<?xml version="1.0" encoding="UTF-8" standalone="no"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v2.3 20070202//EN" "journalpublishing.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" article-type="research-article" dtd-version="2.3" xml:lang="EN">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2023.1208262</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Understanding the oceanographic dynamics of the Isla Cha&#xf1;aral baleen whale feeding ground, (Humboldt Archipelago, Northern Chile) to extend habitat protection</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Buchan</surname>
<given-names>Susannah J.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1737532"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ramos</surname>
<given-names>Marcel</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/518335"/>
</contrib>
<contrib contrib-type="author" equal-contrib="yes" corresp="yes">
<name>
<surname>Oyanadel</surname>
<given-names>Jorge</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2285303"/>
</contrib>
<contrib contrib-type="author" equal-contrib="yes">
<name>
<surname>Santos-Carvallo</surname>
<given-names>Macarena</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/894484"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Bedri&#xf1;ana-Romano</surname>
<given-names>Luis</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="aff" rid="aff8">
<sup>8</sup>
</xref>
<xref ref-type="aff" rid="aff9">
<sup>9</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2199007"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Valladares</surname>
<given-names>Mar&#xed;a</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1911152"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Maldonado</surname>
<given-names>Marinella</given-names>
</name>
<xref ref-type="aff" rid="aff10">
<sup>10</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Astudillo</surname>
<given-names>Orlando</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1657587"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Sep&#xfa;lveda</surname>
<given-names>Maritza</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/119144"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Pearce</surname>
<given-names>Steve</given-names>
</name>
<xref ref-type="aff" rid="aff11">
<sup>11</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2529002"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Olavarr&#xed;a</surname>
<given-names>Carlos</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1281643"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Centro de Estudios Avanzados en Zonas &#xc1;ridas (CEAZA)</institution>, <addr-line>La Serena</addr-line>, <country>Chile</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Center for Oceanographic Research COPAS COASTAL, Universidad de Concepci&#xf3;n</institution>, <addr-line>Concepci&#xf3;n</addr-line>, <country>Chile</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Departamento de Oceanograf&#xed;a, Facultad de Ciencias Naturales y Oceanogr&#xe1;ficas, Universidad de Concepci&#xf3;n</institution>, <addr-line>Concepci&#xf3;n</addr-line>, <country>Chile</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Departamento de Biolog&#xed;a Marina, Facultad de Ciencias del Mar, Universidad Cat&#xf3;lica del Norte</institution>, <addr-line>Coquimbo</addr-line>, <country>Chile</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Centro de Ecolog&#xed;a y Manejo Sustentable de Islas Oce&#xe1;nicas (ESMOI), Departamento de Biolog&#xed;a Marina, Facultad de Ciencias del Mar, Universidad Cat&#xf3;lica del Norte</institution>, <addr-line>Coquimbo</addr-line>, <country>Chile</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Centro de Investigaci&#xf3;n y Gesti&#xf3;n de Recursos Naturales (CIGREN), Instituto de Biolog&#xed;a, Facultad de Ciencias, Universidad de Valpara&#xed;so</institution>, <addr-line>Valpara&#xed;so</addr-line>, <country>Chile</country>
</aff>
<aff id="aff7">
<sup>7</sup>
<institution>Centro de Investigaci&#xf3;n Eutropia</institution>, <addr-line>Santiago</addr-line>, <country>Chile</country>
</aff>
<aff id="aff8">
<sup>8</sup>
<institution>Instituto de Ciencias Marinas y Limnol&#xf3;gicas, Facultad de Ciencias, Universidad Austral de Chile</institution>, <addr-line>Valdivia</addr-line>, <country>Chile</country>
</aff>
<aff id="aff9">
<sup>9</sup>
<institution>Non Governmental Organization (NGO) Centro Ballena Azul</institution>, <addr-line>Valdivia</addr-line>, <country>Chile</country>
</aff>
<aff id="aff10">
<sup>10</sup>
<institution>Corporacion Nacional Forestal (CONAF) Atacama, Oficina caleta Cha&#xf1;aral de Aceituno</institution>, <addr-line>Copiap&#xf3;</addr-line>, <country>Chile</country>
</aff>
<aff id="aff11">
<sup>11</sup>
<institution>ASL Environmental Sciences, BC</institution>, <addr-line>Victoria, BC</addr-line>, <country>Canada</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Mark Meekan, University of Western Australia, Australia</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Salvatore Siciliano, Funda&#xe7;&#xe3;o Oswaldo Cruz (Fiocruz), Brazil</p>
<p>Luciana C. Ferreira, Australian Institute of Marine Science (AIMS), Australia</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Jorge Oyanadel, <email xlink:href="mailto:jorge.oyanadel@ce.ucn.cl">jorge.oyanadel@ce.ucn.cl</email>
</p>
</fn>
<fn fn-type="equal" id="fn003">
<p>&#x2020;These authors have contributed equally to this work</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>04</day>
<month>01</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>10</volume>
<elocation-id>1208262</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>04</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>05</day>
<month>12</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Buchan, Ramos, Oyanadel, Santos-Carvallo, Bedri&#xf1;ana-Romano, Valladares, Maldonado, Astudillo, Sep&#xfa;lveda, Pearce and Olavarr&#xed;a</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Buchan, Ramos, Oyanadel, Santos-Carvallo, Bedri&#xf1;ana-Romano, Valladares, Maldonado, Astudillo, Sep&#xfa;lveda, Pearce and Olavarr&#xed;a</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Off Northern Chile, baleen whales use the area around Isla Cha&#xf1;aral as a feeding ground where they forage on euphausiids. Isla Cha&#xf1;aral is part of the highly productive Humboldt Archipelago (&#x223c; 28&#xb0;S-29&#xb0;S) within the Humboldt Current System (HCS). In this study, we seek to understand the sub-mesoscale spatial distribution of fin and blue whales and their prey around Isla Cha&#xf1;aral using systematic and opportunistic visual sighting data of whales and systematic acoustic backscatter data of zooplankton from an Acoustic Zooplankton and Fish Profiler (AZFP); and to examine the oceanographic dynamics of the wider Humboldt Archipelago area with remote-sensing oceanographic data. We completed a total of 512.6&#xa0;km of survey effort over 20 days in 2018 and 318.3&#xa0;km over 16 days in 2019 collecting systematic whale sighting and backscatter data. A total of 42 fin whales, 0 blue whales and 66 unidentified whales were sighted in 2018, and 7 fin whales, 3 blue whales and 12 unidentified whales were sighted in 2019. Observed spatial distribution of backscatter and whales was strongly associated with a bathymetric feature, i.e., the submarine canyon that curves around Isla Cha&#xf1;aral. Generalized Additive Models showed that fin whale presence was associated with high levels of backscatter and shallow depths similar to those of the canyon. We found that long-term average geostrophic currents form a recirculation system between 28&#xb0;S and 31&#xb0;S that can transport nutrient-rich upwelled surface waters back towards the Humboldt Archipelago and contribute to high biological productivity in this area. However, in summer 2019 geostrophic flow occurred away from the coast and a warm low-productivity spring explained low backscatter and whale sightings around Isla Cha&#xf1;aral. The unique oceanographic features of Isla Cha&#xf1;aral and the Humboldt Archipelago that contribute to high concentrations of Endangered baleen whales and their prey justify the extension of the Isla Cha&#xf1;aral Marine Reserve to include the canyon between the mainland and the island, and the implementation of a Multiple Use Marine Protected Area for the entire Humboldt Archipelago area that explicitly protects this critical feature.</p>
</abstract>
<kwd-group>
<kwd>baleen whale feeding ground</kwd>
<kwd>zooplankton acoustic backscatter</kwd>
<kwd>oceanographic dynamics</kwd>
<kwd>Humboldt archipelago</kwd>
<kwd>Isla Cha&#xf1;aral</kwd>
<kwd>marine protected area</kwd>
</kwd-group>
<counts>
<fig-count count="10"/>
<table-count count="1"/>
<equation-count count="7"/>
<ref-count count="63"/>
<page-count count="16"/>
<word-count count="8065"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Marine Megafauna</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>Fin whales (<italic>Balaenoptera physalus</italic>) and blue whales (<italic>Balaenoptera musculus</italic>) are the largest animals to have ever existed on earth. Off Northern Chile, fin whales use the Humboldt Archipelago and particularly the area around Isla Cha&#xf1;aral as a feeding ground where they forage on euphausiids (<xref ref-type="bibr" rid="B43">P&#xe9;rez et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B59">Toro et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B54">Sep&#xfa;lveda et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B15">Buchan et&#xa0;al., 2021</xref>). Blue whales, although not sighted as frequently as fin whales, are also known to forage on the same prey type in this area (<xref ref-type="bibr" rid="B15">Buchan et&#xa0;al., 2021</xref>).</p>
<p>The Humboldt Archipelago (&#x223c; 28&#xb0;S-29&#xb0;S) lies within the Humboldt Current System (HCS), a highly productive Eastern Boundary Upwelling System (EBUS) (<xref ref-type="bibr" rid="B18">Daneri et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B58">Thiel et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B39">Montecino and Lange, 2009</xref>) off central-northern Chile. The large-scale oceanographic processes, which drive biological productivity associated with wind-driven coastal upwelling, are relatively well-understood in the HCS (e.g., <xref ref-type="bibr" rid="B18">Daneri et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B39">Montecino and Lange, 2009</xref>). The HCS is mainly characterized by a cold surface current that flows towards the equator, and an opposite subsurface current (Peru-Chile Undercurrent: PCUC) that flows south close to the continental slope and deep shelf. The PCUC is the source of the upwelled water that is poor in dissolved oxygen, has a low pH, and is rich in nutrients (e.g., nitrates, phosphates) that fertilize the coastal photic layer favoring primary productivity and maintaining a productive food web (<xref ref-type="bibr" rid="B58">Thiel et&#xa0;al., 2007</xref>). In addition, wind forcing in this region is dominated by the southeast Pacific subtropical anticyclone driving winds with an equatorward component near the coast, which favor upwelling and vary latitudinally and seasonally (<xref ref-type="bibr" rid="B55">Shaffer et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B40">Monteiro et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B33">K&#xe4;mpf and Chapman, 2016</xref>). Permanent intrusions of cold water are present throughout almost the entire year, from coastal upwelling events driven by Ekman transport, with greater seasonal variability in spring. These frequent upwelling events recorded along the coast, made possible by a narrow continental shelf, a strong orographic slope and quasi-weekly wind pulses associated with atmospheric disturbances trapped in the mountain range (i.e. coastal lows) (<xref ref-type="bibr" rid="B49">Rutllant and Montecinos, 2002</xref>). In this sense, coastal topography and the flow over the exposed platform play an important role in upwelling zones since they allow certain bays and promontories to function as larval retention areas and allow primary producers and higher trophic levels to congregate (<xref ref-type="bibr" rid="B35">Largier, 2020</xref>).</p>
<p>The Humboldt Archipelago is a remote stretch of complex coastline with and despite great touristic and scientific interest in this area, its hydrography and coastal dynamics have been poorly studied. The sub-mesoscale oceanographic features that explain the particularly high productivity and the presence of foraging baleen whales in the Humboldt Archipelago has not been examined. It is known that mesoscale (10-100&#xa0;km) and sub-mesoscale (1-10&#xa0;km) oceanographic processes and environmental features, such as bathymetry (<xref ref-type="bibr" rid="B17">Croll et&#xa0;al., 2005</xref>), thermal fronts (<xref ref-type="bibr" rid="B21">Doniol-Valcroze et&#xa0;al., 2007</xref>), water masses (<xref ref-type="bibr" rid="B14">Buchan and Qui&#xf1;ones, 2016</xref>; <xref ref-type="bibr" rid="B13">Buchan et&#xa0;al., 2022</xref>) can drive local whale distribution. Adequate| understanding of the oceanography that drives the distribution of whales and their prey can provide important justification for marine habitat protection. For example, <xref ref-type="bibr" rid="B16">Cox et&#xa0;al. (2018)</xref> identify common key features of marine mammal habitats in coastal areas, which include interactions between bathymetry and tidal currents, and patterns of seasonally stratification and shelf edge upwelling.</p>
<p>Protection of this habitat is necessary given that the Humboldt Archipelago area has been threatened by a series of industrial projects, including the now discarded Barranquilla coal plant project, the proposed Dominga copper and iron mine and mega-port, and the already approved Cruz Grande mega-port. These mega-ports propose to open shipping lanes within this critical feeding habitat for Endangered baleen whales at a time when record numbers of ship strikes are increasing off northern Chile (Toro et&#xa0;al. <italic>In prep</italic>). Currently, only 1 nmi around Isla Cha&#xf1;aral (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>) and two other islands (Isla Damas and Isla Choros) are effectively protected as Marine Reserves for fisheries, although a much larger Multiple Use Marine Protected Area (MU MPA) has recently been approved that would include the entire Humboldt Archipelago. However, it is unclear whether the Humboldt Archipelago MU MPA will explicitly exclude large industrial projects and effectively protect this whale habitat from higher levels of marine traffic.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Map of study area <bold>(A)</bold> with bathymetry around Isla Cha&#xf1;aral (IC) and the Isla Cha&#xf1;aral Marine Reserve (black Line). Systematic survey effort is shown for 2018 <bold>(B)</bold> and 2019 <bold>(C)</bold>. Note: the two other main islands of the Humboldt Archipelago can be seen in panel <bold>(B)</bold>: Isla Damas (D) and Isla Choros (CH).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g001.tif"/>
