134 
Fishery Bulletin 113(2) 
A. personatus 
A. hexapterus 
A. heian n. sp 
A. japonicus 
tobianus 
Figure 2 
Haplotype network of species of Ammodytes . Size of circle indicates number 
of individuals with that haplotype. Length of branch indicates number of 
segregating nucleotides; numbers on the longer branches are the number of 
nucleotide differences between haplotypes. In the clade of A. dubius and A. 
americanus, A. americanus haplotypes are represented by filled circles and A. 
dubius haplotypes by open circles. 
nologies Inc., Warrenton, VA), Spotfire S+ 8.2 (TIBCO 
Software Inc., Palo Alto, CA), and SPSS 11.5.1 (IBM, 
Armonk, NY). Using the Bonferroni method to adjust 
for multiple comparisons, we considered differences 
significant at P<0.05. 
More than 600 specimens were examined morpho- 
logically for meristic and morphometric data or for 
meristic data only. For all specimens identified geneti- 
cally or by geographic range, meristic frequency dis- 
tributions are presented. A reduced data set of 214 
genetically identified specimens that had a complete 
suite of meristic and morphometric character data was 
used to conduct both univariate and multivariate anal- 
yses. Arcsine-transformed morphometric ratios (with 
SL as denominator) were tested to meet the assump- 
tions of normality and equality of variance required 
for analysis of covariance (ANCOVA). A Kruskal- Wallis 
nonparametric test was used to test for differences in 
all characters with heterogenetic variances. Differences 
were considered significant at PcO.OOl. A stepwise dis- 
criminant function analysis was conducted with mor- 
phometric and meristic data to establish the relative 
significance of characters in distinguishing the species 
and to identify specimens preserved in formalin with- 
out tissues available for genetic identification. Morpho- 
metric data were standardized by dividing values by 
SL. The robustness of the discriminant function anal- 
ysis was tested with a leave-one-out cross-validation 
procedure conducted in SPSS 11.5.1. Allometry was 
evaluated by plotting each measurement divided by 
head length (HL) against SL. 
Results 
Genetics 
In the 560-bp COI sequences, 152 haplotypes were 
identified at 176 polymorphic sites, with no insertions 
or deletions. Seven monophyletic clades emerged from 
the sequence analysis of Pacific and Atlantic speci- 
mens. Each clade included many singleton haplotypes 
and up to 4-bp differences radiating from a common 
haplotype (Fig. 2). 
Four clades within the North Pacific region were rep- 
resented in the haplotype network of Ammodytes (Fig. 
2). We refer to these clades as A. personatus (range: 
throughout the eastern North Pacific Ocean from Cali- 
