6
J.T. Eastman and R.R. Eakin
December 15, 2016
Table 1. Notothenioid classification and list of species. The taxonomy follows relationships identified by recent molecular phylogenetic analyses. These revisions have Percophis brasiliensis as the phylogenetically basal notothenioid (Near et al. 2015), and amalgamate the genera Cryothenia and Pagothenia into Trematomus, and Paranotothenia into Notothenia (Sanchez et al. 2007; Kuhn and Near, 2009; Dettai et al. 2012; Near et al. 2015). The polyphyly in the genus Artedidraco (Lecointre et al. 2011) has yet to be resolved so the status quo is retained. We employ the new rank free names for basal clades from Near et al. (2015) but, because notothenioid taxonomy is in a state of flux, we also retain some traditional family names while acknowledging that the relationships implied by these names are no longer tenable as recent work indicates Nototheniidae and Bathydraconidae are paraphyletic (Dettai et al., 2012; Near et al., 2012). Catalog of Fishes (Eschmeyer and Fricke, 2016, online version) and van der Laan et al. (2014) are used for authorities and dates in cases of uncertainty, and for spelling of names of species and families. Catalog of Fishes attempts to resolve nomenclatural problems and also monitors the literature for descriptions of new species, but does not vet or render a judgment on the scientific validity of each of the several hundred new species appearing each year. Thus of the 165 valid species names for notothenioids in the “Species by Family/Subfamily” (Eschmeyer and Fong, 2016) section of the Catalog of Fishes, we recognize only 142 in our list, 14% fewer. Arrangement of list is alphabetical for species within genera. Major changes since Eastman and Eakin (2000) and points of contention are indicated by footnotes. Asterisks (*) denote non-Antarctic species. We recognize 142 species (113 Antarctic and 29 non-Antarctic) and 42 genera.
Percomorpha
Notothenioidei
Percophidae (1)a
Percophis brasilinesis Quoyand and Giamard 1825*
Notothenioidea
Bovichtidae (9)
Bovichtus
angustifrons Regan 1913*
chilensis Regan 1913*b
diacanthus (Carmichael 1819)*
oculus Hardy 1988*c
psychrolutes Günther 1860*
variegatus Richardson 1846*
veneris Sauvage 1879*
Cottoperca trigloides (Forster 1801)*d
Halaphritis platycephala Last, Balushkin and Hutchins 2002*
Pseudaphritioidea (1)
Pseudaphritis urvillii (Valenciennes 1832)*
Eleginopsioidea (1)e
Eleginops maclovinus (Cuvier 1830)*
Cryonotothenioidea (formerly “Antarctic clade”)
Nototheniidae (50)
Aethotaxis mitopteryx DeWitt 1962
Dissostichus
eleginoides Smitt 1898
mawsoni Norman 1937
Gobionotothen
acuta (Günther 1880)
barsukovi Balushkin 1991f
gibberifrons (Lönnberg 1905)
marionensis (Günther 1880)
Gvozdarus svetovidovi Balushkin 1989g
Lepidonotothen
larseni (Lönnberg 1905)
mizops (Günther 1880)
nudifrons (Lönnberg 1905)h
squamifrons (Günther 1880)
Notothenia
angustata Hutton 1875*
coriiceps Richardson 1844
cyanobrancha Richardson 1844
dewitti (Balushkin 1990)
magellanica (Forster 1801)
microlepidota Hutton 1875*
rossii Richardson 1844
Patagonotothen
brevicauda (Lönnberg 1905)*
canina (Smitt 1897)*
cornucola (Richardson 1844)*
elegans (Günther 1880)*
guntheri (Norman 1937)
jordani (Thompson 1916)*
kreffti Balushkin and Stehmann 1993*
longipes (Steindachner 1876)*
ramsayi (Regan 1913)*
sima (Richardson 1844)*
