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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2024.1509455</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Recovery of coral cover at Lizard island Australia 6 years post-disturbance</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Anderson</surname>
<given-names>Gabriel Dax</given-names>
</name>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2697989"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
</contrib-group>
<aff id="aff1">
<institution>California Polytechnic State University San Luis Obispo, Biological Sciences Department</institution>, <addr-line>San Luis Obispo, CA</addr-line>, <country>United States</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Charles Alan Jacoby, University of South Florida St. Petersburg, United States</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Ernesto F. Weil, University of Puerto Rico at Mayag&#xfc;ez, Puerto Rico</p>
<p>Sanqiang Gong, Chinese Academy of Sciences (CAS), China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Gabriel Dax Anderson, <email xlink:href="mailto:gabrieldax38@gmail.com">gabrieldax38@gmail.com</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>21</day>
<month>01</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1509455</elocation-id>
<history>
<date date-type="received">
<day>11</day>
<month>10</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>24</day>
<month>12</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Anderson</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Anderson</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Coral reefs are experiencing more intense and frequent disturbances induced by climate change, such as cyclones and bleaching events. This necessitates a better understanding of the ongoing environmental conditions that stress these systems and the subsequent arc of longer-term reef responses to these stressful conditions. From March of 2014 to May of 2017, the Lizard Island reefs in the northern region of the Great Barrier Reef experienced four consecutive annual disturbances; Cyclone Ita in 2014, Cyclone Nathan in 2015, and two massive bleaching events in 2016 and 2017. Between the concentrated patches of physical damage from the cyclones and the uniform impact of the bleaching events, these reefs were devastated, with none of the eight study sites harboring more than 20% live coral cover by May of 2017. In November of 2023, after six years of relatively calmer conditions with no conspicuous region-wide, large-scale disturbances, I documented the extant coral community on eight previously-monitored reefs around Lizard Island. All reefs showed significant (<italic>p</italic> = 0.0054, F = 3.46, df = 47) improvement from their 2017 immediate post-disturbance degradation. Living coral at my study sites had recovered to between 18.4 &#xb1; 0.6 (mean &#xb1; 1 SE) to 59.9 &#xb1; 5.3% of the reef area per site by 2023, with many sites towards the higher end of that range. Recovery of coral extent appeared to follow a north-south trend in which more Trade Wind-sheltered northerly sites had generally greater recovery and higher live coral cover compared to more exposed southern sites, which experienced significantly less coral recovery. Fast-growing Acroporid corals drove the recovery of coral extent in these more northern sites. While family richness across all sites improved by 2023 (4.0 &#xb1; 0.1; grand mean &#xb1; 1 se), Lizard Island reefs have yet to reach their pre-disturbance diversity (4.8 &#xb1; 0.6 in 2014). Future annual surveys of the study sites as well as others surveyed in 2017 may better clarify the relationship between reef location and the rate of recovery of coral cover post-disturbance.</p>
</abstract>
<kwd-group>
<kwd>disturbance ecology</kwd>
<kwd>biodiversity loss</kwd>
<kwd>coral recovery</kwd>
<kwd>habitat resilience</kwd>
<kwd>coral bleaching</kwd>
</kwd-group>
<counts>
<fig-count count="5"/>
<table-count count="3"/>
<equation-count count="0"/>
<ref-count count="47"/>
<page-count count="9"/>
<word-count count="3968"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Coral Reef Research</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>Coral reefs globally are ecologically (<xref ref-type="bibr" rid="B46">Wilson et&#xa0;al., 2006</xref>), economically (<xref ref-type="bibr" rid="B37">Santavy et&#xa0;al., 2021</xref>), and culturally significant (<xref ref-type="bibr" rid="B6">Cinner et&#xa0;al., 2009</xref>) to developed and developing countries alike (<xref ref-type="bibr" rid="B25">Laurans et&#xa0;al., 2013</xref>). Extants reefs are subject to increasingly frequent and intense disturbances, including climate change-induced factors (<xref ref-type="bibr" rid="B20">Hughes et&#xa0;al., 2017</xref>). Major proximate threats to the world&#x2019;s largest coral reef system&#x2013;the Great Barrier Reef&#x2013;include predation by Crown-of-Thorns starfish (<xref ref-type="bibr" rid="B33">Pratchett, 2010</xref>), unsustainable fishing practices (<xref ref-type="bibr" rid="B37">Santavy et&#xa0;al., 2021</xref>), disease outbreaks (<xref ref-type="bibr" rid="B34">Roff et&#xa0;al., 2011</xref>), and bleaching due to climate-induced high temperature stress (<xref ref-type="bibr" rid="B17">Hoegh-Guldberg, 1999</xref>). As these disturbances occur more often and with greater intensity, heterogenous coral recovery drives changes in reef composition (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). This often reduces the quality of immediate recovery and is correlated with negative long-term trends in biodiversity (<xref ref-type="bibr" rid="B45">Wakeford et&#xa0;al., 2008</xref>). Future bleaching events in coming decades will likely surpass the thermal tolerance of essential corals, impeding their continued growth, development, and ecological function (<xref ref-type="bibr" rid="B17">Hoegh-Guldberg, 1999</xref>). Given that reefs continue to face these intense conditions, it is critical to understand how reef assemblage will be impacted in the long term.</p>
<p>From 2014-2017, the northern Great Barrier Reef faced a harsh sequence of disturbances&#x2013;Cyclone Ita in 2014, Cyclone Nathan in 2015, and two massive bleaching events in 2016 and 2017 (<xref ref-type="bibr" rid="B45">Wakeford et&#xa0;al., 2008</xref>). The physical damage caused by those cyclones coupled with the large-scale bleaching to devastate reefs around the island. Before these stressors, most Lizard Island reefs were coral-rich (typically &gt;40% coral cover; <xref ref-type="bibr" rid="B10">Done et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B23">Johns et&#xa0;al., 2014</xref>), but by late 2017 many were close to 4% of live coral cover (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>).These collective phase shifts away from more structurally secure, complex reefs have had dramatic impacts on fish abundance and species richness (<xref ref-type="bibr" rid="B24">Jones et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B46">Wilson et&#xa0;al., 2006</xref>). Since changes in composition of coral communities tend to be discrete following these episodic stressors, recovery can occur if a reef has sufficient time between perturbations (<xref ref-type="bibr" rid="B12">Gilmour et&#xa0;al., 2013</xref>). However, as the time between these events continues to decrease, phase-shifts become more common, rendering such reefs less able to effectively recover to a pre-disturbance condition (T. P. <xref ref-type="bibr" rid="B19">Hughes and Connell, 1999</xref>). The rising prevalence of phase-shifts, together with increasingly frequent disturbances, hinders and, in some cases, completely inhibits coral recruitment and community succession, leaving these potential habitats in a volatile state (<xref ref-type="bibr" rid="B2">Baird and Hughes, 2000</xref>).</p>
<p>There are extant examples of other reefs undergoing similar conditions. Following a high urchin mortality event in 1983, grazing was greatly reduced across Caribbean reefs where herbivorous fish populations were overfished, leading to the emergence of alternate stable states in which macroalgae made up the majority of cover in previously coral-dominated areas (<xref ref-type="bibr" rid="B32">Mumby et&#xa0;al., 2007</xref>). This weakened the resilience of these reefs against future disturbances that target corals more specifically. Between 1980 and 2001, 286 Caribbean reefs experienced ~17% coral cover reduction in years following hurricanes, with no evidence of recovery to a pre-storm condition for at least eight years post-disturbance (<xref ref-type="bibr" rid="B11">Gardner et&#xa0;al., 2005</xref>). This suggests that not only does reef condition decline with more frequent and damaging impacts, but that reef recovery is also being impaired. Caribbean reefs have continued to experience a decline in structural and biological complexity to this day, with Caribbean-wide homogenization of current structures and species (<xref ref-type="bibr" rid="B1">Alvarez-Filip et&#xa0;al., 2009</xref>). Similar trends of more homogenized, less healthy reefs have been documented across the same time scale in the Philippines (<xref ref-type="bibr" rid="B27">Licuanan et&#xa0;al., 2019</xref>), Indonesia (<xref ref-type="bibr" rid="B47">Wolfe et&#xa0;al., 2020</xref>), Papua New Guinea (<xref ref-type="bibr" rid="B3">Berzunza-Sanchez et&#xa0;al., 2013</xref>). Each of these reefs have also suffered from overexploitation of resources and poor management decisions, but the negative impacts of cyclones and bleaching cannot be mitigated at the source in the same way. This makes it imperative that we better understand how reefs cope with these large-scale disturbances.</p>
