Orr and Hawkins: Species of the rougheye rockfish complex 
129 
to as “menuke” in Japanese, meaning “exploding eye” 
because of the bulging eyeballs seen in fishes brought to 
the surface after being captured at depth. Except for S. 
matsubarai, each of these is easily distinguished from 
both S. melanostictus and S. aleutianus by the absence 
of infraorbital spines. In S. matsubarai, two spines are 
present but each is strong and invariably present in 
the same position: one on the lachrymal and one on 
the base of the suborbital stay of infraorbital 3. In ad- 
dition, the lachrymal has two or three strong spines on 
its ventral margin, in contrast with the two typically 
rounded or flattened lobes, the second with up to four 
small spines in S. melanostictus and S. aleutianus. 
Sebastes flammeus and S. iracundus are very similar 
to one another and were considered synonymous by 
Balanov et al. (2004). However, both Kai et al. (2003) 
and Hyde and Vetter (2007) found differences in genetic 
sequences between the two species, providing additional 
evidence for their validity as separate species. They are 
morphologically most similar to S. borealis, with which 
they appear to be closely related (Hyde and Vetter, 
2007). Both lack infraorbital spines and have reduced 
cranial spines that lack coronal and nuchal spines. 
A recent comprehensive phylogenetic analysis of Se- 
bastes, based on molecular data conducted by Hyde 
and Vetter (2007), revealed notable differences as well 
as similarities between ideas of relationships based on 
overall morphological similarity and detailed genetic 
analyses. Based on maximum parsimony and Bayes- 
ian phylogenetic hypotheses, the subgenus Zalopyr was 
placed in a basal position within the tree and encom- 
passed S. aleutianus and S. melanostictus, as well as 
the western Pacific species S. glaucus Hilgendorf (1880), 
gray rockfish; S. steindachneri Hilgendorf (1880), 
yanaginomai; S. minor Barsukov (1972), akagaya; and 
S. owstoni (Jordan and Thompson, 1914), hatsume. The 
clade comprising Sebastes aleutianus and S. melanost- 
ictus was placed in a terminal position in relation to 
the western Pacific species. All other species of Zalopyr 
of Hyde and Vetter (2007) are readily distinguished 
from S. aleutianus and S. melanostictus by the absence 
of infraorbital spines and reduced or obsolete cranial 
spines. Only S. aleutianus and S. melanostictus are 
deeper-water species — both species commonly occurring 
below 300 m and at least as deep as 500 m; all other 
species within Zalopyr are more shallow-water species 
recorded to a maximum depth of 300 m. 
The purpose of naming elements of phylogenetic trees 
is to communicate information on evolutionary relation- 
ships, primarily the existence of monophyletic groups. 
The subgenus Zalopyr Jordan and Evermann (1898) 
was originally erected for Sebastes aleutianus (the type 
species of the subgenus) and S. atrovirens (Jordan and 
Gilbert, 1880) (Kendall, 2000). Although Zalopyr of 
Hyde and Vetter (2007) comprised a monophyletic group, 
the inclusion of any species other than S. aleutianus 
and S. melanostictus on the basis of molecular evidence 
alone is premature, particularly because only the clade 
including S. aleutianus and S. melanostictus may be 
readily diagnosed morphologically. The name Zalopyr 
should be restricted to the terminal clade of S. aleu- 
tianus and S. melanostictus, a subgenus diagnosed by 
the combination of two character states: the presence 
of 2-10 infraorbital spines and 8 pairs of major cranial 
spines. To avoid naming a paraphyletic group, the spe- 
cies in the more basal portion of the clade should be 1) 
removed from Zalopyr and placed in their own subgen- 
era, 2) the name Emmelas Jordan and Evermann (1898) 
applied to the basalmost member, S. glaucus, and 3) the 
name Hatumeus Matsubara (1943), previously used for 
the monotypic subgenus comprising S. owstoni, applied 
to the western Pacific clade that also includes S. stein- 
dachneri and S. minor. 
This phylogenetic hypothesis indicates the derivation 
of S. melanostictus and S. aleutianus from a western 
North Pacific and shallow-water ancestor that dispersed 
into deeper waters of the eastern North Pacific region. 
Their divergence from other members of the clade has 
been estimated to have occurred at over 4.5 million 
years ago (Hyde and Vetter, 2007) and from one an- 
other at about several hundred thousand to just over 
one million years ago (Gharrett et al., 2005; Hyde and 
Vetter, 2007). 
The zoogeographic pattern exhibited by Sebastes 
aleutianus and S. melanostictus is similar to several 
other species pairs found in waters of Alaska, includ- 
ing Sebastolobus alascanus Bean (1890) and S. altivelis 
Gilbert (1896), the long- and shortspine thornyheads; 
Lepidopsetta polyxystra Orr and Matarese (2000) and 
L. bilineata (Ayres, 1855), the northern and southern 
rock soles; and Lycodes palearis Gilbert (1896) and L. 
brevipes Bean (1890), the wattled and shortfin eelpouts 
(Mecklenburg et al., 2002). In each of these pairs, one 
species ranges across the North Pacific region from 
the Kuril Islands in the west to California in the east. 
The other species is an eastern North Pacific species, 
ranging more or less into the Bering Sea and ending 
to the west in the Aleutian Islands at Samalga Pass, 
the western extent of the ancient and now fragmented 
Alaska Peninsula. Recent studies across a wide spec- 
trum of organisms, from whales and seabirds to fishes 
and zooplankton, across the Aleutian Islands have 
found differences in the nature of oceanographic eco- 
systems on either side of Samalga Pass: to the east the 
environment is more like that of the continental shelf, 
whereas to the west it is more oceanic (Hunt and Sta- 
beno, 2005). Although many of the distribution patterns 
may be related to geologically recent ecological factors, 
a deeper vicariant event may have been responsible for 
affecting species distributions in the region (Logerwell 
et al., 2005) 
Fisheries for rockfish on the U.S. west coast are pres- 
ently managed by species, geographic region, and habi- 
tat (primarily depth) parameters. Although overlap- 
ping broadly in distribution, S. aleutianus is typically 
found at shallower depths than S. melanostictus (Fig. 8; 
Hawkins et al., 2005; Gharrett et al., 2005, 2006), and, 
under present management regimes, S. melanostictus 
may be fished more heavily because the major part of its 
distribution lies outside the more shallow restricted ar- 
