., 2019: 90 (listed).

? Austrophycis marginata (nec Günther, 1878). Cohen in Cohen et al., 1990: 356 (taxonomy of A. megalops).

Diagnosis. First dorsal fin with 8–10 rays, second dorsal fin with 44–52 rays; anal fin with 49–56 rays; total vertebrae 42– 45; nostrils adjacent to anterior margin of orbit, interval between nostril and orbital margin less than diameter of combined nostrils; posterior nostril with raised anterior border but without tubular flap posteriorly; gill rakers of outer arch moderately short, those near angle much shorter than opposing gill filaments; caudal peduncle short 28.8–33.4% HL; chin barbel of moderate length, 23.4–30.5% HL; scales above lateral line in oblique series from base of first dorsal fin ray 7–9; oblique rows of scales intersecting with lateral line 77–93; pyloric caeca 7–8; caudal fin rounded without angular corners, the middle rays equal to or longer than the rays above and below; no distinct dark blotch at base of pectoral fin; underside of head, body and anal fin tan to brown, anal fin with fine black distal margin. A small species reaching at least 151 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 9 (8–10, rarely 8), first ray often minute; second dorsal fin 46 (44–52, rarely less than 46); anal fin 50 (49–56); pectoral fin 21 (20–23); pelvic fin 4 (4–5); caudal fin 26 (26–32, rarely 26); gill rakers 3 (2–3) + 8 (7–8) = 9–12; lateral line pores not associated with individual scales; oblique scale rows intersecting with lateral line 93 (77–93); scales in oblique series above lateral line 7 (7–9), scales in oblique series below lateral line 23 (22–28, rarely 28); vertebrae 13 (11–14, rarely 11) + 31 (30–33) = 42–45, rarely 42; pyloric caeca 7–8.

Body of moderate depth, moderately compressed laterally (fig. 8), greatest depth at or just in advance of anal fin origin 22.8 (20.6–23.3)% SL, tapering gradually from second dorsal fin origin to posterior portion of fin and then more steeply to shallow caudal peduncle; caudal peduncle short, 4.8 (5.7– 8.5)% SL, strongly compressed, depth subequal to orbital diameter. Distance between middle of anus and base of anal fin less than diameter of posterior nostril. Body cavity extending to above origin of anal fin.

Head acute, moderately short, length 25.8 (23.4–26.0)% SL, its width 16.6 (12.4–17.9)% SL and depth 12.1 (10.8–15.2)% SL; snout moderately short 5.9 (5.1–7.0)% SL, not projecting in advance of upper jaw, rounded in dorsal view. Nostrils small, located on posterior half of snout just in front of eye, distance from nostrils to orbital margin less than diameter of combined nostrils; nostrils positioned above horizontal through centre of eye; diameter of posterior opening about 1.5 diameter of anterior, separated from it by raised skin flap; posterior opening without tubular flap encircling posterior margin. Interorbital of moderate width, slightly convex. Eye of moderate size, orbital diameter 27.4 (24.7–35.7)% HL, 0.91 (0.53–1.10) times in interorbital space, 3.72 (2.20–5.78) times suborbital distance, circular, upper edge of eye just below dorsal margin of head in lateral view, skin covering eye unpigmented. Postorbital moderately long, 1.09 (0.97–1.50) times length of upper jaw. Mouth large, terminal, upper jaw terminating just posterior to vertical through posterior margin of eye. Jaw teeth small, caniniform, slightly curved, depressible, band of up to four or five irregular rows in upper jaw with narrow hiatus at symphysis; band tapering near rear of jaw. Teeth of lower jaw similar in form to those of upper jaw; band of several rows with broader patch on either side of symphysis, tapering 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 club-shaped, moderately short, about 0.3–0.8 times length of opposing gill filaments at angle, 15–23 times in head, slightly denticulate. Chin barbel of moderate length, subconical 21.0 (19.6–30.5)% HL.

