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<div class=Section1>
<p class=MsoNormal>Journal of Applied Ecology<o:p></o:p></p>
<p class=MsoNormal style='outline: 0px;orphans: auto;text-align:start;
widows: auto;-webkit-text-stroke-width: 0px;word-spacing:0px'><span
style='color:black'>Article first published online: 6 OCT 2014<o:p></o:p></span></p>
<p class=MsoNormal style='outline: 0px;word-wrap: break-word;orphans: auto;
text-align:start;widows: auto;-webkit-text-stroke-width: 0px;word-spacing:0px'
id=doi><span style='color:black'>DOI: 10.1111/1365-2664.12337<o:p></o:p></span></p>
<p class=earlyview style='margin:0in;margin-bottom:.0001pt;line-height:15.6pt;
background:white;vertical-align:baseline'><b><span style='font-size:14.5pt;
font-family:"Arial","sans-serif";color:#007E8B'><o:p> </o:p></span></b></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:18.0pt;font-family:"Arial","sans-serif";
color:black;border:none windowtext 1.0pt;padding:0in'>Sex differences in risk
perception in deep-diving bottlenose dolphins leads to decreased foraging
efficiency when exposed to human disturbance</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:l1 level1 lfo2;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'>John Symons</span><sup><span style='font-size:7.0pt;font-family:
"Arial","sans-serif";color:black;border:none windowtext 1.0pt;padding:0in'>*</span></sup><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>,<span
class=apple-converted-space> </span>Enrico Pirotta and David Lusseau<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'>Institute of Biological and Environmental Sciences, University of
Aberdeen, Aberdeen, UK. E-mail:<a href="mailto:j.symons.12@aberdeen.ac.uk"
title="Link to email address"><span style='color:#007E8A;border:none windowtext 1.0pt;
padding:0in;text-decoration:none'>j.symons.12@aberdeen.ac.uk</span></a><o:p></o:p></span></p>
<p class=MsoNormal style='mso-line-height-alt:16.2pt;background:white;
vertical-align:baseline'><b><span style='font-size:21.5pt;font-family:"Arial","sans-serif";
color:black'><o:p> </o:p></span></b></p>
<p class=MsoNormal style='line-height:16.2pt;background:white;vertical-align:
baseline'><b><span style='font-size:14.0pt;font-family:"Arial","sans-serif";
color:black'>Summary<o:p></o:p></span></b></p>
<p class=MsoNormal style='margin-left:.45in;text-indent:-.25in;line-height:
18.0pt;mso-list:l0 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'>Individuals make behavioural decisions by weighing potential
advantages and costs (e.g. increased food intake vs. increased risk of
predation). When animals change their activities in response to a perceived
threat, their energetic input may decline. Marine ecotourism, including whale
and dolphin watching, is growing globally and cetaceans perceive interactions
with tour vessels as a form of risk. Observable behavioural changes need to be
linked to bioenergetic effects to determine the potential population consequences
of this disturbance.<o:p></o:p></span></p>
<p class=MsoNormal style='margin-left:.45in;text-indent:-.25in;line-height:
18.0pt;mso-list:l0 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'>We developed a theoretical optimal dive model for bottlenose
dolphins under three potential types of perceived risk resulting from human
interactions at the surface (decreasing instantaneous risk, increasing
instantaneous risk and no risk). We compared the predictions of these
theoretical models to observed dive cycles of foraging male and female dolphins
in the presence and absence of tour vessels. We used mixture models to classify
dive types and mixed effects models to analyse changes in the interbreath
interval of surface and bottom dives and the frequency of estimated bottom
dives.<o:p></o:p></span></p>
<p class=MsoNormal style='margin-left:.45in;text-indent:-.25in;line-height:
18.0pt;mso-list:l0 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'>3.<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'>Males significantly increased bottom time and performed fewer
bottom dives when boats were present, matching predictions of our theoretical model
for perceived decreasing instantaneous risk. In contrast, females significantly
decreased bottom times and increased the frequency of bottom dives, matching
predictions from the model for perceived increasing instantaneous risk.
Therefore, our empirical results suggest differences in the perception of risk
between sexes.<o:p></o:p></span></p>
<p class=MsoNormal style='margin-left:.45in;text-indent:-.25in;line-height:
18.0pt;mso-list:l0 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'>4.<span style='font:7.0pt "Times New Roman"'>
</span></span></span><![endif]><i><span style='font-size:9.0pt;font-family:
"Arial","sans-serif";color:black'>Synthesis and applications</span></i><span
style='font-size:9.0pt;font-family:"Arial","sans-serif";color:black'>. By
comparing theoretical predictions with observed dive data, our study suggests
that boat interactions during foraging can cause decreased net energy gain over
a foraging bout for both sexes, with females being more impacted. The
population under study is currently listed as critically endangered.
Understanding whether these predicted energetic impacts affect an individual's
vital rates will provide a link to the population-level consequences of this
disturbance. Previous analytical approaches have failed to capture the costs
associated with disturbance during foraging, leading to management
recommendations that only protect animals from increased energetic expenditure.
We suggest that the current management scheme should be revised to include
foraging areas in order to secure the energy intake of animals.<o:p></o:p></span></p>
<p class=MsoNormal><o:p> </o:p></p>
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