</fig>
<p>In this study, we seek to understand the submesoscale spatial distribution of fin and blue whales and their prey around Isla Cha&#xf1;aral using systematic and opportunistic whale visual sighting data and zooplankton acoustic backscatter survey data; and to examine the oceanographic dynamics of the wider Humboldt Archipelago area with remote sensing oceanographic data and Acoustic Doppler Current Profiler (ADCP) data.</p>
</sec>
<sec id="s2">
<title>Methods</title>
<sec id="s2_1">
<title>Systematic survey data collection of zooplankton backscatter data and whale sightings</title>
<p>Systematic surveys were carried out between the 23<sup>rd</sup> January and the 19<sup>th</sup> of February 2018, and between the 2<sup>nd</sup> and the 28<sup>th</sup> of February 2019. Surveys were conducted on a 10&#xa0;m vessel moving at a maximum speed of 4 kn (7.41 km/h) in the area surrounding Isla Cha&#xf1;aral (29&#xb0;02&#x2019;3.61&#x201d;S; 71&#xb0;34&#x2019;37.03&#x201d;W) along track lines separated by approximately 1 nmi (<xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1B, C</bold>
</xref>). During these surveys, a side-mounted Acoustic Zooplankton and Fish Profiler (AZFP) was deployed off the side of the vessel, at approximately 1&#xa0;m below the sea surface and facing down, to collect acoustic backscatter data at 38, 125, 200 and 455 kHz center frequencies.</p>
<p>During the AZFP survey, a team of 4 observers conducted visual surveys of whales from the bow of the vessel (elevation 1.6&#xa0;m), where three observers simultaneously performed continuous scans of the horizon with the naked eye covering a 180&#xb0; angle, and a fourth observer acted as data scorer. Each observer performed visual scans for no more than 1.5 hours and then took a 30-min break; observers worked for approximately 6 hours per day. For all marine mammal sightings, the following data were recorded: species (in this case, fin whale, blue whale, humpback whale (<italic>Megaptera novaeangliae</italic>), or unidentified baleen whale), date/time, GPS position, estimated distance from the vessel and estimate angle relative to the bow. Estimates of distance from the vessel were calibrated by taking estimates from observations of objects of known distance determined by a range finder, estimated distances were then corrected according to this calibration. Corrected distances, angle and GPS position were used to calculate whale positions. Observations were made when weather conditions provided good visibility to ensure reliable data collection, i.e., Beaufort Sea state of 3 or less, with no coastal fog or rain.</p>
</sec>
<sec id="s2_2">
<title>Acoustic zooplankton and fish profiler data processing and analysis</title>
<p>The AZFP acoustic backscatter data were converted by AZFPLink software (ASL Environmental Sciences Inc.) into comma separated value format files and then processed as a matrix using custom routines of Matlab software, which allowed estimating the related variable with volume backscattering strength (<italic>Sv</italic>, dB re 1 m<sup>-1</sup>), and all others mentioned below. The <italic>Sv</italic> described as <italic>Sv<sub>meas</sub>
</italic> by (<xref ref-type="bibr" rid="B20">de Robertis and Higginbottom, 2007</xref>) (<xref ref-type="disp-formula" rid="eq1">Equation 1</xref>), can be expressed as the arithmetic sum of the contributions in recorded signal by scatterers and noise namely <italic>Sv<sub>signal</sub>
</italic> and <italic>Sv<sub>noise</sub>
</italic>.</p>
<disp-formula id="eq1">
<label>(1)</label>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>m</mml:mi>
<mml:mi>e</mml:mi>
<mml:mi>a</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mn>10</mml:mn>
<mml:mi>l</mml:mi>
<mml:mi>o</mml:mi>
<mml:msub>
<mml:mi>&#x261;</mml:mi>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2217;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:msup>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
<mml:mrow>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>g</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>a</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">/</mml:mo>
<mml:mn>10</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:msup>
<mml:mo>+</mml:mo>
<mml:msup>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
<mml:mrow>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>n</mml:mi>
<mml:mi>o</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>s</mml:mi>
<mml:mi>e</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo stretchy="false">/</mml:mo>
<mml:mn>10</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
</mml:msup>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>&#xa0;</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>So, to correctly associate the received echo to its source, the acoustic signal was first refined by eliminating background noise, estimated according to the following equation (<xref ref-type="bibr" rid="B20">de Robertis and Higginbottom, 2007</xref>; <xref ref-type="bibr" rid="B62">Ventero et al., 2019</xref>) (<xref ref-type="disp-formula" rid="eq2">Equation 2</xref>):</p>
<disp-formula id="eq2">
<label>(2)</label>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>n</mml:mi>
<mml:mi>o</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>s</mml:mi>
<mml:mi>e</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mn>20</mml:mn>
<mml:mi>l</mml:mi>
<mml:mi>o</mml:mi>
<mml:msub>
<mml:mi>&#x261;</mml:mi>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2217;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mn>2</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>&#x3b1;</mml:mi>
<mml:mo>&#x2217;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mtext>offset</mml:mtext>
</mml:mrow>
</mml:math>
</disp-formula>
<p>Where <italic>&#x3b1;</italic> is the absorption coefficient, <italic>R</italic> is the frequency range respective (m) and offset is the logarithmic form of <italic>sv</italic> (<xref ref-type="disp-formula" rid="eq3">Equation 3</xref>):</p>
<disp-formula id="eq3">
<label>(3)</label>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:mtext>offset</mml:mtext>
<mml:mo>=</mml:mo>
<mml:mn>10</mml:mn>
<mml:mi>l</mml:mi>
<mml:mi>o</mml:mi>
<mml:msub>
<mml:mi>&#x261;</mml:mi>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2217;</mml:mo>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>v</mml:mi>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>&#xa0;</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>The offset was determined by using the backscattering strength recorded in the space immediately below the seabed where the records are considered passive because there is no source of reflection, therefore allowing the estimation of noise values (<xref ref-type="disp-formula" rid="eq3">Equation 3</xref>). For each file, an offset value was estimated using an average of at least five <italic>Sv<sub>meas</sub>
</italic> values &#x200b;&#x200b;selected in that area which was converted to linear form (<italic>sv</italic>), as well as the respective hourly values &#x200b;&#x200b;of the absorption coefficient.</p>
<p>Continuing with the array processing, acoustic records below the seafloor were eliminated from the dataset, resulting in files with only valid records from the water column. Adding to this, with the purpose of obtaining more robust integration of <italic>Sv<sub>signal</sub>
</italic> values, remaining effects of noise still visible in the hydroacoustic signal were eliminated. An SNR indicator was determined which reflects the contribution of noise and scatterers in the captured hydroacoustic signal, according to the methodology applied by (<xref ref-type="bibr" rid="B20">de Robertis and Higginbottom, 2007</xref>) (<xref ref-type="disp-formula" rid="eq4">Equation 4</xref>):</p>
<disp-formula id="eq4">
<label>(4)</label>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:mtext>SNR</mml:mtext>
<mml:mo>=</mml:mo>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>g</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>a</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2013;</mml:mo>
<mml:mi>S</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>n</mml:mi>
<mml:mi>o</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>s</mml:mi>
<mml:mi>e</mml:mi>
</mml:mrow>
</mml:msub>
</mml:mrow>
</mml:math>
</disp-formula>
<p>A value of SNR = 3 dB was determined after a sensitivity analysis, in which it was established that this value was sufficient threshold to eliminate the remaining noise and get a greater percentage of useful data at same time. Cross-sections of all transects were reviewed to examine backscatter. Thus, from each transect, only <italic>Sv<sub>signal</sub>
</italic> records above this SNR threshold value were considered for subsequent analyses.</p>
<p>Thereafter <italic>Sv<sub>signal</sub>
</italic> was analyzed and first processed using a mask (data&lt;-55 dB) to remove possible micronekton&#x2019;s backscatter and thus to obtain acoustic density for the 125, 200 and 455 kHz channels, by means of area backscattering coefficient (<xref ref-type="disp-formula" rid="eq5">Equation 5</xref>) assessment which is defined by (<xref ref-type="bibr" rid="B36">MacLennan et&#xa0;al., 2002</xref>) as the integral of <italic>sv</italic> (<italic>sv<sub>signal</sub>(z, f)</italic> in this study) over a range interval <italic>R</italic>1 to <italic>R</italic>2 (i.e., <italic>z</italic>1 to <italic>z</italic>2) for each frequency (<italic>f</italic>), that is, the integrated backscatter over the entire water column of the linear form of <italic>Sv<sub>signal</sub>
</italic>:</p>
<disp-formula id="eq5">
<label>(5)</label>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>a</mml:mi>
<mml:mo>=</mml:mo>
<mml:munderover>
<mml:mo>&#x222b;</mml:mo>
<mml:mrow>
<mml:mi>z</mml:mi>
<mml:mn>2</mml:mn>
</mml:mrow>
<mml:mrow>
<mml:mi>z</mml:mi>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:munderover>
<mml:mi>s</mml:mi>
<mml:msub>
<mml:mi>v</mml:mi>
<mml:mrow>
<mml:mi>s</mml:mi>
<mml:mi>i</mml:mi>
<mml:mi>g</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>a</mml:mi>
<mml:mi>l</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mi>d</mml:mi>
<mml:mi>z</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>sa</italic> is expressed in linear form (m<sup>2</sup> m<sup>-2</sup>) and then was converted to nautical miles called nautical area scattering coefficient (<italic>NASC</italic>) (m<sup>2</sup> nmi<sup>-2</sup>) (<xref ref-type="disp-formula" rid="eq6">Equation 6</xref>). In this study, the range interval was from five meters above the seafloor up to five meters below the sea surface, which avoids the high reverberation in those layers.</p>
<disp-formula id="eq6">
<label>(6)</label>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>A</mml:mi>
<mml:mi>S</mml:mi>
<mml:mi>C</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>4</mml:mn>
<mml:mtext>&#xa0;&#x3c0;&#xa0;</mml:mtext>
<mml:msup>
<mml:mrow>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mrow>
<mml:mn>1.852</mml:mn>
</mml:mrow>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>s</mml:mi>
<mml:mi>a</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>Then, <italic>NASC</italic> values were transformed to a logarithmic term defined as Nautical area scattering strength (<italic>SA</italic>), expressed in dB re 1(m<sup>2</sup> nmi<sup>-2</sup>) (<xref ref-type="disp-formula" rid="eq7">Equation 7</xref>):</p>
<disp-formula id="eq7">
<label>(7)</label>
<mml:math display="block" id="M7">
<mml:mrow>
<mml:mi>S</mml:mi>
<mml:mi>A</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>10</mml:mn>
<mml:mtext>&#xa0;</mml:mtext>
<mml:mi>l</mml:mi>
<mml:mi>o</mml:mi>
<mml:msub>
<mml:mi>&#x261;</mml:mi>
<mml:mrow>
<mml:mn>10</mml:mn>
</mml:mrow>
</mml:msub>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>N</mml:mi>
<mml:mi>A</mml:mi>
<mml:mi>S</mml:mi>
<mml:mi>C</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>In addition, simultaneous to the AZFP deployments on February 17<sup>th</sup> and 25<sup>th</sup> 2019, to confirm the presence (or absence) of acoustic signal dispersers such as zooplankton groups, diurnal plankton tows were conducted using Bongo nets (0.7&#xa0;m diameter) equipped with 200 &#xb5;m mesh and cod-end buckets, towed from the bottom (110&#xa0;m) to the surface along a total of eleven AZFP survey transects. Also, a Nansen net (200 &#xb5;m mesh) was towed vertically at five stations along survey transects.</p>
<p>Samples were preserved and carried at laboratory of Universidad de Valpara&#xed;so, Chile and then observed through an Olympus model S251 stereoscopic magnifying glass, up to a taxonomic resolution of specie in most of the recorded taxa. The samples were observed without a Folsom sample separator, using a Bogorov type plate and a mechanical counter to quantify the richness of taxa and abundance of organisms obtained in the different stations and sampled transects. The identification was made based on appropriate and specific literature for the zooplanktonic taxa of the coastal area of &#x200b;&#x200b;evaluation (i.e., <xref ref-type="bibr" rid="B12">Boltovskoy, 1981</xref>; <xref ref-type="bibr" rid="B42">Palma and Kaiser, 1993</xref>; <xref ref-type="bibr" rid="B28">Harris et&#xa0;al., 2000</xref>).</p>
<p>It should be noted that the 38 kHz channel is useful mainly for detection of large targets and air bubbles, such as fish with swim bladders (<xref ref-type="bibr" rid="B61">Ventero, 2016</xref>). Based on this and preliminary backscatter results, this channel was used to estimate depth since it had the longest range of the four frequencies (<xref ref-type="bibr" rid="B7">ASL, 2017</xref>). A sudden increase in the count value was observed at a given range due to the reflection of the signal with the seabed. For more accurate depth estimates, all gaps in the raw depth log, as well as those generated by subsequent outlier removal, were filled by interpolating from neighboring values. So, to estimate slope and roughness from the AZFP data, based on the resultant bathymetric layer, roughness (ROU) and slope (SLO) were calculated using the R package raster (<xref ref-type="bibr" rid="B31">Hijmans et&#xa0;al., 2018</xref>).</p>
</sec>
<sec id="s2_3">
<title>Systematic sighting generalized additive model analysis</title>
<p>For the statistical analysis, a grid of 500&#xa0;m x 500&#xa0;m grid-cell size was used to assign whales presence/absence, explanatory variables described above, and effort. Years were considered independent so this variable allocation process was conducted separately for 2018 and 2019. Fin whales and unidentified whales&#x2019; sightings were combined to obtain a sufficient sample size for statistical analyses. Other species were not included in the statistical analysis because of low sample size. More than one sighting per grid-cell occurred on five occasions (8% of the presence data), because of the small number of such records we preferred to model presence/absence per grid-cell for each year instead of using a count model. On-effort track length, expressed in km, was calculated for each one of the grid-cells and for each year. To avoid extreme values in effort allocation, values below the lowest 5% of effort values (0.1&#xa0;km) were replaced by this value, and a similar but inverse procedure was undertaken for the highest 1% values (5&#xa0;km). Grid-cells with missing data (i.e., without effort allocation and/or AZFP backscatter data) were excluded from this analysis and the remaining were grouped for both years to generate the final data base.</p>
