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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.19.204479</article-id>
      <article-id pub-id-type="publisher-id">204479</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Review Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Actinopterygii</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biogeography</subject>
          <subject>Marine</subject>
          <subject>Phylogenetic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Pacific Ocean</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>The historical biogeography of marine ray-finned fishes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class" reg="Actinopterygii">Actinopterygii</tp:taxon-name-part></tp:taxon-name>) in the Southwest Pacific: species richness and endemism patterns</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Samayoa</surname>
            <given-names>André P.</given-names>
          </name>
          <email xlink:type="simple">apsbonin@hotmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-3852-7935</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/formal-analysis/">Formal analysis</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>Liggins</surname>
            <given-names>Libby</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1143-2346</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <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/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">School of Natural Sciences, Massey University, Auckland 0745, New Zealand</addr-line>
        <institution>Natural Sciences, Auckland Museum Tāmaki Paenga Hira</institution>
        <addr-line content-type="city">Auckland</addr-line>
        <country>New Zealand</country>
        <uri content-type="ror">https://ror.org/010na6b76</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">School of Biological Sciences, University of Auckland, Auckland 1010, New Zealand</addr-line>
        <institution>School of Biological Sciences, University of Auckland</institution>
        <addr-line content-type="city">Auckland</addr-line>
        <country>New Zealand</country>
        <uri content-type="ror">https://ror.org/03b94tp07</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Natural Sciences, Auckland Museum Tāmaki Paenga Hira, Auckland 1010, New Zealand</addr-line>
        <institution>School of Natural Sciences, Massey University</institution>
        <addr-line content-type="city">Auckland</addr-line>
        <country>New Zealand</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: André P. Samayoa (<email xlink:type="simple">apsbonin@hotmail.com</email>)</p>
        </fn>
        <fn>
          <p><bold>Editors</bold> Christopher Burridge and Sérgio Ávila</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>12</day>
        <month>08</month>
        <year>2026</year>
      </pub-date>
      <volume>19</volume>
      <elocation-id>e204479</elocation-id>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/14B8FAE5-9616-5DDF-A9BC-CBB9577A2126">14B8FAE5-9616-5DDF-A9BC-CBB9577A2126</uri>
      <history>
        <date date-type="received">
          <day>18</day>
          <month>06</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>03</day>
          <month>08</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>André P. Samayoa, Libby Liggins</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>Inferring how biodiversity is generated and how it responded to past events is fundamental to understanding how biodiversity may respond to future changes. As the largest ubiquitous taxonomic group amongst marine vertebrates, ray-finned fishes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class" reg="Actinopterygii">Actinopterygii</tp:taxon-name-part></tp:taxon-name>) represent a good model to understand the generation of biodiversity and the processes that shaped contemporary biogeographic patterns in the sea. Evidence shows that hotspots of marine fish richness and endemism do not always overlap, due to the distinct processes that contribute to them. In the Pacific Ocean, the tropical western region concentrates the highest records of species richness. In contrast, the peripheral oceanic islands are major endemism centres. Although the evolutionary setting for marine fish endemism in parts of the Central Pacific has been described, processes that generate and maintain biodiversity in other peripheral regions remain unknown. The Southwest Pacific (20°S-55°S and 150°E-165°W) spans subtropical, temperate and subantarctic climates, containing major landmasses such as eastern Australia and New Zealand, as well as oceanic islands. Due to its configuration, the region displays unique biogeographic patterns that can improve our understanding of marine biodiversity evolution. Here, we aim to address this topic by reviewing the latest biogeographic investigations that examine the origin, evolution, and processes shaping patterns of marine fish richness and endemism in the Southwest Pacific. We highlight the utility of specific methodological approaches used in previous investigations, including the inference of molecular phylogenies based on fossil data, probabilistic biogeographic modelling and statistical analysis of phylogenetic measures of richness and endemism. We propose a biogeographic scenario where the subtropical islands of the Southwest Pacific act as biodiversity cradles, with mainland Australia representing the main source of endemic lineages. Empirical data highlight the significance of both rare colonisation events and vicariance within formerly widely distributed lineages in shaping endemism patterns and reveal that the processes shaping patterns of endemism and richness differ at local scales.</p>
        <sec sec-type="Highlights" id="sec1">
          <title>Highlights</title>
          <boxed-text position="float">
            <list list-type="bullet">
              <list-item>
                <p>The Southwest Pacific is a geologically active region that harbours multiple oceanic islands of volcanic origin.</p>
              </list-item>
              <list-item>
                <p>Hotspots of marine fish richness are not geographically concordant with endemism centres along New Zealand, with richness peaking north of the North Island of Aotearoa New Zealand.</p>
              </list-item>
              <list-item>
                <p>Heightened richness in these locations likely originates from an overlap of fish fauna that includes older, distantly related taxa inhabiting warmer waters in the north and closely-related marine fish taxa in colder latitudes in the south.</p>
              </list-item>
              <list-item>
                <p>Within the Southwest Pacific, the subtropical oceanic islands of Lord Howe, Norfolk and Rangitāhua (Kermadec Islands) are characterised by the highest endemism rates in coastal marine ray-finned fishes.</p>
              </list-item>
              <list-item>
                <p>The geographic isolation and unique oceanic, subtropical climate of these islands have likely influenced the establishment of endemism.</p>
              </list-item>
            </list>
          </boxed-text>
        </sec>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>Biodiversity</kwd>
        <kwd>biogeography</kwd>
        <kwd>endemism</kwd>
        <kwd>evolution</kwd>
        <kwd>marine ray-finned fishes</kwd>
        <kwd>molecular phylogenetics</kwd>
        <kwd>Southwest Pacific</kwd>
        <kwd>species richness</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec2">
      <title>Introduction</title>
