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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">127</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:15f175fc-9690-5d13-a2ea-114d8a2e74bd</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Frontiers of Biogeography</journal-title>
        <abbrev-journal-title xml:lang="en">biogeography</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">1948-6596</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.21425/fob.19.199542</article-id>
      <article-id pub-id-type="publisher-id">199542</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Perspectives</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Animalia</subject>
          <subject>Archaea</subject>
          <subject>Bacteria</subject>
          <subject>Chromista</subject>
          <subject>Fungi</subject>
          <subject>Plantae</subject>
          <subject>Protozoa</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Conservation</subject>
          <subject>Global environmental change</subject>
          <subject>Human impact</subject>
          <subject>Methodology</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>World</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>A code of conduct for environmental and biodiversity scientists</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Costello</surname>
            <given-names>Mark John</given-names>
          </name>
          <email xlink:type="simple">mark.j.costello@nord.no</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-2362-0328</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Muller-Karger</surname>
            <given-names>Frank E.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-3159-5011</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Bates</surname>
            <given-names>Amanda E.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-0198-4537</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Faculty of Biosciences and Aquaculture, Nord University, Bodo, Norway</addr-line>
        <institution>Faculty of Biosciences and Aquaculture, Nord University</institution>
        <addr-line content-type="city">Bodo</addr-line>
        <country>Norway</country>
        <uri content-type="ror">https://ror.org/030mwrt98</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">College of Marine Science, University of South Florida, St Petersburg, Florida, USA</addr-line>
        <institution>College of Marine Science, University of South Florida</institution>
        <addr-line content-type="city">St Petersburg</addr-line>
        <country>United States of America</country>
        <uri content-type="ror">https://ror.org/032db5x82</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">University of Victoria, 3800 Finnerty Rd, Victoria, British Columbia V8P 5C2, Canada</addr-line>
        <institution>University of Victoria</institution>
        <addr-line content-type="city">Victoria</addr-line>
        <country>Canada</country>
        <uri content-type="ror">https://ror.org/04s5mat29</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Mark John Costello (<email xlink:type="simple">mark.j.costello@nord.no</email>)</p>
        </fn>
        <fn>
          <p><bold>Editor</bold> Robert Whittaker</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>06</day>
        <month>07</month>
        <year>2026</year>
      </pub-date>
      <volume>19</volume>
      <elocation-id>e199542</elocation-id>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/6152125F-DDF5-5248-A6A5-FF2E9937BDAE">6152125F-DDF5-5248-A6A5-FF2E9937BDAE</uri>
      <history>
        <date date-type="received">
          <day>13</day>
          <month>05</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>25</day>
          <month>06</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Mark John Costello, Frank E. Muller-Karger, Amanda E. Bates</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>Abstract</label>
        <p>There are thousands of codes of conduct in workplaces. However, most are general for all employees and do not address the behaviour of scientists conducting fieldwork and associated activities. Thus, through consultation with many colleagues within and outside the Global Earth Observation Biodiversity Observation Network, we synthesised a proposed code of practice for researchers. This is applicable to people working in environmental sciences, including ecology, biogeography, geology and associated subjects involving fieldwork.</p>
        <sec sec-type="Highlights" id="sec1">
          <title>Highlights</title>
          <boxed-text position="float">
            <list list-type="bullet">
              <list-item>
                <p>This code of conduct recommends that scientists:</p>
              </list-item>
              <list-item>
                <p>Respect all people, nature and laws</p>
              </list-item>
              <list-item>
                <p>Challenge inappropriate behaviour and misinformation</p>
              </list-item>
              <list-item>
