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<div class=Section1>
<p class=MsoNormal><b><span style='font-size:10.0pt;font-family:"Arial","sans-serif"'>Global
Ecology and Biogeography<o:p></o:p></span></b></p>
<p class=MsoNormal><span class=article-headermeta-info-label><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>First published:</span></b></span><span
class=apple-converted-space><b><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'> </span></b></span><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'>27 April 2017 <o:p></o:p></span></p>
<p class=MsoNormal><span class=article-headermeta-info-label><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>DOI:</span></b></span><span class=apple-converted-space><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'> </span></b></span><span class=article-headermeta-info-data><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>10.1111/geb.12588</span></span><span class=apple-converted-space><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'> </span></span><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'> <span
style='color:#414141'><o:p></o:p></span></span></p>
<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:5.05pt;
margin-left:0in;vertical-align:baseline'><span style='font-size:19.0pt;
font-family:"Arial","sans-serif";color:black'><o:p> </o:p></span></h1>
<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:5.05pt;
margin-left:0in;vertical-align:baseline'><span style='font-size:19.0pt;
font-family:"Arial","sans-serif";color:black'>Aggregate patterns of macrofaunal
diversity: An interocean comparison<o:p></o:p></span></h1>
<h3 style='mso-margin-top-alt:0in;margin-right:3.75pt;margin-bottom:0in;
margin-left:0in;margin-bottom:.0001pt;text-indent:-.25in;mso-list:l2 level1 lfo4;
background:white;vertical-align:baseline'><![if !supportLists]><span
style='font-size:10.0pt;font-family:Symbol;font-weight:normal'><span
style='mso-list:Ignore'>·<span style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'>Omar Defeo, </span><span style='font-size:12.5pt;
font-family:"Arial","sans-serif";font-weight:normal'>Carlos A. M. Barboza,</span><span
style='font-size:12.5pt;font-family:"Arial","sans-serif";font-weight:normal'> </span><span
style='font-size:12.5pt;font-family:"Arial","sans-serif";font-weight:normal'>Francisco
R. Barboza,</span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'> </span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'>William H. Aeberhard,</span><span style='font-size:12.5pt;
font-family:"Arial","sans-serif";font-weight:normal'> </span><span
style='font-size:12.5pt;font-family:"Arial","sans-serif";font-weight:normal'>Tatiana
M. B. Cabrini,</span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'> </span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'>Ricardo S. Cardoso,</span><span style='font-size:12.5pt;
font-family:"Arial","sans-serif";font-weight:normal'> </span><span
style='font-size:12.5pt;font-family:"Arial","sans-serif";font-weight:normal'>Leonardo
Ortega,</span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'> </span><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'>Viviane Skinner,Boris Worm<o:p></o:p></span></h3>
<p class=MsoNormal style='mso-margin-top-alt:0in;margin-right:3.75pt;
margin-bottom:6.0pt;margin-left:0in;text-indent:-.25in;mso-list:l2 level2 lfo4;
background:white;vertical-align:baseline'><![if !supportLists]><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'><span
style='mso-list:Ignore'>0.<span style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><span style='font-size:10.0pt;font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<p class=MsoNormal style='margin-left:0in;text-indent:-.25in;mso-list:l0 level1 lfo5;
vertical-align:baseline'><![if !supportLists]><span style='font-size:10.0pt;
font-family:Symbol'><span style='mso-list:Ignore'>·<span style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><b><span style='font-size:14.0pt;font-family:
"Arial","sans-serif"'>Abstract</span></b><span style='font-size:14.0pt;
font-family:"Arial","sans-serif"'><o:p></o:p></span></p>
<p class=MsoNormal style='margin-left:0in;text-indent:-.25in;mso-list:l0 level1 lfo5;
vertical-align:baseline'><![if !supportLists]><span style='font-size:10.0pt;
font-family:Symbol;color:#414141'><span style='mso-list:Ignore'>·<span
style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><span style='font-size:19.0pt;font-family:"Arial","sans-serif";
color:#414141'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Aim<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>While geographical patterns of species richness are reasonably
well explored for single well-studied taxa, less is known about aggregate
patterns of total richness for major biomes and their environmental correlates.
Here we analyse continental-scale aggregate patterns of macrofaunal diversity
for sandy beaches, a dominant habitat along the Atlantic and Pacific coasts of
South America.<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Location<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>South American coastlines.<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Time period<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>Present day (data amassed from studies performed since
1971).<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Major taxa studied<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>Benthic macrofauna, including crustaceans, polychaetes
and molluscs.<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Methods<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>We compiled richness information for all macrofaunal
groups on 263 sandy beaches in South America using standard criteria. We
further matched these data with environmental variables including sea surface
temperature (SST), chlorophyll<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>a</span></em>, grain size, beach slope, tide range and various
morphometrics. We used generalized linear mixed models to relate environmental
factors to observed variation in total macrofaunal richness across all beaches,
testing competing hypotheses about environmental correlates and possible
drivers of latitudinal diversity.<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Results<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>Macrofaunal richness decreased from tropical to temperate
beaches in the Pacific and followed a parabolic trend in the Atlantic, with the
highest biodiversity found at tropical and mid-latitudinal bands. Beach slope,
tidal range and chlorophyll<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>a</span></em><span class=apple-converted-space> </span>mostly
explained latitudinal trends in macrofaunal richness, followed by grain size,
SST and ocean basin.<o:p></o:p></span></p>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline'><span
style='font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>
<h3 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:9.45pt;
margin-left:0in;vertical-align:baseline;outline: 0px;background-position-x:
0px;background-position-y:0px'><span style='font-size:14.5pt;font-family:"Arial","sans-serif";
color:black;font-weight:normal'>Main conclusions<o:p></o:p></span></h3>
<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>This study indicates that richness of macrofaunal species
at a given beach is most closely related to characteristics of the physical
habitat, such as beach slope, area and grain size. At this scale, planktonic
food supply also appeared to be more important than temperature, which is a
dominant explanatory variable of global-scale variation in species richness.<o:p></o:p></span></p>
<p class=MsoNormal><o:p> </o:p></p>
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