<p>Generalized Additive Models (GAMs) were built to examine the relationships between whale occurrence and water column integrated acoustic density from the 125, 200 and 455 kHz channels as previously described. Depth, slope, and bottom roughness were also included as predictors. GAMs were built using the R package mgcv v1.8-31 (<xref ref-type="bibr" rid="B63">Wood, 2017</xref>). The response variable presence/absence was modeled using a binomial distribution with effort used as an offset term. Restricted maximum likelihood (REML) was used to optimize the parameter estimates and a variable selection process that uses a shrinkage approach to modify the smoothing penalty, allowing the smooth to be identically zero and to be removed from the model (<xref ref-type="bibr" rid="B37">Marra and Wood, 2011</xref>). Variables that had p&#x2010;values &gt; 0.05 were removed and the models were refitted to ensure that all remaining variables had p&#x2010;values&lt; 0.05 (<xref ref-type="bibr" rid="B47">Roberts et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B46">Redfern et&#xa0;al., 2017</xref>). Thin-plate regression splines were used for all predictors. Established metrics to compare the performance of the models were used, including Akaike&#x2019;s information criterion (AIC; <xref ref-type="bibr" rid="B2">Akaike, 1973</xref>), Akaike weights (wAIC), REML score, the percentage of explained deviance (ExpDev), the area under curve (AUC), and visual inspection of predicted densities. AUC was calculated using the R package ROCR v.1.0-11 (<xref ref-type="bibr" rid="B56">Sing et&#xa0;al., 2005</xref>). Model fit was assessed through a simulation-based approach and scaled residuals diagnostics using the R package DHARMa v. 0.4.551 (<xref ref-type="bibr" rid="B29">Hartig, 2022</xref>). With this package, quantile dispersion, outliers and deviation tests were implemented.</p>
</sec>
<sec id="s2_4">
<title>Opportunistic whale sighting data</title>
<p>Opportunistic whale sighting data has been compiled by the Corporaci&#xf3;n Nacional Forestal (CONAF) park service since 2015, based on reports from the fleet of 24 whale-watching boats that operate out of the village of Cha&#xf1;aral de Aceituno and provide whale-watching tours around Isla Cha&#xf1;aral during the spring and summer months. These data were analyzed to examine any possible interannual variation in whale presence off Isla Cha&#xf1;aral over a 5-year period (2015-2019). Opportunistic sighting data for each day during January and February were used from this database because these months are the high tourist season, and we could confidently assume that the full fleet of whale-watching boats were operating every day during these two months with negligible differences in sighting effort between both months. For both months, days with presence were determined and the monthly presence of whales (blue, fin, humpback) was calculated as the number of days with presence as a percentage of total days in the month for January and February 2015-2019.&#xa0;A five-year average was also calculated for monthly presence between 2015 and 2019.</p>
<p>Sources of bias in this citizen science dataset are probable, given that boats do not cover the study area systematically or uniformly and will often sight animals closest to port to reduce fuel costs. There is bias in reporting of species, blue whales and humpbacks are less common and arguably more charismatic than fin whales, so although there may be many fin whales in the area, when a single humpback or blue whale is present, boats will prefer these species to fin whales, which results in underreporting fin whales and overreporting blue and humpback whales. To minimize these possible biases, we preferred a presence/absence metric (monthly presence %) versus a count metric of number of sightings.</p>
</sec>
<sec id="s2_5">
<title>Auxiliary oceanographic data</title>
<p>Remote-sensing oceanographic data were obtained to interpret any possible interannual differences in whale sightings (systematic and opportunistic) and to examine the oceanographic dynamics of the wider Humboldt Archipelago area. Thus, satellite sea surface temperature (SST) and chlorophyll-a (Chla) were obtained from the MUR (Global, 0.01 degrees) product and Aqua MODIS (Global, 0.025 degrees), respectively. An average SST and Chla were obtained for the spring (November 2017 and 2018) preceding the systematic data collection in 2018 and 2019, and for the AZFP summer surveys (i.e., January 23rd to February 19th, 2018, and between February 2nd and 28th, 2019). Examining SST and Chla in November can provide information on what the upwelling-favorable conditions were like in the spring preceding the summer sampling campaigns, as November is historically the month with the strongest favorable upwelling winds (Ramos, Unpublished) which increases nutrients available for the development of phytoplankton blooms and the subsequent proliferation of zooplankton (<xref ref-type="bibr" rid="B9">Barlow et&#xa0;al., 2021</xref>).</p>
<p>To estimate the differences in current flow in the study area, mean geostrophic currents (from AVISO-CNES) were calculated for January and February 2018 and 2019. Moreover, mean currents obtained from a moored Acoustic Doppler Current Profiler (ADCP: RDI-Teledyne, Workhorse Sentinel, 300 kHz Model) deployed between Isla Cha&#xf1;aral and the mainland at 100&#xa0;m depth, were also included in this analysis (Ramos, Unpublished). In this case, the mean corresponds to the depth average of the velocity vector for the same periods considered in obtaining the mean geostrophic currents. Additionally, to determine possible recirculation current systems off central-northern Chile, long-term averages (1993-2020) of geostrophic currents (from AVISO-CNES) in a meridional band between 20&#xb0; and 40&#xb0;S were calculated.</p>
<p>During the intensive campaign with the AZFP, no measurements of spatial and hydrographic currents were made around Isla Cha&#xf1;aral. For this reason, results from one of the few oceanographic studies with sufficient spatial coverage (with sampling resolution of ~1.5 km) around the island were used to analyze the oceanographic conditions in the area. This oceanographic study was a FIP project (Fisheries Research Fund: FIP 2006-56, <xref ref-type="bibr" rid="B25">Gaymer et&#xa0;al., 2008)</xref> carried out between January 22 and 25, 2008, in which hydrographic and current measurements were obtained around Isla Cha&#xf1;aral. Thus, temperature, salinity (conductivity) and pressure data were obtained with a CTD (conductivity, Temperature and Depth) Sea-Bird, model SBE 19. A total of 15 CTD stations were carried out in two zonal sections, one in the north and one in the south of the island. On the other hand, measurements of the velocity profiles of the currents were made along four transects. For this, an ADCP (RDI Workhorse Sentinel-300 KHz) with a range of 120&#xa0;m was used. The ADCP was installed on an arm at the side of the vessel (starboard) at an average depth of 1&#xa0;m, the position and speed of the ship were recorded with a Garmin GPS (GPSmap 188C). The setup and acquisition of ADCP measurements were performed with the WinRiver program. The processing and analysis of the current data obtained with the ADCP (in Bottom Tracking mode) was carried out according to the protocol proposed by the manufacturer and the procedure followed by <xref ref-type="bibr" rid="B41">Moraga et&#xa0;al. (2011)</xref>, and <xref ref-type="bibr" rid="B60">Valle-Levinson and Moraga-Opazo (2006)</xref>. This includes the correction proposed by <xref ref-type="bibr" rid="B32">Joyce (1989)</xref> to eliminate the error of the ADCP magnetic compass with respect to the GPS direction.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<title>Results</title>
<sec id="s3_1">
<title>Spatial distribution of whales and prey backscatter</title>
<p>A total of 512.6&#xa0;km of survey effort over 20 days in 2018 (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1B</bold>
</xref>) and 318.3&#xa0;km over 16 days in 2019 (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1C</bold>
</xref>) were achieved. A total of 42 fin whales, 0 blue whales and 66 unidentified whales were sighted in 2018, and 7 fin whales, 3 blue whales and 12 unidentified whales in 2019. No humpback whales were sighted during the systematic surveys. In 2018, whales were sighted predominantly to the south of Isla Cha&#xf1;aral and between the island and the mainland (<xref ref-type="fig" rid="f2">
<bold>Figures&#xa0;2A&#x2013;C</bold>
</xref>). In 2019, due to very low whale sightings in general around Isla Cha&#xf1;aral, survey effort was conducted further offshore where higher densities of fin and blue whales were eventually found (80% of total sightings that year) (<xref ref-type="fig" rid="f2">
<bold>Figures&#xa0;2D&#x2013;F</bold>
</xref>), (but this offshore area was not included in the statistical modeling analysis because this area was only surveyed on 2019). In addition, there were five sightings of odontocetes in 2019: two of Risso&#x2019;s dolphins (<italic>Grampus griseus</italic>), two of pilot whales (<italic>Globicephala</italic> sp.) and one sighting of bottlenose dolphins (<italic>Tursiops truncatus</italic>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Nautical area scattering strength (SA) for the 125 kHz <bold>(A, D)</bold>, 200 kHz <bold>(B, E)</bold> and 455 kHz channels <bold>(C, F)</bold> and whale sightings during 2018 (upper panels) and 2019 (bottom panels). In panel <bold>(F)</bold> the limits of Isla Cha&#xf1;aral Marine Reserve (black Line) are shown. It is worth noting low sightings during 2019, and mainly further offshore to the northwest of the island.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g002.tif"/>
</fig>
<p>Backscatter was much higher overall in 2018 compared with 2019 and was generally patchy, with areas of higher density to the south, to the north and in the canyon between the island and the mainland (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Complex bathymetric features (steep slope and topographic breaks) around the island were apparent on visual inspection of the bathymetric data (<xref ref-type="fig" rid="f1">
<bold>Figures&#xa0;1A</bold>
</xref>, <xref ref-type="fig" rid="f3">
<bold>3A</bold>
</xref>), with a canyon that surrounds the island, starting at 500&#xa0;m depth southwest of the island, shoaling to 200&#xa0;m depth in the channel between the island and the mainland, and then deepening again to 500&#xa0;m to the northwest of the island. In effect, the cross-section of AZFP survey data across the Isla Cha&#xf1;aral submarine canyon shows an example of high backscatter over a topographic break of the canyon (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>) particularly visible in 200 kHz channel (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>). At the right panels of the figure it is possible to observe swarms of zooplankton over the Isla Cha&#xf1;aral submarine canyon. As the frequency increases, the range decreases, so the detection of swarms is closer and appears larger in size, and the intensity of the backscatter depends on the taxonomic group.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Bathymetric map of systematic survey area <bold>(A)</bold> with a cross-section over the canyon (red line) and start location (black dot) carried out on February 18th 2018 at 15:09 (*) on the map. Echograms of the cross-section for frequencies of 125 kHz <bold>(B)</bold>, 200 kHz <bold>(C)</bold> and 455 kHz <bold>(D)</bold> are shown. Seabed in panels (B) and (C) marked by solid line.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g003.tif"/>
</fig>
<p>The dominant species in the zooplankton trawls were copepods and to a lesser extent euphausiids (identified as <italic>Euphausia mucronata</italic>) as well as larvae of crustaceans and mollusks. This was found from both the Bongo net samples and the Nansen net samples (data not shown).</p>
</sec>
<sec id="s3_2">
<title>Statistical modelling of fin whale habitat</title>
<p>The top three ranking models all included 455 kHz backscatter, along with one other variable, i.e., depth, slope, and roughness, respectively. However, the best model clearly outperformed all other models (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) and showed that whale probability of occurrence increased with increasing backscatter in the 455 kHz channel, and shallower depths (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). The 200 kHz channel also gave good model performances, although not as good as 455 kHz (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). Expected versus observed residual plots using scaled residuals and tests for dispersion, outliers and deviation showed good model fit for the best model (<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Material Figure&#xa0;1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Generalized Additive Model results and model selection based on Area Under the Curve (AUC), Deviance explained, Restricted Maximum Likelihood score (REML), Akaike&#x2019;s Information Criteria difference (&#x394;AIC), and Akaike weights (wAIC).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Model</th>
<th valign="middle" align="center">AUC</th>
<th valign="middle" align="center">Deviance Explained</th>
<th valign="middle" align="center">REML</th>
<th valign="middle" align="center">&#x394;AIC</th>
<th valign="middle" align="center">wAIC</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">y~s(SA455)+s(depth)</td>
<td valign="top" align="center">0.76</td>
<td valign="top" align="center">9.03</td>
<td valign="top" align="center">184.68</td>
<td valign="top" align="center">0.00</td>
<td valign="top" align="center">0.31</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA455)+s(slope)</td>
<td valign="top" align="center">0.76</td>
<td valign="top" align="center">9.01</td>
<td valign="top" align="center">184.67</td>
<td valign="top" align="center">0.06</td>
<td valign="top" align="center">0.30</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA455)+s(roughness)</td>
<td valign="top" align="center">0.76</td>
<td valign="top" align="center">8.57</td>
<td valign="top" align="center">185.11</td>
<td valign="top" align="center">1.40</td>
<td valign="top" align="center">0.15</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200_55)+s(slope)+s(depth)</td>
<td valign="top" align="center">0.76</td>
<td valign="top" align="center">8.29</td>
<td valign="top" align="center">184.32</td>
<td valign="top" align="center">2.22</td>