<p>In the immediate wake of these disturbances, researchers characterized the post-impact composition of 13 reef sites around the island in 2017. They found that coral cover across 10 of the 19 sites was below 4%, with only two sites having cover above 40%, the highest cover for most sites prior to the four years&#x2019; disturbances (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). Researchers predicted reef structure and complexity would begin to return. I sought to document any potential coral recovery six years after the 2017 surveys to determine if that recovery had begun on Lizard Island reefs. In the fall of 2023, I sought to test the hypothesis that coral cover and composition were unchanged from 2017.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<title>Materials and methods</title>
<p>I explored coral reefs around the Lizard Island Research Station (14.6688&#xb0; S, 145.4594&#xb0; E), on the northern portion of Australia&#x2019;s Great Barrier Reef. I sampled eight reef sites at Lizard Island from 23 October to 18 November,2023, using the same site name and locations as the previous study (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>)&#x2013;Cook&#x2019;s Path (14.660557&#xb0; S, 145.453396&#xb0; E), Lagoon 1 (14.686888&#xb0; S, 145.456658&#xb0; E), Lagoon 2 (14.698240&#xb0; S, 145.450682&#xb0; E), Horseshoe (14.687851&#xb0; S, 145.444216&#xb0; E), Vicki&#x2019;s (14.685130&#xb0; S, 145.442876&#xb0; E), Osprey (14.667579&#xb0; S, 145.443119&#xb0; E), Mermaid Cove (14.647248&#xb0; S, 145.453939&#xb0; E), and North Reef (14.645044&#xb0; S, 145.453987&#xb0; E). These sites are dispersed across the relatively protected side of Lizard Island, leeward of the prevailing Southeast Trade Winds (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). Risk management concerns (restrictions to remain in range of radio contact and avoid the rougher sea conditions of Lizard Island&#x2019;s east side) prevented me from collecting data on the Northeast to Southeast of the island and as such I could only sample a subset of the representative sites across the island reefs.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Map of Lizard Island oriented to show North-South range of study sites.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1509455-g001.tif"/>
</fig>
<p>At each site, six 10-meter line-intercept transects were randomly laid out over the reef face at one to two meters deep, with each transect no further than 10 meters apart from the last (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). The substrate type beneath the transect tape was recorded to the nearest milimeter, and hard corals were classified by family to determine the percent cover. The categories recorded at each site included soft coral, <italic>Acropora, Porites, Pocillopora</italic>, encrusting coralline algae, hard dead coral, rubble, sand, and algae. In data visualization, these types were put into the following groups: hard coral (<italic>Acropora, Porites, and Pocillopora</italic>), soft coral, algae, encrusting coralline algae, and abiotic substrate (hard dead substrate, rubble, and sand). Transect data were analyzed with JMP Pro (v 17). I compared the proportion of total live coral cover by reef, family proportion of total cover by year, grand means of coral cover proportion by year, and grand means of family richness by year using analysis of variance.</p>
</sec>
<sec id="s3" sec-type="results">
<title>Results</title>
<p>In the wake of the 2014-2017 sequence of disturbances, total live coral cover declined dramatically from pre-disturbance conditions in 2014. Some reefs had single percent coverage of live coral by 2017. In the six years since, total live coral cover on all reefs increased dramatically, with six out of eight reefs harboring more live coral cover than in 2014. The remaining two sites (Lagoon 1 and Lagoon 2) show lower coral cover than pre-disturbance 2014 levels (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Grand means of live coral cover across my eight sites went from 39.6% in 2014 to 12% in 2017 to 51% in 2023.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Proportion of total live coral cover &#xb1; 1 SE at my eight reef sites in 2014 (pre-disturbance), 2017 (post-disturbance), and 2023 (6 years post-disturbance). 2014 and 2017 data sourced from <xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>. Reefs are ordered from northern-most on the left to southernmost on the right.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1509455-g002.tif"/>
</fig>
<p>The proportion of soft coral and <italic>Acropora</italic> cover declined heavily, while the proportion of <italic>Pocillopora</italic> and <italic>Porites</italic> cover fluctuated more moderately (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). By 2023 (six years post-disturbance), all groups recovered by various margins, with the proportion of cover of all groups increasing&#x2013;<italic>Acropora</italic> by 20.1% (p = 0.049, F = 2.03, df = 7), soft coral by 9.5% (p = 0.025, F = 1.97, df = 7), <italic>Pocillopora</italic> by 6.8% (p = 0.015, F = 2.49, df = 7), and <italic>Porites</italic> by 4.2% (p = 0.009, F = 1.42, df = 7). All groups except soft coral recovered to 2014 (pre-disturbance) proportion of cover (<xref ref-type="table" rid="T1">
<bold>Tables&#xa0;1</bold>
</xref>&#x2013;<xref ref-type="table" rid="T3">
<bold>3</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Proportion of soft coral and dominant hard coral families (<italic>Acroporidae</italic>, <italic>Poritidae</italic>, and <italic>Pocilloporidae</italic>) cover &#xb1; 1 SE across Lizard Island reefs from 1995-2023. 1995-2017 data sourced from <xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1509455-g003.tif"/>
</fig>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>ANOVAs of spatial and temporal factors impacting coral cover on Lizard Island reefs.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="4" align="center">Effect on Total Live Coral Cover</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">df</th>
<th valign="middle" align="center">p</th>
</tr>
<tr>
<td valign="middle" align="center">Year</td>
<td valign="middle" align="center">0.67</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="center">Reef</td>
<td valign="middle" align="center">3.31</td>
<td valign="middle" align="center">18</td>
<td valign="middle" align="center">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="center">Year*Reef</td>
<td valign="middle" align="center">1.21</td>
<td valign="middle" align="center">77</td>
<td valign="middle" align="center">0.148</td>
</tr>
</tbody>
</table>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="4" align="center">Effect on Live Acropora Cover</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">df</th>
<th valign="middle" align="center">p</th>
</tr>
<tr>
<td valign="middle" align="center">Year</td>
<td valign="middle" align="center">2.03</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">0.049</td>
</tr>
<tr>
<td valign="middle" align="center">Reef</td>
<td valign="middle" align="center">1.16</td>
<td valign="middle" align="center">18</td>
<td valign="middle" align="center">0.288</td>
</tr>
<tr>
<td valign="middle" align="center">Year*Reef</td>
<td valign="middle" align="center">1.15</td>
<td valign="middle" align="center">77</td>
<td valign="middle" align="center">0.194</td>
</tr>
</tbody>
</table>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="4" align="center">Effect on Live Pocillopora Cover</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">df</th>
<th valign="middle" align="center">p</th>
</tr>
<tr>
<td valign="middle" align="center">Year</td>
<td valign="middle" align="center">2.49</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">0.015</td>
</tr>
<tr>
<td valign="middle" align="center">Reef</td>
<td valign="middle" align="center">1.87</td>
<td valign="middle" align="center">18</td>
<td valign="middle" align="center">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="center">Year*Reef</td>
<td valign="middle" align="center">1.04</td>
<td valign="middle" align="center">77</td>
<td valign="middle" align="center">0.414</td>
</tr>
</tbody>
</table>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="4" align="center">Effect on Live <italic>Porites Cover</italic>
</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">df</th>
<th valign="middle" align="center">p</th>
</tr>
<tr>
<td valign="middle" align="center">Year</td>
<td valign="middle" align="center">1.42</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">0.009</td>
</tr>
<tr>
<td valign="middle" align="center">Reef</td>
<td valign="middle" align="center">2.38</td>
<td valign="middle" align="center">18</td>
<td valign="middle" align="center">&lt;0.001</td>
</tr>
<tr>
<td valign="middle" align="center">Year*Reef</td>
<td valign="middle" align="center">1.32</td>
<td valign="middle" align="center">77</td>
<td valign="middle" align="center">0.234</td>
</tr>
</tbody>
</table>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" colspan="4" align="center">Effect on Live Soft Coral <italic>Cover</italic>
</th>
</tr>
</thead>
<tbody>
<tr>
<th valign="middle" align="center">Variable</th>
<th valign="middle" align="center">F</th>
<th valign="middle" align="center">df</th>
<th valign="middle" align="center">p</th>
</tr>
<tr>
<td valign="middle" align="center">Year</td>
<td valign="middle" align="center">1.97</td>
<td valign="middle" align="center">7</td>
<td valign="middle" align="center">0.025</td>
</tr>
<tr>
<td valign="middle" align="center">Reef</td>
<td valign="middle" align="center">1.33</td>