Moderately 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; very 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 extending from tip of snout posteriorly just above lower edge of suborbital; row of enlarged mandibular pores on underside of lower jaw. Lateral line comprising pores on tubes arising above upper end of gill opening, anterior end curved upwards slightly, then gradually descending to lateral midline below posterior third 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 1.2 times length of 2nd ray at anterior end of fin, last eight or so rays progressively shorter; rays of both dorsal fins mostly unbranched, only about last 14 or 15 rays, except last few, branched. Fleshy basal sheath on base of first dorsal and anterior portion of second dorsal fins not especially prominent with scales apparently confined to basal margin of fin; fin rays interconnected by membranes to tips; sheath more obvious on body below much of combined dorsal fin base, broadest anteriorly, narrowing posteriorly. Profile of anal fin similar to that of second dorsal fin, with comparable unbranched and branched rays. Fleshy sheath on and below anal fin similar to that of dorsal fins. Caudal fin rounded without distinct dorsoposterior and ventroposterior corners, posterior margin distinctly convex; all but anteriormost rays of similar length; base of fin covered by indistinct sheath (sheath demarcated from scales of caudal peduncle). Pectoral fin tip reaching past vertical through anal fin origin, seventh or eighth ray longest. Pelvic fin inserted anterior to vertical through posterior edge of preopercle; outer two rays longer than inner rays; second ray longest, 17.6 (16.6–21.6)% SL, two to four times length of subsequent ray, reaching vertical through first dorsal fin origin.

Fresh colour. (Based on images of non-type material; fig. 8B.) Medium brown to pale brown above, paler below with white chest and belly in smaller individuals, bronze in larger individuals; lateral line not distinctive. Underside of head and jaws brownish; ventral edge of suborbital darker brown; barbel white. Dorsal, anal and caudal fins very pale brown in smaller individuals, darker in large individuals; fins with distinct broad to narrow black margin distally, more diffuse in large individuals. Pectoral fin orange with very faint semi-circular brown blotch basally. Pelvic fin rays white with pink to orange hue.

Preserved colour. Head, body and fins pale dusky to dusky, chest and belly very pale, ventral edge of suborbital darker; distal edges of dorsal, caudal and anal fin with diffuse darker margins.

Etymology. The name breviuscula is Latin for “rather short”, probably in reference to the relatively small maximum size of the species compared with others of the genus, although Richardson separated it from other New Zealand morids simply by fin counts.

Distribution. Endemic to coastal waters of Australia and New Zealand, from Perth, Western Australia (32° 01' S, 115° 30' E), to Tweed Heads, New South Wales (28° 14' S, 153° 50' E), including all of Tasmania and around the North Island of New Zealand (fig. 3C). A demersal reef-dwelling species at 0–273 m, although most often encountered at less than 100 m on rocky and boulder reefs (Struthers et al. in Roberts et al., 2016: 864).

Remarks. P. breviuscula is the smallest of the four species in the genus, reaching less than half the maximum length of its congeners. It also occurs in slightly warmer waters compared with the other three members of the genus, reaching well onto the northern New South Wales coast in eastern Australia and around New Zealand’s North Island.

Paulin (1983: 94) provided a thorough synonymy of P. breviuscula from a New Zealand perspective but failed to mention Austrophycis megalops Ogilby, 1897, which was based on a specimen from New South Wales, Australia. This omission was possibly due to the widespread uncertainty about the identity of Waite’s shrivelled 65 mm type specimen. Some authors regarded Ogilby’s species as congeneric with Günther’s (1878: 19) A. marginatus based on material from the south-eastern Pacific and others thought the two were likely to be conspecific (Cohen in Cohen et al., 1990: 356). Sazonov (2001: 343) reported that he had examined the type specimen and found it to be P. breviuscula relegating the name A. megalops to synonymy with that species.

Material examined. Types. Lota breviuscula BMNH 1855.9.19.1182 (152, holotype) Bay of Islands, New Zealand (fig. 8a); Austrophycis megalops AMS I. 3655 (65, holotype) Maroubra Bay, New South Wales, Australia, 33° 57' S, 151° 16' E, T. Whitelegge, 1897 (after Sazonov, 2001: 343, holotype, not re-examined for this study).