<td valign="top" align="center">0.10</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200_55)+s(depth)</td>
<td valign="top" align="center">0.74</td>
<td valign="top" align="center">7.45</td>
<td valign="top" align="center">184.99</td>
<td valign="top" align="center">4.07</td>
<td valign="top" align="center">0.04</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125_55)+s(slope)+s(depth)</td>
<td valign="top" align="center">0.76</td>
<td valign="top" align="center">7.62</td>
<td valign="top" align="center">184.92</td>
<td valign="top" align="center">4.52</td>
<td valign="top" align="center">0.03</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200)+s(slope)+s(depth)</td>
<td valign="top" align="center">0.75</td>
<td valign="top" align="center">7.35</td>
<td valign="top" align="center">185.85</td>
<td valign="top" align="center">5.73</td>
<td valign="top" align="center">0.02</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125_55)+s(depth)</td>
<td valign="top" align="center">0.75</td>
<td valign="top" align="center">6.94</td>
<td valign="top" align="center">185.43</td>
<td valign="top" align="center">5.83</td>
<td valign="top" align="center">0.02</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200)+s(depth)</td>
<td valign="top" align="center">0.74</td>
<td valign="top" align="center">6.69</td>
<td valign="top" align="center">186.32</td>
<td valign="top" align="center">6.99</td>
<td valign="top" align="center">0.01</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125)+s(depth)</td>
<td valign="top" align="center">0.74</td>
<td valign="top" align="center">6.53</td>
<td valign="top" align="center">186.25</td>
<td valign="top" align="center">7.45</td>
<td valign="top" align="center">0.01</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125_55)+s(slope)</td>
<td valign="top" align="center">0.71</td>
<td valign="top" align="center">6.27</td>
<td valign="top" align="center">186.51</td>
<td valign="top" align="center">8.31</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125)+s(slope)</td>
<td valign="top" align="center">0.71</td>
<td valign="top" align="center">5.93</td>
<td valign="top" align="center">187.19</td>
<td valign="top" align="center">9.65</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125_55)+s(roughness)</td>
<td valign="top" align="center">0.70</td>
<td valign="top" align="center">5.78</td>
<td valign="top" align="center">187.26</td>
<td valign="top" align="center">10.06</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA125)+s(roughness)</td>
<td valign="top" align="center">0.70</td>
<td valign="top" align="center">5.48</td>
<td valign="top" align="center">187.89</td>
<td valign="top" align="center">11.27</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200)+s(slope)</td>
<td valign="top" align="center">0.68</td>
<td valign="top" align="center">5.04</td>
<td valign="top" align="center">188.97</td>
<td valign="top" align="center">13.12</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200_55)+s(slope)</td>
<td valign="top" align="center">0.68</td>
<td valign="top" align="center">4.80</td>
<td valign="top" align="center">189.46</td>
<td valign="top" align="center">14.06</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA455_55)+s(depth)</td>
<td valign="top" align="center">0.73</td>
<td valign="top" align="center">4.75</td>
<td valign="top" align="center">189.35</td>
<td valign="top" align="center">14.17</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200)+s(roughness)</td>
<td valign="top" align="center">0.66</td>
<td valign="top" align="center">4.54</td>
<td valign="top" align="center">189.77</td>
<td valign="top" align="center">14.94</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA455_55)+s(slope)</td>
<td valign="top" align="center">0.71</td>
<td valign="top" align="center">4.39</td>
<td valign="top" align="center">189.86</td>
<td valign="top" align="center">15.43</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA200_55)+s(roughness)</td>
<td valign="top" align="center">0.66</td>
<td valign="top" align="center">4.27</td>
<td valign="top" align="center">190.34</td>
<td valign="top" align="center">16.04</td>
<td valign="top" align="center">0.00</td>
</tr>
<tr>
<td valign="top" align="left">y~s(SA455_55)+s(roughness)</td>
<td valign="top" align="center">0.70</td>
<td valign="top" align="center">4.00</td>
<td valign="top" align="center">190.39</td>
<td valign="top" align="center">16.74</td>
<td valign="top" align="center">0.00</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Response curves for the selected GAM with <bold>(A)</bold> 455kHz backscatter, and <bold>(B)</bold> depth as explanatory variables. Grey shades represent confidence intervals.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g004.tif"/>
</fig>
</sec>
<sec id="s3_3">
<title>Opportunistic whale sightings</title>
<p>A total of 2,571 opportunistic sightings were recorded in January and February between 2015 and 2019 by CONAF, including 1,745 fin whale sightings (67.9% of total), 588 blue whale sightings (22.9% of total) and 238 humpback whale sightings (9.3% of total). No other whale species were sighted. Fin whale presence dominated during all years compared with blue and humpback whales, with a much higher average over the 5 years of data (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). Regarding monthly presence, the years with least fin whale presence were 2018 and 2019 (particularly in February) and with lowest blue whale presence was 2018 (both in January and February) and 2019 (particularly in February) (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). For humpback whales, no sightings were reported in 2018 and only two days with presence were recorded in 2019. The low overall whale presence in February 2019 was also reflected by low systematic sightings in 2019. Additionally, odontocete species were reported: bottlenose dolphins were the most common due to resident and transient populations in this area (<xref ref-type="bibr" rid="B52">Santos Carvallo et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B44">P&#xe9;rez-Alvarez et&#xa0;al., 2018</xref>); Sperm whales (<italic>Physeter macrocephalus</italic>), pilot whales, Risso&#x2019;s dolphins, Dusky dolphins (<italic>Lagenorhynchus obscurus</italic>), and orca (<italic>Orcinus orca</italic>) were also sighted.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Monthly presence of whales in January and February as percentage of days with presence per month using opportunistic sighting data of <bold>(A)</bold> blue whales, <bold>(B)</bold> fin whales and <bold>(C)</bold> humpback whales. Dotted line indicates average over 5 years.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g005.tif"/>
</fig>
</sec>
<sec id="s3_4">
<title>Oceanographic data</title>
<p>Local current measurements (<xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6</bold>
</xref>, <xref ref-type="fig" rid="f7">
<bold>7</bold>
</xref>) and hydrography (<xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7</bold>
</xref>) from 2008 show recirculation around the island and within the canyon. <xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6</bold>
</xref>, <xref ref-type="fig" rid="f7">
<bold>7B</bold>
</xref> show southward flow around the Isla Cha&#xf1;aral especially in the northern part of the channel between the island and the mainland. However, the southern transect (<xref ref-type="fig" rid="f7">
<bold>Figures&#xa0;7D&#x2013;F</bold>
</xref>) across the southern part of the canyon reveals northerly flow along the western bank of the canyon (towards the left hand side of <xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7E</bold>
</xref>), while southerly flow persists on the eastern bank of the canyon (towards the right hand side of <xref ref-type="fig" rid="f7">
<bold>Figure&#xa0;7E</bold>
</xref>). In the southern section of the canyon, we observe colder, higher salinity, higher density water along the bottom of the canyon intruding upwards, indicated by upward bending temperature, salinity, and density isolines (<xref ref-type="fig" rid="f7">
<bold>Figures&#xa0;7J&#x2013;L</bold>
</xref>). Based on the remote sensing data, spring 2017 was markedly cooler and more productive than spring 2018 (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8</bold>
</xref>), with the area of high Chla extending further offshore in 2017 (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8C</bold>
</xref>) compared to 2018 (<xref ref-type="fig" rid="f8">
<bold>Figure&#xa0;8D</bold>
</xref>). Note that spring 2017 and summer 2018 were under La Ni&#xf1;a conditions, while spring 2018 and summer 2019 were under El Ni&#xf1;o conditions (see Oceanic Ni&#xf1;o Index, <ext-link ext-link-type="uri" xlink:href="https://origin.cpc.ncep.noaa.gov/products/analysis_monitoring/ensostuff/ONI_v5.php">https://origin.cpc.ncep.noaa.gov/products/analysis_monitoring/ensostuff/ONI_v5.php</ext-link>).</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Surface current measurements around Isla Cha&#xf1;aral carried out on January 23, 2008 (Fisheries Research Fund grant: FIP 2006-56, 2008), over bathymetry from this study. Current vectors (black arrows) represent the vertical averages of current velocities (from a towed Acoustic Doppler Current Profiler, ADCP) over four transects. The red circles are the positions of the hydrographic (CTD) sampling stations north and south of Isla Cha&#xf1;aral. The cross-sections of the currents and hydrography are shown in <xref ref-type="fig" rid="f9">
<bold>Figure&#xa0;9</bold>
</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g006.tif"/>
</fig>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p>Cross-sections of the current velocities and hydrography around Isla Cha&#xf1;aral during January 23, 2008 (see <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref> for locations). <bold>(A&#x2013;C)</bold> U (East-West) and V (North-South) components of the current velocities, and backscatter (from ADCP) for the northern section, respectively. <bold>(D&#x2013;F)</bold> same as above but for the southern section. <bold>(G&#x2013;I)</bold> Temperature, salinity and density (from the CTD) for the northern section, respectively. <bold>(J&#x2013;L)</bold> same as previous, but for the southern section.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g007.tif"/>
</fig>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Mean sea surface temperature (top panels <bold>A</bold>, <bold>B</bold>)  (&#xb0;C) and chlorophyll a (bottom panels <bold>C</bold>, <bold>D</bold>) (mg m<sup>-3</sup>) during spring (November) 2017 and 2018.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g008.tif"/>
</fig>
<p>Average geostrophic currents near the coast for both survey years showed different patterns, especially south of Isla Cha&#xf1;aral (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9A, B</bold>
</xref>). In summer 2018, currents near the coast flowed towards the coast generating a flow towards Isla Cha&#xf1;aral and the Humboldt Archipelago; while in summer 2019, the currents to the south of Isla Cha&#xf1;aral flowed away from the coast. However, very close to shore, in the channel between Isla Cha&#xf1;aral and the mainland, we observed flow to the southeast during both years (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9A, B</bold>
</xref>) which is in line with southward flow reported in <xref ref-type="fig" rid="f6">
<bold>Figures&#xa0;6</bold>
</xref>, <xref ref-type="fig" rid="f7">
<bold>7B</bold>
</xref>. Backscatter was highest over the canyon as seen in both northern and southern transects, and the edge of the continental shelf (<xref ref-type="fig" rid="f7">
<bold>Figures&#xa0;7C, F</bold>
</xref>).</p>
<fig id="f9" position="float">
<label>Figure&#xa0;9</label>
<caption>
<p>(Top panels <bold>A</bold>, <bold>B</bold>) Mean geostrophic currents during <bold>(A)</bold> January-February 2018, and <bold>(B)</bold> January-February 2019, respectively. In <bold>(A, B)</bold> the average velocity vectors located between Isla Cha&#xf1;aral and the mainland were obtained from an ADCP (Acoustic Doppler Current Profiler) deployed at a depth of 100&#xa0;m. (Bottom panels <bold>C</bold>, <bold>D</bold>) Mean geostrophic currents over 28 years (1993-2020) in <bold>(C)</bold> for the most coastal pixel (+/- STD), and in <bold>(D)</bold> for the considered domain (20&#xb0;-40&#xb0;S), respectively. Note that in <bold>(D)</bold> the color shading is the velocity zonal component (Ug) to highlight the zonal flow (positive and negative) off the north-central zone of Chile. Red box contains the recirculation system that could explain the great biological productivity of the Humboldt Archipelago.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g009.tif"/>
</fig>
<p>The long-term average of the geostrophic currents off central-northern Chile (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9C, D</bold>
</xref>) showed a recirculation system between 28&#xb0; and 31&#xb0;S, with a flow towards the coast in the northern part of that region. This is also consistent with the average geostrophic flows from Isla Cha&#xf1;aral to the north during both surveys (<xref ref-type="fig" rid="f9">
<bold>Figures&#xa0;9A, B</bold>
</xref>).</p>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<title>Discussion</title>
<p>Our results highlight areas of higher backscatter and whale sightings around Isla Cha&#xf1;aral and the submarine canyon; and greater presence of whales and higher backscatter associated with bathymetric features (depth, slope and roughness). There is however, significant variation year-to-year in the presence of whales, as seen in both the systematic and opportunistic sighting datasets, driven by variation in oceanographic conditions. Here we discuss the circulation regime around Isla Cha&#xf1;aral and the Humboldt Archipelago which we summarize in a conceptual schema in <xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref>. We also discuss the dynamics of whales and prey around Isla Cha&#xf1;aral, and make some recommendations for protection of this habitat.</p>
<fig id="f10" position="float">
<label>Figure&#xa0;10</label>
<caption>
<p>Scheme of the circulation and physical processes around Isla Cha&#xf1;aral: 1) Isla Cha&#xf1;aral Submarine Canyon, 2) Topographically caused upwelling associated with the canyon, 3) Upwelling Jet, 4) Peru-Chile Undercurrent (polewards), 5) Ekman Transport, 6) Coastal Upwelling, 7) Recirculation associated with zonal striations*, 8) Coastal circulation*, 9) Flow with changing direction. *Poorly understood connection between geostrophic flow and coastal circulation.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-10-1208262-g010.tif"/>
</fig>
<sec id="s4_1">
<title>Circulation regime around the Humboldt Archipelago</title>