<td valign="middle" align="center">18</td>
<td valign="middle" align="center">0.273</td>
</tr>
<tr>
<td valign="middle" align="center">Year*Reef</td>
<td valign="middle" align="center">1.26</td>
<td valign="middle" align="center">77</td>
<td valign="middle" align="center">0.421</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Transect data was ARCSIN transformed for analysis. 1995-2017 data sourced from <xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Tukey HSD <italic>post-hoc</italic> comparison of effect of year on total live coral cover.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center"/>
<th valign="middle" align="center">1995</th>
<th valign="middle" align="center">1996</th>
<th valign="middle" align="center">1997</th>
<th valign="middle" align="center">2011</th>
<th valign="middle" align="center">2014</th>
<th valign="middle" align="center">2015</th>
<th valign="middle" align="center">2016</th>
<th valign="middle" align="center">2017</th>
<th valign="middle" align="center">2023</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">
<bold>1995</bold>
</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">
<bold>0.002</bold>
</td>
<td valign="middle" align="center">0.147</td>
<td valign="middle" align="center">0.912</td>
<td valign="middle" align="center">0.129</td>
<td valign="middle" align="center">
<bold>0.0001</bold>
</td>
<td valign="middle" align="center">
<bold>0.007</bold>
</td>
<td valign="middle" align="center">
<bold>0.023</bold>
</td>
<td valign="middle" align="center">0.382</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>1996</bold>
</td>
<td valign="middle" align="center">
<bold>0.002</bold>
</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">
<bold>0.005</bold>
</td>
<td valign="middle" align="center">
<bold>0.006</bold>
</td>
<td valign="middle" align="center">
<bold>0.027</bold>
</td>
<td valign="middle" align="center">0.549</td>
<td valign="middle" align="center">
<bold>0.036</bold>
</td>
<td valign="middle" align="center">
<bold>0.048</bold>
</td>
<td valign="middle" align="center">
<bold>0.005</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>1997</bold>
</td>
<td valign="middle" align="center">0.147</td>
<td valign="middle" align="center">
<bold>0.005</bold>
</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0.832</td>
<td valign="middle" align="center">0.0574</td>
<td valign="middle" align="center">
<bold>0.033</bold>
</td>
<td valign="middle" align="center">0.568</td>
<td valign="middle" align="center">
<bold>0.019</bold>
</td>
<td valign="middle" align="center">
<bold>0.023</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2011</bold>
</td>
<td valign="middle" align="center">0.912</td>
<td valign="middle" align="center">
<bold>0.006</bold>
</td>
<td valign="middle" align="center">0.832</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0.326</td>
<td valign="middle" align="center">
<bold>0.047</bold>
</td>
<td valign="middle" align="center">
<bold>0.031</bold>
</td>
<td valign="middle" align="center">
<bold>0.012</bold>
</td>
<td valign="middle" align="center">
<bold>0.006</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2014</bold>
</td>
<td valign="middle" align="center">0.129</td>
<td valign="middle" align="center">
<bold>0.027</bold>
</td>
<td valign="middle" align="center">0.574</td>
<td valign="middle" align="center">0.326</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">
<bold>0.036</bold>
</td>
<td valign="middle" align="center">
<bold>0.025</bold>
</td>
<td valign="middle" align="center">
<bold>0.035</bold>
</td>
<td valign="middle" align="center">
<bold>0.033</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2015</bold>
</td>
<td valign="middle" align="center">
<bold>0.001</bold>
</td>
<td valign="middle" align="center">0.549</td>
<td valign="middle" align="center">
<bold>0.033</bold>
</td>
<td valign="middle" align="center">
<bold>0.047</bold>
</td>
<td valign="middle" align="center">
<bold>0.036</bold>
</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">
<bold>0.038</bold>
</td>
<td valign="middle" align="center">0.62</td>
<td valign="middle" align="center">
<bold>0.024</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2016</bold>
</td>
<td valign="middle" align="center">
<bold>0.007</bold>
</td>
<td valign="middle" align="center">
<bold>0.036</bold>
</td>
<td valign="middle" align="center">0.568</td>
<td valign="middle" align="center">
<bold>0.031</bold>
</td>
<td valign="middle" align="center">
<bold>0.025</bold>
</td>
<td valign="middle" align="center">
<bold>0.038</bold>
</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0.075</td>
<td valign="middle" align="center">
<bold>0.013</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2017</bold>
</td>
<td valign="middle" align="center">
<bold>0.023</bold>
</td>
<td valign="middle" align="center">
<bold>0.048</bold>
</td>
<td valign="middle" align="center">
<bold>0.019</bold>
</td>
<td valign="middle" align="center">
<bold>0.012</bold>
</td>
<td valign="middle" align="center">
<bold>0.035</bold>
</td>
<td valign="middle" align="center">0.062</td>
<td valign="middle" align="center">0.075</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">
<bold>0.016</bold>
</td>
</tr>
<tr>
<td valign="middle" align="center">
<bold>2023</bold>
</td>
<td valign="middle" align="center">0.382</td>
<td valign="middle" align="center">
<bold>0.005</bold>
</td>
<td valign="middle" align="center">
<bold>0.023</bold>
</td>
<td valign="middle" align="center">
<bold>0.006</bold>
</td>
<td valign="middle" align="center">
<bold>0.033</bold>
</td>
<td valign="middle" align="center">
<bold>0.024</bold>
</td>
<td valign="middle" align="center">
<bold>0.013</bold>
</td>
<td valign="middle" align="center">
<bold>0.016</bold>
</td>
<td valign="middle" align="center"/>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Bolded values are statistically significant.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Tukey HSD <italic>post-hoc</italic> comparison of effect of site on total live coral cover.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="bottom" align="center"/>
<th valign="bottom" align="center">North Reef</th>
<th valign="bottom" align="center">Osprey</th>
<th valign="bottom" align="center">Horseshoe</th>
<th valign="bottom" align="center">Lagoon 2</th>
<th valign="bottom" align="center">Cook&#x2019;s Path</th>
<th valign="bottom" align="center">Vicki&#x2019;s</th>
<th valign="bottom" align="center">Lagoon 1</th>
<th valign="bottom" align="center">Mermaid Cove</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="bottom" align="center">
<bold>North Reef</bold>
</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">0.644</td>
<td valign="bottom" align="center">
<bold>0.049</bold>
</td>
<td valign="bottom" align="center">
<bold>0.007</bold>
</td>
<td valign="bottom" align="center">
<bold>0.045</bold>
</td>
<td valign="bottom" align="center">
<bold>0.009</bold>
</td>
<td valign="bottom" align="center">
<bold>0.013</bold>
</td>
<td valign="bottom" align="center">0.717</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Osprey</bold>
</td>
<td valign="bottom" align="center">0.644</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">0.085</td>
<td valign="bottom" align="center">
<bold>0.008</bold>
</td>
<td valign="bottom" align="center">
<bold>0.038</bold>
</td>
<td valign="bottom" align="center">
<bold>0.016</bold>
</td>
<td valign="bottom" align="center">
<bold>0.042</bold>
</td>
<td valign="bottom" align="center">0.273</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Horseshoe</bold>
</td>
<td valign="bottom" align="center">
<bold>0.049</bold>
</td>
<td valign="bottom" align="center">0.085</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">0.057</td>
<td valign="bottom" align="center">0.072</td>
<td valign="bottom" align="center">
<bold>0.034</bold>
</td>
<td valign="bottom" align="center">0.125</td>
<td valign="bottom" align="center">0.106</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Lagoon 2</bold>
</td>
<td valign="bottom" align="center">
<bold>0.007</bold>
</td>
<td valign="bottom" align="center">
<bold>0.008</bold>
</td>
<td valign="bottom" align="center">0.057</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">
<bold>0.011</bold>
</td>
<td valign="bottom" align="center">0.074</td>
<td valign="bottom" align="center">
<bold>0.046</bold>
</td>
<td valign="bottom" align="center">
<bold>0.004</bold>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Cook&#x2019;s Path</bold>
</td>
<td valign="bottom" align="center">
<bold>0.045</bold>
</td>
<td valign="bottom" align="center">
<bold>0.038</bold>
</td>
<td valign="bottom" align="center">0.072</td>
<td valign="bottom" align="center">
<bold>0.011</bold>
</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">0.147</td>
<td valign="bottom" align="center">
<bold>0.018</bold>
</td>
<td valign="bottom" align="center">
<bold>0.015</bold>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Vicki&#x2019;s</bold>
</td>
<td valign="bottom" align="center">
<bold>0.009</bold>
</td>
<td valign="bottom" align="center">
<bold>0.016</bold>
</td>
<td valign="bottom" align="center">
<bold>0.034</bold>
</td>
<td valign="bottom" align="center">0.074</td>
<td valign="bottom" align="center">0.147</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">0.093</td>
<td valign="bottom" align="center">
<bold>0.004</bold>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Lagoon 1</bold>
</td>
<td valign="bottom" align="center">
<bold>0.013</bold>
</td>
<td valign="bottom" align="center">
<bold>0.042</bold>
</td>
<td valign="bottom" align="center">0.125</td>
<td valign="bottom" align="center">
<bold>0.046</bold>
</td>
<td valign="bottom" align="center">
