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<div class=Section1>

<p class=MsoNormal><span style='color:#5D5D5D'>Journal of Biogeography<o:p></o:p></span></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-5/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 5</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
965–976</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'>May 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'>Article first published online:
15 JAN 2014<o:p></o:p></p>

<p class=MsoNormal style='outline: 0px;word-wrap: break-word;background-position:
initial initial;background-repeat:initial initial' id=doi>DOI: 10.1111/jbi.12260<o:p></o:p></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: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'>Phylogeography of a
marine acanthocephalan: lack of cryptic diversity in a cosmopolitan parasite of
mole crabs</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:10.0pt;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:10.0pt;font-family:"Arial","sans-serif";
color:black'>Tricia C. Goulding<sup><span style='border:none windowtext 1.0pt;
padding:0in'>1,2</span></sup><span class=apple-converted-space> </span>and
C. Sarah Cohen<sup><span style='border:none windowtext 1.0pt;padding:0in'>1,*</span></sup><o:p></o:p></span></p>

<p class=MsoNormal style='mso-line-height-alt:7.5pt;vertical-align:baseline;
outline: 0px;background-position:initial initial;background-repeat:initial initial'
id=publishedOnlineDate><sup><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>1</span></sup><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";color:black'>Romberg
Tiburon Center for Environmental Studies, Biology Department, San Francisco
State University, Tiburon, CA, USA<sup><span style='border:none windowtext 1.0pt;
padding:0in'>2</span></sup>Pennsylvania State University, University Park, PA,
USA <sup><span style='border:none windowtext 1.0pt;padding:0in'>*</span></sup><span
class=apple-converted-space> </span>Correspondence: E-mail:<span
class=apple-converted-space> </span><a href="mailto:sarahcoh@sfsu.edu"
title="Link to email address"><span style='color:#007E8A;border:none windowtext 1.0pt;
padding:0in'>sarahcoh@sfsu.edu</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'>Little is known about phylogeography and cryptic diversity of
parasites in the marine environment. The acanthocephalan</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'>Profilicollis
altmani</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'>parasitizes
intermediate hosts that are broadly distributed around the Americas and final
hosts that are highly motile. We investigated the spatial genetic structure of
this acanthocephalan found in three species 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'>Emerita</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'>crabs: (1) to test whether land masses serve as biogeographic barriers
promoting ocean basin divergence among parasite lineages or species; and (2) to
test whether the distribution of parasite species matches the distribution of
different crab host species.<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'>The
Pacific, Atlantic and Gulf coasts of the USA, and the Pacific coast of Panama
and Chile.<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'>Sequences of 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'>c</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'>oxidase subunit I (<em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'>COI</span></em>) and ribosomal
internal transcribed spacers (ITS) were obtained from 204 acanthocephalans.
Parasites were sampled from crabs in 15 sampling localities. These sequences
were analysed with coalescent-based methods and other population genetic
analyses to infer phylogeographic patterns.<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'>Haplotype diversity for</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'>COI</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'>sequences was high (0.96) among parasites sampled, but nucleotide
diversity was low (0.071) and there was no distinct geographic pattern between
regions. Pairwise genetic distances were generally low, although there was a
degree of population structure between oceans. Sequence comparisons showing an
excess of low divergence alleles and a bimodal mismatch distribution provide
evidence of either past selective events or demographic expansions. No
variation was observed in the ITS sequences.<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='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'>The lack of geographic patterning in haplotype diversity of this
parasite indicates that gene flow is probably occurring between ocean basins.
In addition, the low genetic diversity suggests that the acanthocephalan
parasitizing</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. analoga</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'>in Chile is conspecific to the species found parasitizing several</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'>Emerita</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'>species along the coasts of North America, and is thus a
cosmopolitan parasite that is most likely dispersed long distances by marine
birds that serve as definitive hosts.<o:p></o:p></span></p>

<p class=MsoNormal><o:p> </o:p></p>

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