J.T. Eastman and R.R. Eakin
January 15, 2015
Table 1. Notothenioid classification and list of species. Arrangement is phylogenetic for
families, based on Near et al. (2004) and Near and Cheng (2008), and alphabetical for genera
within families and for species within genera. Superfamilies from Voskoboinikova (2004).
Although recent work indicates that the Nototheniidae and Bathydraconidae are paraphyletic
(Near et al., 2012; Dettai et al., 2012), we are provisionally retaining the traditional family
names as relationships and taxonomy of notothenioids, and acanthomorphs in general, are
entering a period of flux (Dettai et al., 2012; Betancur-R et al., 2013; Near et al., 2013). Fishes
of the Southern Ocean (Gon and Heemstra, 1990) is starting point for species list with
subsequent additions noted. Bovichtids from Hardy (1988), Balushkin (1992) and Bravo et al.
(1999); pseudaphritids and eleginopsids from Balushkin (1992, 2000); nototheniids from DeWitt
et al. (1990), but following Schneppenheim et al. (1994) for the Lepidonotothen squamifrons
group, and with non-Antarctic species from DeWitt (1966, 1970); harpagiferids from Hureau
(1990); artedidraconids from Eakin (1990) with revision of Pogonophryne following Balushkin
and Eakin (1998); bathydraconids from Gon (1990) and channichthyids from Iwami and Kock
(1990). Dagger (†) precedes names of species described or resurrected from synonymy after
publication of Fishes of the Southern Ocean in 1990. Species placed in synonymy since 1990
are underlined in the list and are not counted. Other changes since Eastman and Eakin (2000)
indicated by footnotes. Catalog of Fishes (Eschmeyer, 2014, online version) is 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 render judgment on the validity of new species.
In our list we do not recognize all notothenioid species listed in the Catalog of Fishes. Asterisk
(*) indicates non-Antarctic species. Our total count is 139 species (107 Antarctic and 32 nonAntarctic) and 44 genera. Up to date as of January 15, 2015.
Superfamilies, families (with no. of valid species) and species
Bovichtoidea
Bovichtidae (9)
Bovichtus
angustifrons Regan 1913*
argentinus MacDonagh 1931*a
chilensis Regan 1913*
diacanthus (Carmichael 1819)*
elongatus Hureau and Tomo 1977a
oculus Hardy 1988*
2
psychrolutes Günther 1860*
variegatus Richardson 1846*
veneris Sauvage 1879*
Cottoperca trigloides (Forster 1801)*b
†Halaphritis platycephala Last, Balushkin and Hutchins 2002*c
Notothenioidea
Pseudaphritidae (1)
Pseudaphritis urvillii (Valenciennes in Cuvier and Valenciennes 1832)*
Eleginopsidaed (1)
Eleginops maclovinus (Cuvier in Cuvier and Valenciennes 1830)*
Nototheniidae (50)
Aethotaxis mitopteryx DeWitt 1962
Cryothenia
†amphitreta Cziko and Cheng 2006e
peninsulae Daniels 1981
Dissostichus
eleginoides Smitt 1898
mawsoni Norman 1937
Gobionotothen
acuta (Günther 1880)
†barsukovi Balushkin 1991f
gibberifrons (Lönnberg 1905)
marionensis (Günther 1880)
Gvozdarus svetovidovi Balushkin 1989
Lepidonotothen
kempi (Norman 1937)
larseni (Lönnberg 1905)
macrophthalma (Norman 1937)*
mizops (Günther 1880)
nudifrons (Lönnberg 1905)
3
squamifrons (Günther 1880)
Notothenia
angustata Hutton 1875*
coriiceps Richardson 1844
cyanobrancha Richardson 1844
microlepidota Hutton 1875*
rossii Richardson 1844
Pagothenia
borchgrevinki (Boulenger 1902)
brachysoma (Pappenheim 1912)
Paranotothenia
†dewitti Balushkin 1990
magellanica (Forster in Bloch and Schneider 1801)
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)*g
†thompsoni Balushkin 1993*
wiltoni (Regan 1913) *
Pleuragramma antarctica Boulenger 1902h
Trematomus
bernacchii Boulenger 1902
eulepidotus Regan 1914
hansoni Boulenger 1902
lepidorhinus (Pappenheim 1911)i
loennbergii Regan 1913i
newnesi Boulenger 1902
4
nicolai (Boulenger 1902)
pennellii Regan 1914
scotti (Boulenger 1907)
tokarevi Andriashev 1978
vicarius Lönnberg 1905
Harpagiferidae (11)
Harpagifer
†andriashevi Prirodina 2000j
antarcticus Nybelin 1947
bispinis (Schneider in Bloch and Schneider 1801)*
†crozetensis Prirodina 2004k