<bold>0.018</bold>
</td>
<td valign="bottom" align="center">0.093</td>
<td valign="bottom" align="center"/>
<td valign="bottom" align="center">
<bold>0.008</bold>
</td>
</tr>
<tr>
<td valign="bottom" align="center">
<bold>Mermaid Cove</bold>
</td>
<td valign="bottom" align="center">0.717</td>
<td valign="bottom" align="center">0.273</td>
<td valign="bottom" align="center">0.106</td>
<td valign="bottom" align="center">
<bold>0.004</bold>
</td>
<td valign="bottom" align="center">
<bold>0.015</bold>
</td>
<td valign="bottom" align="center">
<bold>0.004</bold>
</td>
<td valign="bottom" align="center">
<bold>0.008</bold>
</td>
<td valign="bottom" align="center"/>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Bolded values are statistically significant.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The grand mean of live coral cover significantly declined from 2014-2017 (mid-disturbance) but rose significantly higher by 2023 (six years post-disturbance) to rise above the pre-disturbance grand mean of live coral cover (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). The grand mean of family richness declined from 2014-2017 (mid-disturbance) and recovered significantly by 2023, but not to the 2011 or 2014 (immediate pre-disturbance) grand means of family richness (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). Family richness across all sites improved by 2023 (4.0 &#xb1; 0.11; grand mean &#xb1; 1 se); however, Lizard Island reefs have yet to reach their pre-disturbance family diversity (4.77 &#xb1; 0.58 in 2014).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Grand means &#xb1; 1 SE of proportion of live coral cover across Lizard Island reefs from 1995-2023. 1995-2017 data sourced from <xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1509455-g004.tif"/>
</fig>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Grand means &#xb1; 1 SE of family richness across Lizard Island reefs from 1995-2023. 1995-2017 data sourced from <xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1509455-g005.tif"/>
</fig>
</sec>
<sec id="s4" sec-type="discussion">
<title>Discussion</title>
<sec id="s4_1">
<title>Limitations: A coral-centric assessment</title>
<p>My study focused on the change of aggregate living coral cover at Lizard Island, and as such it speaks to the dynamics of the foundational ecosystem engineers of coral reef systems. That said, it is important to note that while these 2023 data characterize current reef conditions, I did not sample fish, other invertebrates, or other ecosystem dimensions. My insights about overall reef condition are driven by aggregate coral cover.</p>
</sec>
<sec id="s4_2">
<title>North-south trends</title>
<p>Higher coral recovery occurred in the more northerly reefs sites. This apparent north-south latitudinal gradient of recovery may in part be driven by the various wind and wave conditions across the island. The prevailing Southeast Trade Winds put the southeast-facing portion of Lizard Island under the most constant physical stress. While this study was unable to include previously-surveyed sites on the east side of the Island due to logistical constraints, the physical protection via South Island, Palfrey Island, and the exterior rim of the lagoon provide relatively calmer conditions for marine growth. The difference in wave action and nutrient availability between each site, influenced by physical barriers, also plays a role in which marine growth is best-adapted to each site. Wave action on a better-protected reef is relatively gentle and ideal for less rigid/robust life forms to settle and grow. Partially protected reefs with moderate wave action have more potential to be damaged but also are afforded a suite of potential benefits. First, there can be better circulation of sediment in the water column (<xref ref-type="bibr" rid="B16">Hench et&#xa0;al., 2008</xref>), which allows for more sunlight availability. Second, such sites allow sub-optimal conditions for potential hard coral competitors (i.e. macroalgae, soft corals) and provide hard corals a greater chance of successfully recruiting and establishing upon that reef. The most protected reef sites experience less water column circulation and more nutrient buildup, favoring competitors of hard corals better adapted to rapid growth who often quickly block otherwise accessible sunlight incident on the reef surface (<xref ref-type="bibr" rid="B8">D&#x2019;Angelo and Wiedenmann, 2014</xref>). This is pivotal in creating a phase shift away from a primarily coral-based substrate and greatly obstructs further coral growth.</p>
<p>It seems that these gross differences in oceanographic conditions (moderate vs strong protections) may be encouraging potential phase shifts of these reefs. Sites with stronger protection, particularly lagoon sites, see lower rates of coral recovery (<xref ref-type="bibr" rid="B2">Baird and Hughes, 2000</xref>). Lagoon 1 and Lagoon 2 were in some of the calmest wave conditions I observed across Lizard Island in 2023. These quiescent sites appear to afford more ideal conditions for macroalgae to outcompete hard corals for space and ultimately for light once their thalli grow tall enough to overstory surrounding reef surfaces, reducing incident sunlight in their shadows. Vicki&#x2019;s and Horseshoe, despite being protected by Palfrey Island, are still on the edge of the shallow reef habitat, and as such experience much more wave action. Osprey and Cook&#x2019;s Path face similar conditions. Mermaid Cove and North Reef sit on the northern tip of Lizard Island where they receive a heavy amount of wave action and, despite being very protected from the southeast by the main body of the island, had the lowest percent cover of macroalgae and soft corals. In short, the degree of gross oceanographic protection seems to explain why Mermaid Cove and North Reef were the most recovered, and Lagoon 1 and Lagoon 2 the least-recovered, and the trend in the intermediary sites.</p>
</sec>
<sec id="s4_3">
<title>Coral composition</title>
<p>Equally noteworthy is that the bulk of the hard coral cover recorded was of the family Acroporidae. Acroporids are cosmopolitan reef-building corals, often the most common stony coral group across the expanse of Indo-Pacific reefs (<xref ref-type="bibr" rid="B2">Baird and Hughes, 2000</xref>). Past surveys (from 1995&#x2013;2017) found the genus <italic>Acropora</italic> remained the most abundant taxon from 1995 through to 2014, then declined by 95% from 2014 to 2017, being one of the least abundant taxa post-disturbance (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). The two most well-recovered sites, Mermaid Cove and North Reef, had <italic>Acropora</italic> consisting of 80% of the hard corals identified. For reasons to be determined, <italic>Acropora</italic> seems to be on track to again become the most abundant hard coral taxon at Lizard Island, as it was in the 1995&#x2013;2014 pre-disturbance area. The less recovered reefs, however, did not experience the same dominance of <italic>Acropora</italic> species. Instead, <italic>Porites</italic> was most dominant at Lagoon 2 and <italic>Pocillopora</italic> was most dominant at Lagoon 1 in 2023.</p>
<p>Acroporids tend to be a dominant family on shallow reefs due to their fast growth rate and structural advantage that allows them to outcompete understory growth (<xref ref-type="bibr" rid="B2">Baird and Hughes, 2000</xref>). However, the overabundance of <italic>Acropora</italic> on a reef can also have potentially harmful impacts on the resilience of that reef going forward. <italic>Acropora</italic> are highly susceptible to bleaching events (<xref ref-type="bibr" rid="B29">Marshall and Baird, 2000</xref>). Past bleaching events of 1998 and 2002 did not diminish the intensity of bleaching events in 2016 (<xref ref-type="bibr" rid="B20">Hughes et&#xa0;al., 2017</xref>), and those bleaching events saw a huge die-off of tabular and staghorn corals (<xref ref-type="bibr" rid="B18">Hughes et&#xa0;al., 2018</xref>), some of which belong to the <italic>Acropora</italic> family. Acroporids have also been documented to be among the preferred prey of Crown-of-Thorns starfish at Lizard Island (<xref ref-type="bibr" rid="B33">Pratchett, 2010</xref>). In a more general sense, less diverse reefs can suffer greatly from disease outbreaks&#x2013;in the case of <italic>Acropora</italic>, Acroporid White Syndrome (<xref ref-type="bibr" rid="B34">Roff et&#xa0;al., 2011</xref>) could culminate in massive disturbance events due to the prevalence of <italic>Acropora</italic> at these recovering sites.</p>
</sec>
<sec id="s4_4">
<title>Reef resilience</title>
<p>With coral reefs declining globally (<xref ref-type="bibr" rid="B46">Wilson et&#xa0;al., 2006</xref>), many extant reef-dwelling species are at risk of becoming endangered or locally or regionally extinct (<xref ref-type="bibr" rid="B39">Sherman et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B7">Coker et&#xa0;al., 2009</xref>). With climate change bringing more frequent and intense disturbance events (<xref ref-type="bibr" rid="B22">Jentsch and Beierkuhnlein, 2008</xref>), communities of reef-dwelling species are likely to continue declining without updating current management decisions (<xref ref-type="bibr" rid="B43">van Woesik et&#xa0;al., 2022</xref>). Ideal management decisions should focus on an &#x201c;ecosystem-based approach&#x201d; (<xref ref-type="bibr" rid="B15">Harvey et&#xa0;al., 2018</xref>) that takes a more holistic perspective of all aspects of coral reef communities.</p>
