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<div class=Section1>
<p class=MsoNormal>Global Ecology and Biogeography<o:p></o:p></p>
<p class=MsoNormal style='outline: 0px'><span style='color:#007E8B'><a
href="http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1466-8238/earlyview"><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:#007E8A;
border:none windowtext 1.0pt;padding:0in'>Early View (Online Version of Record
published before inclusion in an issue)</span></a><o:p></o:p></span></p>
<p class=MsoNormal><span style='color:black'>Article first published online: 25
MAY 2015<o:p></o:p></span></p>
<p class=MsoNormal style='outline: 0px;word-wrap: break-word' id=doi><span
style='color:black'>DOI: 10.1111/geb.12328<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:16.0pt;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:16.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>Geographic patterns of
diversification and the latitudinal gradient of richness of rocky intertidal
gastropods: the ‘into the tropical museum’ hypothesis</span></span><span
style='font-size:16.0pt;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:l1 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 lang=ES style='font-size:9.0pt;font-family:
"Arial","sans-serif";color:black'>Marcelo M. Rivadeneira</span><sup><span
lang=ES style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>1,*</span></sup><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Alex H. Alballay</span><sup><span
lang=ES style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>1</span></sup><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Jaime A. Villafaña</span><sup><span
lang=ES style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>1</span></sup><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Peter T. Raimondi</span><sup><span
lang=ES style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>2</span></sup><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,</span><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Carol A.
Blanchette</span><sup><span style='font-size:7.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>3</span></sup><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'>and Phillip B. Fenberg</span><sup><span style='font-size:7.0pt;
font-family:"Arial","sans-serif";color:black;border:none windowtext 1.0pt;
padding:0in'>4</span></sup><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'><o:p></o:p></span></p>
<p class=MsoNormal style='mso-line-height-alt:7.5pt;vertical-align:baseline;
outline: 0px' id=publishedOnlineDate><span class=affiliationnumber><span
lang=ES style='font-size:8.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>1 </span></span><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Laboratorio
de Paleobiología, Centro de Estudios Avanzados en Zonas Áridas (CEAZA) & Universidad
Católica del Norte, Coquimbo, Chile </span><span class=affiliationnumber><span
lang=ES style='font-size:8.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>2 </span></span><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Department
of Ecology and Evolutionary Biology, Long Marine Laboratory, University of
California, Santa Cruz, CA, USA </span><span class=affiliationnumber><span
lang=ES style='font-size:8.0pt;font-family:"Arial","sans-serif";color:black;
border:none windowtext 1.0pt;padding:0in'>3 </span></span><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Marine
Science Institute, University of California, Santa Barbara, CA, USA </span><span
class=affiliationnumber><span lang=ES style='font-size:8.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>4</span></span><span
lang=ES 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'><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Ocean and Earth Science, National Oceanography Centre, University
of Southampton, Southampton, UK. </span><sup><span lang=ES style='font-size:
9.0pt;font-family:"Arial","sans-serif";color:black;border:none windowtext 1.0pt;
padding:0in'>*</span></sup><span class=apple-converted-space><span lang=ES
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'> </span></span><span
lang=ES style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Correspondence:
E-mail:</span><span class=apple-converted-space><span lang=ES
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'><a
href="mailto:marcelo.rivadeneira@ceaza.cl" title="Link to email address"><span
lang=ES style='color:#007E8A;border:none windowtext 1.0pt;padding:0in'>marcelo.rivadeneira@ceaza.cl</span></a></span><span
lang=ES style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'><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'><span
style='font-family:"Arial","sans-serif";color:black'>Aim<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'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>To
evaluate the existence of a latitudinal gradient of richness (LGR) in rocky
intertidal gastropods and the role of evolutionary processes in shaping the
LGR.<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'><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'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>The entire
eastern Pacific coast, from Alaska to Tierra del Fuego, encompassing ca. 120°
latitude.<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'><span
style='font-family:"Arial","sans-serif";color:black'>Methods<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'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>The LGR
was assessed using field surveys and a comprehensive literature dataset from
308 rocky intertidal sites, representing 328 species and 159 genera. The
importance of geographic patterns of diversification in shaping the LGR was assessed
using three complementary approaches. First, we used the fossil record to
evaluate the region of origination of genera (tropical vs. extra-tropical).
Secondly, we used a nestedness analysis to compare present-day biogeographic
distributions of genera with their region of origin. Finally, we evaluated the
importance of biogeographic dynamics for shaping the LGR using a projection
matrix and estimating transition probabilities among different biogeographic
states.<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'><span
style='font-family:"Arial","sans-serif";color:black'>Results<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'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Eastern
Pacific rocky intertidal gastropods follow the canonical latitudinal gradient
of richness (higher richness in the tropics). Originations were similar in the
tropics and extra-tropics, a pattern that was robust to geographic differences
in fossil sampling. The biogeographic distribution of genera was significantly
nested, irrespective of the region of origin of genera. The distribution
dynamics model accurately reproduced the LGR and showed that key-transitions
are low extinction in the tropics and high dispersal of taxa from the
extra-tropics to the tropics.<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'><span
style='font-family:"Arial","sans-serif";color:black'>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'><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>The
existence of a canonical LGR in rocky intertidal gastropods can be explained by
the combined effect of reduced extinction rates in the tropics and the range
expansion of taxa from the extra-tropics toward the tropics, in what we have
called the ‘into the tropical museum’ hypothesis.<o:p></o:p></span></p>
<p class=MsoNormal><o:p> </o:p></p>
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