Other material. (47 specimens examined for meristic or morphometric values, 70.2–139 mm SL; see Appendix 2 for additional material in Australasian collections.) Australia, Victoria: NMV A2261-002 (2, 106–111) 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 A8882 (2, 84.0–127) Bass Strait, 80 km south-east of Loch Sport, 38° 34.3' S, 148° 18.2' E, 86 m, Victorian Marine Sciences Laboratory, 6 June 1984. Tasmania: CSIRO H 7698-09 † (66) Huon Commonwealth Marine Reserve, 43° 42.72' S, 147° 11.32' E – 43° 42.97' S, 147° 10.35' E, 122 m, A. Graham and J. Pogonoski, 8 April 2015. Western Australia: WAM P.25342.004 (112) Cape Naturaliste, 33°32' S, 115°01' E, J. Scott, 16 February 1964; WAM P.25343.017 (2, 108–119) Fremantle, 32°02' S, 115°40' E, L.M. Marsh et al, 24 June 1975; WAM P.26616.005 (125) Point Clune, 32°00' S, 115°30' E, 8 m, J.B. Hutchins et al, 29 March 1979; WAM P.28297.006 (5, 59.4–139) Lucky Bay, 34°08' S, 122°15' E, 8–10 m, J.B. Hutchins, 13 April 1984; WAM P.28300.005 (5, 63.8–131) Lucky Bay, 34°05' S, 122°15' E, 11– 12 m, J.B. Hutchins et al, 16 April 1984. New Zealand, North Island : NMNZ P.028044 (9, 82–127) Matatuahu Point, Tawharanui Peninsula, Hauraki Gulf, 36° 23' S, 174° 49' E, 0–5 m, A.L Stewart and C.D. Paulin, 8 April 1992; NMNZ P.028118 (7, 62–119) Onepoto Bay, Hicks Bay, East Cape, 37° 35.25' S, 178° 18.00' E, 0–3 m, NMNZ East Cape 1992 field team, 4 May 1992; NMNZ P.029805 (3, 55–135) Waihau Bay, Bay of Plenty, 37° 36.8' S, 177° 54.6' E, 4–6 m, NMNZ East Cape 1993 field team, 27 January 1993; NMNZ P.030036 (4, 103–135) inside of Tuamotu Island, Gisborne Harbour, 38° 42.2' S, 178° 2.4' E, 4–7 m, NMNZ East Cape 1993 field team, 21 January 1993; NMNZ P.046263 (103) north of Tom Bowling Bay; North Cape, 34° 22.56' S, 172° 55.28' E, 61–72 m, RV Tangaroa, 14 July 2009; NMNZ P.046285 † (2, 72.2–81.4) east of Purerua Peninsula, Bay of Islands, 35° 6.36' S, 174° 17.10' E, 119–121 m, RV Tangaroa, 7 July 2009; NMNZ P.048310 (70.4) western end of Omapere wharf, 35° 32.05' S, 173° 23.14' E, 4 m, NMNZ & AIM Northland 2011 field team, 13 February 2011; NMNZ P.048311 (70.5) western end of Omapere wharf, 35° 32.05' S, 173° 23.14' E, 4 m, NMNZ & AIM Northland 2011 field team, 13 February 2011; NMNZ P.048312 (82.6) western end of Omapere wharf, 35° 32.05' S, 173° 23.14' E, 4 m, NMNZ & AIM Northland 2011 field team, 13 February 2011; NMNZ P.048380 (124), NMNZ P.048381 (96.8) and NMNZ P.048382 (3, 59.9–101) reef inside southern headland of Hokianga Harbour, 35° 32.01' S, 173° 22.10' E, 5–8 m, WCN 11/09, NMNZ & AIM Northland 2011 field team, 15 February 2011; NMNZ P.049600 (84.8) and NMNZ P.049703 (93) Tauroa Point, Ahipara, 35° 10.48' S, 173° 2.73' E, 21 m, WCN 11/35, NMNZ & AIM Northland 2011 field team, 22 February 2011; NMNZ P.049708 (100) Tauroa Point, Ahipara, 35° 9.96' S, 173° 3.12' E, 14 m, NMNZ & AIM Northland 2011 field team, 22 February 2011; NMNZ P.051786 (119) Tauroa Point, Ahipara, 35° 10.48' S, 173° 2.73' E, 21 m, NMNZ & AIM Northland 2011 field team, 22 February 2011; NMNZ P.057218 † (78.5) and NMNZ P.052719 † (82.3) Ranfurly Bank, 37° 32.78' S, 178° 53.42' E, 68–70 m, RV Tangaroa, 30 May 2011.