<p>Species diversity has been found to be 5.7x more essential to maximizing biomass compared to other ecological and environmental influences (<xref ref-type="bibr" rid="B26">Lefcheck et&#xa0;al., 2021</xref>). This suggests coral reefs require a variety of reef-building corals for continued recovery of habitat health and biological diversity. In terms of coral cover, even the most well-recovered reefs on Lizard Island need to contain a relatively more diverse range of reef-building corals to support the benthic and mid-water species that not only rely on available coral reef habitat but could beneficially augment their recovery (<xref ref-type="bibr" rid="B38">Schneider et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B15">Harvey et&#xa0;al., 2018</xref>).</p>
<p>As coral reef habitats continue to degrade in the long term, many species that rely on such habitats are likely to suffer heavy population decline (<xref ref-type="bibr" rid="B7">Coker et&#xa0;al., 2009</xref>). Limited habitat complexity&#x2013;combined with smaller, less reproductively-successful adult populations of reef-dwelling species&#x2013;also inhibits recruitment to other reef habitats, which reduces adult replenishment and hinders reef habitat recovery overall (<xref ref-type="bibr" rid="B36">Roth et&#xa0;al., 2018</xref>). Recent research has found that among dominant reef-building corals, <italic>Acropora</italic> recovery and growth is heavily influenced by larval connectivity (<xref ref-type="bibr" rid="B13">Gouezo et&#xa0;al., 2019</xref>). Future studies at my Lizard Island sites could provide a more in-depth understanding of how significant a role larval connectivity plays in overall reef recovery.</p>
</sec>
<sec id="s4_5">
<title>Climate-exacerbated disturbances</title>
<p>Disturbance events are growing in frequency and intensity across many of the world&#x2019;s ecosystems (<xref ref-type="bibr" rid="B4">Buma and Schultz, 2020</xref>; <xref ref-type="bibr" rid="B21">Iwasaki and Noda, 2018</xref>). A better understanding of the impact of such disturbances on various ecosystems not only enhances understanding of the particular system in question, but also contributes to understanding larger global-scale trends and multi-factor processes (<xref ref-type="bibr" rid="B14">Gough et&#xa0;al., 2024</xref>). For example, climate-exacerbated disturbances have altered fire return rates in temperate forests (<xref ref-type="bibr" rid="B9">Dale et&#xa0;al., 2001</xref>), frequency of drought in tropical rainforests (<xref ref-type="bibr" rid="B42">Tao et&#xa0;al., 2022</xref>), rate of sea level rise in mangrove forests (<xref ref-type="bibr" rid="B5">Carugati et&#xa0;al., 2018</xref>), and apparent phase shifts in kelp forests (<xref ref-type="bibr" rid="B41">Smith et&#xa0;al., 2016</xref>).</p>
<p>Coral reefs are among the most productive ecosystems on Earth (<xref ref-type="bibr" rid="B31">Moberg and Folke, 1999</xref>), and their decline is likely to have direct and indirect effects on species diversity, standing biomass, and an array of ecosystem services globally relied on by coastal communities (<xref ref-type="bibr" rid="B17">Hoegh-Guldberg, 1999</xref>). There is particular concern around decreased biodiversity driving reduced ecosystem productivity (<xref ref-type="bibr" rid="B35">Rogers et&#xa0;al., 2018</xref>) as well as ecosystem resistance to future disturbances (<xref ref-type="bibr" rid="B40">Shin et&#xa0;al., 2022</xref>). As coral reefs and other essential ecosystems continue to experience the pressures of climate change, it is crucial that we continue to monitor these changes, especially in the case of disturbances. Changing climate conditions, in combination with other anthropogenic disturbances, are gradually changing the assemblage of coral reefs and their inhabitants (<xref ref-type="bibr" rid="B44">Vanwonterghem and Webster, 2020</xref>). While some species or groups are more resistant under such new conditions, others struggle to recover at the same rate (<xref ref-type="bibr" rid="B30">Miller et&#xa0;al., 2011</xref>). The impacts of such changes will remain uncertain without further monitoring of coral reefs or other productive systems, as described above, to provide more clarity on these community-wide shifts and how they may change conservation or management strategies in the future.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<title>Conclusion</title>
<p>
<italic>Acropora</italic> dominance relative to total live coral cover may turn out to be a useful metric for forecasting future near-term condition of impacted reefs on Lizard Island. To the extent reef managers can influence reef communities, they should consider emphasizing efforts which promote restoration or recruitment of Acroporids. Recovering these key reef builders seems necessary to recover historic conditions as fast as possible. Before the four-year period of consecutive disturbance events, most Lizard Island reefs were relatively live-coral rich, typically with &gt;40% coral cover (<xref ref-type="bibr" rid="B10">Done et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B23">Johns et&#xa0;al., 2014</xref>), but by late 2017 many were close to 4% (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). Between the intense patches of physical damage from the cyclones and the more uniform impact of the bleaching events, these reefs were devastated, with none of the eight study sites reaching more than 20% total coral cover (<xref ref-type="bibr" rid="B28">Madin et&#xa0;al., 2018</xref>). After six years of relatively calmer conditions with fewer conspicuous disturbances, living coral cover at the study sites improved by various margins between 18.4 &#xb1; 0.6 to 59.9 &#xb1; 5.3%, with many sites on the higher end of that spectrum. Aggregate coral cover across reefs at Lizard Island has recovered since the wake of disturbances ending in 2017, with their rate of recovery closely related to the physical environment they inhabit. More regular (annual or biennial) monitoring of each of these sites could also better inform scientific understanding of the long-term recovery of Lizard Island reefs in the face of continual disturbances at the hands of cyclones, bleaching, Crown-of-Thorns Starfish, and other environmental pressures.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>GA: Conceptualization, Data curation, Formal Analysis, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>Thank you to Lizard Island Research Station staff for providing facilities and transportation integral to data collection for this project. In addition, thank you to Dr. Joshua S. Madin for providing essential coral cover data from 1995-2017 at Lizard Island. My undergraduate research was generously supported by a Frost Scholarship via the William &amp; Linda Frost Fund in the Bailey College of Science and Mathematics at California Polytechnic State University San Luis Obispo.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<ref-list>
<title>References</title>
<ref id="B1">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Alvarez-Filip</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Dulvy</surname> <given-names>N. K.</given-names>
</name>
<name>
<surname>Gill</surname> <given-names>J. A.</given-names>
</name>
<name>
<surname>C&#xf4;t&#xe9;</surname> <given-names>I. M.</given-names>
</name>
<name>
<surname>Watkinson</surname> <given-names>A. R.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>Flattening of Caribbean coral reefs: region-wide declines in architectural complexity</article-title>. <source>Proc. R. Soc. B: Biol. Sci.</source> <volume>276(1669)</volume>, <fpage>3019</fpage>&#x2013;<lpage>3025</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1098/rspb.2009.0339</pub-id>
</citation>
</ref>
<ref id="B2">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Baird</surname> <given-names>A. H.</given-names>
</name>
<name>
<surname>Hughes</surname> <given-names>T. P.</given-names>
</name>
</person-group> (<year>2000</year>). <article-title>Competitive dominance by tabular corals: an experimental analysis of recruitment and survival of understorey assemblages</article-title>. <source>J. Exp. Mar. Biol. Ecol.</source> <volume>251</volume>, <fpage>117</fpage>&#x2013;<lpage>132</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/S0022-0981(00)00209-4</pub-id>
</citation>
</ref>
<ref id="B3">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Berzunza-Sanchez</surname> <given-names>M. M.</given-names>
</name>
<name>
<surname>del Carmen Gomez Cabrera</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Pandolfi</surname> <given-names>J. M.</given-names>
</name>
</person-group> (<year>2013</year>). <article-title>Historical patterns of resource exploitation and the status of Papua New Guinea coral reefs1</article-title>. <source>Pacific Sci.</source> <volume>67</volume>, <fpage>425</fpage>&#x2013;<lpage>405</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.2984/67.3.9</pub-id>
</citation>
</ref>
<ref id="B4">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Buma</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Schultz</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Disturbances as opportunities: learning from disturbance-response parallels in social and ecological systems to better adapt to climate change</article-title>. <source>J. Appl. Ecol.</source> <volume>57</volume>, <fpage>1113</fpage>&#x2013;<lpage>1235</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1365-2664.13606</pub-id>
</citation>
</ref>
<ref id="B5">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Carugati</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Gatto</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Rastelli</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Lo Martire</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Coral</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Greco</surname> <given-names>S.</given-names>