<p>The long-term average geostrophic currents over almost three decades revealed a recirculation system between 28&#xb0; and 31&#xb0;S showing flow back toward the coast in the northern part of that area. This recirculation could be part of a near zonal (east-west) pattern of the average geostrophic currents along the Chilean coast that is associated with so-called striations (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref>), i.e., bands interspersed with positive and negative zonal flows (<xref ref-type="bibr" rid="B11">Belmadani et&#xa0;al., 2017</xref>). These striations have been observed in different regions of the global ocean through remote-sensing altimetry and derived geostrophic currents (e.g., <xref ref-type="bibr" rid="B38">Maximenko et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B11">Belmadani et&#xa0;al., 2017</xref>). The presence of eastward and westward jets seems to result from the eddy field generated in the presence of planetary vorticity gradient and they efficiently develop near eastern boundaries related to particular coastal features (<xref ref-type="bibr" rid="B19">Davis et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B11">Belmadani et&#xa0;al., 2017</xref>). This circulation pattern may explain the high biological productivity in the area because productivity gets trapped nearer the coast in the Humboldt Archipelago area.</p>
<p>In the northern part of this study area we observed the mean geostrophic flow towards the coast (with an eastern component) (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref>) which should favor the transport of planktonic organisms towards the coast, especially those species that display diel vertical migration below of the superficial Ekman layer, like euphausiids, taking advantage of the opposite flow at depth. In general, selective vertical migration whether diel, seasonal or ontogenetic helps the species to conserve energy, locate food, retain a certain location or to move to other locations (e.g., <xref ref-type="bibr" rid="B33">K&#xe4;mpf and Chapman, 2016</xref>). A recent study about lateral transport of zooplankton explains trophic and taxonomic similarities over the zonal gradient of central Chile showing the importance of geostrophic current on the zooplankton distribution and suggesting that cross-shelf advection is a key process promoting zooplankton export to the deep-water ecosystem (<xref ref-type="bibr" rid="B26">Gonz&#xe1;lez et&#xa0;al., 2023</xref>). Although this study focused on offshore transport over a short observation period and did not consider the onshore component of transport, its results could show offshore zonal flow as part of the oceanic striations reported along the coast for long-term average flows, but without considering the onshore transport reported here. This onshore flow could in part explain why this area is so biologically productive and biodiverse and provides a good feeding habitat for whales that feed on euphausiids. During our study period, there was however variability in the flow of currents in the Humboldt Archipelago, mainly at south of Isla Cha&#xf1;aral i.e. 2018 the flow was toward coast and away from coast in 2019. This demonstrate interannual variability in the circulation regime around the Humboldt Archipelago, and may explain the low productivity, and consequently the low whale sightings observed that year.</p>
</sec>
<sec id="s4_2">
<title>Circulation around Isla Cha&#xf1;aral and the submarine canyon</title>
<p>Local current velocities measured in 2008 around Isla Cha&#xf1;aral clearly show the effects of the island and the submarine canyon on flow regime, with southeast flow coming from the northern part of the channel between the island and the mainland (<xref ref-type="fig" rid="f10">
<bold>Figure&#xa0;10</bold>
</xref>), which is consistent with the ADCP measurements made during this study. The intrusion of colder, more saline, denser bottom water (elevations of the temperature, salinity, and density isolines) into the southern part of submarine canyon suggests the entrance of locally upwelled water into the canyon from the south.</p>
<p>The geostrophic currents closest to the coast in the northern part of the island show a similar flow towards the coast for both measured periods (2018 and 2019) and for the long-term average. We hypothesize that this southeastward flow (towards the coast) is the dominant flow to the north of the island, but more observations of currents in the region are needed to verify this. The currents to the south of the island are more complex and show a cyclonic rotation over the canyon. This has been reported in several studies on the hydrodynamic effects of canyons in EBUS (e.g., <xref ref-type="bibr" rid="B30">Hickey, 1997</xref>; <xref ref-type="bibr" rid="B57">Sobarzo et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B50">Sald&#xed;as and Allen, 2020</xref>). The presence of canyons in the continental shelf produces a geostrophic flow imbalance along the isobaths because of the abrupt change in the bottom topography, generating ageostrophic flows transverse to the shelf within and over the canyon (<xref ref-type="bibr" rid="B5">Allen et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B3">Allen and Durrieu de Madron, 2009</xref>), inducing local upwelling and even cyclonic eddies (<xref ref-type="bibr" rid="B34">Klinck, 1996</xref>; <xref ref-type="bibr" rid="B4">Allen and Hickey, 2010</xref>; <xref ref-type="bibr" rid="B45">Ramos-Musalem and Allen, 2019</xref>), depending on the direction of regional flow and stratification. However, it is not clear whether the pattern of currents detected in the canyon is due solely to the presence of the canyon or is also caused by the joint effect of the canyon and the island; this requires a more detailed study of the currents in the zone. Despite the above, the pattern of currents in the region can explain the concentration of zooplankton around Isla Cha&#xf1;aral and in the canyon, as found for other canyons elsewhere (e.g., <xref ref-type="bibr" rid="B5">Allen et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B4">Allen et&#xa0;al., 2010</xref>).</p>
</sec>
<sec id="s4_3">
<title>Spatial distribution of whales and prey around Isla Cha&#xf1;aral</title>
<p>Based on both observations and studies of fecal plumes from whales around Isla Cha&#xf1;aral, we know that fin whales and blue whales feed exclusively on the euphausiid species <italic>E. mucronata</italic> in this area (<xref ref-type="bibr" rid="B43">P&#xe9;rez et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B15">Buchan et&#xa0;al., 2021</xref>). <italic>E. mucronata</italic> is a keystone species in the HCS, and sustains higher trophic levels (<xref ref-type="bibr" rid="B6">Antezana, 2010</xref>) and is strongly associated with wind-driven coastal upwelling centers (<xref ref-type="bibr" rid="B23">Escribano et&#xa0;al., 2000</xref>). Planktonic prey like euphausiids are known to concentrate in areas where physical processes, such as the interaction between bathymetric and local circulation, promote their aggregation (e.g., <xref ref-type="bibr" rid="B17">Croll et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B22">Doniol-Valcroze et&#xa0;al., 2012</xref>). Our results indicate that the circulation regime in the wider Humboldt Archipelago area and the local circulation around Isla Cha&#xf1;aral and the submarine canyon, concentrate planktonic prey and therefore provide feeding habitat for baleen whales.</p>
<p>During the 2018 surveys, fin whales were spatially associated with the submarine canyon between Isla Cha&#xf1;aral and the mainland and the slope to the south of the canyon, features which are apparent in the bathymetric data presented here. Zooplankton backscatter was also highest in the canyon, as seen in the AZFP survey data from 2018 and the towed ADCP from 2008. The association of whales and their prey with bathymetric features and submarine canyons has been described before for blue whale feeding grounds, such as the Monterey Bay (<xref ref-type="bibr" rid="B17">Croll et&#xa0;al., 2005</xref>) and the St. Lawrence Gulf Estuary (<xref ref-type="bibr" rid="B22">Doniol-Valcroze et&#xa0;al., 2012</xref>), and has been suggested in Chilean Patagonia (<xref ref-type="bibr" rid="B14">Buchan and Qui&#xf1;ones, 2016</xref>). Complex bathymetry has a known effect on the aggregation of zooplankton (e.g., <xref ref-type="bibr" rid="B17">Croll et&#xa0;al., 2005</xref>) and retention of fish larval as described by <xref ref-type="bibr" rid="B48">Rojas and Landaeta (2014)</xref> for Mejillones Bay northern of Chile. A review of canyons in the California Current System (also an EBUS) by <xref ref-type="bibr" rid="B51">Santora et&#xa0;al. (2018)</xref> showed that 76% of krill hotspots occurred within or near submarine canyons, including smaller canyons (25 km<sup>2</sup>) like the Isla Cha&#xf1;aral canyon. <xref ref-type="bibr" rid="B5">Allen et&#xa0;al. (2001)</xref> showed eddy formation at the head of a shelf-break canyon during an upwelling event that concentrated euphausiids; it is not clear if this mechanism also applies to a much smaller canyon like the Isla Cha&#xf1;aral canyon.</p>
<p>Spatial distribution of whales and backscatter was in line with the results of the GAMs, where higher presence was associated with shallower depths (like those found near-shore and in the canyon) and high backscatter, particularly in the 455 kHz and 200 kHz channels. Similarly, a distribution model of fin whales in the study area demonstrated that shallow depths was one of the variables that better explained the presence and habitat use of this species (<xref ref-type="bibr" rid="B8">Barilari, 2022</xref>). Future research should include further surveys and should examine the physical and biological dynamics of the submarine canyon, particularly the links between circulation and euphausiid density in space and time.</p>
</sec>
<sec id="s4_4">
<title>Interannual variation in whale presence and oceanographic conditions</title>
<p>Although opportunistic sightings of whales off Isla Cha&#xf1;aral have been consistent in the summer months since at least 2015, interannual variability is likely caused by interannual variation in oceanographic conditions and therefore prey availability for whales. In effect, February 2019 was a particularly poor year for fin whale sightings, as seen by both the systematic surveys and the CONAF opportunistic sighting data. The differences in geostrophic flow and remote-sensing oceanographic data can explain differences between 2018 and 2019, where geostrophic flow was towards the coast in summer 2018 and the summer of 2018 was preceded by a cooler and more productive spring (September to December 2017). In contrast, during summer 2019, geostrophic flow was away from the coast and a warmer and less productive spring preceded this summer. The low near-shore sightings of whales during 2019, due to the low-productivity conditions around Isla Cha&#xf1;aral, suggests that in years where productivity is low around Isla Cha&#xf1;aral, whales may move offshore. Offshore scientific surveys of fisheries during summer 2019 did in fact report offshore sightings of whales (<xref ref-type="bibr" rid="B62">Bedri&#xf1;ana-Romano et al., 2022</xref>).</p>
</sec>
<sec id="s4_5">
<title>Expansion of the Isla Cha&#xf1;aral marine reserve and implementation of the Humboldt Archipelago marine protected area</title>
<p>Isla Cha&#xf1;aral has become increasingly well known nationally and internationally as a feeding ground for baleen whales, particularly fin whales, and an area where small-scale community-based whale-watching has flourished (<xref ref-type="bibr" rid="B54">Sep&#xfa;lveda et&#xa0;al., 2018</xref>). The Humboldt Archipelago and Isla Cha&#xf1;aral is critical feeding habitat for fin, blue and humpback whales, and a host of other marine species, including marine birds and other marine mammals. Fin whales were the most hunted mysticete whales in the HCS (<xref ref-type="bibr" rid="B1">Aguayo-Lobo et&#xa0;al., 1998</xref>) and are considered Endangered by the Chilean Ministry of the Environment, as are blue whales and humpback whales (<ext-link ext-link-type="uri" xlink:href="https://clasificacionespecies.mma.gob.cl/">https://clasificacionespecies.mma.gob.cl/</ext-link>). Despite the recent population estimate for fin whales in the HCS of 2511.5 ind. (2.5-97.5% C.I. = 1387.9 &#x2013; 5068.5) (<xref ref-type="bibr" rid="B10">Bedri&#xf1;ana-Romano et&#xa0;al., 2022</xref>) there is currently no information on the population trend of this species or the other baleen whales in the HCS, and thus no knowledge of whether these populations are recovering or not.</p>
<p>Given the unique oceanographic and bathymetric features that lead to high prey availability for baleen whales around Isla Cha&#xf1;aral, and also due to recent studies have shown that a large proportion of the sightings of cetacean species occur outside the current protected area (<xref ref-type="bibr" rid="B53">Sep&#xfa;lveda et&#xa0;al., 2016</xref>, <xref ref-type="bibr" rid="B27">Guti&#xe9;rrez et&#xa0;al. In press</xref>) we recommend that the Isla Cha&#xf1;aral Marine Reserve needs to be expanded. The Humboldt Archipelago has recently received international recognition as an Important Marine Mammal Area (IMMA) (<ext-link ext-link-type="uri" xlink:href="https://www.marinemammalhabitat.org/portfolio-item/humboldt-archipelago-imma/">https://www.marinemammalhabitat.org/portfolio-item/humboldt-archipelago-imma/</ext-link>). IMMAs are &#x201c;<italic>discrete portions of habitat, important to marine mammal species, that have the potential to be delineated and managed for conservation&#x201d;.</italic> Also, during 2023&#xa0;a MU MPA was approved by the Chilean Council of Ministers for Sustainability. This MPA could strengthen the conservation of marine mammals and their habitat, by including special zoning for critical features like the canyon described here. The protection of the canyon is likely to have benefits beyond the protection of critical habitat for whales, given the important ecosystem services provided by submarine canyons (<xref ref-type="bibr" rid="B24">Fern&#xe1;ndez-Arcaya et&#xa0;al., 2017</xref>). Lastly, we recommend that the MU MPA administration plan excludes the development of large industrial projects that lead to high levels of marine traffic and other environmental threats for whales and other species.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<title>Conclusions</title>
<list list-type="simple">
<list-item>
<p>1) We observed overlapping spatial distribution of whales, backscatter and the submarine canyon that is located to the south of the island and in the channel between the island and the mainland. This is in line with whale presence being associated with high levels of backscatter and shallow depths according to the GAM analysis. Higher sightings in summer 2018 vs. summer 2019 coincided with higher backscatter, indicating that these differences in whale presence were likely driven by differences in prey availability.</p>
</list-item>
<list-item>
<p>2) In summer 2018, geostrophic flow towards the coast and a cold productive spring could explain high backscatter and whale sightings around Isla Cha&#xf1;aral. In summer 2019, geostrophic flow away from the coast and a warm spring with low productivity could explain low backscatter levels and whale sightings around Isla Cha&#xf1;aral, highlighting the interannual variability of the area.</p>
