<META HTTP-EQUIV="Content-Type" CONTENT="text/html; charset=iso-2022-jp">
<html xmlns:v="urn:schemas-microsoft-com:vml" xmlns:o="urn:schemas-microsoft-com:office:office" xmlns:w="urn:schemas-microsoft-com:office:word" xmlns:m="http://schemas.microsoft.com/office/2004/12/omml" xmlns="http://www.w3.org/TR/REC-html40"><head><meta name=Generator content="Microsoft Word 15 (filtered medium)"><style><!--
/* Font Definitions */
@font-face
{font-family:"Cambria Math";
panose-1:2 4 5 3 5 4 6 3 2 4;}
@font-face
{font-family:Calibri;
panose-1:2 15 5 2 2 2 4 3 2 4;}
/* Style Definitions */
p.MsoNormal, li.MsoNormal, div.MsoNormal
{margin:0cm;
margin-bottom:.0001pt;
font-size:11.0pt;
font-family:"Calibri","sans-serif";
mso-fareast-language:EN-US;}
a:link, span.MsoHyperlink
{mso-style-priority:99;
color:#0563C1;
text-decoration:underline;}
a:visited, span.MsoHyperlinkFollowed
{mso-style-priority:99;
color:#954F72;
text-decoration:underline;}
span.EstiloCorreo17
{mso-style-type:personal-compose;
font-family:"Calibri","sans-serif";
color:windowtext;}
.MsoChpDefault
{mso-style-type:export-only;
font-family:"Calibri","sans-serif";
mso-fareast-language:EN-US;}
@page WordSection1
{size:612.0pt 792.0pt;
margin:70.85pt 3.0cm 70.85pt 3.0cm;}
div.WordSection1
{page:WordSection1;}
--></style><!--[if gte mso 9]><xml>
<o:shapedefaults v:ext="edit" spidmax="1026" />
</xml><![endif]--><!--[if gte mso 9]><xml>
<o:shapelayout v:ext="edit">
<o:idmap v:ext="edit" data="1" />
</o:shapelayout></xml><![endif]--></head><body lang=ES link="#0563C1" vlink="#954F72"><div class=WordSection1><p class=MsoNormal><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Global Ecology and Biogeography<o:p></o:p></span></p><p class=MsoNormal><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>First published: 06 November 2019<o:p></o:p></span></p><p class=MsoNormal><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'><a href="https://doi.org/10.1111/geb.13023"><span lang=EN-US style='color:windowtext;text-decoration:none'>https://doi.org/10.1111/geb.13023</span></a></span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'><o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:24.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Global meta</span></b><b><span lang=EN-US style='font-size:24.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span></b><b><span lang=EN-US style='font-size:24.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>analysis of how marine upwelling affects herbivory<o:p></o:p></span></b></p><p class=MsoNormal><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'><a href="https://onlinelibrary.wiley.com/action/doSearch?ContribAuthorStored=Sellers%2C+Andrew+J"><span lang=EN-US style='color:windowtext;text-decoration:none'>Andrew J. Sellers</span></a></span><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'> </span><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'><a href="https://onlinelibrary.wiley.com/action/doSearch?ContribAuthorStored=Leung%2C+Brian"><span lang=EN-US style='color:windowtext;text-decoration:none'>Brian Leung</span></a></span><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'> </span><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'><a href="https://onlinelibrary.wiley.com/action/doSearch?ContribAuthorStored=Torchin%2C+Mark+E"><span lang=EN-US style='color:windowtext;text-decoration:none'>Mark E. Torchin</span></a></span><span style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'> <span lang=EN-US><o:p></o:p></span></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:16.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Abstract<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Aim<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Nutrient subsidies support high primary productivity, increasing herbivore abundance and influencing their top</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>down control of producers. Wind</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>driven upwelling events deliver cold nutrient</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>rich water to coastlines, supporting highly productive marine environments. Results from studies comparing ecological processes across upwelling regimes are mixed: some reveal weaker herbivory in upwelling regions, while others report a positive relationship between upwelling and herbivory. In this synthesis we examine the influence of upwelling on top</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>down control of producers across the globe.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Location<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Global; marine ecosystems.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Time period<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>1978–2017.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Major taxa studied<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Marine herbivores and algae.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Methods<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>We used data from herbivory studies focusing specifically on the influence of upwelling activity (upwelling studies), and a broader collection of herbivore exclusion studies dating back four decades. For the upwelling studies we compared herbivore effects between experiments replicated across sites for which upwelling conditions were described by the authors. Meanwhile, for the broader collection of experiments we used externally sourced oceanographic data to characterize upwelling activity, and examined how herbivory changed along a gradient of upwelling activity.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Results<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Our results consistently reveal that upwelling weakens herbivore effects on producers. Herbivory was, on average, four times weaker in upwelling sites relative to sites under weak upwelling or downwelling regimes in studies that specifically examined upwelling. The analysis of the broader herbivory literature revealed a similar weakening influence of upwelling on herbivory; however, the effect size was smaller and varied across producer functional groups.<o:p></o:p></span></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><b><span lang=EN-US style='font-size:13.5pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Main conclusions<o:p></o:p></span></b></p><p class=MsoNormal style='mso-margin-top-alt:auto;mso-margin-bottom-alt:auto'><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>Nutrient subsidies from upwelling events reduce top</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>down control by herbivores in coastal ecosystems; however, the negative relationship between upwelling intensity and herbivory is likely the result of a combination of co</span><span lang=EN-US style='font-size:12.0pt;font-family:"Cambria Math","serif";mso-fareast-language:ES'>$B!>(J</span><span lang=EN-US style='font-size:12.0pt;font-family:"Arial","sans-serif";mso-fareast-language:ES'>occurring processes. First, increased primary production overwhelms consumption by herbivores. Second, cold water reduces herbivore metabolism and activity. Finally, surface currents associated with upwelling activity transport herbivore larvae offshore, decoupling secondary production from herbivory.<o:p></o:p></span></p><p class=MsoNormal><span lang=EN-US><o:p> </o:p></span></p></div></body></html>