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<div class=Section1>

<p class=MsoNormal>Journal of Biogeography<o:p></o:p></p>

<p class=MsoNormal style='outline: 0px;background-position:initial initial;
background-repeat:initial initial'><span style='color:#A3A2A2'><a
href="http://onlinelibrary.wiley.com/doi/10.1111/jbi.2014.41.issue-6/issuetoc"><b><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:#007E8B;
border:none windowtext 1.0pt;padding:0in'>Volume 41</span></b><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:#007E8A;
border:none windowtext 1.0pt;padding:0in'>,</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:#007E8A;border:none windowtext 1.0pt;padding:0in;text-decoration:none'> </span></span><b><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:#007E8B;
border:none windowtext 1.0pt;padding:0in'>Issue 6</span></b><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:#007E8A;
border:none windowtext 1.0pt;padding:0in'>,</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:#007E8A;border:none windowtext 1.0pt;padding:0in;text-decoration:none'> </span></span></a></span><b><span
style='color:#007E8B;border:none windowtext 1.0pt;padding:0in'>pages
1171–1182</span></b><span style='color:#A3A2A2'>,</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:#A3A2A2'> </span></span><b><span style='color:#007E8B;border:none windowtext 1.0pt;
padding:0in'>June 2014</span></b><span style='color:#A3A2A2'><o:p></o:p></span></p>

<p class=MsoNormal style='outline: 0px;background-position:initial initial;
background-repeat:initial initial;float:left'><span style='color:black'>Article
first published online: 23 JAN 2014<o:p></o:p></span></p>

<p class=MsoNormal style='outline: 0px;word-wrap: break-word;background-position:
initial initial;background-repeat:initial initial' id=doi><span
style='color:black'>DOI: 10.1111/jbi.12275<o:p></o:p></span></p>

<h1 style='mso-margin-top-alt:0in;margin-right:24.0pt;margin-bottom:0in;
margin-left:0in;margin-bottom:.0001pt;mso-line-height-alt:7.5pt;vertical-align:
baseline'><span class=maintitle><span style='font-size:10.5pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'><o:p> </o:p></span></span></h1>

<h1 style='mso-margin-top-alt:0in;margin-right:24.0pt;margin-bottom:0in;
margin-left:0in;margin-bottom:.0001pt;mso-line-height-alt:7.5pt;vertical-align:
baseline'><span class=maintitle><span style='font-size:18.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'><o:p> </o:p></span></span></h1>

<h1 style='mso-margin-top-alt:0in;margin-right:24.0pt;margin-bottom:0in;
margin-left:0in;margin-bottom:.0001pt;mso-line-height-alt:7.5pt;vertical-align:
baseline'><span class=maintitle><span style='font-size:18.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>Anchovies go north and
west without losing diversity: post-glacial range expansions in a small pelagic
fish</span></span><span style='font-size:18.0pt;font-family:"Arial","sans-serif";
color:black'><o:p></o:p></span></h1>

<p class=MsoNormal style='mso-margin-top-alt:0in;margin-right:3.75pt;
margin-bottom:0in;margin-left:0in;margin-bottom:.0001pt;text-indent:-.25in;
line-height:18.0pt;mso-list:l0 level1 lfo1;vertical-align:baseline'><![if !supportLists]><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'><span
style='mso-list:Ignore'>1.<span style='font:7.0pt "Times New Roman"'>    
</span></span></span><![endif]><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Gonçalo Silva,<span class=apple-converted-space> </span>John
B. Horne and Rita Castilho</span><sup><span style='font-size:7.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>*</span></sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'><o:p></o:p></span></p>

<p style='margin:0in;margin-bottom:.0001pt;line-height:15.6pt;vertical-align:
baseline;outline: 0px;background-position:initial initial;background-repeat:
initial initial' id=publishedOnlineDate><span style='font-size:9.0pt;
font-family:"Arial","sans-serif";color:black'>Centre of Marine Sciences,
CCMAR/CIMAR Associate Laboratory, University of Algarve, Gambelas, Faro,
Portugal. </span><sup><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>*</span></sup><span
class=apple-converted-space><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
color:black'> </span></span><span style='font-size:10.0pt;font-family:
"Arial","sans-serif";color:black'>Correspondence: E-mail:<span
class=apple-converted-space> </span><a href="mailto:rcastil@ualg.pt"
title="Link to email address"><span style='color:#007E8A;border:none windowtext 1.0pt;
padding:0in'>rcastil@ualg.pt</span></a><o:p></o:p></span></p>

<h3 style='margin:0in;margin-bottom:.0001pt;line-height:16.2pt;vertical-align:
baseline'><span style='font-family:"Arial","sans-serif";color:black'><o:p> </o:p></span></h3>

<h3 style='margin:0in;margin-bottom:.0001pt;line-height:16.2pt;vertical-align:
baseline'><span style='font-family:"Arial","sans-serif";color:black'>Abstract<o:p></o:p></span></h3>