                <p>Maintain their integrity and be accountable</p>
              </list-item>
              <list-item>
                <p>Minimise impacts on wildlife and habitats</p>
              </list-item>
              <list-item>
                <p>Publish their findings with full acknowledgements</p>
              </list-item>
            </list>
          </boxed-text>
        </sec>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>Best practice</kwd>
        <kwd>environmental impact</kwd>
        <kwd>ethics</kwd>
        <kwd>fieldwork</kwd>
        <kwd>field work conduct</kwd>
        <kwd>health and safety</kwd>
        <kwd>individual behaviours</kwd>
        <kwd>law abiding</kwd>
        <kwd>publishing</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Preamble" id="sec2">
      <title>Preamble</title>
      <p>Science advances through critical thinking and the testing of ideas and practices, and discoveries may confirm and challenge existing paradigms. Science benefits from the diverse perspectives of people, with knowledge gained from oral and written history, philosophy, observations, experiments, data analysis, and practical skills. The diversity of beliefs, practices, values and thinking can create tensions, which can be constructive but may upset people and the status quo. As scientists we need to be sensitive and respectful of these tensions and their consequences, encourage collaboration, discussion and not imposition of ideas, and strive to do the best possible science that will benefit people and nature in present and future generations. Here, we consider “scientists” to cover all researchers and practitioners who are engaged in discovery, including consultants and others in the private sector, government policy advisors, and social and natural scientists spanning engineering and biomedicine to humanities and the arts. We reflect on the behaviours and practices that enable a science that benefits society. Therefore, the code is applicable to the field of biogeography, which is based on field observations of wildlife in all environments and parts of the world, benefits from a diverse community of scientists (including amateur naturalists), and has a history of challenging prevailing paradigms and controversies (e.g., assisted colonisation, de-extinction, new conservation, conservation trade-offs, excluding people from protected areas). As with other codes, it aims to be a moral compass for individual and group behaviour.</p>
      <p>Effective science practices and good citizenship require that we hold ourselves to high ethical and moral standards (<xref ref-type="bibr" rid="B21">Institute of Medicine 2009</xref>; <xref ref-type="bibr" rid="B15">Barbier et al. 2018</xref>). Most existing scientific codes of ethics and conduct tend to focus on employer-employee relationships (e.g., <xref ref-type="bibr" rid="B22">NOAA 2021</xref>), legal obligations (e.g., in conducting experiments on animals), publication ethics (e.g., <xref ref-type="bibr" rid="B1">Albert and Wager 2010</xref>; <xref ref-type="bibr" rid="B7">Anonymous 2019a</xref>), and data ethics and best practices (e.g., <xref ref-type="bibr" rid="B18">Costello and Wieczorek 2014</xref>; <xref ref-type="bibr" rid="B32">Wilkinson et al. 2016</xref>; <xref ref-type="bibr" rid="B25">Moher et al. 2020</xref>; <xref ref-type="bibr" rid="B26">OECD 2021</xref>). Respect for the diversity of people in the scientific community and society (<xref ref-type="bibr" rid="B31">TRUST 2018</xref>; <xref ref-type="bibr" rid="B28">Popp et al. 2020</xref>, e.g., CARE principles regarding Indigenous Peoples) and equity are also highlighted as guiding principles (e.g., <xref ref-type="bibr" rid="B20">Giakoumi et al. 2021</xref>; <xref ref-type="bibr" rid="B29">Research Data Alliance International Indigenous Data Sovereignty Interest Group 2019</xref>). Online resources for teaching good ethical practice have also emerged (e.g., <xref ref-type="bibr" rid="B11">Anonymous 2021a</xref>, <xref ref-type="bibr" rid="B12">2021b</xref>). Many national academies, scientific societies and research programmes have codes of ethics for their members such as guidelines for institutional practice regarding research ethics, promotion and hiring (e.g., <xref ref-type="bibr" rid="B4">Anonymous 2007</xref>, <xref ref-type="bibr" rid="B5">2008</xref>, <xref ref-type="bibr" rid="B8">2019b</xref>, <xref ref-type="bibr" rid="B9">2020a</xref>, <xref ref-type="bibr" rid="B10">2020b</xref>; <xref ref-type="bibr" rid="B2">ALLEA 2017</xref>). More than 2,500 codes and guidelines from over 1,500 organisations around the world, including current and historical codes dating to 1887, have been archived in the Ethics Codes Collection (<xref ref-type="bibr" rid="B11">Anonymous 2021a</xref>, <xref ref-type="bibr" rid="B13">2021c</xref>, <xref ref-type="bibr" rid="B14">2021d</xref>). These codes have been developed through committees at research institutions, national academies, and other organisations. However, most existing codes of scientific conduct have gaps. For example, most do not explicitly provide guidance on how field studies should be conducted (<xref ref-type="bibr" rid="B3">Anonymous 2004</xref>, <xref ref-type="bibr" rid="B9">2020a</xref>; <xref ref-type="bibr" rid="B23">Minteer and Collins 2005a</xref>, <xref ref-type="bibr" rid="B24">2005b</xref>; <xref ref-type="bibr" rid="B17">Costello et al. 2016</xref>; <xref ref-type="bibr" rid="B19">Eichhorn et al. (2020)</xref> discuss the need to work across communities in collaborative ways, publishing data in open access channels and through proactive engagement with local researchers in developing countries before and after any research is conducted together.</p>