      <p>Marine organisms are not uniformly distributed in the ocean realm, a phenomenon resulting from the effect of evolutionary and ecological mechanisms (<xref ref-type="bibr" rid="B17">Costello and Chaudhary 2017</xref>). Biogeography studies the distribution of living organisms and the mechanisms influencing their distribution, often aiming to identify “hotspots” - geographic locations with high levels of biodiversity - given the insight they can provide regarding the processes that generate and maintain biodiversity, as well as the value in protecting such places in the face of the ongoing biodiversity loss (<xref ref-type="bibr" rid="B90">Norman 2003</xref>). We currently have an advanced understanding of the geographic distribution of the most diverse terrestrial (<xref ref-type="bibr" rid="B85">Myers et al. 2000</xref>) and marine (<xref ref-type="bibr" rid="B105">Roberts et al. 2002</xref>) regions, as well as hotspots within these regions that can be defined either by their number of total species (<xref ref-type="bibr" rid="B103">Reid 1998</xref>) or of range-restricted species only (<xref ref-type="bibr" rid="B84">Myers 1990</xref>).</p>
      <p>Ray-finned fishes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class" reg="Actinopterygii">Actinopterygii</tp:taxon-name-part></tp:taxon-name>) belong to one of the largest taxonomic groups in the animal kingdom with over 36,900 valid species and 5,283 valid genera (<xref ref-type="bibr" rid="B40">Fricke et al. 2024</xref>). Actinopterygians are evenly distributed between freshwater and marine environments (<xref ref-type="bibr" rid="B115">Seehausen and Wagner 2014</xref>), with marine ray-finned fishes (hereby “marine fishes”) spreading from tropical to polar latitudes (<xref ref-type="bibr" rid="B64">Lin et al. 2021</xref>) and reaching depths up to 8,336 m (<xref ref-type="bibr" rid="B54">Jamieson et al. 2023</xref>). Marine fishes are fundamental for ecosystem function, occupying a wide array of trophic positions and dietary niches (<xref ref-type="bibr" rid="B48">Hayden et al. 2019</xref>) and for human consumption (<xref ref-type="bibr" rid="B86">Natale et al. 2013</xref>). Given that highly diverse marine fish communities seem more resilient to current climatic pressures (<xref ref-type="bibr" rid="B30">Duffy et al. 2016</xref>), marine fishes represent an interesting model to understand how marine biodiversity originates and persists and how future biodiversity patterns might be determined.</p>
      <p>In the face of biodiversity loss in the sea, intense anthropogenic pressure on fish communities and the immediate changes in biodiversity patterns due to climate change, it is imperative to keep paying attention to the spatial patterns of marine fishes and to understand the causal processes that have played a role in their varied distributions (<xref ref-type="bibr" rid="B22">Cowman et al. 2017</xref>). The Pacific Ocean, the world’s largest marine basin, is an actively studied area that includes regions of distinct biogeographic features (<xref ref-type="bibr" rid="B119">Springer 1982</xref>; <xref ref-type="bibr" rid="B118">Spalding et al. 2007</xref>; <xref ref-type="bibr" rid="B58">Kulbicki et al. 2013</xref>; <xref ref-type="bibr" rid="B6">Bowen et al. 2016</xref>). While the western tropical region that sits between the Indian and the Pacific Oceans represents a major area for biogeographic research (<xref ref-type="bibr" rid="B65">Lohman et al. 2011</xref>), few investigations have focused on the Southwest Pacific, a region characterised by a mixture of major landmasses and oceanic islands that cross subtropical, temperate and subantarctic latitudes (<xref ref-type="bibr" rid="B73">McClymont et al. 2016</xref>). As evidence shows that climate change drives latitudinal, distributional shifts of marine biodiversity (<xref ref-type="bibr" rid="B95">Poloczanska et al. 2016</xref>), a potentially higher climatic vulnerability and significant distributional changes for marine organisms are expected in regions that span multiple climatic latitudes, such as the Southwest Pacific.</p>
      <p>This review summarises the state of knowledge for the biogeography of shallow (up to 500 metres) marine ray-finned fishes in the Southwest Pacific, based on the latest contributions of historical biogeographic studies in the region. Here, (i) we describe general patterns of species richness and endemism for marine biodiversity; (ii) we provide a concise review of the biogeographic patterns and the search for underlying historical causes for actinopterygians in the broader Pacific Ocean; (iii) we present the major geological, oceanographic and biological features of the Southwest Pacific; (iv) we outline the emerging biogeographic setting for marine actinopterygians in the region, based on recent studies and (v) we propose future lines of work to fill current knowledge gaps towards a better understanding of the geographic distribution of marine fish biodiversity in the Southwest Pacific.</p>
    </sec>
    <sec sec-type="Key biogeographic patterns of marine biodiversity in the Pacific Ocean" id="sec3">
      <title>Key biogeographic patterns of marine biodiversity in the Pacific Ocean</title>
      <sec sec-type="Theoretical overview of patterns" id="sec4">
        <title>Theoretical overview of patterns</title>
        <p>Historically, biogeographers have focused on two major distributional patterns in the sea, areas of high species richness and endemism (<xref ref-type="bibr" rid="B17">Costello and Chaudhary 2017</xref>). In the marine environment, cross-taxon species richness concentrates in tropical ecosystems, peaks at the edges of warm waters and dips at the equator following a bimodal latitudinal distribution (<xref ref-type="bibr" rid="B14">Chaudhary et al. 2016</xref>). This latitudinal gradient seems shaped by the strong influence of temperature, as shown by the symmetrical thermal gradient along latitude (<xref ref-type="bibr" rid="B15">Chaudhary et al. 2017</xref>) and the ongoing poleward range shifts of marine species caused by equatorial warming conditions (<xref ref-type="bibr" rid="B16">Chaudhary et al. 2021</xref>). Within tropical ecosystems, it is largely accepted that the diversity of marine coastal species is higher at the intersection of the East Indian and West Pacific Oceans (<xref ref-type="bibr" rid="B120">Tittensor et al. 2010</xref>), a region known as the Indo-Australian Archipelago of long-standing interest for biogeographers (<xref ref-type="bibr" rid="B65">Lohman et al. 2011</xref>). There are three major hypotheses regarding the mechanisms underpinning this tropical hotspot (<xref ref-type="bibr" rid="B4">Bellwood et al. 2012</xref>): the Centre of Origin – species originate in the tropical Indo-Pacific and spread outwards (<xref ref-type="bibr" rid="B7">Briggs 2003</xref>); the Centre of Overlap – the Indo-Pacific is the meeting area where the ranges of nearby species overlap (<xref ref-type="bibr" rid="B128">Woodland 1983</xref>); and the Centre of Accumulation/Survival – species originate at the periphery of the Indo-Pacific and spread inwards (<xref ref-type="bibr" rid="B59">Ladd 1960</xref>; <xref ref-type="bibr" rid="B1">Barber and Bellwood 2005</xref>). A review of historical biogeographic data suggests that the processes referred to in each hypothesis have interacted over multiple spatio-temporal scales to generate the biodiversity hotspot (<xref ref-type="bibr" rid="B20">Cowman 2014</xref>).</p>