Notes

Published as part of 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, pp. 59-99 in Memoirs of Museum Victoria 80 on pages 80-82, DOI: 10.24199/j.mmv.2021.80.04, http://zenodo.org/record/8065377

Files

Files (15.9 kB)

Name Size Download all
md5:fbb2228a4598e5dee74bc8c04d4c2df8
15.9 kB Download

System files (114.8 kB)

Name Size Download all
md5:cfec38ac8b4c6fb0206149bc5d7f8dc9
114.8 kB Download

Linked records

Additional details

References

  • Richardson, J. 1846. Ichthyology of the voyage of H. M. S. Erebus & Terror, under the command of Captain Sir James Clark Ross, R. N., F. R. S. Pp. 53 - 74 in: J. Richardson, and J. E. Gray (eds.), The zoology of the voyage of H. M. S. Erebus & Terror, under the command of Captain Sir J. C. Ross, R. N., F. R. S., during the years 1839 to 1843. E. W. Janson: London. 139 pp.
  • Gunther, A. 1862. Catalogue of the fishes in the British Museum. Catalogue of the Acanthopterygii, Pharyngognathi and Anacanthini in the collection of the British Muesum. 4. Department of Zoology, British Museum (Natural History): London. 534 pp.
  • Gunther, A. 1863. On new species of fishes from Victoria, South Australia. Annals and Magazine of Natural History (Series 3) 11 (62): 114 - 117.
  • Hutton, F. W., and Hector J. 1872. Fishes of New Zealand. Colonial Museum and Geological Survey Department: Wellington. 133 pp.
  • Dambeck, K. 1879. Appendix A. Geographical distribution of the Gadidae or the cod family, in its relation to fisheries and commerce in II. Pp. 531 - 557 in: Part IV: Report of the Commissioner for 1877. United States Commission of Fish and Fisheries: Washington, DC. 981 pp.
  • Hector, J. 1884. The fisheries of New Zealand in Handbook of New Zealand, 1883. Bulletin of the United States Fisheries Commission 4: 53 - 55.
  • Hector, J. 1886. Fisheries. Pp. 26 - 28 in: J. Hector (ed.), Handbook of New Zealand. Government Printer: Wellington. 120 pp.
  • Sherrin, R. A. A. 1886. Handbook of the fishes of New Zealand. Wilson and Horton: Auckland. 307 pp.
  • Gill, T. 1893. A comparison of antipodal faunas. Memoirs of the National Academy of Sciences 6 (5): 91 - 121.
  • Hutton, F. W. 1904. Index Faunae Novae Zealandiae. Dulau and Company: London. 372 pp.
  • Thomson, G. M. 1906. The Portobello Marine Fish-hatchery and Biological station. Transactions and Proceedings of the New Zealand Institute 38: 529 - 558.
  • Young, M. W. 1925. Physiculus (Lotella) rhacinus (Forster). A record of its capture in Otago waters; with some notes on its range and vernacular nomenclature. New Zealand Journal of Science and Technology 7 (5): 369 - 371.
  • Graham, D. H. 1938. Fishes of Otago Harbour and adjacent seas with additions to previous records. Transactions and Proceedings of the Royal Society of New Zealand 68 (3): 399 - 419.
  • Graham, D. H. 1939. Breeding habits of the fishes of Otago Harbour and adjacent seas. Transactions and Proceedings of the Royal Society of New Zealand 69: 361 - 372.
  • Ayling, T., and Cox, G. J. 1982. Collins guide to the sea fishes of New Zealand. William Collins Publishers Ltd: Auckland. 343 pp.
