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Fishery Bulletin 113(2) 
cally with A. hexapterus. Ammodytes hexapterus exhib- 
its an Arctic-Kuril distribution (Allen and Smith, 1988; 
Briggs, 1995; Logerwell et al., 2005), extending from 
the southern Sea of Okhotsk in the Soya Strait to the 
western Bering Sea, the northeastern Bering Sea, and 
into the Chukchi Sea. It may be a circum-Arctic spe- 
cies; sequence data in GenBank indicates that its range 
extends at least to the Beaufort Sea and Hudson Bay 
(Table 4). Ammodytes japonicus exhibits a distribution 
that encompasses the Sea of Japan and the Central 
Kuroshio Current area (Spalding et al., 2007), and this 
species is found on both coasts of Japan, in the Soya 
Strait, in the Sea of Japan, in the Seto Inland Sea, in 
the Yellow and East China seas (Kim et al., 2006, 2008, 
2010), and on the southern Pacific coast of Japan. Am- 
modytes heian exhibits a distribution that consists of 
the northern Sea of Japan and northeastern Honshu 
(Spalding et al., 2007), having been found in the Soya 
Strait and on the northeastern Pacific coast of Japan. 
A collection in the Soya Strait at Wakkanai, Japan, 
contained 3 species: A. hexapterus A. japonicus, and 
A. heian n. sp. This distributional pattern is similar 
to the one based on allozyme data of Okamoto (1989), 
who found 3 “genetically distinct” groups around Ja- 
pan, including 2 in the Soya Strait: a southern species 
ranging from Kyushu and the Seto Inland Sea to Sen- 
dai Bay (=A. japonicus), a more northern species from 
Iwate Prefecture to the northern Sea of Japan and Soya 
Strait (=A. heian n. sp.), and a third species on the 
Pacific Ocean side of Hokkaido off Kushiro and Soya 
Strait (=A. hexapterus). This strong geographic pattern 
may indicate that present-day, large-scale movement of 
sand lances is limited. 
The genetic information presented in this article is 
meant to support the morphological data and describe 
the species of Ammodytes in the North Pacific region. It 
is not meant to provide a complete phylogenetic recon- 
struction of the genus Ammodytes. A productive look at 
phylogeny will require a more detailed study of Ammo- 
dytes of the Atlantic Ocean, as well as other members 
of the group. Although each clade of species is robust, 
the relationships among the Atlantic species are weak, 
as indicated by the low bootstrap branch support and 
alternative positions of A. personatus among Atlantic 
species in phylogenetic trees (Figs. 3 and 4). Addition- 
ally, Tajima D values (Tajima, 1989) indicate that the 
COI gene is likely under selection in Ammodytes. 
Because selection can obscure phylogenetic pat- 
terns and because no literature is available to suggest 
a mutation rate in species of Ammodytes, we are re- 
luctant to provide estimates of times of divergence be- 
tween these species. Ammodytes hexapterus is nearly 
homogenous at COI and, because bottlenecks decrease 
molecular diversity, was likely derived from a founder 
population followed by rapid expansion, as indicated 
by the broad range over which it is homogenous. The 
lack of genetic divergence across haplotypes is typical 
for marine fishes that have experienced fluctuations 
in population size (Grant and Bowen, 1998). More in- 
formation about the Atlantic species, and members of 
other genera, may allow for a more robust examination 
of phylogeny in Ammodytes. 
Our taxonomic revision of the sand lances of the 
genus Ammodytes in the North Pacific region resulted 
in the recognition of 4 species: A. hexapterus, A. per- 
sonatus, A. japonicus, and the new species A. heian. 
Only 2 species, with a confused nomenclature, have 
been recognized as valid for the past several decades. 
Without a clear knowledge of the number of species 
present in the area, each with its unique life history 
and biology, the contributions of this genus to the eco- 
system will be inadequately understood and successful 
management of these species will be impossible. 
This study underscores the importance of increas- 
ing our knowledge of forage species. As scientists move 
to manage marine resources with an ecosystem-based 
approach, species at the base of the food web should 
be examined at least as rigorously as species of com- 
mercial interest. In this study, we uncovered aspects of 
the evolutionary history of the genus Ammodytes and 
its current-day isolation into 4 species in the ecosys- 
tem of the North Pacific Ocean and surrounding seas. 
Ecosystem models would benefit from further studies of 
the population structure within each species, as well as 
studies of the unusual life history of these sand- and 
water-dwelling species. 
Acknowledgments 
We thank B. Armstrong for the initial inspiration to 
conduct this project, and P. Mqller and T. Menne (Uni- 
versity of Copenhagen); A. Abookire, R. Armstrong, E. 
Brown, C. Kondzela, C. Munk, D. Nichol, S. Paine, M. 
Paquin, J. Thedinga, S. Johnson, and L. Britt (Alas- 
ka Fisheries Science Center ); J. Blaine, D. Gunder- 
son, and K. Maslenikov (University of Washington); 
A. Kinziger (Humboldt State University); A. Lopez 
(University of Alaska Fairbanks); E. Hilton (Virginia 
Institute of Marine Sciences); A. Kagley (Northwest 
Fisheries Science Center), and the Northeast Fisher- 
ies Science Center Ecosystems Surveys Branch for 
samples. We also thank T. Pietsch and K. Maslenikov 
(University of Washington Fish Collection) for provid- 
ing access to collections, archiving new specimens, and 
facilitating loans from other institutions; P. Bartsch, C. 
Lamour (Museum fur Naturkunde), and R. Thiel (Zoo- 
logical Museum Hamburg) for access to type material; 
and L. Park, P. Schwenke, A. Elz, S. Howard, and J. 
Hemplemann (Northwest Fisheries Science Center) for 
assistance with assays. D. Stevenson (Alaska Fisheries 
Science Center) and W. Grant (Alaska Department of 
Fish and Game) provided critical reviews at the manu- 
script stage. 
Literature cited 
Allen, M., and Smith, G. 
1988. Atlas and zoogeography of common fishes in the 
Bering Sea and northeastern Pacific. NOAA Tech. Rep. 
NMFS 66, 151 p. 
