Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals
Identifieur interne : 000930 ( Pmc/Checkpoint ); précédent : 000929; suivant : 000931Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals
Auteurs : Tiphaine Jeanniard-Du-Dot [Canada, France] ; Andrew W. Trites [Canada] ; John P. Y. Arnould [Australie] ; John R. Speakman [Royaume-Uni] ; Christophe Guinet [France]Source :
- Scientific Reports [ 2045-2322 ] ; 2016.
Abstract
Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 ± 0.39 J/kg per stroke and had a cost of transport of ~1.6–1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R2 = 0.76) and to a lesser extent while transiting (R2 = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R2 = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude × number of strokes) predicted total energy expenditure (R2 = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.
Url:
DOI: 10.1038/srep33912
PubMed: 27658718
PubMed Central: 5034273
Affiliations:
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Trites</name><affiliation wicri:level="1"><nlm:aff id="a1"><institution>Department of Zoology and Marine Mammal Research Unit, Institute for the Oceans and Fisheries, 2202 Main Mall, AERL, University of British Columbia</institution>, Vancouver, BC, V6T1Z4,<country>Canada</country></nlm:aff><country xml:lang="fr">Canada</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Arnould, John P Y" sort="Arnould, John P Y" uniqKey="Arnould J" first="John P. Y." last="Arnould">John P. Y. Arnould</name><affiliation wicri:level="1"><nlm:aff id="a3"><institution>Deakin University, School of Life and Environmental Sciences (Burwood Campus)</institution>, Geelong,<country>Australia</country></nlm:aff><country xml:lang="fr">Australie</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Speakman, John R" sort="Speakman, John R" uniqKey="Speakman J" first="John R." last="Speakman">John R. Speakman</name><affiliation wicri:level="1"><nlm:aff id="a4"><institution>The Institute of Biological and Environmental Sciences, Zoology Bldg</institution>, Tillydrone Avenue, Aberdeen, AB24 2TZ,<country>UK</country></nlm:aff><country xml:lang="fr">Royaume-Uni</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author><author><name sortKey="Guinet, Christophe" sort="Guinet, Christophe" uniqKey="Guinet C" first="Christophe" last="Guinet">Christophe Guinet</name><affiliation wicri:level="1"><nlm:aff id="a2"><institution>Centre d’Etudes Biologiques de Chizé</institution>, CNRS, 79360 Villiers en Bois,<country>France</country></nlm:aff><country xml:lang="fr">France</country><wicri:regionArea># see nlm:aff country strict</wicri:regionArea></affiliation></author></analytic><series><title level="j">Scientific Reports</title><idno type="eISSN">2045-2322</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 ± 0.39 J/kg per stroke and had a cost of transport of ~1.6–1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R<sup>2</sup> = 0.76) and to a lesser extent while transiting (R<sup>2</sup> = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R<sup>2</sup> = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude × number of strokes) predicted total energy expenditure (R<sup>2</sup> = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Le Boeuf, B J" uniqKey="Le Boeuf B">B. J. Le Boeuf</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Boyd, I L" uniqKey="Boyd I">I. L. Boyd</name></author><author><name sortKey="Woakes, A J" uniqKey="Woakes A">A. J. Woakes</name></author><author><name sortKey="Butler, P J" uniqKey="Butler P">P. J. Butler</name></author><author><name sortKey="Davis, R W" uniqKey="Davis R">R. W. Davis</name></author><author><name sortKey="Williams, T M" uniqKey="Williams T">T. M. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Sci Rep</journal-id><journal-id journal-id-type="iso-abbrev">Sci Rep</journal-id><journal-title-group><journal-title>Scientific Reports</journal-title></journal-title-group><issn pub-type="epub">2045-2322</issn><publisher><publisher-name>Nature Publishing Group</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">27658718</article-id><article-id pub-id-type="pmc">5034273</article-id><article-id pub-id-type="pii">srep33912</article-id><article-id pub-id-type="doi">10.1038/srep33912</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Flipper strokes can predict energy expenditure and locomotion costs in free-ranging northern and Antarctic fur seals</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Jeanniard-du-Dot</surname><given-names>Tiphaine</given-names></name><xref ref-type="corresp" rid="c1">a</xref><xref ref-type="aff" rid="a1">1</xref><xref ref-type="aff" rid="a2">2</xref></contrib><contrib contrib-type="author"><name><surname>Trites</surname><given-names>Andrew W.