  • Ogilby, J. D. 1897. New genera and species of Australian fishes. Proceedings of the Linnean Society of New South Wales 22 (1): 62 - 95.
  • McCulloch, A. R. 1927. The fishes and fish-like animals of New South Wales. Sydney: Royal Zoological Society of New South Wales: Sydney. 104 pp.
  • McCulloch, A. R. 1929 - 30. A check-list of the fishes recorded from Australia. Parts I - IV. Memoirs of the Australian Museum, Sydney 5: 1 - 534 (Pt I: 1 - 144, II: 145 - 329, III: 329 - 436, IV: 437 - 534).
  • Munro, I. S. R. 1938. Handbook of Australian fishes. Fishery Newsletter 16 (11): 1 - 172.
  • Whitley, G. P. 1964. A survey of Australian Ichthyology. Proceedings of the Linnean Society of New South Wales 89 (1): 11 - 127.
  • Paxton, J. R., Hoese, D. F., Allen, G. R., and Hanley, J. E. 1989. Zoological Catalogue of Australia. Vol. 7. Pisces: Petromyzontidae to Carangidae. Australian Government Publishing Service: Canberra, Australia. 665 pp.
  • Graham, D. H. 1956. A treasury of New Zealand fishes. Second edition. A. H. and A. W. Reed: Wellington. 424 pp.
  • Whitley, G. P. 1956. Name-list of New Zealand fishes. Pp. 397 - 415 in Graham, D. H. (ed.) A treasury of New Zealand fishes. 2 nd Edition. Reed: Wellington. 424 pp.
  • Svetovidov, A. N. 1967. K poznaniyu semeistva Moridae (Pisces, Gadiformes), Zoologicheskii Zhurnal, 46 (11): 1684 - 1693, figs 1 - 3.
  • Whitley, G. P. 1968. A checklist of the fishes recorded from the New Zealand Region. Australian Zoologist 15 (1): 1 - 102.
  • Paulin, C. D. 1983. A revision of the family Moridae (Pisces: Anacanthini) within the New Zealand region. National Museum of New Zealand Records 2 (9): 81 - 126.
  • Francis, M. P. 1988. Coastal fishes of New Zealand: a diver's identification guide. Heinemann Reed: Auckland. 63 pp.
  • Paul, L. J. 1986. NewZealandfishes: anidentificationguide. Auckland, Reed Methuen Publishers Ltd. 184 pp.
  • Paulin, C. D. 1988. Swimbladder structure in Morid cods (Pisces: Gadiformes). Copeia 2: 450 - 454.
  • Paulin, C. D., Stewart, A. L., Roberts, C. D., and McMillan, P. J. 1989. New Zealand fish a complete guide. National Museum of New Zealand Miscellaneous Series. No. 19. National Museum of New Zealand: Wellington. 279 pp, 8 pls. [Also reprinted in 2001].
  • Cohen, D. M., Inada, T., Iwamoto, T., and Scialabba, N. 1990. FAO Species Catalogue. Vol 10. Gadiform fishes of the world. FAO: Rome. 442 pp.
  • Paulin, C. D., and Roberts, C. D. 1992. The rockpool fishes of New Zealand. Te ika aaria o Aotearoa. Museum of New Zealand Te Papa Tongarewa: Wellington. 177 pp.
  • Kuiter, R. H. 1993. Coastal fishes of south-eastern Australia. University of Hawaii Press: Honolulu. 437 pp.
  • Gomon, M. F., Glover, C. J. L., and Kuiter, R. H. 1994. The fishes of Australia's south coast. Flora and Fauna of South Australia Handbooks Committee: Adelaide, Australia. 992 pp.
  • Francis, M. P. 1996. Coastal fishes of New Zealand: an identification guide. Revised edition. Reed Publishing: Auckland. 72 pp.