      <p>The code presented below originated within the Group on Earth Observations Biodiversity Observation Network (<abbrev xlink:title="Group on Earth Observations Biodiversity Observation Network">GEO BON</abbrev>) community to provide guidance to individual scientists practising in the fields of biodiversity and environmental sciences and was made available on its website to the community (<ext-link xlink:href="https://geobon.org/about-code-of-conduct)-" ext-link-type="uri">https://geobon.org/about-code-of-conduct)-</ext-link> yet it is more widely applicable and so being published here. These guidelines were developed through review of existing literature and by consulting with a wide range of individual scientists representing the diversity of our community around the world to provide perspectives across ages, gender, languages, cultures, and ethnicities. Following a review of existing codes, a draft was circulated within the Marine Biodiversity Observation Network (<abbrev xlink:title="Marine Biodiversity Observation Network">MBON</abbrev>) and the International Association of Biological Oceanography (<abbrev xlink:title="International Association of Biological Oceanography">IABO</abbrev>) for feedback. Following revisions, it was then circulated within <abbrev xlink:title="Group on Earth Observations Biodiversity Observation Network">GEO BON</abbrev>. A finally revised version was then deliberated on by the <abbrev xlink:title="Group on Earth Observations Biodiversity Observation Network">GEO BON</abbrev> Steering Committee before its acceptance. Some feedback suggested expanding particular parts of this code. However, there was consensus that the code should be concise and short to make it more likely to be widely read, digested and applicable to a broad community of scientists. Thus, we encourage users to expand on this code to fit their needs. We don’t claim special moral authority to dictate such a code, nor that this code is definitive. Rather, its value will be judged by who adopts it, and whether and how it affects the practice of science and the well-being of people and nature.</p>
      <p>We hope this code advances changes in behaviour and thought-provoking discussion that move us toward those goals. We encourage organisations and institutions to adopt any and all parts of this code for their purposes, and to help review and update it as appropriate. People building on this code may wish to add consideration of emerging issues, like the use of Artificial Intelligence, as they develop codes for their communities.</p>
      <p>The guidelines below summarise values and principles regarding how we should work with others, engage with society, and how our work and methods impact life and nature. We encourage all to delve into more detailed guidelines developed by others. We start with three overarching principles, focusing on conduct in relation to people, nature, and scientific practice. Further details are divided into personal behaviours, practices, and publication strategies.</p>
    </sec>
    <sec sec-type="Overarching principles: We…" id="sec3">
      <title>Overarching principles: We…</title>
      <list list-type="simple">
        <list-item>
          <p>1. <italic>respect everyone</italic> and accept diversity of age, gender, sexuality, race, origin, religion, ancestry and culture, social and economic status, and physical and mental ability. This includes promoting and facilitating equity in all transactions.
                </p>
        </list-item>
        <list-item>
          <p>2. <italic>respect nature</italic> by practising and promoting sustainable use of natural resources and minimising disturbance of living organisms and natural environments.
                </p>
        </list-item>
        <list-item>
          <p>3. <italic>conduct our science</italic> with honesty and transparency of evidence, data, knowledge, know-how and/or ideas (<xref ref-type="bibr" rid="B30">Resnik and Shamoo 2011</xref>; <xref ref-type="bibr" rid="B6">Anonymous 2013</xref>).
                </p>
        </list-item>
      </list>
    </sec>
    <sec sec-type="Behaviours: We…" id="sec4">
      <title>Behaviours: We…</title>
      <list list-type="simple">
        <list-item>
          <p>4. <italic>take time to be mindful</italic>, considerate, and supportive of others and their circumstances.