</name>
<etal/>
</person-group>. (<year>2018</year>). <article-title>Impact of mangrove forests degradation on biodiversity and ecosystem functioning</article-title>. <source>Sci. Rep.</source> <volume>8</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-018-31683-0</pub-id>
</citation>
</ref>
<ref id="B6">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Cinner</surname> <given-names>J. E.</given-names>
</name>
<name>
<surname>McClanahan</surname> <given-names>T. R.</given-names>
</name>
<name>
<surname>Daw</surname> <given-names>T. M.</given-names>
</name>
<name>
<surname>Graham</surname> <given-names>N. A.J.</given-names>
</name>
<name>
<surname>Maina</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Wilson</surname> <given-names>S. K.</given-names>
</name>
<etal/>
</person-group>. (<year>2009</year>). <article-title>Linking social and ecological systems to sustain coral reef fisheries</article-title>. <source>Curr. Biol.</source> <volume>19</volume>, <fpage>206</fpage>&#x2013;<lpage>125</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.cub.2008.11.055</pub-id>
</citation>
</ref>
<ref id="B7">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Coker</surname> <given-names>D. J.</given-names>
</name>
<name>
<surname>Pratchett</surname> <given-names>M. S.</given-names>
</name>
<name>
<surname>Munday</surname> <given-names>P. L.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>Coral bleaching and habitat degradation increase susceptibility to predation for coral-dwelling fishes</article-title>. <source>Behav. Ecol.</source> <volume>20</volume>, <fpage>1204</fpage>&#x2013;<lpage>1105</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/beheco/arp113</pub-id>
</citation>
</ref>
<ref id="B8">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>D&#x2019;Angelo</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Wiedenmann</surname> <given-names>J.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Impacts of nutrient enrichment on coral reefs: new perspectives and implications for coastal management and reef survival</article-title>. <source>Curr. Opin. Environ. Sustainability</source> <volume>7</volume>, <fpage>82</fpage>&#x2013;<lpage>93</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.cosust.2013.11.029</pub-id>
</citation>
</ref>
<ref id="B9">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Dale</surname> <given-names>V. H.</given-names>
</name>
<name>
<surname>Joyce</surname> <given-names>L. A.</given-names>
</name>
<name>
<surname>McNulty</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Neilson</surname> <given-names>R. P.</given-names>
</name>
<name>
<surname>Ayres</surname> <given-names>M. P.</given-names>
</name>
<name>
<surname>Flannigan</surname> <given-names>M. D.</given-names>
</name>
<etal/>
</person-group>. (<year>2001</year>). <article-title>Climate change and forest disturbances: climate change can affect forests by altering the frequency, intensity, duration, and timing of fire, drought, introduced species, insect and pathogen outbreaks, hurricanes, windstorms, ice storms, or landslides</article-title>. <source>BioScience</source> <volume>51</volume>, <fpage>723</fpage>&#x2013;<lpage>734</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1641/0006-3568(2001)051[0723:CCAFD]2.0.CO;2</pub-id>
</citation>
</ref>
<ref id="B10">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Done</surname> <given-names>T. J.</given-names>
</name>
<name>
<surname>DeVantier</surname> <given-names>L. M.</given-names>
</name>
<name>
<surname>Turak</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Fisk</surname> <given-names>D. A.</given-names>
</name>
<name>
<surname>Wakeford</surname> <given-names>M.</given-names>
</name>
<name>
<surname>van Woesik</surname> <given-names>R.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Coral growth on three reefs: development of recovery benchmarks using a space for time approach</article-title>. <source>Coral Reefs</source> <volume>29</volume>, <fpage>815</fpage>&#x2013;<lpage>833</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00338-010-0637-y</pub-id>
</citation>
</ref>
<ref id="B11">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gardner</surname> <given-names>T. A.</given-names>
</name>
<name>
<surname>C&#xf4;t&#xe9;</surname> <given-names>I. M.</given-names>
</name>
<name>
<surname>Gill</surname> <given-names>J. A.</given-names>
</name>
<name>
<surname>Grant</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Watkinson</surname> <given-names>A. R.</given-names>
</name>
</person-group> (<year>2005</year>). <article-title>Hurricanes and Caribbean coral reefs: impacts, recovery patterns, and role in long-term decline</article-title>. <source>Ecology</source> <volume>86</volume>, <fpage>174</fpage>&#x2013;<lpage>845</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1890/04-0141</pub-id>
</citation>
</ref>
<ref id="B12">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gilmour</surname> <given-names>J. P.</given-names>
</name>
<name>
<surname>Smith</surname> <given-names>L. D.</given-names>
</name>
<name>
<surname>Heyward</surname> <given-names>A. J.</given-names>
</name>
<name>
<surname>Baird</surname> <given-names>A. H.</given-names>
</name>
<name>
<surname>Pratchett</surname> <given-names>M. S.</given-names>
</name>
</person-group> (<year>2013</year>). <article-title>Recovery of an isolated coral reef system following severe disturbance</article-title>. <source>Sci. (New York N.Y.)</source> <volume>340(6128)</volume>, <fpage>69</fpage>&#x2013;<lpage>71</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1126/science.1232310</pub-id>
</citation>
</ref>
<ref id="B13">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gouezo</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Golbuu</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Fabricius</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Olsudong</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Mereb</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Nestor</surname> <given-names>V.</given-names>
</name>
<etal/>
</person-group>. (<year>2019</year>). <article-title>Drivers of recovery and reassembly of coral reef communities</article-title>. <source>Proc. R. Soc. B: Biol. Sci.</source> <volume>286(1897)</volume>, <fpage>20182908</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1098/rspb.2018.2908</pub-id>
</citation>
</ref>
<ref id="B14">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gough</surname> <given-names>C. M.</given-names>
</name>
<name>
<surname>Buma</surname> <given-names>B.</given-names>
</name>
<name>
<surname>Jentsch</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Mathes</surname> <given-names>K. C.</given-names>
</name>
<name>
<surname>Fahey</surname> <given-names>R. T.</given-names>
</name>
</person-group> (<year>2024</year>). <article-title>Disturbance theory for ecosystem ecologists: A primer</article-title>. <source>Ecol. Evol.</source> <volume>14</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/ece3.11403</pub-id>
</citation>
</ref>
<ref id="B15">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Harvey</surname> <given-names>B. J.</given-names>
</name>
<name>
<surname>Nash</surname> <given-names>K. L.</given-names>
</name>
<name>
<surname>Blanchard</surname> <given-names>J. L.</given-names>
</name>
<name>
<surname>Edwards</surname> <given-names>D. P.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Ecosystem-based management of coral reefs under climate change</article-title>. <source>Ecol. Evol.</source> <volume>8</volume>, <fpage>6354</fpage>&#x2013;<lpage>6685</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/ece3.4146</pub-id>
</citation>
</ref>
<ref id="B16">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hench</surname> <given-names>J. L.</given-names>
</name>
<name>
<surname>Leichter</surname> <given-names>J. J.</given-names>
</name>
<name>
<surname>Monismith</surname> <given-names>S. G.</given-names>
</name>
</person-group> (<year>2008</year>). <article-title>Episodic circulation and exchange in a wave-driven coral reef and lagoon system</article-title>. <source>Limnology Oceanography</source> <volume>53</volume>, <fpage>2681</fpage>&#x2013;<lpage>2694</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.4319/lo.2008.53.6.2681</pub-id>
</citation>
</ref>
<ref id="B17">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hoegh-Guldberg</surname> <given-names>O.</given-names>
</name>
</person-group> (<year>1999</year>). <article-title>Climate change, coral bleaching and the future of the world&#x2019;s coral reefs</article-title>. <source>Mar. Freshw. Res.</source> <volume>50</volume>, <fpage>839</fpage>&#x2013;<lpage>866</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1071/mf99078</pub-id>
</citation>
</ref>
<ref id="B18">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hughes</surname> <given-names>T. P.</given-names>
</name>
<name>
<surname>Anderson</surname> <given-names>K. D.</given-names>
</name>
<name>
<surname>Connolly</surname> <given-names>S. R.</given-names>
</name>
<name>
<surname>Heron</surname> <given-names>S. F.</given-names>
</name>
<name>
<surname>Kerry</surname> <given-names>J. T.</given-names>
</name>
<name>
<surname>Lough</surname> <given-names>J. M.</given-names>
</name>
<etal/>
</person-group>. (<year>2018</year>). <article-title>Spatial and temporal patterns of mass bleaching of corals in the anthropocene</article-title>. <source>Science</source> <volume>359</volume>, <fpage>80</fpage>&#x2013;<lpage>83</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1126/science.aan8048</pub-id>