</list-item>
<list-item>
<p>3) Long-term average geostrophic currents form a recirculation system between 28&#xb0;S and 31&#xb0;S that could transport nutrient-rich upwelled surface waters back towards the Humboldt Archipelago and contribute to high biological productivity in this area.</p>
</list-item>
<list-item>
<p>4) The unique oceanographic features of the Isla Cha&#xf1;aral and the Humboldt Archipelago area contribute to successful foraging conditions for Endangered baleen whales. Based on this evidence, we recommend the expansion of the Isla Cha&#xf1;aral Marine Reserve to include the submarine canyon between the mainland and the island, and the implementation of a MU MPA for the entire Humboldt Archipelago area that explicitly protects this critical habitat from large industrial projects (including mega-ports and associated marine traffic routes).</p>
</list-item>
</list>
</sec>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation. Further inquiries can be directed to the corresponding author. Requests to access the datasets should be directed to jorge.oyanadel@ce.ucn.cl.</p>
</sec>
<sec id="s7" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>Ethical approval was not required for the study involving animals in accordance with the local legislation and institutional requirements because no animals were manipulated and the regulations for sighting marine mammals established by the National Fisheries Service and Undersecretary of Fisheries of Chile (D.S. N&#xb0; 38-2011) were strictly followed.</p>
</sec>
<sec id="s8" sec-type="author-contributions">
<title>Author contributions</title>
<p>SB contributed to study conception, design, data collection and analysis, manuscript preparation, including writing the first draft of this manuscript, and securing funding. MR contributed to study design, oceanographic and hydroacoustic data collection and analysis, securing funding, and writing and preparation of manuscript. JO contributed to hydroacoustic data collection, processing and analysis, oceanographic data analysis, and writing and preparation of manuscript. MS contributed to sighting data collection and analysis, writing and preparation of manuscript. LB-R contributed to study design, sighting data collection and analysis, and writing and preparation of manuscript. MV contributed to acoustic data collection, processing and data analysis. MM carried out CONAF sighting data collection. OA contributed to processing and analysis of satellite data. MS and CO contributed to study design, manuscript preparation and securing funding. SP contributed to acoustic data collection, processing and analysis. All authors contributed to the article and approved the submitted version.</p>
</sec>
</body>
<back>
<sec id="s9" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. Financial support was provided by Agencia Nacional de Investigaci&#xf3;n y Desarrollo (<underline>ANID</underline>) grant R16A10003 to CO and SB. Partial funding was provided by Centro COPAS Sur-Austral (ANID PFB31 and ANID AFB170006), and Centro COPAS Coastal (ANID FB210021). This study was also made possible with an Early Career Scientist award to SB from ASL Environmental Sciences. Support was also provided by ANID project ANILLO ATE220044 (BiodUCCT) awarded to MR and FONDECYT project #11190999 awarded to OA, also OA, SB, CO, MR and MV are grateful for ANID&#x2019;s funding through the R20F0008 CLAP project. We thank the Fisheries Research Fund FIP 2006-56 for auxiliary oceanographic data. All instrument deployments were carried out under SHOA resolutions 13270-24-74, 13270-24-136, 13270-24-137, 13270-24-155, 13270-24-161.&#xa0;A MATLAB, 2022a Academic License to the Universidad Cat&#xf3;lica del Norte was used in this research.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>Our thanks to our captains Rafael Gonz&#xe1;lez and Rafael Gonz&#xe1;lez Jr. The authors thank Dr. Ricardo Bravo for help with analysis of zooplankton samples. Our thanks to the entire fleet of community whale-watching adventure tourism vessels that report opportunistic sighting data on a daily basis and the Corporaci&#xf3;n Nacional Forestal (CONAF) Atacama and their team of park rangers for collecting and compiling the opportunistic sighting data. Our thanks to Dr. Ana Ventero for help with acoustic data analysis and, finally, to all the anonymous reviewers.</p>
</ack>
<sec id="s10" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2023.1208262/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2023.1208262/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Image_1.jpeg" id="SF1" mimetype="image/jpeg">
<label>Supplementary Figure&#xa0;1</label>
<caption>
<p>Results for simulation-based and scaled residuals diagnostics for model-fit assessment on the best model retaining SA455 and depth. Left top panel shows expected versus observed residual plot using scaled residuals. Top right panel shows residual dispersion for observed data (red line) and simulated data (histogram). Left bottom panel show quantile deviation test, indicating no deviations detected. Right bottom panel shows result for outlier test, indicating now significant outliers detected.</p>
</caption>
</supplementary-material>
</sec>
<ref-list>
<title>References</title>
<ref id="B1">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Aguayo-Lobo</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Torres.</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Acevedo.</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>1998</year>). <article-title>Los mam&#xed;feros marinos de Chile</article-title>. <source>Ser&#xed;e Cient&#xed;fica. INACH</source> <volume>48</volume> (<issue>January</issue>), <fpage>19</fpage>&#x2013;<lpage>159</lpage>.</citation>
</ref>
<ref id="B2">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Akaike</surname> <given-names>H.</given-names>
</name>
</person-group> (<year>1973</year>). <source>Information theory and an extension of the maximum likelihood principle</source>. Eds. <person-group person-group-type="editor">
<name>
<surname>Petrov</surname> <given-names>B. N.</given-names>
</name>
<name>
<surname>Csaki</surname> <given-names>F.</given-names>
</name>
</person-group> (<publisher-loc>Akademiai Kiado, Budapest</publisher-loc>: <publisher-name>Proceedings of the 2nd International Symposium on Information Theory</publisher-name>), <fpage>267</fpage>&#x2013;<lpage>281</lpage>.</citation>
</ref>
<ref id="B3">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Allen</surname> <given-names>S. E.</given-names>
</name>
<name>
<surname>Durrieu de Madron</surname> <given-names>X.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>A review of the role of submarine canyons in deep-ocean exchange with the shelf</article-title>. <source>Ocean Sci.</source> <volume>5</volume> (<issue>4</issue>), <fpage>607</fpage>&#x2013;<lpage>620</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5194/os-5-607-2009</pub-id>
</citation>
</ref>
<ref id="B4">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Allen</surname> <given-names>S. E.</given-names>
</name>
<name>
<surname>Hickey</surname> <given-names>B.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Dynamics of advection-driven upwelling over a shelf break submarine canyon</article-title>. <source>J. Geophysical Res.</source> <volume>115</volume> (<issue>C8</issue>). doi:&#xa0;<pub-id pub-id-type="doi">10.1029/2009JC005731</pub-id>
</citation>
</ref>
<ref id="B5">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Allen</surname> <given-names>S. E.</given-names>
</name>
<name>
<surname>Vindeirinho</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Thomson</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Foreman</surname> <given-names>M. G.</given-names>
</name>
<name>
<surname>Mackas</surname> <given-names>D.</given-names>
</name>
</person-group> (<year>2001</year>). <article-title>Physical and biological processes over a submarine canyon during an upwelling event</article-title>. <source>Can. J. Fisheries Aquat. Sci.</source> <volume>58</volume> (<issue>4</issue>), <fpage>671</fpage>&#x2013;<lpage>684</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1139/f01-008</pub-id>
</citation>
</ref>
<ref id="B6">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Antezana</surname> <given-names>T.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Euphausia mucronata: A keystone herbivore and prey of the Humboldt Current System</article-title>. <source>Deep Sea Res. Part II: Topical Stud. Oceanography</source> <volume>57</volume> (<issue>7&#x2013;8</issue>), <fpage>652</fpage>&#x2013;<lpage>662</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.dsr2.2009.10.014</pub-id>
</citation>
</ref>
<ref id="B7">
<citation citation-type="journal">
<person-group person-group-type="author">
<collab>ASL</collab></person-group> (<year>2017</year>). <source>AZFP (Acoustic Zooplankton Fish Profiler) operator&#x2019;s manual</source> (<publisher-loc>Victoria, Canada</publisher-loc>: <publisher-name>ASL Editor Environmental Sciences Inc.</publisher-name>), <fpage>108</fpage>.</citation>
</ref>
<ref id="B8">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Barilari</surname> <given-names>F.</given-names>
</name>
</person-group> (<year>2022</year>). <source>Distribuci&#xf3;n potencial de la ballena fin (Balaenoptera physalus) en la zona norte y centro de Chile</source> (<publisher-loc>Universidad de Valpara&#xed;so</publisher-loc>: <publisher-name>Tesis de Magister en Ciencias Biol&#xf3;gicas menci&#xf3;n Biodiversidad y Conservaci&#xf3;n</publisher-name>).</citation>
</ref>
<ref id="B9">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Barlow</surname> <given-names>D. R.</given-names>
</name>
<name>
<surname>Klinck</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Ponirakis</surname> <given-names>D. W.</given-names>
</name>
<name>
<surname>Garvey</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Torres</surname> <given-names>L.G.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Fashionably late: Temporal and spatial lags between wind and blue whales in a coastal upwelling system</article-title>. <source>Sci. Rep.</source> <volume>1&#x2013;10</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-021-86403-y</pub-id>
</citation>
</ref>
<ref id="B10">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bedri&#xf1;ana-Romano</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Zarate</surname> <given-names>P. M.</given-names>
</name>
<name>
<surname>Hucke-Gaete</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Viddi</surname> <given-names>F. A.</given-names>
</name>
<name>
<surname>Buchan</surname> <given-names>S. J.</given-names>
</name>
<name>
<surname>Cari</surname> <given-names>I.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Abundance and distribution patterns of cetaceans and their overlap with vessel traffic in the Humboldt Current Ecosystem, Chile</article-title>. <source>Sci. Rep.</source> <volume>12</volume> (<issue>1</issue>), <fpage>1</fpage>&#x2013;<lpage>15</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-022-14465-7</pub-id>
</citation>
</ref>
<ref id="B11">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Belmadani</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Concha</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Donoso</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Chaigneau</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Colas</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Maximenko</surname> <given-names>N.</given-names>
</name>
<etal/>
</person-group>. (<year>2017</year>). <article-title>Striations and preferred eddy tracks triggered by topographic steering of the background flow in the eastern South Pacific</article-title>. <source>J. Geophysical Research: Oceans</source> <volume>122</volume> (<issue>4</issue>), <fpage>2847</fpage>&#x2013;<lpage>2870</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/2016JC012348</pub-id>
</citation>
</ref>
<ref id="B12">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Boltovskoy</surname> <given-names>D.</given-names>
</name>
</person-group> (<year>1981</year>). <source>Radiolaria: en Atlas del zooplancton del Atl&#xe1;ntico Sudoccidental y m&#xe9;todos de trabajo con el zooplancton marino</source> (<publisher-loc>Mar del Plata, Argentina</publisher-loc>: <publisher-name>Mar del Plata: Instituto Nacional de Investigaci&#xf3;n y Desarrollo Pesquero INIDEP</publisher-name>), <fpage>261</fpage>&#x2013;<lpage>316</lpage>.</citation>
</ref>
<ref id="B13">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Buchan</surname> <given-names>S. J.</given-names>
</name>
<name>
<surname>Guti&#xe9;rrez</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Baumgartner</surname> <given-names>M. F.</given-names>
</name>
<name>
<surname>Stafford</surname> <given-names>K. M.</given-names>
</name>
<name>
<surname>Ramirez</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Pizarro</surname> <given-names>O.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Distribution of blue and sei whale vocalizations, and temperature - salinity characteristics from glider surveys in the Northern Chilean Patagonia mega-estuarine system</article-title>. <source>Front. Mar. Sci.</source> <volume>9</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fmars.2022.903964</pub-id>
</citation>
</ref>
<ref id="B14">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Buchan</surname> <given-names>S. J.</given-names>
</name>
<name>
<surname>Qui&#xf1;ones</surname> <given-names>R. A.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>First insights into the oceanographic characteristics of a blue whale feeding ground in northern Patagonia, Chile</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>554</volume>, <page-range>183&#x2013;199</page-range>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps11762}</pub-id>
</citation>
</ref>
<ref id="B15">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Buchan</surname> <given-names>S. J.</given-names>
</name>
<name>
<surname>V&#xe1;squez</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Olavarr&#xed;a</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Castro</surname> <given-names>L. R.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Prey items of baleen whale species off the coast of Chile from fecal plume analysis</article-title>. <source>Mar. Mammal Sci.</source> <volume>37</volume> (<issue>3</issue>), <fpage>1116</fpage>&#x2013;<lpage>1127</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/mms.12782</pub-id>
</citation>
</ref>
<ref id="B16">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Cox</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Embling</surname> <given-names>C. B.</given-names>
</name>
<name>
<surname>Hosegood</surname> <given-names>P. J.</given-names>
</name>
<name>
<surname>Votier</surname> <given-names>S. C.</given-names>
</name>
<name>