georgianus Nybelin 1947
kerguelensis Nybelin 1947
†macquariensis Prirodina 2000l
†nybelini Prirodina 2002m
†permitini Neyelov and Prirodina 2006n
palliolatus Richardson 1845
spinosus Hureau, Louis, Tomo and Ozouf 1980
Artedidraconidae (35)
Artedidraco
†glareobarbatus Eastman and Eakin 1999
loennbergi Roule 1913
mirus Lönnberg 1905
orianae Regan 1914
shackletoni Waite 1911
skottsbergi Lönnberg 1905
Dolloidraco longedorsalis Roule 1913
Histiodraco velifer (Regan 1914)
Pogonophryne
albipinna Eakin 1981
barsukovi Andriashev 1967
†bellingshausenensis Eakin, Eastman and Matallanas 2008o
†brevibarbata Balushkin, Petrov and Prutko 2010p
†cerebropogon Eakin and Eastman 1998
curtilemma Balushkin 1988
dewitti Eakin 1988
5
dolichobranchiata Andriashev 1967
†eakini Balushkin 1999
†favosa Balushkin and Korolkova 2013q
†fusca Balushkin and Eakin 1998
immaculata Eakin 1981
lanceobarbata Eakin 1987
macropogon Eakin 1981
†maculiventrata Spodareva and Balushkin 2014r
marmorata Norman 1938
mentella Andriashev 1967
†minor Balushkin and Spodareva 2013s
†neyelovi Shandikov and Eakin 2013t
†orangiensis Eakin and Balushkin 1998
orcadensis Tomo 1981
†pavlovi Balushkin 2013u
permitini Andriashev 1967
phyllopogon Andriashev 1967
platypogon Eakin 1988
†sarmentifera Balushkin and Spodareva 2013v
scotti Regan 1914
†skorai Balushkin and Spodareva 2013w
†stewarti Eakin, Eastman and Near 2009x
†squamibarbata Eakin and Balushkin 2000
†tronio Shandivov, Eakin and Usachev 2013y
velifera Eakin 1981
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
6
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 1844z
Chionobathyscus dewitti Andriashev and Neelov 1978
Chionodraco
hamatus (Lönnberg 1905)
myersi DeWitt and Tyler 1960
rastrospinosus DeWitt and Hureau 1979
Cryodraco
antarcticus Dollo 1900
†atkinsoni Regan 1914aa
Dacodraco hunteri Waite 1916
Neopagetopsis ionah Nybelin 1947
Pagetopsis
macropterus (Boulenger 1907)
7
maculatus Barsukov and Permitin 1958
Pseudochaenichthys georgianus Norman 1937
aBravo
et al. (1999) placed these two species in the synonymy of Bovichtus chilensis. Eastman
and Eakin (2000) were unaware of this paper when compiling their 2000 list.
bThis
is a change from Eastman and Eakin (2000). We now follow Balushkin (1992, 2000) and
Eschmeyer (2013) in considering Cottoperca gobio (Günther 1861) as a junior synonym of C.
trigloides (Forster in Bloch and Schneider 1801).
cLast
et al. (2002)
dWe
follow Catalog of Fishes in spelling this family name by including the final “s” in the stem
inops-. 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.
eCziko
and Cheng (2006)
fDuhamel
et al. (2005, pp. 327 and 334), for example, 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 remain skeptical about the validity of this
species.
gNot
included in previous lists; the status of this species is uncertain but it is treated here as
valid; it is widely recognized by South American ichthyologists, e.g. García and Menni (1996)
and Ojeda et al. (2000), and has been collected off West Falkland Island (Paul Brickle, Fisheries
Department, Falkland Islands Government, personal communication, 2005).
hThis
change is necessary because Pleuragramma is feminine, based on agramma meaning
absence of a lateral line. Thus the correct species spelling is antarctica (Eschmeyer 2013).
iUntil
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).
8
jPrirodina
(2000)
kPrirodina
(2004)
lPrirodina
(2000). This is the same reference as in footnote j.
mPrirodina
nNeyelov
oEakin
(2002)
and Prirodina (2006)
et al. (2008)
pBalushkin
et al. (2010)
qBalushkin
and Korolkova (2013)
rSpodareva
and Balushkin (2014)
sBalushkin
and Spodareva (2013a). The validity of this species is doubtful. See Eakin’s critique
in Appendix A following the References.
tShandikov
and Eakin (2013)
uBalushkin
and Spodareva (2013b)
vBalushkin
and Spodareva (2013c)
wShandikov
xEakin
and Eakin (2013)
et al. (2009)
yShandikov
et al. (2013)
zThere
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, Shandivov (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.