  • Paul, L. J. 2000. New Zealand fishes: identification, natural history and fisheries. Revised edition. Reed Publishing: Auckland. 253 pp.
  • Francis, M. P. 2001. Coastal fishes of New Zealand: An identification guide. Third edition. Reed Publishing: Auckland. 103 pp.
  • Hutchins, J. B. 2001. Checklist of the fishes of Western Australia. Records of the Western Australian Museum Supplement 63: 9 - 50.
  • Sazonov, Y. I. 2001. Notophycis - a new genus of fish of the family Moridae (Gadiformes) with the description of a new species and two new subspecies, and notes on the state of the genus Austrophycis. Voprosy Ikhtiologii 41 (3): 293 - 303. [In Russian. English translation in Journal of Ichthyology 41 (5): 343 - 353.]
  • Paxton, J. R., Gates, J. E., Bray, D. J., and Hoese, D. F. 2006. Moridae, Pp. 608 - 617 in: Hoese, D. F. Bray, D. J. Paxton, J. R. and Allen, G. R. (eds), Fishes, Pp i - xxiv, 1 - 670. In, Beesley, P. L. and Wells, A. (eds) Zoological catalogue of Australia. ABRS and CSIRO Publishing, Australia Volume 35, Part 1.
  • Gomon, M. F., Bray, D. J., and Kuiter, R. H. 2008. Fishes of Australia's southern coast. New Holland Publishers: Melbourne, Australia. 928 pp.
  • McMillan P. J., Francis, M. P., Paul, L. J., Marriott, P. J., Mackay, E., Baird, S. - J., Griggs, L. H., Sui, H., and Wei, F. 2011 a. New Zealand fishes. Vol. 1: A field guide to common species caught by bottom and midwater fishing. New Zealand Aquatic Environment and Biodiversity Report No. 68. Ministry of Fisheries: Wellington. 329 pp.
  • McMillan P. J., Francis, M. P., James, G. D., Paul, L. J., Marriott, P. J., Mackay, E., Wood, B. A., Griggs, L. H., Sui, H., and Wei, F. 2011 b. New Zealand fishes. Vol. 2: A field guide to less common species caught by bottom and midwater fishing. New Zealand Aquatic Environment and Biodiversity Report No. 78. Ministry of Fisheries: Wellington. 181 pp.
  • Francis M. P. 2012. Coastal fishes of New Zealand: identification, biology, behaviour. Fourth edition. Craig Potton Publishing: Nelson. 268 pp.
  • Roberts, C. D, Barker, J. J, Stewart A. L, Struthers, C. D, and Kortet, S. 2015. Checklist of the fishes of New Zealand. Pp. S 147 - S 178 in: Roberts, CD, Stewart, A. L., and Struthers, C. D. (eds), The fishes of New Zealand. Volume 1. Te Papa Press: Wellington. 2000 pp.
  • Roberts, C. D, Stewart, A. L., Struthers, C. D., Barker, J. J., and Kortet, S. 2017. Checklist of the Fishes of New Zealand. Online Version 1.0. Museum of New Zealand Te Papa Tongarewa. Accessed at: https: // collections. tepapa. govt. nz / document / 10564 on 10 July 2018.
  • Roberts, C. D, Stewart, A. L., Struthers, C. D., Barker, J. J., and Kortet, S. 2019. Checklist of the Fishes of New Zealand. Online Version 1.1. Museum of New Zealand Te Papa Tongarewa. Accessed at: https: // collections. tepapa. govt. nz / document / 10564 on 10 July 2019.
  • Gunther, A. 1878. Preliminary notices of deep-sea fishes collected during the voyage of H. M. S. ' Challenger'. Annals and Magazine of Natural History (Series 5) 2 (7, 8, 9): 17 - 28; 179 - 187; 248 - 251.