<surname>Ingram</surname> <given-names>S. N.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Oceanographic drivers of marine mammal and seabird habitat-use across shelf-seas: a guide to key features and recommendations for future research and conservation management</article-title>. <source>Estuarine Coast. Shelf Sci.</source> <volume>212</volume>, <fpage>294</fpage>&#x2013;<lpage>310</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ecss.2018.06.022</pub-id>
</citation>
</ref>
<ref id="B17">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Croll</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Marinovic</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Benson</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Chavez</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Black</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Ternullo</surname> <given-names>R.</given-names>
</name>
<etal/>
</person-group>. (<year>2005</year>). <article-title>From wind to whales: trophic links in a coastal upwelling system</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>289</volume>, <fpage>117</fpage>&#x2013;<lpage>130</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps289117</pub-id>
</citation>
</ref>
<ref id="B18">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Daneri</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Dellarossa</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Qui&#xf1;ones</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Jacob</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Montero</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Ulloa</surname> <given-names>O.</given-names>
</name>
</person-group> (<year>2000</year>). <article-title>Primary production and community respiration in the Humboldt Current System off Chile and associated oceanic areas</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>197</volume>, <fpage>41</fpage>&#x2013;<lpage>49</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps197041</pub-id>
</citation>
</ref>
<ref id="B19">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Davis</surname> <given-names>A.</given-names>
</name>
<name>
<surname>di Lorenzo</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Luo</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Belmadani</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Maximenko</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Melnichenko</surname> <given-names>O.</given-names>
</name>
<etal/>
</person-group>. (<year>2014</year>). <article-title>Mechanisms for the emergence of ocean striations in the North Pacific</article-title>. <source>Geophysical Res. Lett.</source> <volume>41</volume> (<issue>3</issue>), <fpage>948</fpage>&#x2013;<lpage>953</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/2013GL057956</pub-id>
</citation>
</ref>
<ref id="B20">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>de Robertis</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Higginbottom</surname> <given-names>I.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>A post-processing technique to estimate the signal-to-noise ratio and remove echosounder background noise</article-title>. <source>ICES J. Mar. Sci.</source> <volume>64</volume> (<issue>6</issue>), <fpage>1282</fpage>&#x2013;<lpage>1291</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsm112</pub-id>
</citation>
</ref>
<ref id="B21">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Doniol-Valcroze</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Berteaux</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Larouche</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Sears</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Influence of thermal fronts on habitat selection by four rorqual whale species in the Gulf of St. Lawrence</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>335</volume>, <fpage>207</fpage>&#x2013;<lpage>216</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps335207</pub-id>
</citation>
</ref>
<ref id="B22">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Doniol-Valcroze</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Lesage</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Giard</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Michaud</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2012</year>). <article-title>Challenges in marine mammal habitat modelling: Evidence of multiple foraging habitats from the identification of feeding events in blue whales</article-title>. <source>Endangered Species Res.</source> <volume>17</volume> (<issue>3</issue>), <fpage>255</fpage>&#x2013;<lpage>268</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/esr00427</pub-id>
</citation>
</ref>
<ref id="B23">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Escribano</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Marin</surname> <given-names>V. H.</given-names>
</name>
<name>
<surname>Irribarren</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>2000</year>). <article-title>Distribution of Euphausia mucronata at the upwelling area of Peninsula Mejillones , northern Chile: the influence of the oxygen minimum layer</article-title>. <source>Scientia Marina</source> <volume>64</volume> (<issue>1</issue>), <fpage>69</fpage>&#x2013;<lpage>77</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3989/scimar.2000.64n169</pub-id>
</citation>
</ref>
<ref id="B24">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Fern&#xe1;ndez-Arcaya</surname> <given-names>U.</given-names>
</name>
<name>
<surname>Ram&#xed;rez-Llodra</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Aguzzi</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Allcock</surname> <given-names>A. L.</given-names>
</name>
<name>
<surname>Davies</surname> <given-names>J. S.</given-names>
</name>
<name>
<surname>Dissanayake</surname> <given-names>A.</given-names>
</name>
<etal/>
</person-group>. (<year>2017</year>). <article-title>Ecological role of submarine canyons and need for canyon conservation: a review</article-title>. <source>Front. Mar. Sci.</source> <volume>4</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fmars.2017.00005</pub-id>
</citation>
</ref>
<ref id="B25">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Gaymer</surname> <given-names>C. F.</given-names>
</name>
<name>
<surname>Stotz</surname> <given-names>W.</given-names>
</name>
<name>
<surname>Garay Fl&#xfc;hmann</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Luna-Jorquera</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Ramos</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2008</year>). <source>Evaluaci&#xf3;n de l&#xed;nea base de Las Reservas Marinas "Isla Cha&#xf1;aral" e "Isla Choros-Damas"</source> (<publisher-name>Informe Final Proyecto FIP 2006-56, Subsecretar&#xed;a de Pesca, Chile</publisher-name>), <fpage>532</fpage>.</citation>
</ref>
<ref id="B26">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gonz&#xe1;lez</surname> <given-names>C. E.</given-names>
</name>
<name>
<surname>Bode</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Fern&#xe1;ndez-Urruzola</surname> <given-names>I.</given-names>
</name>
<name>
<surname>Hidalgo</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Oerder</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Escribano</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2023</year>). <article-title>The lateral transport of zooplankton explains trophic and taxonomic similarities over the zonal gradient of central Chile</article-title>. <source>J. Mar. Syst.</source> <volume>238</volume>, <elocation-id>103840</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jmarsys.2022.103840</pub-id>
</citation>
</ref>
<ref id="B27">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Guti&#xe9;rrez</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Letelier</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Santos-Carvallo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Barilari</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Guti&#xe9;rrez</surname> <given-names>L.</given-names>
</name>
<name>
<surname>P&#xe9;rez-Alvarez</surname> <given-names>M. J.</given-names>
</name>
<etal/>
</person-group>. <article-title>Zoning proposal for a Marine Protected Area in Chile: a conservation tool for large cetaceans</article-title>. <source>Ocean Coast. Management</source>. [Epub ahead of print].</citation>
</ref>
<ref id="B28">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Harris</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Wiebe</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Lenz</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Skjoldal</surname> <given-names>H. R.</given-names>
</name>
<name>
<surname>Huntley</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2000</year>). <source>ICES Zooplankton methodology manual</source> (<publisher-loc>London</publisher-loc>: <publisher-name>Academic Press</publisher-name>).</citation>
</ref>
<ref id="B29">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Hartig</surname> <given-names>F.</given-names>
</name>
</person-group> (<year>2022</year>). <source>DHARMa: residual diagnostics for hierarchical (Multi-level / mixed) regression models</source>.</citation>
</ref>
<ref id="B30">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hickey</surname> <given-names>B.</given-names>
</name>
</person-group> (<year>1997</year>). <article-title>The response of a steep-sided, narrow canyon to time-variable wind forcing</article-title>. <source>J. Phys. Oceanography</source> <volume>27</volume> (<issue>5</issue>), <fpage>697</fpage>&#x2013;<lpage>726</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1175/1520-0485(1997)027&lt;0697:TROASS&gt;2.0.CO;2</pub-id>
</citation>
</ref>
<ref id="B31">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Hijmans</surname> <given-names>R. J.</given-names>
</name>
<name>
<surname>van Etten</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Cheng</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Sumner</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Mattiuzzi</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Greenberg</surname> <given-names>J. A.</given-names>
</name>
</person-group> (<year>2018</year>). <source>Raster: geographic data analysis and modeling</source>.</citation>
</ref>
<ref id="B32">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Joyce</surname> <given-names>T. M.</given-names>
</name>
</person-group> (<year>1989</year>). <article-title>On in <italic>situ</italic> &#x201c;calibration&#x201d; of shipboard ADCPs</article-title>. <source>J. Atmospheric Oceanic Technol.</source> <volume>6</volume>, <fpage>169</fpage>&#x2013;<lpage>172</lpage>. doi: <pub-id pub-id-type="doi">10.1175/1520-0426(1989)006&lt;0169:OISOSA&gt;2.0.CO;2</pub-id>
</citation>
</ref>
<ref id="B33">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>K&#xe4;mpf</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Chapman</surname> <given-names>P.</given-names>
</name>
</person-group> (<year>2016</year>). <source>Upwelling systems of the world A scientific journey to the most productive marine Ecosystems</source> (<publisher-loc>Switzerland</publisher-loc>: <publisher-name>Springer International Publishing</publisher-name>), <fpage>443</fpage>.</citation>
</ref>
<ref id="B34">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Klinck</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>1996</year>). <article-title>Circulation near submarine canyons: A modeling study</article-title>. <source>J. Geophysical Res.</source> <volume>101</volume> (<issue>C1</issue>), <fpage>1211</fpage>&#x2013;<lpage>1223</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1029/95JC02901</pub-id>
</citation>
</ref>
<ref id="B35">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Largier</surname> <given-names>J. L.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Upwelling bays: How coastal upwelling controls circulation, habitat and productivity in bays</article-title>. <source>Annu. Rev. of Mar. Sci.</source> <volume>12</volume>, <fpage>415</fpage>&#x2013;<lpage>447</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1146/annurev-marine-010419-011020</pub-id>
</citation>
</ref>
<ref id="B36">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>MacLennan</surname> <given-names>D. N.</given-names>
</name>
<name>
<surname>Fernandes</surname> <given-names>P. G.</given-names>
</name>
<name>
<surname>Dalen</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2002</year>). <article-title>A consistent approach to definitions and symbols in fisheries acoustics</article-title>. <source>ICES J. Mar. Sci.</source> <volume>59</volume> (<issue>2</issue>), <fpage>365</fpage>&#x2013;<lpage>369</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1006/jmsc.2001.1158</pub-id>
</citation>
</ref>
<ref id="B37">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Marra</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Wood</surname> <given-names>S. N.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Practical variable selection for generalized additive models</article-title>. <source>Comput. Stat Data Anal.</source> <volume>55</volume> (<issue>7</issue>), <fpage>2372</fpage>&#x2013;<lpage>2387</lpage>. doi: <pub-id pub-id-type="doi">10.1016/j.csda.2011.02.004</pub-id>
</citation>
</ref>
<ref id="B38">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Maximenko</surname> <given-names>N. A.</given-names>
</name>
<name>
<surname>Bang</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Sasaki</surname> <given-names>H.</given-names>
</name>
</person-group> (<year>2005</year>). <article-title>Observational evidence of alternating zonal jets in the world ocean</article-title>. <source>Geophysical Res. Lett.</source> <volume>32</volume> (<issue>12</issue>). doi:&#xa0;<pub-id pub-id-type="doi">10.1029/2005GL022728</pub-id>
</citation>
</ref>
<ref id="B39">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Montecino</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Lange</surname> <given-names>C. B.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>The Humboldt Current System: Ecosystem components and processes, fisheries, and sediment studies</article-title>. <source>Prog. Oceanography</source> <volume>83</volume> (<issue>1&#x2013;4</issue>), <fpage>65</fpage>&#x2013;<lpage>79</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.pocean.2009.07.041</pub-id>
</citation>
</ref>
<ref id="B40">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Monteiro</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Dewitte</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Scranton</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Palmier</surname> <given-names>A.</given-names>
</name>
<name>
<surname>van der Plas</surname> <given-names>A. K.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>The role of open ocean boundary forcing on seasonal to decadal-scale variability and long-term change of natural shelf hypoxia</article-title>. <source>Environ. Res. Letters.</source> <volume>6</volume>, <elocation-id>25002</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1088/1748-9326/6/2/025002</pub-id>
</citation>
</ref>
