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<div class=Section1>
<p class=MsoNormal>Marine Ecology<o:p></o:p></p>
<p class=MsoNormal style='outline: 0px'><span style='color:black'>Article first
published online: 19 JUL 2014<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/maec.12173<o:p></o:p></span></p>
<p class=MsoNormal style='line-height:7.5pt;vertical-align:baseline;outline: 0px'
id=copyright><span style='font-size:7.5pt;font-family:"Arial","sans-serif";
color:#A3A2A2'><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:16.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>Spatio-temporal
variability of zooplankton biomass and environmental control in the Northern
Benguela Upwelling System: field investigations and model simulation</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 style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Bettina Martin</span><sup><span style='font-size:7.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>1,*</span></sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Anja Eggert</span><sup><span
style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>2,*</span></sup><span style='font-size:9.0pt;
font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Rolf Koppelmann</span><sup><span
style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>1</span></sup><span style='font-size:9.0pt;
font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Rabea Diekmann</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 style='font-size:9.0pt;
font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Volker Mohrholz</span><sup><span
style='font-size:7.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>2</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 Martin Schmidt</span><sup><span style='font-size:7.0pt;
font-family:"Arial","sans-serif";color:black;border:none windowtext 1.0pt;
padding:0in'>2</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><sup><span style='font-size:8.0pt;
font-family:"Arial","sans-serif";color:black;border:none windowtext 1.0pt;
padding:0in'>1</span></sup><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Institut für Hydrobiologie und Fischereiwissenschaft, Universität
Hamburg, Hamburg, Germany</span><sup><span style='font-size:8.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>2</span></sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>Leibniz-Institut
für Ostseeforschung Warnemünde, Rostock, Germany</span><sup><span
style='font-size:8.0pt;font-family:"Arial","sans-serif";color:black;border:
none windowtext 1.0pt;padding:0in'>3</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'><span style='font-size:9.0pt;font-family:"Arial","sans-serif";
color:black'>Thünen Institut für Seefischerei, Hamburg, Germany<sup><span
style='border:none windowtext 1.0pt;padding:0in'>*</span></sup><span
class=apple-converted-space> </span><span style='border:none windowtext 1.0pt;
padding:0in'>Correspondence</span><span class=apple-converted-space> </span><a
href="mailto:bmartin@uni-hamburg.de" title="Link to email address"><span
style='color:#007E8A;border:none windowtext 1.0pt;padding:0in'>bmartin@uni-hamburg.de</span></a>
and<span class=apple-converted-space> </span><a
href="mailto:anja.eggert@io-warnemuende.de" title="Link to email address"><span
style='color:#007E8A;border:none windowtext 1.0pt;padding:0in'>anja.eggert@io-warnemuende.de</span></a><o:p></o:p></span></p>
<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'>2.<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'><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'>Abstract<o:p></o:p></span></h3>
<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'>Spatial
and temporal distribution patterns of zooplankton are highly variable in the
Northern Benguela Upwelling System. We studied the distribution of zooplankton
(size class ≥ 0.33 mm) and used field data from four
cruises that took place between March 2008 and February 2011, as well as
simulation results of a regional ecosystem model. Remotely sensed sea surface
temperatures (SST) and surface chlorophyll concentrations were analysed to
investigate environmental influences on zooplankton biomass. The Intense
Benguela Upwelling Index showed a distinct seasonal signal throughout the years
and the highest upwelling peaks in August/September. Even though surface
chlorophyll concentrations were very variable throughout the year, the highest
concentrations were always detected in September, following the upwelling of
nutrient-rich water. In field catches, zooplankton biomass concentration in the
upper 200 m was highest above the outer shelf and shelf-break in December
2010 and February 2011, i.e. 6 months after the upwelling peaks. In
contrast, zooplankton biomass simulated by the model in the surface water was
highest in September. In March/April, biomass maxima were typically measured in
the field at intermediate water depths, but the vertical distribution was also
affected by extensive oxygen minimum zones. The ecosystem model reproduced this
vertical pattern. Although general trends were similar, simulation data of
zooplankton standing stocks overestimated the field data by a factor of 3. In
upwelling systems, food webs are generally considered to be short and dominated
by large cells. However, our field data indicate more small-sized zooplankton
organisms above the shelf than offshore.<o:p></o:p></span></p>
<p class=MsoNormal><o:p> </o:p></p>
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