        <p>Centres of marine endemism concentrate species found nowhere else (<xref ref-type="bibr" rid="B85">Myers et al. 2000</xref>) and are of conservation interest due to the higher extinction risks of range-restricted species (<xref ref-type="bibr" rid="B69">Manes et al. 2021</xref>), but also for evolutionary biologists, given that endemism sheds light on processes generating biodiversity (<xref ref-type="bibr" rid="B3">Bellwood and Meyer 2009</xref>). In marine (<xref ref-type="bibr" rid="B22">Cowman et al. 2017</xref>) and terrestrial (<xref ref-type="bibr" rid="B91">Orme et al. 2005</xref>) species, evidence shows that hotspots of species richness and endemism do not always overlap, an indication that these patterns originate from different processes. Centres of endemism are often located in oceanic islands characterised by relatively low species richness (<xref ref-type="bibr" rid="B57">Kier et al. 2009</xref>), a pattern explained by models that derive from early island biogeographic theory that predicts species richness as a function of immigration, speciation and extinction (<xref ref-type="bibr" rid="B67">MacArthur and Wilson 1963</xref>, <xref ref-type="bibr" rid="B68">1967</xref>). Despite numerous oceanic island models proposed by theoreticians and empiricists, most models follow the principle that island endemism arises from highly dispersing colonisers, subsequent diversification events driven by geographic and/or ecological isolation and eventual increase in endemism richness (<xref ref-type="bibr" rid="B125">Whittaker et al. 2008</xref>, <xref ref-type="bibr" rid="B126">2010</xref>; <xref ref-type="bibr" rid="B47">Hachich et al. 2015</xref>; <xref ref-type="bibr" rid="B26">Dawson 2016</xref>; <xref ref-type="bibr" rid="B94">Pinheiro et al. 2017</xref>). This theoretical framework brings together two processes belonging to two traditionally opposed views as to the main mechanism shaping biogeographic patterns (<xref ref-type="bibr" rid="B127">Wiens and Donoghue 2004</xref>): rare long-distance dispersal (jump-dispersal) and the geographic separation of two populations by a land or marine barrier (vicariance; see <xref ref-type="bibr" rid="B21">Cowman and Bellwood (2013)</xref> for a review of barriers in the sea). In the last few decades, methodological progress has generated empirical data that supports the interaction of both processes over time in the generation of contemporary biogeographic patterns (<xref ref-type="bibr" rid="B113">Sanmartín 2012</xref>), demonstrating how technological advances facilitate the study of historical processes that shape the distribution of marine organisms, contributing to the expansion of biogeographic theory.</p>
      </sec>
      <sec sec-type="Searching for historical processes acting in the Pacific Ocean" id="sec5">
        <title>Searching for historical processes acting in the Pacific Ocean</title>
        <p>The Pacific Ocean is the largest body of seawater on Earth and includes part of the Indo-Australian Archipelago which hosts the highest diversity of coastal marine fishes in the world (<xref ref-type="bibr" rid="B20">Cowman 2014</xref>). The biogeography of marine fishes in the Pacific is characterised by high species richness in the western tropical “bullseye” of which a relatively small proportion are endemic species and low species richness, but relatively high endemism in the oceanic islands at the periphery of the richness hotspot (<xref ref-type="bibr" rid="B22">Cowman et al. 2017</xref>). Modern works investigating the historical biogeography of marine fishes (<xref ref-type="bibr" rid="B93">Pellissier et al. 2014</xref>; <xref ref-type="bibr" rid="B117">Siqueira et al. 2016</xref>; <xref ref-type="bibr" rid="B42">Gaboriau et al. 2018</xref>; <xref ref-type="bibr" rid="B77">Miller et al. 2018</xref>) suggest that the geographic distribution of actinopterygians in the Indo-Pacific emerged from: the stability of tropical ecosystems during glaciations within the last 34 million years ago (Ma), providing sufficient time for speciation and persistence of old lineages over time; elevated diversification driven by the high topographic variability of reef habitats; the interaction of mechanisms invoked in each of the three “Centre-of” hypotheses (<xref ref-type="bibr" rid="B4">Bellwood et al. 2012</xref>) over distinct geological timings; and the contribution of peripheral regions in generating and maintaining biodiversity (<xref ref-type="bibr" rid="B5">Bowen et al. 2013</xref>).</p>
        <p>The significant role that peripheral oceanic islands play in the dynamics of marine fish biodiversity is explicitly acknowledged in the “biodiversity feedback” model (<xref ref-type="bibr" rid="B5">Bowen et al. 2013</xref>), a biogeographic hypothesis in which the tropical Indo-Pacific exports biodiversity, but also imports biodiversity produced in peripheral islands (Fig. <xref ref-type="fig" rid="F1">1</xref>). More regional evidence supports the model by highlighting the role of peripheral islands as engines for biodiversity origination, as reported for Hawaii (<xref ref-type="bibr" rid="B51">Hodge et al. 2014</xref>) and Easter Island (Rapa Nui) (<xref ref-type="bibr" rid="B29">Delrieu‐Trottin et al. 2019</xref>). In both cases, biogeographic analyses suggest that the conjunct action of vicariant and colonisation events shaped contemporary patterns of insular biodiversity and endemism, a conclusion congruent with the current historical biogeographic philosophy that recognises the relevance of both vicariant and dispersal arguments in the explanation of biogeographic patterns (<xref ref-type="bibr" rid="B113">Sanmartín 2012</xref>). However, despite significant progress in understanding the role of oceanic islands in shaping spatial patterns of marine fish biodiversity, many remote islands that lie at the periphery of the Pacific Ocean remain understudied by biogeographers.</p>
        <fig id="F1">
          <object-id content-type="doi">10.21425/fob.19.204479.figure1</object-id>
          <object-id content-type="arpha">9829CC53-04ED-5D0C-ABD7-30C79E83EDCB</object-id>
          <label>Figure 1.</label>
          <caption>
            <p>Map of the Pacific Ocean showing the position of the Southwest Pacific (green square) defined in this review by latitudes 20°-55°S and longitudes 150°E-165°W as in <xref ref-type="bibr" rid="B110">Samayoa et al. (2022)</xref>. The red area depicts the Indo-Australian Archipelago (IAA), based on <xref ref-type="bibr" rid="B5">Bowen et al. (2013)</xref>. Blue triangles indicate the location of volcanic islands mentioned in the text and that lie at the periphery of the IAA: Lord Howe Island (1); Norfolk Island (2); Rangitāhua/Kermadec Islands (3); Hawaii (4); Marquesas Islands (5); Rapa Nui/Easter Island (6); and Juan Fernández and Desventuradas (7). Arrows illustrate the “biodiversity feedback” model from <xref ref-type="bibr" rid="B5">Bowen et al. (2013)</xref> where the IAA exports biodiversity into the Pacific Ocean (black arrows) and intakes biodiversity generated in peripheral islands (blue arrows). Outgoing blue arrows are positioned in islands where phylogenetic evidence supports the generation of young endemic lineages postdating the emergence of islands (neoendemism): Samayoa et al. (<xref ref-type="bibr" rid="B110">2022</xref>, <xref ref-type="bibr" rid="B111">2023</xref>) for the Southwest Pacific islands (1, 2 and 3); <xref ref-type="bibr" rid="B51">Hodge et al. (2014)</xref> for Hawaii (4); and <xref ref-type="bibr" rid="B28">Delrieu-Trottin et al. (2018)</xref> for Rapa Nui (6). Map projection: WGS84.</p>
          </caption>
          <graphic xlink:href="biogeography-19-001_article-204479__-g001.jpg" id="oo_1744159.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1744159</uri>
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        </fig>
      </sec>
    </sec>