</citation>
</ref>
<ref id="B19">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hughes</surname> <given-names>T. P.</given-names>
</name>
<name>
<surname>Connell</surname> <given-names>J. H.</given-names>
</name>
</person-group> (<year>1999</year>). <article-title>Multiple stressors on coral reefs: A long -term perspective</article-title>. <source>Limnology Oceanography</source> <volume>44</volume>, <fpage>932</fpage>&#x2013;<lpage>940</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.4319/lo.1999.44.3_part_2.0932</pub-id>
</citation>
</ref>
<ref id="B20">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Hughes</surname> <given-names>T. P.</given-names>
</name>
<name>
<surname>Kerry</surname> <given-names>J. T.</given-names>
</name>
<name>
<surname>&#xc1;lvarez-Noriega</surname> <given-names>M.</given-names>
</name>
<name>
<surname>&#xc1;lvarez-Romero</surname> <given-names>J. G.</given-names>
</name>
<name>
<surname>Anderson</surname> <given-names>K. D.</given-names>
</name>
<name>
<surname>Baird</surname> <given-names>A. H.</given-names>
</name>
<etal/>
</person-group>. (<year>2017</year>). <article-title>Global warming and recurrent mass bleaching of corals</article-title>. <source>Nature</source> <volume>543</volume>, <fpage>373</fpage>&#x2013;<lpage>377</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/nature21707</pub-id>
</citation>
</ref>
<ref id="B21">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Iwasaki</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Noda</surname> <given-names>T.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>A framework for quantifying the relationship between intensity and severity of impact of disturbance across types of events and species</article-title>. <source>Sci. Rep.</source> <volume>8</volume>, <fpage>7955</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-017-19048-5</pub-id>
</citation>
</ref>
<ref id="B22">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jentsch</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Beierkuhnlein</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>2008</year>). <article-title>Research frontiers in climate change: effects of extreme meteorological events on ecosystems</article-title>. <source>Comptes Rendus Geoscience Ecosyst&#xe8;mes &#xe9;v&#xe9;nements climatiques extr&#xea;mes</source> <volume>340</volume>, <fpage>621</fpage>&#x2013;<lpage>628</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.crte.2008.07.002</pub-id>
</citation>
</ref>
<ref id="B23">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Johns</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Osborne</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Logan</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Contrasting rates of coral recovery and reassembly in coral communities on the great barrier reef</article-title>. <source>Coral Reefs</source> <volume>33</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00338-014-1148-z</pub-id>
</citation>
</ref>
<ref id="B24">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Jones</surname> <given-names>G. P.</given-names>
</name>
<name>
<surname>McCormick</surname> <given-names>M. I.</given-names>
</name>
<name>
<surname>Srinivasan</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Eagle</surname> <given-names>J. V.</given-names>
</name>
</person-group> (<year>2004</year>). <article-title>Coral decline threatens fish biodiversity in marine reserves</article-title>. <source>Proc. Natl. Acad. Sci. U S A</source> <volume>101</volume>, <fpage>8251</fpage>&#x2013;<lpage>8535</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.0401277101</pub-id>
</citation>
</ref>
<ref id="B25">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Laurans</surname> <given-names>Y.</given-names>
</name>
<name>
<surname>Pascal</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Binet</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Brander</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Clua</surname> <given-names>E.</given-names>
</name>
<name>
<surname>David</surname> <given-names>G.</given-names>
</name>
<etal/>
</person-group>. (<year>2013</year>). <article-title>Economic valuation of ecosystem services from coral reefs in the South Pacific: taking stock of recent experience</article-title>. <source>J. Environ. Manage.</source> <volume>116</volume>, <fpage>135</fpage>&#x2013;<lpage>144</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.jenvman.2012.11.031</pub-id>
</citation>
</ref>
<ref id="B26">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lefcheck</surname> <given-names>J. S.</given-names>
</name>
<name>
<surname>Edgar</surname> <given-names>G. J.</given-names>
</name>
<name>
<surname>Stuart-Smith</surname> <given-names>R. D.</given-names>
</name>
<name>
<surname>Bates</surname> <given-names>A. E.</given-names>
</name>
<name>
<surname>Waldock</surname> <given-names>C.</given-names>
</name>
<name>
<surname>Brandl</surname> <given-names>S. J.</given-names>
</name>
<etal/>
</person-group>. (<year>2021</year>). <article-title>Species richness and identity both determine the biomass of global reef fish communities</article-title>. <source>Nat. Commun.</source> <volume>12</volume>, <fpage>6875</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41467-021-27212-9</pub-id>
</citation>
</ref>
<ref id="B27">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Licuanan</surname> <given-names>W. Y.</given-names>
</name>
<name>
<surname>Robles</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Reyes</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Status and recent trends in coral reefs of the Philippines</article-title>. <source>Mar. pollut. Bull.</source> <volume>142</volume>, <fpage>544</fpage>&#x2013;<lpage>550</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.marpolbul.2019.04.013</pub-id>
</citation>
</ref>
<ref id="B28">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Madin</surname> <given-names>J. S.</given-names>
</name>
<name>
<surname>Baird</surname> <given-names>A. H.</given-names>
</name>
<name>
<surname>Bridge</surname> <given-names>T. C.L.</given-names>
</name>
<name>
<surname>Connolly</surname> <given-names>S. R.</given-names>
</name>
<name>
<surname>Zawada</surname> <given-names>K. J.A.</given-names>
</name>
<name>
<surname>Dornelas</surname> <given-names>M.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Cumulative effects of cyclones and bleaching on coral cover and species richness at Lizard Island</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>604</volume>, <fpage>263</fpage>&#x2013;<lpage>268</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps12735</pub-id>
</citation>
</ref>
<ref id="B29">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Marshall</surname> <given-names>P. A.</given-names>
</name>
<name>
<surname>Baird</surname> <given-names>A. H.</given-names>
</name>
</person-group> (<year>2000</year>). <article-title>Bleaching of corals on the Great Barrier reef: differential susceptibilities among taxa</article-title>. <source>Coral Reefs</source> <volume>19</volume>, <fpage>155</fpage>&#x2013;<lpage>163</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s003380000086</pub-id>
</citation>
</ref>
<ref id="B30">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Miller</surname> <given-names>A. D.</given-names>
</name>
<name>
<surname>Roxburgh</surname> <given-names>S. H.</given-names>
</name>
<name>
<surname>Shea</surname> <given-names>K.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>How frequency and intensity shape diversity&#x2013;disturbance relationships</article-title>. <source>Proc. Natl. Acad. Sci.</source> <volume>108</volume>, <fpage>5643</fpage>&#x2013;<lpage>5485</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.1018594108</pub-id>
</citation>
</ref>
<ref id="B31">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Moberg</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Folke</surname> <given-names>C.</given-names>
</name>
</person-group> (<year>1999</year>). <article-title>Ecological goods and services of coral reef ecosystems</article-title>. <source>Ecol. Economics</source> <volume>29</volume>, <fpage>215</fpage>&#x2013;<lpage>335</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/S0921-8009(99)00009-9</pub-id>
</citation>
</ref>
<ref id="B32">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mumby</surname> <given-names>P. J.</given-names>
</name>
<name>
<surname>Hastings</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Edwards</surname> <given-names>H. J.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Thresholds and the resilience of Caribbean coral reefs</article-title>. <source>Nature</source> <volume>450(7166)</volume>, <fpage>98</fpage>&#x2013;<lpage>101</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/nature06252</pub-id>
</citation>
</ref>
<ref id="B33">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Pratchett</surname> <given-names>M. S.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Changes in Coral Assemblages during an Outbreak of Acanthaster Planci at Lizard Island, Northern Great Barrier Reef, (1995&#x2013;1999)</article-title>. <source>Coral Reefs</source> <volume>29</volume>, <fpage>717</fpage>&#x2013;<lpage>725</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00338-010-0602-9</pub-id>
</citation>
</ref>
<ref id="B34">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Roff</surname> <given-names>G.</given-names>
</name>
<name>
<surname>Kvennefors</surname> <given-names>E.C. E.</given-names>
</name>
<name>
<surname>Fine</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Ortiz</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Davy</surname> <given-names>J. E.</given-names>