</given-names></name><xref ref-type="aff" rid="a1">1</xref></contrib><contrib contrib-type="author"><name><surname>Arnould</surname><given-names>John P. Y.</given-names></name><xref ref-type="aff" rid="a3">3</xref></contrib><contrib contrib-type="author"><name><surname>Speakman</surname><given-names>John R.</given-names></name><xref ref-type="aff" rid="a4">4</xref></contrib><contrib contrib-type="author"><name><surname>Guinet</surname><given-names>Christophe</given-names></name><xref ref-type="aff" rid="a2">2</xref></contrib><aff id="a1"><label>1</label><institution>Department of Zoology and Marine Mammal Research Unit, Institute for the Oceans and Fisheries, 2202 Main Mall, AERL, University of British Columbia</institution>, Vancouver, BC, V6T1Z4,<country>Canada</country></aff><aff id="a2"><label>2</label><institution>Centre d’Etudes Biologiques de Chizé</institution>, CNRS, 79360 Villiers en Bois,<country>France</country></aff><aff id="a3"><label>3</label><institution>Deakin University, School of Life and Environmental Sciences (Burwood Campus)</institution>, Geelong,<country>Australia</country></aff><aff id="a4"><label>4</label><institution>The Institute of Biological and Environmental Sciences, Zoology Bldg</institution>, Tillydrone Avenue, Aberdeen, AB24 2TZ,<country>UK</country></aff></contrib-group><author-notes><corresp id="c1"><label>a</label><email>tiphainejdd@gmail.com</email></corresp></author-notes><pub-date pub-type="epub"><day>23</day><month>09</month><year>2016</year></pub-date><pub-date pub-type="collection"><year>2016</year></pub-date><volume>6</volume><elocation-id>33912</elocation-id><history><date date-type="received"><day>02</day><month>10</month><year>2015</year></date><date date-type="accepted"><day>06</day><month>09</month><year>2016</year></date></history><permissions><copyright-statement>Copyright © 2016, The Author(s)</copyright-statement><copyright-year>2016</copyright-year><copyright-holder>The Author(s)</copyright-holder><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/4.0/"><pmc-comment>author-paid</pmc-comment>
<license-p>This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">http://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><abstract><p>Flipper strokes have been proposed as proxies to estimate the energy expended by marine vertebrates while foraging at sea, but this has never been validated on free-ranging otariids (fur seals and sea lions). Our goal was to investigate how well flipper strokes correlate with energy expenditure in 33 foraging northern and Antarctic fur seals equipped with accelerometers, GPS, and time-depth recorders. We concomitantly measured field metabolic rates with the doubly-labelled water method and derived activity-specific energy expenditures using fine-scale time-activity budgets for each seal. Flipper strokes were detected while diving or surface transiting using dynamic acceleration. Despite some inter-species differences in flipper stroke dynamics or frequencies, both species of fur seals spent 3.79 ± 0.39 J/kg per stroke and had a cost of transport of ~1.6–1.9 J/kg/m while diving. Also, flipper stroke counts were good predictors of energy spent while diving (R<sup>2</sup> = 0.76) and to a lesser extent while transiting (R<sup>2</sup> = 0.63). However, flipper stroke count was a poor predictor overall of total energy spent during a full foraging trip (R<sup>2</sup> = 0.50). Amplitude of flipper strokes (i.e., acceleration amplitude × number of strokes) predicted total energy expenditure (R<sup>2</sup> = 0.63) better than flipper stroke counts, but was not as accurate as other acceleration-based proxies, i.e. Overall Dynamic Body Acceleration.