    <sec sec-type="The Southwest Pacific: geological, oceanographic and biological features" id="sec6">
      <title>The Southwest Pacific: geological, oceanographic and biological features</title>
      <p>In this review, the “Southwest Pacific” is defined as the south-western quadrant of the Pacific Ocean, bounded by latitudes 20°S-55°S and longitudes 150°E-165°W (Fig. <xref ref-type="fig" rid="F1">1</xref>). It is a geologically active region (<xref ref-type="bibr" rid="B87">Neall and Trewick 2008</xref>), characterised by two continental fragments of Gondwanan origin, Australia and Zealandia and by the Australian-Pacific plate boundary that crosses Aotearoa New Zealand between the North and South Islands. Consequently, major landmasses include the continental islands of mainland Australia (eastern segment) and the aerial sections of Zealandia that comprise New Zealand (North Island, South Island, Stewart Island, the Chatham Islands and several smaller islands) and New Caledonia (<xref ref-type="bibr" rid="B121">Trewick et al. 2007</xref>) (Fig. <xref ref-type="fig" rid="F2">2</xref>). Additionally, the Southwest Pacific harbours recently emerged oceanic islands of volcanic origin, with estimated ages of 23.8–1 Ma for subantarctic islands (<xref ref-type="bibr" rid="B98">Quilty 2007</xref>; <xref ref-type="bibr" rid="B114">Scott and Turnbull 2019</xref>) and 6.9–2.58 Ma for subtropical islands (<xref ref-type="bibr" rid="B55">Jones and McDougall 1973</xref>; <xref ref-type="bibr" rid="B74">McDougall et al. 1981</xref>; <xref ref-type="bibr" rid="B11">Brook 1998b</xref>). Most outlying islands sit on Zealandia (<xref ref-type="bibr" rid="B80">Mortimer et al. 2017</xref>), except the Kermadec Archipelago/Rangitāhua on the Kermadec Ridge (<xref ref-type="bibr" rid="B11">Brook 1998b</xref>) and Macquarie Island on the Macquarie Ridge (<xref ref-type="bibr" rid="B98">Quilty 2007</xref>). Politically, Australia administers the subtropical Lord Howe Island and Norfolk Island and the subantarctic Macquarie Island. New Zealand’s outlying territories include subtropical Rangitāhua, the eastern Chatham Islands/Rēkohu/Wharekauri, as well as the subantarctic Bounty Islands/Moutere Hauriri, Antipodes Islands/Moutere Mahue, Snares Islands/Tini Heke, Auckland Islands/Motu Maha and Campbell Island/Motu Ihupuku.</p>
      <fig id="F2">
        <object-id content-type="doi">10.21425/fob.19.204479.figure2</object-id>
        <object-id content-type="arpha">598114C6-52DA-597B-896A-3422A871AF92</object-id>
        <label>Figure 2.</label>
        <caption>
          <p>Map of the Southwest Pacific showing the geographic location of the subtropical islands of Lord Howe Island (1), Norfolk Island (2) and Rangitāhua/Kermadec Islands (3). Major oceanic currents are illustrated as arrows: red represents warm water flow from the East Australian Current (EAC), the Tasman Front (TF) and the East Auckland Current (EAuC); blue represents cold water flow from the Subantarctic Front (SAF); and purple represents mid-temperature water flow from the Subtropical Front (STF). The inner area from the dashed line belongs to the almost submerged continental crust of Zealandia. Map projection: WGS84.</p>
        </caption>
        <graphic xlink:href="biogeography-19-001_article-204479__-g002.jpg" id="oo_1744160.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1744160</uri>
        </graphic>
      </fig>
      <p>Surface currents in the Southwest Pacific follow the anti-clockwise movement of the South Pacific Gyre (<xref ref-type="bibr" rid="B43">Ganachaud et al. 2014</xref>): the main flow moves west at low latitudes as the South Equatorial Current, reaches mainland Australia through the Coral Sea, becomes the East Australian Current at 15°S to then flow southwards along the eastern Australian coastline (Fig. <xref ref-type="fig" rid="F2">2</xref>), branches eastwards between 32°S-34°S as the Tasman Front, crosses Norfolk and reaches the North Island of New Zealand, becoming the East Auckland Current. Additionally, two independent currents from the East Australian Current flow east across the high latitudes of the Southwest Pacific (<xref ref-type="bibr" rid="B88">Nelson and Cooke 2001</xref>): the Subtropical Front (ca. 43°S-45°S) that follows a convoluted path around the southern tip of New Zealand’s South Island to then reach the Chatham Islands; and the Subantarctic Front (ca. 52°S) that flows around the southernmost continental segment of Zealandia to meet the Subtropical Front south of the Chathams. The three major eastward currents of the Southwest Pacific are distinguished by their physicochemical characteristics (<xref ref-type="bibr" rid="B66">Lorrey et al. 2012</xref>): the Tasman Front carries highly saline warm water which is poor in nutrients; the Subantarctic Front is a nutrient-rich, cold-water mass with lower salinity; and the Subtropical Front separates the Tasman and Subantarctic fronts.</p>
      <p>The Southwest Pacific hosts an exceptionally rich marine biodiversity resulting from the high species richness and endemism in New Zealand and to a lesser extent, Australia (<xref ref-type="bibr" rid="B8">Briggs and Bowen 2012</xref>; <xref ref-type="bibr" rid="B19">Costello et al. 2017</xref>). Both countries are considered the best taxonomically studied regions in the West Pacific (<xref ref-type="bibr" rid="B18">Costello et al. 2010</xref>), exemplified by the elevated completeness of New Zealand’s inventory of marine species (<xref ref-type="bibr" rid="B79">Mora et al. 2008</xref>) which include 17,135 taxa and 4,315 undescribed species according to <xref ref-type="bibr" rid="B46">Gordon et al. (2010)</xref>. For fish diversity, <xref ref-type="bibr" rid="B104">Roberts et al. (2020)</xref> report 1,296 species in New Zealand, of which 1,085 are marine actinopterygians. As high marine fish biodiversity seems linked to habitat complexity (<xref ref-type="bibr" rid="B62">Leprieur et al. 2016</xref>), New Zealand’s marine richness might result from the extensive Zealandian continental shelf which likely provides abundant habitats for diversification (<xref ref-type="bibr" rid="B46">Gordon et al. 2010</xref>). Within the Southwest Pacific, taxonomic data point to the subtropical islands of Lord Howe, Norfolk and Rangitāhua as centres of endemism, with an overall 4.6% endemism rate for coastal marine fishes (<xref ref-type="bibr" rid="B37">Francis 1993</xref>), although more recent surveys report the same figure for Rangitāhua alone (<xref ref-type="bibr" rid="B123">Trnski et al. 2015</xref>). The marine fish communities in the three island groups are a mixture of tropical, subtropical and temperate species (<xref ref-type="bibr" rid="B38">Francis and Duffy 2015</xref>), a similarity likely explained by the predominant eastward oceanic currents of the region (<xref ref-type="bibr" rid="B122">Trnski and de Lange 2015</xref>). The recent compilation of marine fish taxa occurring in Lord Howe Island, Norfolk Island and Rangitāhua (<xref ref-type="bibr" rid="B39">Francis and Hoese 2025</xref>) illustrates the significance of marine fish biodiversity around the islands, as well as the relevance of distributional data to understand biogeographic patterns in the region.</p>
      <p>Overall, the contemporary spatial patterns of marine fish biodiversity and endemism are broadly understood in the Southwest Pacific. However, the evolutionary value of regional actinopterygians and key biogeographic locations, have just started to be revealed (<xref ref-type="bibr" rid="B28">Delrieu-Trottin et al. 2018</xref>; <xref ref-type="bibr" rid="B33">Eme et al. 2020</xref>; <xref ref-type="bibr" rid="B63">Liggins et al. 2022</xref>). A comprehensive historical framework is needed to inform biogeographers, evolutionary biologists, ecologists and conservationists about the relevance of the processes shaping the biogeography of marine fishes in the Southwest Pacific, if we are to determine how marine biodiversity will evolve in times of heavy biodiversity loss and intense anthropogenic pressure.</p>