squamiceps (Peters 1876)*
tessellata (Richardson 1845)*
trigramma (Regan 1913)*
thompsoni Balushkin 1993*
wiltoni (Regan 1913)*
Pleuragramma antarctica Boulenger 1902i
Trematomus
amphitreta (Cziko and Cheng 2006)
bernacchii Boulenger 1902
borchgrevinki Boulenger 1902
brachysoma Pappenheim 1912
eulepidotus Regan 1914
hansoni Boulenger 1902
lepidorhinus (Pappenheim 1911)j
loennbergii Regan 1913j
newnesi Boulenger 1902
nicolai (Boulenger 1902)
peninsulae (Daniels 1981)
pennellii Regan 1914
scotti (Boulenger 1907)
tokarevi Andriashev 1978
vicarius Lönnberg 1905
Harpagiferidae (11)
Harpagifer
andriashevi Prirodina 2000
antarcticus Nybelin 1947
bispinis (Forster 1801)*
crozetensis Prirodina 2004
georgianus Nybelin 1947
kerguelensis Nybelin 1947
macquariensis Prirodina 2000
nybelini Prirodina 2002
permitini Neyelov and Prirodina 2006
palliolatus Richardson 1845
spinosus Hureau, Louis, Tomo and Ozouf 1980
Artedidraconidae (37)
Artedidraco
glareobarbatus Eastman and Eakin 1999
longibarbatus Eakin, Riginella and La Mesa 2015
loennbergi Roule 1913
mirus Lönnberg 1905
orianae Regan 1914
shackletoni Waite 1911
skottsbergi Lönnberg 1905
Dolloidraco longedorsalis Roule 1913
Histiodraco velifer (Regan 1914)
Pogonophrynek
albipinna Eakin 1981
barsukovi Andriashev 1967
bellingshausenensis Eakin, Eastman and Matallanas 2008
brevibarbata Balushkin, Petrov and Prutko 2010
cerebropogon Eakin and Eastman 1998
dewitti Eakin 1988
eakini Balushkin 1999
favosa Balushkin and Korolkova 2013
fusca Balushkin and Eakin 1998
immaculata Eakin 1981
lanceobarbata Eakin 1987
macropogon Eakin 1981
maculiventrata Spodareva and Balushkin 2014
marmorata Norman 1938
mentella Andriashev 1967
neyelovi Shandikov and Eakin 2013
orangiensis Eakin and Balushkin 1998
pallida Balushkin and Spodareva 2015
pavlovi Balushkin 2013
permitini Andriashev 1967
platypogon Eakin 1988
sarmentifera Balushkin and Spodareva 2013
scotti Regan 1914
skorai Balushkin and Spodareva 2013
stewarti Eakin, Eastman and Near 2009
squamibarbata Eakin and Balushkin 2000
tronio Shandivov, Eakin and Usachev 2013
ventrimaculata Eakin, 1987
Bathydraconidae (16)
Acanthodraco dewitti Skóra 1995
Akarotaxis nudiceps (Waite 1916)
Bathydraco
antarcticus Günther 1878
joannae DeWitt 1985
macrolepis Boulenger 1907
marri Norman 1938
scotiae Dollo 1906
Cygnodraco mawsoni Waite 1916
Gerlachea australis Dollo 1900
Gymnodraco acuticeps Boulenger 1902
Parachaenichthys
charcoti (Vaillant 1906)
georgianus (Fischer 1885)
Prionodraco evansii Regan 1914
Psilodraco breviceps Norman 1937
Racovitzia glacialis Dollo 1900
Vomeridens infuscipinnis (DeWitt 1964)
Channichthyidae (16)
Chaenocephalus aceratus (Lönnberg 1906)
Chaenodraco wilsoni Regan 1914
Champsocephalus
esox (Günther 1861)
gunnari Lönnberg 1905
Channichthys rhinoceratus Richardson 1844l
Chionobathyscus dewitti Andriashev and Neelov 1978
Chionodraco
hamatus (Lönnberg 1905)
myersi DeWitt and Tyler 1960
rastrospinosus DeWitt and Hureau 1979
Cryodracom
antarcticus Dollo 1900
atkinsoni Regan 1914
Dacodraco hunteri Waite 1916
Neopagetopsis ionah Nybelin 1947
Pagetopsis
macropterus (Boulenger 1907)
maculatus Barsukov and Permitin 1958
Pseudochaenichthys georgianus Norman 1937
aInclusion of percophids in the Notothenioidei is based on Near et al. (2015).
bBravo et al. (1999) have placed Bovichtus elongatus and B. argentinus in the synonymy of Bovichtus chilensis.