</p></abstract></article-meta></front><floats-group><fig id="f1"><label>Figure 1</label><caption><title>Relationships between number of flipper strokes during diving or transiting at the surface (2 left panels) or cumulative swimming intensity (from cumulative acceleration amplitude per stroke in m/s<sup>2</sup>, 2 right panels) and activity-specific energy expenditure (in MJ) for northern fur seals (orange triangles, N = 15) and Antarctic fur seals (green squares N = 15).</title><p>Solid lines show the results of linear models (with species included as an independent variable), and dotted lines are the 95% CI on the predicted values.</p></caption><graphic xlink:href="srep33912-f1"></graphic></fig><fig id="f2"><label>Figure 2</label><caption><p>Relationships between cumulative swimming intensity (from cumulative acceleration amplitude per stroke in m/s<sup>2</sup>, graph (<bold>A</bold>) model 5 from <xref ref-type="table" rid="t2">Table 2</xref>, R<sup>2</sup> = 0.63) or total number of flipper stroke during time at sea ((<bold>B</bold>) model 10 from <xref ref-type="table" rid="t2">Table 2</xref>, R<sup>2</sup> = 0.50) over both diving and transiting periods and total energy expenditure (MJ/kg) for the foraging trip for female northern fur seals (N = 13, orange triangles) and Antarctic fur seals (N = 13, green squares). Solid lines are results of the general linear model and dotted lines are the 95% CI on the predicted values.</p></caption><graphic xlink:href="srep33912-f2"></graphic></fig><fig id="f3"><label>Figure 3</label><caption><title>Examples of detection of fur seal flipper strokes from the dynamic acceleration on the X and Y accelerometer axes while diving, while transiting at the surface at speeds > 1 m/s, or while performing slower movements at the surface at speeds < 1 m/s.</title></caption><graphic xlink:href="srep33912-f3"></graphic></fig><fig id="f4"><label>Figure 4</label><caption><title>Number of flipper strokes detected per distance traveled vertically while diving (red triangles) and horizontally while transiting near the surface (blue circles) for 2 northern fur seals (NFS 2 and NFS10) and 2 Antarctic fur seals (AFS 4 and AFS 9) between 2 GPS locations during a foraging trip.</title><p>Transiting at the surface is defined as periods of time when animals travel at speeds > 1 m/s between 2 GPS locations.</p></caption><graphic xlink:href="srep33912-f4"></graphic></fig><table-wrap position="float" id="t1"><label>Table 1</label><caption><title>Morphometric, metabolic and foraging parameters for 13 female northern and 13 female Antarctic fur seals during a single foraging trip at sea.</title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="left" valign="top" charoff="50">Parameters</th><th colspan="2" align="center" valign="top" charoff="50">Northern fur seals</th><th colspan="2" align="center" valign="top" charoff="50">Antarctic fur seals</th></tr></thead><tbody valign="top"><tr><td align="left" valign="top" charoff="50">Mass (kg)</td><td colspan="2" align="center" valign="top" charoff="50">37.9 ± 1.3<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">31.1 ± 0.8<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Length (cm)</td><td colspan="2" align="center" valign="top" charoff="50">128.1 ± 1.6<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">114.5 ± 1.3<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Axillary girth (cm)</td><td colspan="2" align="center" valign="top" charoff="50">78.7 ± 1.2</td><td colspan="2" align="center" valign="top" charoff="50">81 ± 1.2</td></tr><tr><td align="left" valign="top" charoff="50">Fore-flipper length (cm)</td><td colspan="2" align="center" valign="top" charoff="50">39.4 ± 0.9<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">34.3 ± 0.3<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Fore-flipper width (cm)</td><td colspan="2" align="center" valign="top" charoff="50">12.5 ± 0.2<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">11.1 ± 0.1<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Hind-flipper length (cm)</td><td colspan="2" align="center" valign="top" charoff="50">34.6 ± 0.6<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">24.4 ± 0.4<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Hind-flipper width (cm)</td><td colspan="2" align="center" valign="top" charoff="50">14.4 ± 0.2<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">10.8 ± 0.2<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Trip duration (d)</td><td