    </sec>
    <sec sec-type="Historical biogeography of marine fishes in the Southwest Pacific" id="sec7">
      <title>Historical biogeography of marine fishes in the Southwest Pacific</title>
      <sec sec-type="Why historical biogeography?" id="sec8">
        <title>Why historical biogeography?</title>
        <p>Historical biogeography focuses on evolutionary processes (e.g. speciation, extinction) over geological time to explain the geographic distribution of organisms through time and space (<xref ref-type="bibr" rid="B23">Crisci 2001</xref>), requiring a sound understanding of the evolutionary history of biodiversity and contemporary range information (<xref ref-type="bibr" rid="B107">Ronquist and Sanmartín 2011</xref>). The technological innovations of the 20<sup>th</sup> century have improved methods to infer the evolutionary trajectories of living organisms using molecular sequences, resulting in the substantial integration of phylogenetic trees in historical biogeographic methods (<xref ref-type="bibr" rid="B96">Posadas et al. 2006</xref>). For marine biodiversity, the incorporation of range information has been slow, given the sampling bias towards terrestrial ecosystems (<xref ref-type="bibr" rid="B53">Hughes et al. 2021</xref>) and coastal areas (<xref ref-type="bibr" rid="B52">Hortal et al. 2015</xref>), leaving vast open-water regions unsampled. To fill this gap, recent modelling techniques generate probabilistic maps for benthic and pelagic taxa (<xref ref-type="bibr" rid="B56">Kaschner et al. 2019</xref>) to gain marine range information, combining environmental variables with occurrence data from web-based repositories (e.g. Ocean Biogeographic Information System, <ext-link ext-link-type="uri" xlink:href="http://www.obis.org">www.obis.org</ext-link>; and Global Biodiversity Information Facility, <ext-link ext-link-type="uri" xlink:href="http://www.gbif.org">www.gbif.org</ext-link>).</p>
        <p>Phylogenetic and range information are the basis of countless historical biogeographic methods (<xref ref-type="bibr" rid="B23">Crisci 2001</xref>), which, until recent decades, excluded ecological explanations (e.g. dispersal ability, competition) for contemporary biogeographic patterns. To address calls to unify historical and ecological perspectives (<xref ref-type="bibr" rid="B32">Ebach 2004</xref>; <xref ref-type="bibr" rid="B127">Wiens and Donoghue 2004</xref>; <xref ref-type="bibr" rid="B25">Crisp et al. 2011</xref>), more recent approaches (e.g. <xref ref-type="bibr" rid="B70">Matzke (2013)</xref>) integrate evolutionary and ecological processes, leading to a more dynamic methodological approach in biogeographic research (<xref ref-type="bibr" rid="B113">Sanmartín 2012</xref>). In this review, we acknowledge the plurality of historical biogeographic methods, referring the reader to the reviews by <xref ref-type="bibr" rid="B23">Crisci (2001)</xref>, <xref ref-type="bibr" rid="B96">Posadas et al. (2006)</xref> and <xref ref-type="bibr" rid="B113">Sanmartín (2012)</xref> for detailed information on each methodological approach. Hereby, we refer to methods and tools relevant for the inference of historical processes shaping contemporary patterns of marine fish richness and endemism in the Southwest Pacific.</p>
      </sec>
      <sec sec-type="Richness hotspots" id="sec9">
        <title>Richness hotspots</title>
        <p>A recent phylogenetic study examined the biogeography of New Zealand’s marine actinopterygians across a marine section (25°S-55°S, 160°E-170°W) of the Southwest Pacific, finding that richness hotspots and endemism centres do not co-occur geographically at the study’s spatial scale (<xref ref-type="bibr" rid="B112">Samayoa et al. 2025</xref>; Fig. <xref ref-type="fig" rid="F3">3</xref>), a similar biogeographic pattern observed in the same taxonomic group at the larger Indo-Pacific scale (<xref ref-type="bibr" rid="B20">Cowman 2014</xref>; <xref ref-type="bibr" rid="B22">Cowman et al. 2017</xref>). To characterise the biogeography of marine fishes, the investigators used phylogenetic metrics (see <xref ref-type="bibr" rid="B124">Tucker et al. (2017)</xref> for a thorough description of each metric), such as “phylogenetic diversity”, the sum of branch lengths within a phylogenetic tree spanning all occurring taxa at a given location (<xref ref-type="bibr" rid="B34">Faith 1992</xref>) and “phylogenetic endemism”, the sum of branch lengths spanning taxa present in an assemblage, weighted by the range of the clade that includes those taxa (<xref ref-type="bibr" rid="B108">Rosauer et al. 2009</xref>). Phylogenetic metrics are developed to measure evolutionary differences between lineages within a phylogenetic tree, translating historical aspects of biodiversity (<xref ref-type="bibr" rid="B124">Tucker et al. 2017</xref>). Contrasting the spatial distribution of phylogenetic measures of diversity and endemism with species richness is a proven approach to infer the evolutionary and ecological processes shaping richness hotspots (<xref ref-type="bibr" rid="B13">Cadotte et al. 2010</xref>; <xref ref-type="bibr" rid="B92">Pavoine and Bonsall 2011</xref>).</p>
        <fig id="F3">
          <object-id content-type="doi">10.21425/fob.19.204479.figure3</object-id>
          <object-id content-type="arpha">2F29D618-ADC2-58EC-A6D5-4A6747728C6E</object-id>
          <label>Figure 3.</label>
          <caption>
            <p>The larger map illustrates the estimated geographic origin (coloured circles) of marine fish lineages endemic to the subtropical islands of Lord Howe (1), Norfolk (2) and Rangitāhua (3) (numbers apply to both maps). The surface area of each circle is proportionate to the percentage of endemic lineages originating at the indicated location. The greyed circle illustrates an undefined origin. The arrows indicate the hypothesised colonisation routes. The inset map enlarges the Southwest Pacific, as defined in this review, to show the area investigated in a recent phylogenetic study (dashed square ranging 25°-55°S and 160°E-170°W; <xref ref-type="bibr" rid="B112">Samayoa et al. (2025)</xref>) that revealed the latitudinal variation of biodiversity metrics (right graph) for marine ray-finned fishes occurring in New Zealand’s waters. The blue line shows species richness (SR), defined as the number of species per latitudinal degree. The red line represents weighted endemism (WE), defined as the number of taxa per latitude, weighted by the number of geographic grid cells each taxon occurs at (<xref ref-type="bibr" rid="B24">Crisp et al. 2001</xref>). Map projections: WGS84.</p>
          </caption>
          <graphic xlink:href="biogeography-19-001_article-204479__-g003.jpg" id="oo_1744161.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1744161</uri>
          </graphic>
        </fig>
        <p>For marine fish richness distribution along New Zealand’s waters, data showed higher taxonomic and phylogenetic diversities in warm, low-latitude waters, peaking at around 35°S (<xref ref-type="bibr" rid="B112">Samayoa et al. 2025</xref>), in congruence with previous phylogenetic and ecological research in the region (<xref ref-type="bibr" rid="B33">Eme et al. 2020</xref>; <xref ref-type="bibr" rid="B82">Myers et al. 2021</xref>; <xref ref-type="bibr" rid="B76">Middleton et al. 2023</xref>) and the distribution of marine fish biodiversity in the Southern Hemisphere (<xref ref-type="bibr" rid="B15">Chaudhary et al. 2017</xref>). Additionally, phylogenetic results showed that the warmer waters of the North Island of New Zealand and northern subtropical oceanic islands act as “museums” for old marine fish lineages. Altogether, these results could reflect a range overlap of tropical and mid-latitude species (<xref ref-type="bibr" rid="B14">Chaudhary et al. 2016</xref>).</p>
