Orr et al.: Systematics of North Pacific sand lances of the genu s Ammodytes and description of a new species 
149 
Distribution 
On the basis of material examined, we determined 
that Ammodytes heian n. sp. ranges from the southern 
Sea of Okhotsk to the Pacific Ocean off the northern 
part of Japan (Fig. 1). In the southern Sea of Okhotsk, 
its range overlaps with that of A. hexapterus and A. 
japonicus in the Soya Strait off Wakkanai, Hokkaido 
(Fig. 1). Published sequences in GenBank (FJ666915, 
FJ666916) indicate that A. heian n. sp. also is found in 
the Sea of Japan (Table 4). 
Etymology 
The specific name heian is a transliteration of the 
Japanese word for “peace.” The holotype and paratypes 
were collected from waters off Fukushima and Iwate 
prefectures, before the earthquake and tsunami of 
March 2011. 
Remarks 
Early surveys of vertebral counts in Ammodytes by 
Kawamura (1940), Ohshima (1950), and Hashimoto 
and Kawasaki (1981) around Japan revealed signifi- 
cant variation in different regions. Populations from 
off Onagawa on the northeastern Pacific coast and in 
Moura Bay at the northern tip of Honshu (Ohshima, 
1950, table 6, 61-66) are clearly different from popula- 
tions in the Sea of Japan and farther south along the 
Pacific coast and match our data for A. heian n. sp. 
(Tables 5 and 7). Two populations examined by Hashi- 
moto and Kawasaki (1981) had significantly different 
vertebral counts, as well as differences in life history 
characteristics. In a more comprehensive work, Hashi- 
moto (1984) surveyed additional meristic characters, 
classifying populations around Japan into 3 groups: 
a northern group (“I”;=A. hexapterus) found in Japan 
only in Soya Strait; a northcentral group (“H”;=A. he- 
ian n. sp.) found on the northeastern Pacific Ocean 
side of Honshu and in the Sea of Japan from Tottori 
to the Soya Strait; and a southcentral group (“IH”;=A. 
japonicus) found in the southern Sea of Japan, on the 
southcentral side of Honshu in the Pacific Ocean, and 
in the Seto Inland Sea. 
Genetic research has consistently shown strong het- 
erogeneity among populations around Japan (all origi- 
nally identified as A. personatus), whether they are 
strictly considered as one species or potentially as more 
than one. Distributions of allozyme genotypes present- 
ed by Okamoto (1989) revealed 3 genetically isolated 
groups in Japan concordant with our results: a north- 
ern Japan group (likely representing A. heian n. sp., on 
the basis of the distribution of our material), a south- 
ern Japan group (=A. japonicus), a Kushiro and Wak- 
kanai group (=A. hexapterus)-, also shown was a fourth 
group from Kodiak, Alaska (=A. personatus). Later 
work by Kim et al. (2006, 2008, 2010) with mtDNA also 
revealed genetic differences among populations around 
Korea. Through comparisons between our sequence 
data and data in GenBank and BOLD, we identified 2 
individuals of A. heian n. sp. from the Sea of Japan in 
the material of Kim et al. (2006, 2008, 2010; Table 4); 
all other individuals were identified by us as A. japoni- 
cus. Most recently, Han et al. (2012) found 2 lineages 
around Japan and in the Yellow Sea on the basis of 
analysis of the mitochondrial control region, conclud- 
ing that further study with nuclear DNA markers is 
needed to clarify their taxonomic status. However, the 
morphological and COI sequence differences disclosed 
here as well as the isozyme analysis of Hashimoto and 
Kawasaki (1981) strongly support the conclusion that 
the lineages represent 2 species. Lineage A appears to 
represent A. heian n. sp., and Lineage B, A. japonicus. 
Discussion 
Our primary objective was to taxonomically re- 
vise the genus Ammodytes in the North Pacific Ocean 
and its fringing seas by examining morphological and 
mtDNA variation. Morphological and genetic analyses 
show that the genus Ammodytes comprises 4 species in 
the North Pacific region: Ammodytes personatus, rang- 
ing from California to the Aleutian Islands and eastern 
Bering Sea; A. hexapterus, ranging from the Hudson 
Bay in the east, through the Chukchi and Bering seas, 
to the western Pacific region in the Sea of Okhotsk and 
to Soya Strait between Hokkaido and Sakhalin islands; 
A. japonicus, found in the Sea of Japan, the Seto Inland 
Sea, the Pacific Ocean side of southern Honshu, and 
the Yellow Sea; and a new species, A. heian, herein de- 
scribed and found on the Pacific Ocean side of northern 
Honshu, in the Soya Strait, and in the Sea of Japan. 
Although clearly distinct genetically, some species 
are morphologically similar to one another and, despite 
significant differences found among several characters, 
only combinations of morphometric and meristic char- 
acters serve to distinguish the species from one anoth- 
er. The species of the eastern or northern areas of the 
North Pacific region (A. personatus and A. hexapterus), 
together, can readily be distinguished from the species 
of the western North Pacific region (A. japonicus and 
A. heian n. sp.) on the basis of vertebral counts, but 
species within these pairs are nearly indistinguish- 
able morphologically. To add to the challenge of species 
identification, ranges of A. personatus and A. hexapter- 
us overlap in the northern Bering Sea, and A. japoni- 
cus and A. heian n. sp. were found together in our Soya 
Strait material; data from other authors strongly indi- 
cate that the latter 2 species are also found together 
in the Sea of Japan (Hashimoto, 1984; Kim et al., 2006, 
2008, 2010; Han et al., 2012). 
The distribution of each species generally corre- 
sponds well with general zoogeographic regions (Fig. 
1). Ammodytes personatus exhibits an Aleutian-Or- 
egonian distribution (Allen and Smith, 1988; Briggs, 
1995; Logerwell et al., 2005) and is found in the east- 
ern North Pacific Ocean from California to the Gulf 
of Alaska, the Aleutian Islands, and into the eastern 
Bering Sea, where to the north it is found sympatri- 