colspan="2" align="center" valign="top" charoff="50">7.96 ± 2.17</td><td colspan="2" align="center" valign="top" charoff="50">7.65 ± 3.88</td></tr><tr><td align="left" valign="top" charoff="50">Distance traveled (km)</td><td colspan="2" align="center" valign="top" charoff="50">750 ± 50</td><td colspan="2" align="center" valign="top" charoff="50">635 ± 77</td></tr><tr><td align="left" valign="top" charoff="50">Number of dives</td><td colspan="2" align="center" valign="top" charoff="50">2551 ± 323<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">3949 ± 597<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Dive duration (s)</td><td colspan="2" align="center" valign="top" charoff="50">62.8 ± 7.3<xref ref-type="fn" rid="t1-fn1">*</xref></td><td colspan="2" align="center" valign="top" charoff="50">42.6 ± 4.5<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Time spend diving (%)</td><td colspan="2" align="center" valign="top" charoff="50">28.6 ± 2.0</td><td colspan="2" align="center" valign="top" charoff="50">29.0 ± 0.7</td></tr><tr><td align="left" valign="top" charoff="50">Time spent transiting (%)</td><td colspan="2" align="center" valign="top" charoff="50">30.5 ± 1.8</td><td colspan="2" align="center" valign="top" charoff="50">26.4 ± 1.6</td></tr><tr><td align="left" valign="top" charoff="50">Time spend diving (h)</td><td colspan="2" align="center" valign="top" charoff="50">53.9 ± 4.7</td><td colspan="2" align="center" valign="top" charoff="50">47.7 ± 6.0</td></tr><tr><td align="left" valign="top" charoff="50">Time spent transiting (h)</td><td colspan="2" align="center" valign="top" charoff="50">59.9 ± 5.9</td><td colspan="2" align="center" valign="top" charoff="50">45.9 ± 7.7</td></tr><tr><td align="left" valign="top" charoff="50">At-sea metabolic rate (MJ/d/kg)</td><td colspan="2" align="center" valign="top" charoff="50">0.56 ± 0.04</td><td colspan="2" align="center" valign="top" charoff="50">0.59 ± 0.04</td></tr><tr><td align="left" valign="top" charoff="50">Total energy expenditure (MJ/kg)</td><td colspan="2" align="center" valign="top" charoff="50">4.2 ± 0.4</td><td colspan="2" align="center" valign="top" charoff="50">4.1 ± 0.6</td></tr><tr><td rowspan="2" align="left" valign="top" charoff="50"> </td><td colspan="2" align="center" valign="top" charoff="50"><bold>Northern fur seals</bold></td><td colspan="2" align="center" valign="top" charoff="50"><bold>Antarctic fur seals</bold></td></tr><tr><td align="center" valign="top" charoff="50"><bold>Diving</bold></td><td align="center" valign="top" charoff="50"><bold>Transit</bold></td><td align="center" valign="top" charoff="50"><bold>Diving</bold></td><td align="center" valign="top" charoff="50"><bold>Transit</bold></td></tr><tr><td align="left" valign="top" charoff="50">Energy expenditure (MJ/kg)</td><td align="center" valign="top" charoff="50">1.89 ± 0.81</td><td align="center" valign="top" charoff="50">1.24 ± 0.51</td><td align="center" valign="top" charoff="50">2.00 ± 1.03</td><td align="center" valign="top" charoff="50">1.15 ± 0.75</td></tr><tr><td align="left" valign="top" charoff="50">Flipper stroke Frequency (Hz)</td><td align="center" valign="top" charoff="50">0.37 ± 0.03<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">0.35 ± 0.02<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">0.44 ± 0.06<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">0.53 ± 0.03<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr><tr><td align="left" valign="top" charoff="50">Swim. intensity/stroke (m/s<sup>2</sup>)</td><td align="center" valign="top" charoff="50">0.46 ± 0.01*</td><td align="center" valign="top" charoff="50">0.43 ± 0.01*</td><td align="center" valign="top" charoff="50">0.75 ± 0.02*</td><td align="center" valign="top" charoff="50">0.48 ± 0.01*</td></tr><tr><td align="left" valign="top" charoff="50">Swim. intensity/min (m/s<sup>2</sup>/min)</td><td align="center" valign="top" charoff="50">10.26 ± 1.0<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">8.70 ± 0.52<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">22.60 ± 1.51<xref ref-type="fn" rid="t1-fn1">*</xref></td><td align="center" valign="top" charoff="50">15.01 ± 1.13<xref ref-type="fn" rid="t1-fn1">*</xref></td></tr></tbody></table><table-wrap-foot><fn id="t1-fn1"><p>Values are mean ± SE and <sup>*</sup> show between species significant differences (<italic>p</italic> < 0.05). Swimming intensity refers to acceleration amplitude per stroke (m/s<sup>2</sup>) or per min of stroking (m/s<sup>2</sup>/min).