<h4 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.05in;
margin-left:0in;line-height:19.2pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-family:"Arial","sans-serif";color:black'>Aim<o:p></o:p></span></h4>

<p style='margin:0in;margin-bottom:.0001pt;line-height:18.0pt;vertical-align:
baseline;outline: 0px;background-position:initial initial;background-repeat:
initial initial'><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>As part of an emerging effort to understand the role played by
climatic fluctuations in shaping the geographical distributions and abundances
of marine organisms, we examined the genetic patterns of leading-edge
populations in the European anchovy,</span><span class=apple-converted-space><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'> </span></span><em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>Engraulis encrasicolus</span></em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>, and its
American counterpart, the morphologically similar silver anchovy,</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'> </span></span><em><span style='font-size:9.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>Engraulis
eurystole</span></em><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>, in the North Atlantic Ocean.<o:p></o:p></span></p>

<h4 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.05in;
margin-left:0in;line-height:19.2pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-family:"Arial","sans-serif";color:black'>Location<o:p></o:p></span></h4>

<p style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:12.0pt;
margin-left:0in;line-height:18.0pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Adults
were collected from the western Atlantic, eastern Atlantic (from Norway to
Ghana) and western Mediterranean.<o:p></o:p></span></p>

<h4 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.05in;
margin-left:0in;line-height:19.2pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-family:"Arial","sans-serif";color:black'>Methods<o:p></o:p></span></h4>

<p style='margin:0in;margin-bottom:.0001pt;line-height:18.0pt;vertical-align:
baseline;outline: 0px;background-position:initial initial;background-repeat:
initial initial'><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>A 1045 bp fragment of the mtDNA cytochrome</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'> </span></span><em><span style='font-size:9.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>b</span></em><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'> </span></span><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>gene was sequenced (<em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'>n </span></em>=<em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'> </span></em>312) and nine microsatellite loci were genotyped
(<em><span style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>n </span></em>=<em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'> </span></em>462) for anchovies
from 13 locations across the temperate North Atlantic. Populations were
surveyed for diversity and differentiation with a range of summary statistics.
Multivariate discriminant analysis of principal components was employed to
detect the number of genetic clusters in the data and assign individuals to
populations based on their microsatellite genotypes. Historical demographic
inferences – mismatch distributions and Bayesian skyline plots –
were used to observe population size changes relating to climatic oscillations.<o:p></o:p></span></p>

<h4 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.05in;
margin-left:0in;line-height:19.2pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-family:"Arial","sans-serif";color:black'>Results<o:p></o:p></span></h4>

<p style='margin:0in;margin-bottom:.0001pt;line-height:18.0pt;vertical-align:
baseline;outline: 0px;background-position:initial initial;background-repeat:
initial initial'><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Two mitochondrial clades were recovered, consistent with previous
studies of</span><span class=apple-converted-space><span style='font-size:9.0pt;
font-family:"Arial","sans-serif";color:black'> </span></span><em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>E. encrasicolus</span></em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>, in which
the frequency of each clade varied by latitude. Four genetic clusters
corresponding loosely to large geographical regions were identified with
microsatellite data. The north-western Atlantic</span><span
class=apple-converted-space><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'> </span></span><em><span style='font-size:9.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>E.
eurystole</span></em><span class=apple-converted-space><span style='font-size:
9.0pt;font-family:"Arial","sans-serif";color:black'> </span></span><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>was not
reciprocally monophyletic for either mtDNA or microsatellite analyses and is
probably conspecific with</span><span class=apple-converted-space><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'> </span></span><em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>E. encrasicolus</span></em><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>. Genetic
diversity peaked in Iberian populations, but differences in genetic diversity
were only statistically significant for the least diverse population, Tangier.
The indications of demographic expansion were more pronounced in the southern
clade and both mtDNA clades exhibited genetic diversity and expansion imprints
that are likely to be older than climatic oscillations of the recent
Pleistocene.<o:p></o:p></span></p>

<h4 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.05in;
margin-left:0in;line-height:19.2pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-family:"Arial","sans-serif";color:black'>Main conclusions<o:p></o:p></span></h4>

<p style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:12.0pt;
margin-left:0in;line-height:18.0pt;vertical-align:baseline;outline: 0px;
background-position:initial initial;background-repeat:initial initial'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>The highly
mobile nature of anchovies has allowed them to track their optimal thermal
physiological conditions during the extreme climate shifts of the Last Glacial
Maximum and avoid wholesale population reductions and genetic bottlenecks. Both
north-eastern and north-western Atlantic were probably rapidly recolonized
after the Last Glacial Maximum by large numbers of anchovies, such that
leading-edge populations retained the genetic diversity of parent populations.<o:p></o:p></span></p>

<p class=MsoNormal><o:p> </o:p></p>

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