aaLa
Mesa et al. (2002)
9
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14
Appendix A
Critique of Pogonophryne minor Balushkin and Spodareva, 2013
Richard R. Eakin
Balushkin and Spodareva (2013) state that the paratype (ZIN no. 55238) of Pogonophryne minor
sp. n. (“small specimen,” 70 mm TL juvenile P. marmorata, Andriashev, 1967) differs from the
adult (173 mm TL female) P. marmorata caught at the same locality (Ob’ station 164, Davis Sea,
January 17, 1957, depth 540-430 m) in having a narrower interorbital space, a larger eye and a
longer mental barbel – character differences Andriashev attributed to age. The authors
compare this specimen with another (holotype: ZIN no. 55237, 103.4 mm TL female; R/V
Chatyr-Dag stn. 56, cruise 18, trawl 53, Mawson Sea, March 14, 1983, depth 420 m) and
conclude that the two represent a new species -- the “dwarf toad plunderfish” (so named
because the holotype is supposedly mature at a smaller-than-normal size for the genus). They
calculate, based on Kock and Kellerman (1991), that P. minor “can not exceed 150 mm” TL.
Dwarfism has not been previously reported in notothenioid fishes and this purported example
may simply reflect the range of variation in a genus that does not exceed about 340 mm TL
(Eakin, 1990). More material is needed to determine if this is a true example of dwarfism.
Dwarfism aside, P. minor differs in no significant way from P. marmorata, based on the
evidence provided by the authors from seven specimens (106-173 mm TL) of P. marmorata and
two type specimens of P. minor. They state that their “measurements based on large samples
of two species” indicate that P. marmorata differs from the new species in having a short
barbel (8.9-11.3 vs. 14.9-15.5% SL) with a short (3.4-5.3 vs. 5.5-6.4% SL) and narrow (1.5-2.2 vs.
2.6-2.8% SL) terminal expansion, a high and wide head (20.5-24.5 vs. 19.9-20.2% SL; 26.6-31.7
vs. 24.2-25.5% SL), and a wide interorbital space (4.7-5.8 vs. 4.3-4.6% SL).
Data from 32 specimens (84-227 mm TL) of P. marmorata from four different collections
(Weddell Sea, 1985: ISH 50/85, 109/85, 122/85; South Orkney Islands, 1999) indicate a wider
range of variation in the above characters than the authors present. A range of 8.9-15.9% SL
(mean 12.9% SL) shows that there is no difference in barbel length between the two species. A
range of 4.2-7.8% SL (mean 5.7% SL) shows that there is no difference in the length of the
terminal expansion between the two species. (Width of the terminal expansion is variable in
the “P. marmorata” group and therefore of no taxonomic significance, in my opinion. Far more
important is the similarity, noted by the authors, in both “shape and structure” of the terminal
expansion in both species.) A range of 3.9-6.1% SL (mean 4.8% SL) shows that there is no
difference in interorbital width between the two species. A range of 19.4-25.7% SL (mean
22.8% SL) shows that there is no difference in head depth between the two species. A range of
26.0-35.9% SL (mean 29.9% SL) in head width does not overlap with P. minor, but this slight
difference (head width greater than 26% SL for P. marmorata and less than 26% SL for P.
minor) is likely not of significance as a diagnostic character.
Other purported differences between P. marmorata and P. minor noted by the authors include
caudal-fin pattern (which in the above samples examined varies from dark and unstriped to
15
dark and striped to light and striped), dark color (“almost black in holotype” of P.minor) of the
peritoneum, and length of pelvic fins (19.9-23.1 vs. 24.2-24.5% SL). Data from 24 specimens of
P. marmorata (ISH collections referred to above) show a range in pelvic-fin length of 17.523.7% SL which supports the difference, albeit slightly.
In conclusion, the two type specimens of P. minor differ from P. marmorata in having slightly
narrower heads, slightly longer pelvic fins, slightly narrower terminal expansions on their
mental barbels and darker peritoneums. Given the overall similarity between these two
specimens (in more critical characters such as barbel structure, head shape and meristics) and
P. marmorata, however, it seems unwarranted to suggest that they represent a new species
based on relatively insignificant differences. P. minor is therefore best considered a junior
synonym of P. marmorata.
Andriashev, A. P. 1967. Review of the plunder fishes of genus Pogonophryne Regan
(Harpagiferidae) with descriptions of five new species from the East Antarctic and South
Orkney Islands (pp. 389-412). In: A. P. Andriashev & P. V. Ushakov (eds), Biol. Res. Sov.
Antarc. Exped. (1955-1958) 3.
Balushkin, A. V. and V. V. Spodareva. 2013. 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):1-6.
Eakin, R. R. 1990. Artedidraconidae (pp. 332-356). In: O. Gon & P. C. Heemstra (eds), Fishes of
the Southern Ocean. Grahamstown, South Africa. J. L. B. Smith Institute of Ichthyology.
Kock, K.-H. and A. Kellermann. 1991. Reproduction in Antarctic notothenioid fish: a review.
Antarct. Sci., 3(2):125-150.