S171
Journal of Ichthyology, Vol. 45, Suppl. 1, 2005, pp. S171–S182.
Original English Text Copyright © 2005 by Stankovic, Spalik, Golik, Balushkin, Borsuk, Koper, Rakusa-Suszczewski, Weglenski.
1
INTRODUCTION
Notothenioids are a suborder of perciform fishes
comprising 8 families, 48 genera, and 139 species (Bal-
ushkin, 2000). While the majority of the species of six
of the notothenoid families live within the Antarctic
Region, one monotypic family, Pseudaphritidae, and
most species of the family Bovichtidae have a non-Ant-
arctic distribution. Notothenioidei is the most abundant
suborder in number, biomass, and taxonomic diversity
among the vertebrates of the southern regions of the
Earth.
The notothenioid ancestor was probably benthic,
like the most primitive members of the group Hal-
aphrithis platycephala (Bovichtidae) (Last et al.,
2002). Many extant taxa are also benthic or demersal in
habit. However, half of all living species have moved
1
This article was submitted by the authors in English.
into non-benthic niches becoming epibenthic, pelagic,
semipelagic or cryopelagic (Andriashev, 1970; East-
man, 1993). The progressive loss of bone tissue, along
with certain other morpho-physiological changes lead-
ing to a decrease in body weight (these changes
included the accumulation of fat in the subdermal layer,
muscles, and special fat deposits) were the important
morphological pre-adaptations which enabled notothe-
nioids to colonize the pelagic biotopes (Balushkin,
2000). Notothenioids are characterized by a reduced
haematocrit and haemoglobin concentration resulting
from low metabolic demand and high oxygen solubility
at low temperatures. In the family Channichthyidae
(icefish), this trend has proceeded as far as the loss of
functional erythrocytes. The synthesis of antifreeze
glycopeptides is an evolutionary novelty of Antarctic
notothenioids which has enabled them to successfully
colonize ice-cold waters. The absence of these glyco-
Polyphyly of Scorpaeniformes and Perciformes:
New Evidence from the Study of Notothenioid’s Mitochondrial
and Nuclear rDNA Sequence Data
1
A. Stankovic
1,4
*, K. Spalik
2
, P. Golik
3,4
, A. V. Balushkin
5
, P. Borsuk
3,4
, M. Koper
3
,
S. Rakusa-Suszczewski
6
, and P. Weglenski
3,4
1
Novae Center for Archaeological Research, Warsaw University, Krakowskie Przedmiescie 32, 00–927 Warsaw, Poland
2
Department of Plant Systematics and Geography, Warsaw University, Aleje Ujazdowskie 4, 00–478 Warsaw, Poland
3
Department of Genetics, Warsaw University, Pawinskiego 5a, 02–106 Warsaw, Poland
4
Institute of Biochemistry and Biophysics, Polish Academy of Sciences, Pawinskiego 5a, 02–106 Warsaw, Poland
5
Laboratory of Ichthyology, Zoological Institute, Russian Academy of Sciences,
Universitetskaia nab. 1, St. Petersburg, 199034 Russia
6
Department of Antarctic Biology, Polish Academy of Sciences, Ustrzycka 10/12, 02–141 Warsaw, Poland
*E-mail: anams@poezta.ibb.waw.pl
Received March 9, 2005
Abstract—Partial sequences of nuclear 18S and mitochondrial 12S and 16S rDNA genes from 21 species were
analyzed using parsimony, maximum likelihood, and Bayesian inference in order to establish the phylogenetic
position of notothenioids, a major group of Antarctic fishes. The outgroup included members of the orders Per-
ciformes and Scorpaeniformes, which are both placed in the superorder Percomorpha. In all analyzes, the
notothenioids containing Pseudaphritidae, Eleginopidae, Nototheniidae, Harpagiferidae, Artedidraconidae, and
Channichthyidae were monophyletic and this result received very strong internal support from decay, bootstrap,
and Bayesian analyzes. However, the orders Perciformes and Scorpaeniformes, as well as the perciform subor-
der Trachinoidei and the scorpaeniform suborder Scorpaenoidei were shown to be polyphyletic. Among the
species included in this study, the closest relatives of the monopyletic clade of notothenioids appeared to be two
species of Sebastidae, Sebastes brevispinis and S. marinus. The sister relationship between notothenioids (with
the exception of bovichtids) and S. brevispinis was also confirmed in a large analysis using 125 partial mito-
chondrial 12S and 16S rDNA sequences. This phylogeny is in serious disagreement with current opinions on
the timing of Sebastes and notothenioid radiations. The oldest separations in Sebastes are estimated at 18 mil-
lion years ago, while the divergence of the Antarctic notothenioid lineage from its temperate pseudaphitid cous-
ins is associated with the separation of Australia from Antarctica, which took place 45–55 million years ago.
Molecular data suggest that the origin and diversification of notothenioids occurred much more recently than
previously believed.