cAlthough Balushkin (2016) has placed Bovichtus oculus in the synonymy of B. psychrolutes, we continue to recognize B. oculus as valid. B. oculus is known only from the holotype in the Museum of New Zealand Te Papa Tongarewa (NMNZ P. 18510). Although Balushkin (pages 632 and 637) indicates he examined a radiograph of this specimen and measured its eye diameter, it is doubtful that the specimen he examined was the holotype. Records at the Museum of New Zealand provide no evidence that the specimen was ever radiographed or that it was ever sent out on loan. Bovichtus oculus has large eyes and a horizontal eye diameter 9.0 times SL is the most important character in distinguishing it from other species of bovichtids including B. psychrolutes (Hardy, 1988). Balushkin reports that in his radiograph this value was 9.2–9.4 times in SL, not the 8.2 in SL reported by Hardy in his Table 1. He claims that his measurements place the holotytpe in the range of eye diameters for B. psychrolutes and therefore that it is this latter species. However re-measurement of the NMNZ type specimen of B. oculus using two different calipers showed that the eye was a little smaller than recorded by Hardy, but still < 9 times in SL. The actual measurements were: bony eye diameter = 13.04 mm; SL = 109.15 mm = 8.37 in SL or 11.94% of SL (Andrew Stewart, Collection Manager: Science (Fishes), NMNZ, personal communication to J.T. Eastman, 12 December 2016.) Given this information, we continue to recognize B. oculus as a valid species.
dThis is a change from Eastman and Eakin (2000). We now follow Balushkin (1992, 2000) and Eschmeyer and Fricke (2016) in considering Cottoperca gobio (Günther 1861) as a junior synonym of C. trigloides (Forster 1801).
eWe follow Catalog of Fishes (Eschmeyer and Fricke, 2016) and Van Der Laan et al. (2014, page 116) in spelling this family name by including the final “s” in the stem eleginops-. We asked the authors of the Catalog to review the Eleginopsidae/Eleginopidae issue and they indicated that, because Eleginopsidae is the most familiar name and used in prevailing recent practice, it should stand.
fDuhamel et al. (2005, pp. 327 and 334) are not convinced that there is sufficient evidence to prove that Gobionotothen barsukovi is distinct from G. acuta. Although Balushkin (2014) has supplemented his original description of G. barsukovi with meristic data from additional specimens, there is no genetic data and we also remain skeptical about the validity of this species. A recent molecular phylogenetic study supported the validity of G. acuta and G. marionensis, but unfortunately did not included specimens of G. barsukovi (Miya et al., 2016).
gVoskoboinikova and Kellermann (1993) described Gvozdarus balushkini from the Weddell Sea on the basis if a single 30 mm SL specimen. This seems premature given the dearth of information about G. svetovidovi, a species known from only two specimens.
hUsing a combination of morphological and molecular genetic data, Dornburg et al. (2016a) have convincingly shown that Lepidonotothen nudifrons consists of two cryptic species: L. nudifrons around South Georgia and the South Sandwich Islands, and L. cf. nudifrons elsewhere in the Antarctic Peninsula area. That latter species will not be included in this list and counted until it formally named.
iThis is a necessary change in spelling of the scientific name of this species because Pleuragramma is feminine, based on agramma meaning absence of a lateral line and hus the correct species spelling is antarctica rather than antarcticum (Eschmeyer and Fricke, 2016).
jUntil recently, there had never been any question about the validity of the sister species Trematomus lepidorhinus and T. loennbergii. However there have been reports that investigators have been unable to distinguish Indian Ocean specimens of these species by either morphological or/or molecular methodology (Lautrédou et al. 2010, 2012; Causse et al. 2011; Dettai et al. 2011a, 2011b). In addition, Ross Sea specimens of these species cannot be distinguished by DNA barcodes (Smith et al. 2012), although they are easily separated morphologically (DeWitt et al. 1990; Eastman, unpublished data). Recent karyological studies support the existence of two species (Ghigliotti et al. 2015).
kWe view an additional species of Pogonophryne, Pogonophryne minor Balushkin and Spodareva 2013a, as doubtful, and it is not included in the list. See Eakin’s critique in Appendix A following the References.
lThere are differing opinions about the number of species of Channichthys, a genus endemic to the Kerguelen Plateau. Iwami and Kock (1990) and Kock (2005) recognize only C. rhinoceratus, while Duhamel et al. (2005) recognize C. rhinoceratus, C. velifer and suggest the existence of a single undescribed species. However Shandikov and Balushkin have described additional species and, in his most recent paper, Shandikov (2011) recognizes a total of nine species of Channichthys. Given the existence of phenotypic plasticity in C. rhinoceratus and many other notothenioid species, we think it is best to maintain a conservative approach by recognizing only C. rhinoceratus until genetic data are available to clarify this situation.
mDornburg et al. (2016b) have provided molecular data to supplement previous morphological data (La Mesa et al., 2002) indicating that these two species of Cryodraco are valid.