      </sec>
      <sec sec-type="Centres of endemism" id="sec10">
        <title>Centres of endemism</title>
        <p>Recent research has engaged in the historical biogeography of marine fish endemism in the Southwest Pacific (<xref ref-type="bibr" rid="B110">Samayoa et al. 2022</xref>; <xref ref-type="bibr" rid="B111">Samayoa et al. 2023</xref>; <xref ref-type="bibr" rid="B112">Samayoa et al. 2025</xref>) highlighting the subtropical islands of Lord Howe, Norfolk Island and Rangitāhua (Kermadec Islands) as the main centres of endemism in the region. Despite local differences, the three island groups share similar geographic positions (between 29 and 32°S; Fig. <xref ref-type="fig" rid="F2">2</xref>) and are characterised by their relatively young geological age (6.92–2.58 Ma) originating as volcanic oceanic islands. Former taxonomic surveys estimated an endemism rate of 4.6% for coastal fishes around these subtropical islands (<xref ref-type="bibr" rid="B37">Francis 1993</xref>), a significant, but lower rate than those found in other volcanic islands of the Pacific: 13.7% in the Marquesas islands (<xref ref-type="bibr" rid="B27">Delrieu-Trottin et al. 2015</xref>); 25% in Hawaii (<xref ref-type="bibr" rid="B99">Randall 2007</xref>); and 61.5% in Juan Fernández and Desventuradas (<xref ref-type="bibr" rid="B41">Friedlander et al. 2016</xref>).</p>
        <sec sec-type="Geographic origins" id="sec11">
          <title>Geographic origins</title>
          <p>Phylogenetic investigations have shown that the geographical origin of lineages endemic to the subtropical islands of the Southwest Pacific is mainly located in Australia (Fig. <xref ref-type="fig" rid="F3">3</xref>), which is responsible for 53–67% of endemic lineages (<xref ref-type="bibr" rid="B110">Samayoa et al. 2022</xref>; <xref ref-type="bibr" rid="B111">Samayoa et al. 2023</xref>). Interestingly, most (67%) of these Australia-linked endemic taxa are estimated to have originated in temperate areas (south of Bermagui, New South Wales, at 36.4°S), while the rest in tropical waters (north of Bermagui). The East Pacific appears as the second most relevant origin albeit with a far lesser percentage (7–15%; numbers vary across studies). A minority of endemic lineages (3–7%) are individually linked to other areas of the Pacific Ocean (e.g. Northwest Pacific, Hawaii and the larger Indo-Pacific). As a small percentage of endemics are closely related to taxa distributed in distant Pacific locations, this result highlights the likelihood of transpacific dispersal events to colonise islands across the Pacific (<xref ref-type="bibr" rid="B106">Robertson et al. 2004</xref>). Although the geographic origin has yet to be resolved for around a quarter of endemic lineages (17–26%), the phylogenetic approach of historical biogeography brought clear evidence of the significant influence of Australian fauna as a source of endemism to the eastern subtropical volcanic islands. These updated studies have clarified previous taxonomic data suggesting a geographic affinity of subtropical endemism with Australia (<xref ref-type="bibr" rid="B37">Francis 1993</xref>; <xref ref-type="bibr" rid="B122">Trnski and de Lange 2015</xref>). This connection to mainland Australia is likely facilitated by the predominant eastward oceanic currents in the region (Figs <xref ref-type="fig" rid="F2">2</xref>, <xref ref-type="fig" rid="F3">3</xref>), a similar pattern reported in other marine taxa (<xref ref-type="bibr" rid="B37">Francis 1993</xref>; <xref ref-type="bibr" rid="B10">Brook 1998a</xref>, <xref ref-type="bibr" rid="B12">1999</xref>; <xref ref-type="bibr" rid="B9">Bronstein et al. 2019</xref>).</p>
        </sec>
        <sec sec-type="Temporal evolution" id="sec12">
          <title>Temporal evolution</title>
          <p>The study of the historical biogeography of marine fishes of the Southwest Pacific has additionally provided clues about the evolution of endemism since its inception. From a temporal perspective, the contrast of the geological age of the subtropical islands with the estimated time of lineage splitting within the time-calibrated phylogeny of focal taxa showed that two-thirds of endemic lineages diverged after the emergence of the islands (<xref ref-type="bibr" rid="B110">Samayoa et al. 2022</xref>; <xref ref-type="bibr" rid="B111">Samayoa et al. 2023</xref>). These “neoendemics” are characterised by their relatively young geological age and are likely the evolutionary result of favouring local, ecological conditions for adaptive radiation (<xref ref-type="bibr" rid="B45">Gillespie and Roderick 2002</xref>). Additionally, the role of the subtropical islands as neodendemism centres was reinforced by the phylogenetic study of <xref ref-type="bibr" rid="B112">Samayoa et al. (2025)</xref> who statistically contrasted patterns of phylogenetic endemism with species richness, a demonstrated method to quantitatively distinguish centres of neo- and palaeoendemism (i.e. range-restricted remnant populations of a previously widespread ancestor) (<xref ref-type="bibr" rid="B78">Mishler et al. 2014</xref>). Despite recognising the potential influence of taxa with broader ranges than the study area (Fig. <xref ref-type="fig" rid="F3">3</xref>) on inferred endemism patterns, the authors identified the islands of Norfolk and Rangitāhua as centres of neoendemism - where results highlighted a significant concentration of short, range-restricted phylogenetic branches – in agreement with taxonomic evidence for marine fish endemism in Norfolk (<xref ref-type="bibr" rid="B39">Francis and Hoese 2025</xref>) and Rangitāhua (<xref ref-type="bibr" rid="B123">Trnski et al. 2015</xref>). The subtropical islands of the Southwest Pacific represent a major area of biodiversity origination (e.g. <xref ref-type="bibr" rid="B63">Liggins et al. (2022)</xref>), similar to other volcanic islands within the Pacific (<xref ref-type="bibr" rid="B22">Cowman et al. 2017</xref>; Fig. <xref ref-type="fig" rid="F1">1</xref>), supporting biogeographic models that assume the peripheral islands are engines of marine fish biodiversity in the Pacific (<xref ref-type="bibr" rid="B5">Bowen et al. 2013</xref>; <xref ref-type="bibr" rid="B42">Gaboriau et al. 2018</xref>). Interestingly, the establishment of new, young endemic taxa seems to have occurred as independent events throughout the geological life of the subtropical islands (<xref ref-type="bibr" rid="B111">Samayoa et al. 2023</xref>), similar to the independent founder-events for marine fish endemics of Hawaii (<xref ref-type="bibr" rid="B51">Hodge et al. 2014</xref>) and Rapa Nui (Easter Island) (<xref ref-type="bibr" rid="B28">Delrieu-Trottin et al. 2018</xref>). These results highlight the diversity of historical events shaping modern-day patterns.</p>
        </sec>
        <sec sec-type="Spatial evolution: underpinning mechanisms" id="sec13">
          <title>Spatial evolution: underpinning mechanisms</title>