</name>
<name>
<surname>Hoegh-Guldberg</surname> <given-names>O.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>The ecology of &#x2018;Acroporid white syndrome&#x2019;, a coral disease from the southern great barrier reef</article-title>. <source>PLoS One</source> <volume>6</volume>, <elocation-id>e268295</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1371/journal.pone.0026829</pub-id>
</citation>
</ref>
<ref id="B35">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rogers</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Blanchard</surname> <given-names>J. L.</given-names>
</name>
<name>
<surname>Mumby</surname> <given-names>P. J.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Fisheries productivity under progressive coral reef degradation</article-title>. <source>J. Appl. Ecol.</source> <volume>55</volume>, <fpage>1041</fpage>&#x2013;<lpage>1495</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1365-2664.13051</pub-id>
</citation>
</ref>
<ref id="B36">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Roth</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Saalmann</surname> <given-names>F.</given-names>
</name>
<name>
<surname>Thomson</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Coker</surname> <given-names>D. J.</given-names>
</name>
<name>
<surname>Villalobos</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Jones</surname> <given-names>B. H.</given-names>
</name>
<etal/>
</person-group>. (<year>2018</year>). <article-title>Coral reef degradation affects the potential for reef recovery after disturbance</article-title>. <source>Mar. Environ. Res.</source> <volume>142</volume>, <fpage>48</fpage>&#x2013;<lpage>58</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.marenvres.2018.09.022</pub-id>
</citation>
</ref>
<ref id="B37">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Santavy</surname> <given-names>D. L.</given-names>
</name>
<name>
<surname>Horstmann</surname> <given-names>C. L.</given-names>
</name>
<name>
<surname>Sharpe</surname> <given-names>L. M.</given-names>
</name>
<name>
<surname>Yee</surname> <given-names>S. H.</given-names>
</name>
<name>
<surname>Ringold</surname> <given-names>P.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>What is it about coral reefs? Translation of ecosystem goods and services relevant to people and their well-being</article-title>. <source>Ecosphere (Washington D.C)</source> <volume>12</volume>, <fpage>1</fpage>&#x2013;<lpage>275</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/ecs2.3639</pub-id>
</citation>
</ref>
<ref id="B38">
<citation citation-type="web">
<person-group person-group-type="author">
<name>
<surname>Schneider</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Silverman</surname> <given-names>J.</given-names>
</name>
<name>
<surname>Woolsey</surname> <given-names>E.</given-names>
</name>
<name>
<surname>Eriksson</surname> <given-names>H.</given-names>
</name>
<name>
<surname>Byrne</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Caldeira</surname> <given-names>K.</given-names>
</name>
</person-group> <article-title>Potential influence of sea cucumbers on coral reef CaCO3 budget: A case study at one tree reef</article-title> (Accessed <access-date>September 3, 2024</access-date>).</citation>
</ref>
<ref id="B39">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Sherman</surname> <given-names>C.S.</given-names>
</name>
<name>
<surname>Simpfendorfer</surname> <given-names>C. A.</given-names>
</name>
<name>
<surname>Pacoureau</surname> <given-names>N.</given-names>
</name>
<name>
<surname>Matsushiba</surname> <given-names>J. H.</given-names>
</name>
<name>
<surname>Yan</surname> <given-names>H. F.</given-names>
</name>
<name>
<surname>Walls</surname> <given-names>R. H.L.</given-names>
</name>
<etal/>
</person-group>. (<year>2023</year>). <article-title>Half a century of rising extinction risk of coral reef sharks and rays</article-title>. <source>Nat. Commun.</source> <volume>14</volume>, <fpage>15</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41467-022-35091-x</pub-id>
</citation>
</ref>
<ref id="B40">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Shin</surname> <given-names>Y.-J.</given-names>
</name>
<name>
<surname>Midgley</surname> <given-names>G. F.</given-names>
</name>
<name>
<surname>Archer</surname> <given-names>E. R.M.</given-names>
</name>
<name>
<surname>Arneth</surname> <given-names>A.</given-names>
</name>
<name>
<surname>Barnes</surname> <given-names>D. K.A.</given-names>
</name>
<name>
<surname>Chan</surname> <given-names>L.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Actions to halt biodiversity loss generally benefit the climate</article-title>. <source>Global Change Biol.</source> <volume>28</volume>, <fpage>2846</fpage>&#x2013;<lpage>2874</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/gcb.16109</pub-id>
</citation>
</ref>
<ref id="B41">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Smith</surname> <given-names>J. G.</given-names>
</name>
<name>
<surname>Malone</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Carr</surname> <given-names>M. H.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>Consequences of kelp forest ecosystem shifts and predictors of persistence through multiple stressors</article-title>. <source>Proc. R. Soc. B: Biol. Sci.</source> <volume>291</volume>, <fpage>20232749</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1098/rspb.2023.2749</pub-id>
</citation>
</ref>
<ref id="B42">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Tao</surname> <given-names>S.</given-names>
</name>
<name>
<surname>Chave</surname> <given-names>J&#xe9;r&#xf4;me</given-names>
</name>
<name>
<surname>Frison</surname> <given-names>P.-L.</given-names>
</name>
<name>
<surname>Le Toan</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Ciais</surname> <given-names>P.</given-names>
</name>
<name>
<surname>Fang</surname> <given-names>J.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Increasing and widespread vulnerability of intact tropical rainforests to repeated droughts</article-title>. <source>Proc. Natl. Acad. Sci. U S A</source> <volume>119</volume>, <elocation-id>e2116626119</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.2116626119</pub-id>
</citation>
</ref>
<ref id="B43">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>van Woesik</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Shlesinger</surname> <given-names>T.</given-names>
</name>
<name>
<surname>Grottoli</surname> <given-names>Andr&#xe9;aG.</given-names>
</name>
<name>
<surname>Toonen</surname> <given-names>R. J.</given-names>
</name>
<name>
<surname>Vega Thurber</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Warner</surname> <given-names>M. E.</given-names>
</name>
<etal/>
</person-group>. (<year>2022</year>). <article-title>Coral-bleaching responses to climate change across biological scales</article-title>. <source>Global Change Biol.</source> <volume>28</volume>, <fpage>4229</fpage>&#x2013;<lpage>4250</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/gcb.16192</pub-id>
</citation>
</ref>
<ref id="B44">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Vanwonterghem</surname> <given-names>I.</given-names>
</name>
<name>
<surname>Webster</surname> <given-names>N. S.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Coral reef microorganisms in a changing climate</article-title>. <source>iScience</source> <volume>23</volume>, <fpage>1009725</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.isci.2020.100972</pub-id>
</citation>
</ref>
<ref id="B45">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wakeford</surname> <given-names>M.</given-names>
</name>
<name>
<surname>Done</surname> <given-names>T. J.</given-names>
</name>
<name>
<surname>Johnson</surname> <given-names>C. R.</given-names>
</name>
</person-group> (<year>2008</year>). <article-title>Decadal trends in a coral community and evidence of changed disturbance regime</article-title>. <source>Coral Reefs</source> <volume>27</volume>, <fpage>1</fpage>&#x2013;<lpage>13</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/s00338-007-0284-0</pub-id>
</citation>
</ref>
<ref id="B46">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wilson</surname> <given-names>S. K.</given-names>
</name>
<name>
<surname>Graham</surname> <given-names>N. a. J.</given-names>
</name>
<name>
<surname>Pratchett</surname> <given-names>M. S.</given-names>
</name>
<name>
<surname>Jones</surname> <given-names>G. P.</given-names>
</name>
<name>
<surname>Polunin</surname> <given-names>N. V.C.</given-names>
</name>
</person-group> (<year>2006</year>). <article-title>Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient</article-title>? <source>Global Change Biol.</source> <volume>12</volume>, <fpage>2220</fpage>&#x2013;<lpage>2345</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/j.1365-2486.2006.01252.x</pub-id>
</citation>
</ref>
<ref id="B47">
<citation citation-type="other">
<person-group person-group-type="author">
<name>
<surname>Wolfe</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Anthony</surname> <given-names>K.</given-names>
</name>
<name>
<surname>Babcock</surname> <given-names>R.</given-names>
</name>
<name>
<surname>Bay</surname> <given-names>L.</given-names>
</name>
<name>
<surname>Bourne</surname> <given-names>D.</given-names>
</name>
<name>
<surname>Burrows</surname> <given-names>D.</given-names>
</name>
<etal/>
</person-group>. (<year>2020</year>), <fpage>179</fpage>&#x2013;<lpage>318</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1201/9780429351495-5</pub-id>
</citation>
</ref>
</ref-list>
</back>
</article>