                </p>
        </list-item>
        <list-item>
          <p>5. <italic>respect confidentiality, cultural and indigenous sensitivities</italic>, and do not impose ideologies (e.g., political, religious) on others.
                </p>
        </list-item>
        <list-item>
          <p>6. <italic>refrain from and actively oppose discrimination</italic>, harassment, bullying, and misuse of power.
                </p>
        </list-item>
        <list-item>
          <p>7. <italic>respect the spirit and letter of laws</italic>, regulations, agreements, and institutional requirements where these are appropriately designed for the benefit of a broader society, and otherwise work with these institutions for change.
                </p>
        </list-item>
        <list-item>
          <p>8. <italic>practice integrity</italic>, honesty, impartiality, objectivity, and intellectual rigour in our work.
                </p>
        </list-item>
        <list-item>
          <p>9. <italic>are accountable for our research</italic>, including mistakes and oversights in professional practice, publications, peer review, data collection and interpretation.
                </p>
        </list-item>
        <list-item>
          <p>10. <italic>oppose the spread of misinformation</italic> and misleading presentation of evidence, such as selective reporting to conceal evidence of governmental, industrial, scientific, social and/or environmental issues that society should be aware of; and actively correct the public record, including retraction of flawed work.
                </p>
        </list-item>
        <list-item>
          <p>11. <italic>declare conflicts of interest</italic>, both actual and potential, that may be perceived to influence our research and its interpretation.
                </p>
        </list-item>
        <list-item>
          <p>12. <italic>support the profession and colleagues</italic>, such as by contributing as an editor and peer reviewer, organising meetings, mentoring, educating, and sponsoring as opportunities arise, and by supporting those disadvantaged by breaches of these guidelines.
                </p>
        </list-item>
        <list-item>
          <p>13. <italic>acknowledge our backgrounds</italic> and limitations of expertise in discussions and applications.
                </p>
        </list-item>
        <list-item>
          <p>14. <italic>foster the ideals of this Code of Conduct</italic> in others, including training and mentoring staff and students.
                </p>
        </list-item>
      </list>
    </sec>
    <sec sec-type="Practices: We…" id="sec5">
      <title>Practices: We…</title>
      <list list-type="simple">
        <list-item>
          <p>15. <italic>consider the consequences</italic> our behaviour and work have on people and nature, including wildlife, climate change, and the environment, and minimise detrimental impacts.
                </p>
        </list-item>
        <list-item>
          <p>16. <italic>minimise health and safety risks</italic> to ourselves, other people, and nature.
                </p>
        </list-item>
        <list-item>
          <p>17. <italic>prioritise research</italic> methods and practices that have <italic>low impact</italic> on the well-being of people and nature, in the laboratory and natural environment, including non-lethal methods of collecting data on species and their environment.
                </p>
        </list-item>
        <list-item>
          <p>18. <italic>obtain permission</italic> from landowners, local communities, Indigenous People, and any other relevant authorities in any fieldwork.
                </p>
        </list-item>
        <list-item>
          <p>19. <italic>make it clear that people are free to refuse</italic> to participate in particular research. This includes explaining health and safety risks, and the effects the research may have on participants, other people, and the environment.
                </p>
        </list-item>
        <list-item>
          <p>20. <italic>avoid spreading species</italic>, including microbial pathogens, beyond their natural ranges.
                </p>
        </list-item>
        <list-item>
          <p>21. <italic>deposit physical samples where appropriate</italic>, including specimens, in herbaria, museums, or other appropriate national archives with rigorous metadata and all appropriate permits so they are maximally accessible to other users.
                </p>
        </list-item>
      </list>
    </sec>
    <sec sec-type="Publication: We…" id="sec6">
      <title>Publication: We…</title>
      <list list-type="simple">
        <list-item>
          <p>22. <italic>publish research findings and supporting primary data</italic> in an accurate, timely manner to contribute to scientific knowledge, and enable transparency, reproducibility, and replicability of the work.
                </p>
        </list-item>
        <list-item>
          <p>23. <italic>acknowledge people</italic> who have helped our research, including funding sources, and cite prior work and ideas which our work incorporates.
                </p>
        </list-item>
        <list-item>
          <p>24. <italic>respect the intellectual and material property of others</italic>. Where we use data from third parties, we seek their permission to use and release data as appropriate.