          <p>The combination of time-calibrated phylogenies with range information for terminal taxa is key for estimating ancestral ranges and the geographic origin of biodiversity (<xref ref-type="bibr" rid="B113">Sanmartín 2012</xref>). To this effect, novel biogeographic techniques parametrise processes shaping range evolution (e.g. vicariance and jump-dispersal) along the phylogenetic tree (<xref ref-type="bibr" rid="B70">Matzke 2013</xref>, <xref ref-type="bibr" rid="B71">2014</xref>), estimate parameters under a maximum-likelihood framework (<xref ref-type="bibr" rid="B102">Ree et al. 2005</xref>) and infer the most likely range at each ancestral node (<xref ref-type="bibr" rid="B101">Ree and Smith 2008</xref>). Probabilistic biogeography allows the explicit testing of evolutionary and ecological hypotheses regarding the processes shaping modern-day biogeographic patterns (<xref ref-type="bibr" rid="B61">Lamm and Redelings 2009</xref>), significantly clarifying the prevalence of jump-dispersal and vicariant events during range evolution (<xref ref-type="bibr" rid="B71">Matzke 2014</xref>). <xref ref-type="bibr" rid="B111">Samayoa et al. (2023)</xref> followed this approach to infer the processes that influence range evolution in endemic actinopterygian lineages to the subtropical islands of the Southwest Pacific. Probabilistic biogeographic models suggested that both jump-dispersal and vicariance are significant drivers of endemism, where jump-dispersal is a relevant process for lineages restricted to volcanic islands. This inference supports historical biogeographic models that emphasise the synergy between vicariant and dispersal events over time and space (<xref ref-type="bibr" rid="B113">Sanmartín 2012</xref>), as well as models of marine island biogeography that underline the relevance of colonisation after the emergence of an island (<xref ref-type="bibr" rid="B125">Whittaker et al. 2008</xref>; <xref ref-type="bibr" rid="B47">Hachich et al. 2015</xref>; <xref ref-type="bibr" rid="B26">Dawson 2016</xref>; <xref ref-type="bibr" rid="B94">Pinheiro et al. 2017</xref>). The study from <xref ref-type="bibr" rid="B111">Samayoa et al. (2023)</xref> shows the utility of probabilistic models that incorporate a jump-dispersal parameter (<xref ref-type="bibr" rid="B71">Matzke 2014</xref>), given the congruence between what the models suggested and their ecological expectations. Their work highlights the importance of integrating jump-dispersal in probabilistic models as it is a valid approach (<xref ref-type="bibr" rid="B72">Matzke 2022</xref>) despite previous criticism (<xref ref-type="bibr" rid="B100">Ree and Sanmartín 2018</xref>).</p>
        </sec>
      </sec>
    </sec>
    <sec sec-type="Future directions" id="sec14">
      <title>Future directions</title>
      <p>Constructive research relies on the conscious identification of what we know and what we do not know (<xref ref-type="bibr" rid="B52">Hortal et al. 2015</xref>). Here, we identify five major grounds of investigation that would enrich the inferences of recent biogeographic studies on marine fish biodiversity in the Southwest Pacific.</p>
      <p>First, phylogenetic inferences would improve by adding taxa within poorly-sampled genera, clarifying the evolutionary relationships amongst all extant congeners and avoiding the overestimation of divergence timings (<xref ref-type="bibr" rid="B50">Hodge and Bellwood 2016</xref>). Such addition would derive from an improvement of specimen preservation for DNA analysis and field surveys. Furthermore, the generation and inclusion of novel sequences for additional gene regions would improve topological resolution (<xref ref-type="bibr" rid="B109">Rubinoff and Holland 2005</xref>) and benefit species identification (<xref ref-type="bibr" rid="B31">Dupuis et al. 2012</xref>). The exploration of phylogenomics is a promising, relatively inexpensive alternative to molecular phylogenetics by using genome-scale data to infer evolutionary relationships amongst taxa, estimate divergence times and reconstruct ancestral ranges in biogeography (<xref ref-type="bibr" rid="B129">Young and Gillung 2020</xref>). The inference of molecular phylogenies at the tips of the Fish Tree of Life is a necessary approach that also addresses biological shortfalls (<xref ref-type="bibr" rid="B52">Hortal et al. 2015</xref>). For instance, <xref ref-type="bibr" rid="B110">Samayoa et al. (2022)</xref> clarified the evolutionary relationships within 21 genera, eight of which were fully sampled and four missed only one taxon. Moreover, the investigators built the first comprehensively sampled phylogenies for four genera (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Enneapterygius">Enneapterygius</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lepidoperca">Lepidoperca</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hypoplectrodes">Hypoplectrodes</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Upeneus">Upeneus</tp:taxon-name-part></tp:taxon-name></italic>) and confirmed the species status for 34 endemic taxa, 11 of which are positioned for the first time within a published molecular phylogeny, including three undescribed taxa (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hypoplectrodes">Hypoplectrodes</tp:taxon-name-part></tp:taxon-name></italic> sp. A, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hypoplectrodes">Hypoplectrodes</tp:taxon-name-part></tp:taxon-name></italic> sp. B and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Nemadactylus">Nemadactylus</tp:taxon-name-part></tp:taxon-name></italic> n.sp.). Additionally, novel molecular sequences for six endemics were added to open-access repositories.</p>
      <p>Second, the full extension of the phylogenetic landscape of marine fishes in the shallow waters around New Zealand remains to be examined. Recent investigations focused on richness and endemism metrics, which are a small proportion of the numerous indices measurable with a phylogenetic tree (<xref ref-type="bibr" rid="B124">Tucker et al. 2017</xref>). Future investigations of measures of distance and evenness could provide complementary understanding (e.g. <xref ref-type="bibr" rid="B33">Eme et al. (2020)</xref>). Furthermore, the estimation of divergence rates, based on the phylogenies from <xref ref-type="bibr" rid="B110">Samayoa et al. (2022)</xref>, would enhance our understanding of the evolutionary and ecological processes behind the biogeographic patterns of marine fish fauna, as diversification rates bridge questions raised in macroevolution and macroecology (<xref ref-type="bibr" rid="B75">McGill et al. 2019</xref>). These lines of investigation will benefit from the ongoing molecular extension of New Zealand’s marine fish phylogeny (Liggins unpublished results).</p>
      <p>Third, the ecological aspects of the actinopterygian biogeography in the Southwest Pacific are yet to be fully explored. Fieldwork in subtropical islands, particularly Lord Howe and Norfolk, would improve taxonomic inventories and provide biological/ecological information on endemic taxa (e.g. dispersal capacities and reproductive success), offering an ecological background to contrast with the evolutionary inferences presented in this review. A short-term objective is the formal description of undescribed endemic taxa included in <xref ref-type="bibr" rid="B110">Samayoa et al. (2022)</xref>, an ongoing work for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Nemadactylus">Nemadactylus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hypoplectrodes">Hypoplectrodes</tp:taxon-name-part></tp:taxon-name></italic> (Roberts unpublished results).</p>