</p></fn></table-wrap-foot></table-wrap><table-wrap position="float" id="t2"><label>Table 2</label><caption><title>Best models explaining at-sea energy expenditure in MJ/kg of northern (N = 13) and/or Antarctic fur seals (N = 13) as a function of flipper stroke count or of swimming intensity, i.e., cumulative acceleration amplitude of strokes during diving and transiting combined (Stroke.Count<sub>Total</sub> and Swim.Intens.<sub>Total</sub>), or while diving or transiting at the surface (Stroke.Count<sub>Dive,</sub> Stroke.Count<sub>Transit,</sub> Swim.Intens.<sub>Dive</sub> and Swim.Intens.<sub>Transit</sub>).</title></caption><table frame="hsides" rules="groups" border="1"><colgroup><col align="left"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col><col align="center"></col></colgroup><thead valign="bottom"><tr><th align="left" valign="top" charoff="50">Model</th><th align="center" valign="top" charoff="50">Parameters</th><th align="center" valign="top" charoff="50">Estimates</th><th align="center" valign="top" charoff="50">SE</th><th align="center" valign="top" charoff="50"><italic>p</italic></th><th align="center" valign="top" charoff="50">R<sup>2</sup></th><th align="center" valign="top" charoff="50">ΔAICc</th><th align="center" valign="top" charoff="50">AICc weight</th></tr></thead><tbody valign="top"><tr><td rowspan="3" align="left" valign="middle" charoff="50">5</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">0.16</td><td align="center" valign="top" charoff="50">0.69</td><td align="center" valign="top" charoff="50">0.820</td><td align="center" valign="top" charoff="50">0.63</td><td align="center" valign="top" charoff="50">0.00</td><td align="center" valign="top" charoff="50">0.185</td></tr><tr><td align="center" valign="top" charoff="50">Swim.Intens.<sub>Total</sub></td><td align="center" valign="top" charoff="50">2.5 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">4.1 10<sup>−6</sup></td><td align="center" valign="top" charoff="50"> < 10<sup>−5</sup></td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Species (NFS)</td><td align="center" valign="top" charoff="50">1.33</td><td align="center" valign="top" charoff="50">0.54</td><td align="center" valign="top" charoff="50">0.022</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td rowspan="4" align="left" valign="middle" charoff="50">6</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">0.15</td><td align="center" valign="top" charoff="50">0.67</td><td align="center" valign="top" charoff="50">0.824</td><td align="center" valign="top" charoff="50">0.67</td><td align="center" valign="top" charoff="50">0.48</td><td align="center" valign="top" charoff="50">0.146</td></tr><tr><td align="center" valign="top" charoff="50">Swim.Intens.<sub>Total</sub></td><td align="center" valign="top" charoff="50">5.0 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">1.7 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">0.007</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Stroke.Count<sub>Total</sub></td><td align="center" valign="top" charoff="50">−1.4 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">8.9 10<sup>−6</sup></td><td align="center" valign="top" charoff="50">0.138</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Species (NFS)</td><td align="center" valign="top" charoff="50">1.97</td><td align="center" valign="top" charoff="50">0.67</td><td align="center" valign="top" charoff="50">0.008</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td rowspan="4" align="left" valign="middle" charoff="50">7</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">−0.20</td><td align="center" valign="top" charoff="50">0.71</td><td align="center" valign="top" charoff="50">0.775</td><td align="center" valign="top" charoff="50">0.66</td><td align="center" valign="top" charoff="50">0.82</td><td align="center" valign="top" charoff="50">0.123</td></tr><tr><td align="center" valign="top" charoff="50">Stroke.Count<sub>Dive</sub></td><td align="center" valign="top" charoff="50">4.3 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">1.5 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">0.008</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Swim.Intens.