References (although not indicated by superscripts and footnotes in the species list, we have also included references for new species described since Eastman and Eakin, 2000)
Balushkin AV (1991) Review of green notothenias, Gobionotothen, Balushkin (Nototheniidae) of the Antarctic and Subantarctic. J Ichthyol 31(8): 42-55
Balushkin AV (1992) Classification, phylogenetic relationships, and origins of the families of the suborder Notothenioidei (Perciformes). J. Ichthyol. 32: 90-110
Balushkin AV (2000) Morphology, classification, and evolution of notothenioid fishes of the Southern Ocean (Notothenioidei, Perciformes). J. Ichthyol. 40, Suppl. 1: S74-S109
Balushkin AV (2013) A new species of Pogonophryne (Perciformes: Notothenioidei: Artedidraconidae) from the deep Ross Sea, Antarctica. Trudy Zool Inst 317: 119-124 [In Russian; diagnosis in English]
Balushkin AV (2016) Systematics of the Antarctic thornfishes of the genus Bovichtus (Bovichtidae) of the seamounts of the New Zealand Basin. J Ichthyol 56: 499-507
Balushkin AV, Eakin RR (1998) A new toad plunderfish Pogonophryne fusca sp. nova (Fam. Artedidraconidae: Notothenioidei) with notes on species composition and species groups in the genus Pogonophryne Regan. J. Ichthyol. 38: 574-579
Balushkin AV, Korolkova ED (2013) New species of plunderfish Pogonophryne favosa sp. n. (Artedidraconidae, Notothenioidei, Perciformes) from the Cosmonauts Sea (Antarctica) with the description in artedidraconids of unusual anatomical structures—convexitas superaxillaris. J Ichthyol 53: 562-574
Balushkin AV, Spodareva VV (2013a) Dwarf toad plunderfish Pogonophryne minor sp. n. (Artedidraconidae; Notothenioidei; Perciformes)—a new species and one of the smallest species of autochthonous ichthyofauna of marginal seas of the Antarctic continent. J Ichthyol 53: 1-6
Balushkin AV, Spodareva VV (2013b) Pogonophryne sarmentifera sp. nov. (Artedidraconidae; Notothenioidei; Perciformes) — The deep-water species of Antarctic plunderfishes from the Ross Sea (Southern Ocean). Trudy Zool Inst 317: 275-281 [In Russian; diagnosis in English]
Balushkin AV, Spodareva VV (2013c) Pogonophryne skorai sp. n. (Perciformes: Artedidraconidae), a new species of toadlike plunderfish from the Bransfield Strait and coastal waters of the South Shetland Islands, Antarctica. Russian J Mar Biol 39: 190-196
Balushkin AV, Spodareva VV (2015) New species of the toad plunderfish of the "albipinna" group, genus Pogonophryne (Artedidraconidae) from the Ross Sea (Antarctica). J Ichthyol 55: 757-764
Balushkin AV, Petrov AF, Prutko VG (2010) Pogonophryne brevibarbata sp. nov. (Artedidraconidae, Notothenioidei, Perciformes) — A new species of toadlike plunderfish from the Ross Sea, Antarctica. Proc Zool Inst Acad Sci USSR 314: 381-386 [In Russian; English abstract]
Betancur-R R, Broughton RE, Wiley EO, Carpenter K, López JA, Li C, Holcroft NI, Arcila D, Sanciangco M, Cureton II JC, Zhang F, Buser T, Campbell MA, Ballesteros JA, Roa-Varon A, Willis S, Borden WC, Rowley T, Reneau PC, Hough DJ, Lu G, Grande T, Arratia G, Ortí G (2013) The Tree of Life and a New Classification of Bony Fishes. PLOS Currents Tree of Life. 2013 Apr 18. Edition 1. doi: 10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288. PLOS Currents Tree of Life 2013 Apr 18 Edition 1 doi: 101371/currentstol53ba26640df0ccaee75bb165c8c26288
Bravo R, Lloris D, Pequeño G, Rucabado J (1999) Revisión de las distintas especies del género Bovichtus (Perciformes, Bovichtidae) citadas para el cono sur americano y península Antártica. Rev. Biol. Mar. Oceanogr. 34: 123-137 [In Spanish; English abstract]
Causse R, Ozouf-Costaz C, Koubbi P, Lamy D, Eléaume M, Dettaï A, Duhamel G, Busson F, Pruvost P, Post A, Beaman RJ, Riddle MJ (2011) Demersal ichthyofaunal shelf communities from the Dumont d'Urville Sea (East Antarctica). Polar Sci 5: 272-285