129 
NOAA 
National Marine 
Fisheries Service 
Fishery Bulletin 
rtr established 1881 -<?. 
Spencer F. Baird 
First U.S. Commissioner 
of Fisheries and founder 
of Fishery Bulletin 
Systematics of North Pacific sand lances of 
the genus Ammodytes based on molecular and 
morphological evidence, with the description of 
a new species from Japan 
James W. Orr (contact author ) 1 
Sharon Wildes 2 
Yoshiaki Kai 3 
Nate Raring 1 
T. Nakabo 4 
Oleg Katugin 5 
Jeff Guyon 2 
Email address for contact author: iames.orr@noaa.gov 
Abstract— The systematic status 
of North Pacific sand lances (ge- 
nus Ammodytes) was assessed from 
mitochondrial DNA (cytochrome 
oxidase c subunit 1) sequence data 
and morphological data to identify 
the number of species in the North 
Pacific Ocean and its fringing seas. 
Although only 2 species, Ammodytes 
hexapterus and A. personatus, have 
been considered valid in the region, 
haplotype networks and trees con- 
structed with maximum parsimony 
and genetic distance (neighbor- 
joining) methods revealed 4 highly 
divergent monophyletic clades that 
clearly represent 4 species of Ammo- 
dytes in the North Pacific region. On 
the basis of our material and com- 
parisons with sequence data report- 
ed in online databases, A. personatus 
is found throughout the eastern 
North Pacific Ocean, Gulf of Alaska, 
Aleutian Islands, and the eastern 
Bering Sea where it co-occurs with 
a northwestern Arctic species, A. 
hexapterus, that is found through- 
out the North American Arctic from 
Hudson Bay, Canada, in the east, 
through the Beaufort and Chukchi 
seas, into the northern and western 
Bering Sea, and to the southern Sea 
of Okhotsk in the Soya Strait off 
Hokkaido, Japan. Two other species 
reside in waters around Japan: A. 
japonicus throughout the Sea of Ja- 
pan and the Seto Inland Sea and a 
new species in the Sea of Japan and 
the North Pacific Ocean off north- 
ern Honshu. We designate neotypes 
for A. hexapterus and A. personatus 
because of the absence of type mate- 
rial and the close similarity of these 
2 species. Ammodytes aleutensis is a 
junior synonym of A. japonicus, and 
A. alascanus is a junior synonym of 
A. personatus. 
Manuscript submitted 6 February 2014. 
Manuscript accepted 30 January 2015. 
Fish. Bull. 113:129-156 (2015). 
Electronic publication date: 10 March 2015. 
doi: 10. 7755/FB. 113.2.3 
http://zoobank.org/References/29FEAF57- 
8178-45DB-8A23-76BE33BB2B0D 
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. 
1 Resource Assessment and Conservation 
Engineering Division 
Alaska Fisheries Science Center 
National Marine Fisheries Service, NOAA 
7600 Sand Point Way NE 
Seattle, Washington 98115-6349 
2 Auke Bay Laboratories 
Alaska Fisheries Science Center 
National Marine Fisheries Service, NOAA 
17109 Point Lena Loop Road 
Juneau, Alaska 99801-8344 
Sand lances of the genus Ammodytes 
(family Ammodytidae) are small 
silvery fishes that inhabit marine 
and adjacent brackish waters of the 
Northern Hemisphere. They reside 
primarily over sandy substrates, 
where they are able to quickly bury 
themselves to avoid predators, but 
are found also over bedrock, eelgrass, 
and kelp. Species of Ammodytes are 
cornerstone prey items for more 
than 100 species of marine mam- 
mals, birds, and other fishes in the 
region of the North Pacific Ocean 
(Field, 1988; Willson et al. 1 ). Despite 
1 Willson, M. F., R. H. Armstrong, M. D. 
Robards, and J. F. Piatt. 1999. Sand 
lance as cornerstone prey for predator 
populations. In Sand lance: a review of 
biology and predator relations and anno- 
tated bibliography (M. D. Robards, M. F. 
3 Maizuru Fisheries Research Station 
Field Science Education and Research Center 
Kyoto University 
Nagahama, Maizuru 
Kyoto 625-0086, Japan 
4 The Kyoto University Museum 
Kyoto University 
Yoshida, Sakyo 
Kyoto 606-8501, Japan 
5 Pacific Research Institute of 
Fisheries and Oceanography (TINRO-Center) 
4 Shevchenko Alley 
Vladivostok, Primorsky Kray, Russia, 690950 
their tremendous importance in eco- 
systems and as targets in fisheries 
off Japan (Hamada, 1985; Han et al., 
2012) and in the North Sea (Sher- 
man et al., 1981; Furness, 2002), lit- 
tle is known about their distribution 
or life history. 
The family Ammodytidae com- 
prises about 31 species in 7 gen- 
era and 2 subfamilies found in the 
Arctic, Atlantic, Indian, and Pacific 
oceans (Nelson, 2006; Randall and 
Heemstra, 2008; Shibukawa and Ida, 
2013; Randall and Ida, 2014). Placed 
within the subfamily Ammodytinae, 
Willson, R. H. Armstrong, and J. F. Piatt, 
eds.), p. 17-44. Research Paper PNW- 
RP-521. U.S. Department of Agricul- 
ture, Forest Service, Pacific Northwest 
Research Station. Portland OR. [http:// 
www.fs.fed.us/pnw/pubs/rp_521a.pdfl 