<ref id="B41">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Moraga</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Valle-Levinson</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Ramos</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Pizarro</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Upwelling-triggered near-geostrophic recirculation in an equatorward facing embayment</article-title>. <source>Continental Shelf Res.</source> <volume>31</volume>, <fpage>1991</fpage>&#x2013;<lpage>1999</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.csr.2011.10.002</pub-id>
</citation>
</ref>
<ref id="B42">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Palma</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Kaiser</surname> <given-names>K.</given-names>
</name>
</person-group> (<year>1993</year>). <source>Plancton marino de aguas Chilenas</source> (<publisher-loc>Chile</publisher-loc>: <publisher-name>Ediciones Universitarias de Valpara&#xed;so, Universidad Cat&#xf3;lica de Valpara&#xed;so</publisher-name>), <fpage>151</fpage>.</citation>
</ref>
<ref id="B43">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>P&#xe9;rez</surname> <given-names>M. J.</given-names>
</name>
<name>
<surname>Thomas</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Uribe</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Sep&#xfa;lveda</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Flores</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Moraga</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2006</year>). <article-title>Fin Whales (Balaenoptera physalus) Feeding on Euphausia mucronate in Nearshore Waters off North-Central Chile</article-title>. <source>Aquat. Mammals</source> <volume>32</volume> (<issue>1</issue>), <fpage>109</fpage>&#x2013;<lpage>113</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1578/am.32.1.2006.109</pub-id>
</citation>
</ref>
<ref id="B44">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>P&#xe9;rez-Alvarez</surname> <given-names>M. J.</given-names>
</name>
<name>
<surname>V&#xe1;squez</surname> <given-names>R. A.</given-names>
</name>
<name>
<surname>Moraga</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Santos-Carvallo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Kraft</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Sabaj</surname> <given-names>V.</given-names>
</name>
<etal/>
</person-group>. (<year>2018</year>). <article-title>Home sweet home: social dynamics and genetic variation of a long-term resident bottlenose dolphin population off the Chilean coast</article-title>. <source>Anim. Behav.</source> <volume>139</volume>, <fpage>81</fpage>&#x2013;<lpage>89</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.anbehav.2018.03.009</pub-id>
</citation>
</ref>
<ref id="B45">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ramos-Musalem</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Allen</surname> <given-names>S. E.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>The impact of locally-enhanced vertical diffusivity on the cross-shelf transport of tracers induced by a submarine canyon</article-title>. <source>J. Phys. Oceanography</source> <volume>49</volume>, <fpage>561</fpage>&#x2013;<lpage>584</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1175/jpo-d-18-0174.1</pub-id>
</citation>
</ref>
<ref id="B46">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Redfern</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Moore</surname> <given-names>T. J.</given-names>
</name>
<name>
<surname>Fiedler</surname> <given-names>P. C.</given-names>
</name>
<name>
<surname>de Vos</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Brownell</surname> <given-names>R. L.</given-names>
<suffix>Jr.</suffix>
</name>
<name>
<surname>Forney</surname> <given-names>K. A.</given-names>
</name>
<etal/>
</person-group>. (<year>2017</year>). <article-title>Predicting cetacean distributions in data-poor marine ecosystems</article-title>. <source>Diversity Distributions</source> <volume>23</volume> (<issue>4</issue>), <fpage>394</fpage>&#x2013;<lpage>408</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/ddi.12537</pub-id>
</citation>
</ref>
<ref id="B47">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Roberts</surname> <given-names>J. J.</given-names>
</name>
<name>
<surname>Best</surname> <given-names>B. D.</given-names>
</name>
<name>
<surname>Mannocci</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Fujioka</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Halpin</surname> <given-names>P. N.</given-names>
</name>
<name>
<surname>Palka</surname> <given-names>D. L.</given-names>
</name>
<etal/>
</person-group>. (<year>2016</year>). <article-title>Habitat-based cetacean density models for the U.S. Atlantic and Gulf of Mexico</article-title>. <source>Sci. Rep.</source> <volume>6</volume> (<issue>1</issue>), <elocation-id>22615</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/srep22615</pub-id>
</citation>
</ref>
<ref id="B48">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rojas</surname> <given-names>P. M.</given-names>
</name>
<name>
<surname>Landaeta</surname> <given-names>M. F.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Fish larvae retention linked to abrupt bathymetry at Mejillones Bay (northern Chile) during coastal upwelling events</article-title>. <source>Latin Am. J. Aquat. Res.</source> <volume>42</volume>, <fpage>989</fpage>&#x2013;<lpage>1008</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3856/vol42-issue5-fulltext-6</pub-id>
</citation>
</ref>
<ref id="B49">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rutllant</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Montecinos</surname> <given-names>V.</given-names>
</name>
</person-group> (<year>2002</year>). <article-title>Multiscale upwelling forcing cycles and biological response off north-central Chile</article-title>. <source>Rev. Chil. Hist. Natural</source> <volume>75</volume>, <fpage>231</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.4067/S0716-078X2002000100020</pub-id>
</citation>
</ref>
<ref id="B50">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sald&#xed;as</surname> <given-names>G. S.</given-names>
</name>
<name>
<surname>Allen</surname> <given-names>S. E.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>The influence of a submarine canyon on the circulation and cross-shore exchanges around an upwelling front</article-title>. <source>J. Phys. Oceanography</source> <volume>50</volume> (<issue>6</issue>), <fpage>1677</fpage>&#x2013;<lpage>1698</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1175/jpo-d-19-0130.1</pub-id>
</citation>
</ref>
<ref id="B51">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Santora</surname> <given-names>J. A.</given-names>
</name>
<name>
<surname>Zeno</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Dorman</surname> <given-names>J. G.</given-names>
</name>
<name>
<surname>Sydeman</surname> <given-names>W. J.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Submarine canyons represent an essential habitat network for krill hotspots in a Large Marine Ecosystem</article-title>. <source>Sci. Rep.</source> <volume>8</volume> (<issue>1</issue>), <fpage>7579</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-018-25742-9</pub-id>
</citation>
</ref>
<ref id="B52">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Santos Carvallo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>P&#xe9;rez Alvarez</surname> <given-names>M. J.</given-names>
</name>
<name>
<surname>Muniain</surname> <given-names>V.</given-names>
</name>
<name>
<surname>Moraga</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Oliva</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Sep&#xfa;lveda</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Trophic niche overlap between sympatric resident and transient populations of bottlenose dolphins in the Humboldt Current System off north-central Chile</article-title>. <source>Mar. Mammal Science.</source> <volume>31</volume> (<issue>2</issue>), <fpage>790</fpage>&#x2013;<lpage>799</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/mms.12185</pub-id>
</citation>
</ref>
<ref id="B53">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Sep&#xfa;lveda</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Oliva</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Pavez</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Santos-Carvallo</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2016</year>). <source>Caleta Cha&#xf1;aral de Aceituno: Destino tur&#xed;stico de alta calidad para el avistamiento de cet&#xe1;ceos, otros mam&#xed;feros y aves marinas</source> (<publisher-loc>Chile</publisher-loc>: <publisher-name>Servicio Nacional de Pesca y Acuicultura</publisher-name>).</citation>
</ref>
<ref id="B54">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sep&#xfa;lveda</surname> <given-names>M.</given-names>
</name>
<name>
<surname>P&#xe9;rez-&#xc1;lvarez</surname> <given-names>M. J.</given-names>
</name>
<name>
<surname>Santos-Carvallo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Pavez</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Olavarr&#xed;a</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Moraga</surname> <given-names>R.</given-names>
</name>
<etal/>
</person-group>. (<year>2018</year>). <article-title>From whaling to whale watching: Identifying fin whale critical foraging habitats off the Chilean coast</article-title>. <source>Aquat. Conserv. Mar. Freshw. Ecosystem</source> <volume>28</volume> (<issue>4</issue>), <fpage>821</fpage>&#x2013;<lpage>829</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/aqc.2899</pub-id>
</citation>
</ref>
<ref id="B55">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Shaffer</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Horm&#xe1;zabal</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Pizarro</surname> <given-names>O.</given-names>
</name>
<name>
<surname>Salinas</surname> <given-names>S.</given-names>
</name>
</person-group> (<year>1999</year>). <article-title>Seasonal and interannual variability of currents and temperature off central Chile</article-title>. <source>J. Geophysical Research: Oceans.</source> <volume>104</volume>, <fpage>29951</fpage>&#x2013;<lpage>29961</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1029/1999JC900253</pub-id>
</citation>
</ref>
<ref id="B56">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sing</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Sander</surname> <given-names>O.</given-names>
</name>
<name>
<surname>Beerenwinkel</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Lengauer</surname> <given-names>T.</given-names>
</name>
</person-group> (<year>2005</year>). <article-title>ROCR: visualizing classifier performance in R</article-title>. <source>Bioinformatics</source> <volume>21</volume> (<issue>20</issue>), <fpage>3940</fpage>&#x2013;<lpage>3941</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/bioinformatics/bti623</pub-id>
</citation>
</ref>
<ref id="B57">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sobarzo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Sald&#xed;as</surname> <given-names>G. S.</given-names>
</name>
<name>
<surname>Tapia</surname> <given-names>F. J.</given-names>
</name>
<name>
<surname>Bravo</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Moffat</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Largier</surname> <given-names>J. L.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>On subsurface cooling associated with the Biobio River canyon (Chile)</article-title>. <source>J. Geophysical Research: Oceans</source> <volume>121</volume>, <fpage>4568</fpage>&#x2013;<lpage>4584</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/2016JC011796</pub-id>
</citation>
</ref>
<ref id="B58">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Thiel</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Macaya</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Acu&#xf1;a</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Amtz</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Bast&#xed;as</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Brokordt</surname> <given-names>K.</given-names>
</name>
<etal/>
</person-group>. (<year>2007</year>). <article-title>The Humboldt current system of northern and central Chile: Oceanographic processes, ecological interactions and socio-economic feedback</article-title>. <source>Oceanography Mar. Biology: Annu. Rev.</source> <volume>45</volume>, <fpage>195</fpage>&#x2013;<lpage>344</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1201/9781420050943</pub-id>
</citation>
</ref>
<ref id="B59">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Toro</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Vilina</surname> <given-names>Y. A.</given-names>
</name>
<name>
<surname>Capella</surname> <given-names>J. J.</given-names>
</name>
<name>
<surname>Gibbons</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>Novel coastal feeding area for eastern South Pacific fin whales (Balaenoptera physalus) in mid-latitude humboldt current waters off Chile</article-title>. <source>Aquat. Mammals</source> <volume>42</volume> (<issue>1</issue>), <fpage>47</fpage>&#x2013;<lpage>55</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1578/AM.42.1.2016.47</pub-id>
</citation>
</ref>
<ref id="B60">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Valle-Levinson</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Moraga-Opazo</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2006</year>). <article-title>Bipolar residual circulation in two equatorward facing bays</article-title>. <source>Continental Shelf Res.</source> <volume>26</volume>, <fpage>179</fpage>&#x2013;<lpage>193</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.csr.2005.10.002</pub-id>
</citation>
</ref>
<ref id="B61">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Ventero</surname> <given-names>A.</given-names>
</name>
</person-group> (<year>2016</year>). <source>Estudio biol&#xf3;gico de la capa de dispersi&#xf3;n ac&#xfa;stica estival en el mar de Albor&#xe1;n y su implicaci&#xf3;n en la evaluaci&#xf3;n de pel&#xe1;gicos costeros</source> (<publisher-name>Doctoral dissertation, Universitat de les Illes Balears</publisher-name>).</citation>
</ref>
<ref id="B62">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ventero</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Iglesias</surname> <given-names>M.</given-names>
</name>
<name>
<surname>C&#xf3;rdoba</surname> <given-names>P.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Krill spatial distribution in the Spanish Mediterranean Sea in summer time</article-title>. <source>J. Plankton Res.</source> <volume>41</volume> (<issue>4</issue>), <fpage>491</fpage>&#x2013;<lpage>505</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/plankt/fbz030</pub-id>
</citation>
</ref>
<ref id="B63">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Wood</surname> <given-names>S. N.</given-names>
</name>
</person-group> (<year>2017</year>). <source>Generalized additive models: an introduction with R</source>, <edition>2nd ed</edition>. (<publisher-loc>Boca Raton, USA</publisher-loc>: <publisher-name>CRC Press</publisher-name>).</citation>
</ref>
</ref-list>
</back>
</article>