<sub>Transit</sub></td><td align="center" valign="top" charoff="50">1.9 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">5.7 10<sup>−6</sup></td><td align="center" valign="top" charoff="50">0.003</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Species (NFS)</td><td align="center" valign="top" charoff="50">0.96</td><td align="center" valign="top" charoff="50">0.51</td><td align="center" valign="top" charoff="50">0.075</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td rowspan="3" align="left" valign="middle" charoff="50">8</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">0.46</td><td align="center" valign="top" charoff="50">0.65</td><td align="center" valign="top" charoff="50">0.487</td><td align="center" valign="top" charoff="50">0.61</td><td align="center" valign="top" charoff="50">1.52</td><td align="center" valign="top" charoff="50">0.087</td></tr><tr><td align="center" valign="top" charoff="50">Stroke.Count<sub>Dive</sub></td><td align="center" valign="top" charoff="50">4.2 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">1.5 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">0.011</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Swim.Intens.<sub>Transit</sub></td><td align="center" valign="top" charoff="50">1.7 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">5.9 10<sup>−6</sup></td><td align="center" valign="top" charoff="50">0.008</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td rowspan="4" align="left" valign="middle" charoff="50">9</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">−0.02</td><td align="center" valign="top" charoff="50">0.71</td><td align="center" valign="top" charoff="50">0.920</td><td align="center" valign="top" charoff="50">0.65</td><td align="center" valign="top" charoff="50">1.78</td><td align="center" valign="top" charoff="50">0.076</td></tr><tr><td align="center" valign="top" charoff="50">Stroke.Count<sub>Transit</sub></td><td align="center" valign="top" charoff="50">7.7 10<sup>−6</sup></td><td align="center" valign="top" charoff="50">2.7 10<sup>−6</sup></td><td align="center" valign="top" charoff="50">0.010</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Swim.Intens.<sub>Dive</sub></td><td align="center" valign="top" charoff="50">6.0 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">1.7 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">0.002</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td align="center" valign="top" charoff="50">Species (NFS)</td><td align="center" valign="top" charoff="50">1.53</td><td align="center" valign="top" charoff="50">0.58</td><td align="center" valign="top" charoff="50">0.016</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr><tr><td rowspan="2" align="left" valign="middle" charoff="50">10</td><td align="center" valign="top" charoff="50">Intercept</td><td align="center" valign="top" charoff="50">0.95</td><td align="center" valign="top" charoff="50">0.67</td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50">0.50</td><td align="center" valign="top" charoff="50">4.63</td><td align="center" valign="top" charoff="50">0.045</td></tr><tr><td align="center" valign="top" charoff="50">Stroke.Count<sub>Total</sub></td><td align="center" valign="top" charoff="50">1.2 10<sup>−5</sup></td><td align="center" valign="top" charoff="50">2.510<sup>−6</sup></td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td><td align="center" valign="top" charoff="50"> </td></tr></tbody></table><table-wrap-foot><fn id="t2-fn1"><p>Only models with ΔAICc < 2 and the model with only Stroke.Count<sub>Total</sub> as an explanatory variable (simplest model for comparison) are presented.</p></fn></table-wrap-foot></table-wrap></floats-group></pmc><affiliations><list><country><li>Australie</li><li>Canada</li><li>France</li><li>Royaume-Uni</li></country></list><tree><country name="Canada"><noRegion><name sortKey="Jeanniard Du Dot, Tiphaine" sort="Jeanniard Du Dot, Tiphaine" uniqKey="Jeanniard Du Dot T" first="Tiphaine" last="Jeanniard-Du-Dot">Tiphaine Jeanniard-Du-Dot</name></noRegion><name sortKey="Trites, Andrew W" sort="Trites, Andrew W" uniqKey="Trites A" first="Andrew W." last="Trites">Andrew W. 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