                </p>
        </list-item>
        <list-item>
          <p>25. <italic>respect the sensitivity of data</italic> that may enable the identification of individual people, or aid criminal or unethical behaviour; such as, collection of threatened species, spread of pests, or infringement of others’ patents or copyrights.
                </p>
        </list-item>
        <list-item>
          <p>26. <italic>support publication</italic> of ethical scientific research, as an editor or reviewer, regardless of whether it conflicts with our beliefs or interests, or those of our employers or group consensus.
                </p>
        </list-item>
        <list-item>
          <p>27. <italic>never fabricate or falsify research</italic> findings, plagiarise the work of others, or condone such misconduct by others.
                </p>
        </list-item>
        <list-item>
          <p>28. <italic>follow scholarly publishing guidelines</italic> for authorship and acknowledgements in scientific research papers (e.g., the “Vancouver Guidelines” <xref ref-type="bibr" rid="B7">Anonymous 2019a</xref>; <xref ref-type="bibr" rid="B27">Parker and Berman 1998</xref>; <xref ref-type="bibr" rid="B16">CASRAI 2021</xref>).
                </p>
        </list-item>
      </list>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>This document originated from discussion within the Marine Biodiversity Observation Network (<abbrev xlink:title="Marine Biodiversity Observation Network">MBON</abbrev>) of the Group on Earth Observations (<abbrev xlink:title="Group on Earth Observations">GEO</abbrev>). We thank Tri Arfianti, Michèle Barbier, Eeshan Bhatt, Gilianne Brodie, Chhaya Chaudhary, Carlos Duarte, Emmett Duffy, Graham Edgar, Jens Kattge, Áine Kelly-Costello, Liam Kelly-Costello, Suzan Kholeif, Daniel Lauretta, Yvan Le Bras, Shobha Maharaj, Edem Mahu, Joana Soares, Jonathan Quetzal Tritter, Gautam Hirak Talukdar, ‘Aulani Wilhelm, Dawn Wright, and members of the Executive Committees of the Scientific Committee on Ocean Research, International Association of Biological Oceanography (<abbrev xlink:title="International Association of Biological Oceanography">IABO</abbrev>), Marine Biodiversity Observation Network (<abbrev xlink:title="Marine Biodiversity Observation Network">MBON</abbrev>), and Group on Earth Observations Biodiversity Observation Network (<abbrev xlink:title="Group on Earth Observations Biodiversity Observation Network">GEO BON</abbrev>) for helpful discussion and comments that improved the original draft. We thank Mark Lomolino, an anonymous referee and the editor for helpful comments on the paper. This code has been endorsed by <abbrev xlink:title="Marine Biodiversity Observation Network">MBON</abbrev>, <abbrev xlink:title="International Association of Biological Oceanography">IABO</abbrev>, and <abbrev xlink:title="Group on Earth Observations Biodiversity Observation Network">GEO BON</abbrev>.</p>
    </ack>
    <ref-list>
      <title>References</title>
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      </ref>
      <ref id="B2">
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      </ref>
      <ref id="B3">
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    <sec sec-type="Additional information" id="sec7">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec8">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec9">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec10">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec11">
        <title>Funding</title>
        <p>No funding was reported.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec12">
        <title>Author contributions</title>
        <p>Conceptualization: MJC, AEB, FEMK. Investigation: MJC. Methodology: MJC, AEB, FEMK. Project administration: MJC. Writing – original draft: MJC. Writing – review and editing: FEMK, AEB, MJC.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec13">
        <title>Author ORCIDs</title>
        <p>M.J. Costello <ext-link xlink:href="https://orcid.org/0000-0003-2362-0328" ext-link-type="uri">https://orcid.org/0000-0003-2362-0328</ext-link></p>
        <p>F.E. Muller-Karger <ext-link xlink:href="https://orcid.org/0000-0003-3159-5011" ext-link-type="uri">https://orcid.org/0000-0003-3159-5011</ext-link></p>
        <p>A.E. Bates <ext-link xlink:href="https://orcid.org/0000-0002-0198-4537" ext-link-type="uri">https://orcid.org/0000-0002-0198-4537</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec14">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
  </back>
</article>