      <p>Fourth, the continuing curation of open-access repositories is a substantial work that provides a valuable amount of biological, ecological and biogeographic data. The biogeographic studies presented in this review exemplify the capacity of using contemporary analytical tools to synthesise an ever-growing amount of data and to improve our understanding of natural processes. Similar works should provide constructive inferences if web-based datasets (e.g. occurrence and range information) are maintained and updated by expert opinions, as this is useful information for evolutionary, ecological and biogeographic research.</p>
      <p>Finally, the present review emphasises the relevance of phylogenetic methods in clarifying biogeographic patterns and inferring underlying evolutionary and ecological processes. We confirm the usefulness of fossils to calibrate molecular phylogenies and to estimate divergence timings (<xref ref-type="bibr" rid="B97">Powell et al. 2020</xref>), despite taxonomic and age uncertainties of some fossils (<xref ref-type="bibr" rid="B49">Heads 2005</xref>). Additionally, we highight the efficacy of probabilistic models in tracing back the evolutionary trajectory of biodiversity, particularly those that incorporate a jump-dispersal parameter (<xref ref-type="bibr" rid="B71">Matzke 2014</xref>). Lastly, we stress the value of phylogenetic metrics in determining evolutionary processes uncaptured by species richness alone (<xref ref-type="bibr" rid="B35">Faith 2013</xref>) and in quantitatively delineating centres of endemism (<xref ref-type="bibr" rid="B78">Mishler et al. 2014</xref>). We believe that the development and application of novel phylogenetic-based tools represent a suitable approach to better understand the historical biogeography of marine fishes in the Pacific.</p>
    </sec>
    <sec sec-type="Concluding remarks" id="sec15">
      <title>Concluding remarks</title>
      <p>Climate change is impacting the current geographic distribution of marine biodiversity at a global scale (<xref ref-type="bibr" rid="B16">Chaudhary et al. 2021</xref>) and is expected to aggravate the loss of marine life already stressed by anthropogenic pressure (<xref ref-type="bibr" rid="B2">Beaugrand et al. 2015</xref>). Areas of high biodiversity have been recognised as conservation priorities (<xref ref-type="bibr" rid="B83">Myers 1988</xref>; <xref ref-type="bibr" rid="B85">Myers et al. 2000</xref>; <xref ref-type="bibr" rid="B105">Roberts et al. 2002</xref>), with recent research showing that the evolutionary processes that shape biodiversity “hotspots” are equally valuable if we are to understand how life will cope with the current environmental pressures (<xref ref-type="bibr" rid="B116">Sgrò et al. 2011</xref>; <xref ref-type="bibr" rid="B81">Munday et al. 2013</xref>; <xref ref-type="bibr" rid="B89">Nielsen et al. 2017</xref>). As marine actinopterygians are highly diverse (<xref ref-type="bibr" rid="B40">Fricke et al. 2024</xref>), play critical ecological functions (<xref ref-type="bibr" rid="B48">Hayden et al. 2019</xref>), are economically valuable (<xref ref-type="bibr" rid="B86">Natale et al. 2013</xref>) and display responsiveness to climate change (<xref ref-type="bibr" rid="B30">Duffy et al. 2016</xref>), marine fishes are a good model to understand how marine biodiversity has evolved through time and space. To enhance our understanding of the evolutionary role that regions at the periphery of the Pacific Ocean play in the generation and maintenance of marine biodiversity, we focused on the Southwest Pacific, a highly diverse region with significant marine fish endemism at a global scale (<xref ref-type="bibr" rid="B19">Costello et al. 2017</xref>), but where evolutionary research has been in its infancy (<xref ref-type="bibr" rid="B28">Delrieu-Trottin et al. 2018</xref>; <xref ref-type="bibr" rid="B33">Eme et al. 2020</xref>; <xref ref-type="bibr" rid="B63">Liggins et al. 2022</xref>). To inform and better understand the evolutionary processes that shaped contemporary patterns of marine fish endemism and richness in the Southwest Pacific, we reviewed the latest generated empirical data, based on a variety of historical biogeographic approaches.</p>
      <p>Overall, the current state of knowledge highlights a historical biogeographic scenario for marine fishes of the Southwest Pacific where centres of endemism are not concordant with richness hotspots, suggesting that the patterns originated from distinct historical processes. Biodiversity richness peaks at 35°S to decrease with increasing latitude, a pattern likely driven by the range overlap of tropical, subtropical and temperate species and expected to intensify with current poleward range shifts of tropical marine fish taxa as detected along northern New Zealand (e.g. <xref ref-type="bibr" rid="B76">Middleton et al. (2023)</xref>) and south-eastern Australia (e.g. <xref ref-type="bibr" rid="B36">Fowler et al. (2018)</xref>; <xref ref-type="bibr" rid="B44">Gervais et al. (2021)</xref>). Endemism concentrates in subtropical oceanic islands, which act as peripheral islands that generate neoendemism in the Pacific and originates principally from mainland Australia lineages that occupied the recently emerged islands through rare colonisation events – following the predominant eastward oceanographic currents – to eventually diverge into endemic taxa. To conclude, it is considered that achieving a complete understanding of any natural process is highly unlikely (<xref ref-type="bibr" rid="B60">Ladle and Hortal 2013</xref>). However, we believe that the latest evolutionary and biogeographic research in the Southwest Pacific contributes to a better understanding of the historical biogeography of marine fishes in the Southwest Pacific, representing a knowledge bridge for future work within the marine evolutionary ecology field.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We thank J. David Aguirre (University of Auckland) and Adam Smith (Seathrough Science) for their constructive comments during the early stages of this work. We are also grateful for the positive insights from Prof. Mary Morgan-Richards (Massey University) and Prof. Giacomo Bernardi (University of California Santa Cruz), as well from Dr. Peter Cowman (James Cook University) who particularly acknowledged the relevance in publishing our work. This review derives from the doctoral thesis of André P. Samayoa who was supported by a Massey University Doctoral Scholarship. Libby Liggins is supported by a Rutherford Discovery Fellowship (RDF-UOA2001).</p>
    </ack>
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    <sec sec-type="Additional information" id="sec16">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec17">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec18">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec19">
        <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="sec20">
        <title>Funding</title>
        <p>Libby Liggins is supported by a Rutherford Discovery Fellowship (RDF-UOA2001).</p>
      </sec>
      <sec sec-type="Author contributions" id="sec21">
        <title>Author contributions</title>
        <p>Conceptualization: APS, LL. Formal analysis: APS. Funding acquisition: LL. Methodology: APS. Project administration: LL. Resources: LL. Supervision: LL. Writing - original draft: APS. Writing - review and editing: APS, LL.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec22">
        <title>Author ORCIDs</title>
        <p>A.P. Samayoa <ext-link xlink:href="https://orcid.org/0000-0003-3852-7935" ext-link-type="uri">https://orcid.org/0000-0003-3852-7935</ext-link></p>
        <p>L. Liggins <ext-link xlink:href="https://orcid.org/0000-0003-1143-2346" ext-link-type="uri">https://orcid.org/0000-0003-1143-2346</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec23">
        <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>
