429 
Genetic and morphological differences 
between Sebastes vulpes and S. zonatus 
(Teleostei: Scorpaeniformes: Scorpaenidae) 
Nozomu Muto 1 
Yoshiaki Kai 2 
Tetsuji Nakabo 3 
Email address for contact author: muto@inet. museum. kyoto-u. ac.jp 
1 Division of Applied Biosciences 
Graduate School of Agriculture 
Kyoto University 
do The Kyoto University Museum 
Kyoto University 
Yoshida, Sakyo, Kyoto, 606-8501 Japan 
2 Maizuru Fisheries Research Station 
Field Science Education and Research Center 
Kyoto University 
Nagahama, Maizuru, Kyoto, 625-0086 Japan 
3 The Kyoto University Museum 
Kyoto University 
Yoshida, Sakyo, Kyoto, 606-8501 Japan 
Abstract — The taxonomic status of 
Sebastes vulpes and S. zonatus were 
clarified by comprehensive genetic 
(amplified fragment length poly- 
morphisms [AFLP] and mitochon- 
drial DNA [mtDNA] variation) and 
morphological analyses on a total of 
65 specimens collected from a single 
locality. A principal coordinate analy- 
sis based on 364 AFLP loci separated 
the specimens completely into two 
genetically distinct groups that cor- 
responded to S. vulpes and S. zonatus 
according to body coloration and that 
indicated that they are reproductively 
isolated species. Significant morpho- 
logical differences were also evident 
between the two groups; 1) separa- 
tion by principal component analysis 
based on 31 measurements, and 2) 
separation according to differences 
in counts of gill rakers and dorsal-fin 
spines without basal scales, and in the 
frequencies of specimens with small 
scales on the lower jaw. Restriction 
of gene flow between the two groups 
was also indicated by the pairwise 
<P ST values estimated from variations 
in partial sequences from the mtDNA 
control region although the minimum 
spanning network did not result in 
separation into distinct clades. The 
latter was likely due to incomplete 
lineage sorting between S. vulpes 
and S. zonatus owing to their recent 
speciation. 
Manuscript submitted 21 May 2011. 
Manuscript accepted 22 July 2011. 
Fish. Bull. 109:429-439 (2011). 
The views and opinions expressed 
or implied in this article are those of the 
author (or authors) and do not necessarily 
reflect the position of the National Marine 
Fisheries Service, NOAA. 
The genus Sebastes, containing the 
live-bearing rockfishes, is the most 
species-rich scorpaenid genus, com- 
prising over 110 species worldwide, 
over 30 of which are known from 
the western North Pacific (Kai et 
al., 2003; Nelson, 2006; Hyde and 
Vetter, 2007). Exhibiting a relatively 
high diversity in number of closely 
related species in contrast to other 
genera of marine fishes, the genus 
has long attracted the attention of 
evolutionary biologists (e.g.. Love et 
ah, 2002). The greater part of such 
diversity has been ascribed to an 
ancient explosive speciation event 
and subsequent adaptive radiation. 
These species have been interpreted 
as representing an “ancient species 
flock” — an occurrence rarely seen in 
marine fishes (Johns and Avise, 1998; 
Ruber and Zardoya, 2005). However, 
an increasing number of recent stud- 
ies have documented recently diverged 
sibling species pairs that are indica- 
tive of ongoing speciation within the 
genus (Kai et ah, 2002a; Narum et ah, 
2004; Hawkins et ah, 2005; Hyde et 
ah, 2008; Burford, 2009; Stefansson 
et ah, 2009). These events present 
a series of “snapshots” of the specia- 
tion process, providing us with unique 
insights into evolutionary processes in 
the marine realm (Sobel et ah, 2009). 
Sebastes vulpes, S. ijimae, and S. 
zonatus are closely related and mor- 
phologically similar western North 
Pacific species (Chen and Barsukov, 
1976; Nakabo, 2002b; Kai et ah, 
2003; Hyde and Vetter, 2007) that 
are subject to some taxonomic con- 
fusion. Doderlein in Steindachner 
and Doderlein (1884) first described 
Sebastes vulpes; Jordan and Metz 
(1913) subsequently listed S. vulpes 
as valid and described a new species, 
S. ijimae. Two color variants within 
<S. vulpes recognized by Matsubara 
(1943) were later considered to repre- 
sent separate species (S. vulpes and a 
new species, S. zonatus) by Chen and 
Barsukov (1976). They characterized 
S. vulpes as having a dark gray body 
with distinct dense white spots and 
S. zonatus as having a white to pink- 
ish body with three distinct vertical 
dark bands. In addition, Chen and 
Barsukov (1976) also recognized S. 
ijimae as a distinct species, noting 
that the three species shared almost 
the same distributional range, be- 
ing known from southern Hokkaido 
