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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2022.895906</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>The Impact of Climate Change on Potential Invasion Risk of <italic>Oryctes monoceros</italic> Worldwide</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Aidoo</surname> <given-names>Owusu Fordjour</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="author-notes" rid="fn002"><sup>&#x2020;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1719979/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Hao</surname> <given-names>Mengmeng</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="author-notes" rid="fn002"><sup>&#x2020;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1743529/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Ding</surname> <given-names>Fangyu</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1416002/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Wang</surname> <given-names>Di</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Jiang</surname> <given-names>Dong</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/116224/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Ma</surname> <given-names>Tian</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="corresp" rid="c002"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1724880/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Qian</surname> <given-names>Yushu</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Tettey</surname> <given-names>Elizabeth</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Yankey</surname> <given-names>Ndede</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Dadzie Ninsin</surname> <given-names>Kodwo</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Borgemeister</surname> <given-names>Christian</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Department of Biological, Physical and Mathematical Sciences, School of Natural and Environmental Sciences, University of Environment and Sustainable Development</institution>, <addr-line>Somanya</addr-line>, <country>Ghana</country></aff>
<aff id="aff2"><sup>2</sup><institution>State Key Laboratory of Resources and Environmental Information Systems, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>College of Resources and Environment, University of Chinese Academy of Sciences</institution>, <addr-line>Beijing</addr-line>, <country>China</country></aff>
<aff id="aff4"><sup>4</sup><institution>Council for Scientific and Industrial Research (CSIR), Oil Palm Research Institute, Coconut Research Programme</institution>, <addr-line>Sekondi</addr-line>, <country>Ghana</country></aff>
<aff id="aff5"><sup>5</sup><institution>Centre for Development Research (ZEF), University of Bonn</institution>, <addr-line>Bonn</addr-line>, <country>Germany</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Mark Maboeta, North-West University, South Africa</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Ji-Zhong Wan, Qinghai University, China; Matan Shelomi, National Taiwan University, Taiwan</p></fn>
<corresp id="c001">&#x002A;Correspondence: Fangyu Ding, <email>dingfy@igsnrr.ac.cn</email></corresp>
<corresp id="c002">Tian Ma, <email>mat.19b@igsnrr.ac.cn</email></corresp>
<fn fn-type="equal" id="fn002"><p><sup>&#x2020;</sup>These authors have contributed equally to this work</p></fn>
<fn fn-type="other" id="fn004"><p>This article was submitted to Conservation and Restoration Ecology, a section of the journal Frontiers in Ecology and Evolution</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>06</day>
<month>07</month>
<year>2022</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>10</volume>
<elocation-id>895906</elocation-id>
<history>
<date date-type="received">
<day>14</day>
<month>03</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>15</day>
<month>06</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2022 Aidoo, Hao, Ding, Wang, Jiang, Ma, Qian, Tettey, Yankey, Dadzie Ninsin and Borgemeister.</copyright-statement>
<copyright-year>2022</copyright-year>
<copyright-holder>Aidoo, Hao, Ding, Wang, Jiang, Ma, Qian, Tettey, Yankey, Dadzie Ninsin and Borgemeister</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>As a significant threat to agriculture, pests have caused a great disservice to crop production and food security. Understanding the mechanisms of pests&#x2019; outbreaks and invasion is critical in giving sound suggestions on their control and prevention strategies. The African rhinoceros beetle, <italic>Oryctes monoceros</italic> (Olivier), as the most damaging pest of palms, banana, sugarcane, and pineapple, severely threatens their production due to its ability to kill both young and matured hosts. Analyzing the effect of climate change on major parameters of <italic>O. monoceros</italic> life history has been an important issue recently, given its sensitivity to thermal conditions. However, information on how climate change alters geographical distribution of <italic>O. monoceros</italic> is poorly understood. By combining environmental variables and occurrence records, we were able to assess environmental risk factors for <italic>O. monoceros</italic> and create risk maps for the pest using the Boosted Regression Tree model. Our results significance of environmental variables showed that the annual temperature variation (39.45%), seasonality of temperature (23.00%), the isothermality (18.76%), precipitation of the hottest quarter months (6.07%), average variation of day time temperature (3.27%), were relatively important environmental factors that affected the distribution <italic>O. monoceros</italic>. We also found that the projected potential distributions of the pest&#x2019;s habitats in all future global warming scenarios exceeded its present known distribution. The model predicts that habitat suitability for <italic>O. monoceros</italic> is predominantly concentrated along Africa&#x2019;s west and east coastlines, Asia&#x2019;s south coasts, South America&#x2019;s north and east coasts, and a few locations spread over North America&#x2019;s southern coasts and coastal regions. These outputs provide a solid theoretical foundation for <italic>O. monoceros</italic> risk evaluations and control.</p>
</abstract>
<kwd-group>
<kwd>African rhinoceros beetle</kwd>
<kwd>Boosted Regression Trees</kwd>
<kwd>BRT</kwd>
<kwd>climate change</kwd>
<kwd><italic>Oryctes monoceros</italic></kwd>
</kwd-group>
<counts>
<fig-count count="3"/>
<table-count count="3"/>
<equation-count count="0"/>
<ref-count count="85"/>
<page-count count="10"/>
<word-count count="7353"/>
</counts>
</article-meta>
</front>
<body>
<sec id="S1" sec-type="intro">
<title>Introduction</title>
<p>The African rhinoceros beetle, <italic>Oryctes monoceros</italic> (Olivier; Coleoptera: Scarabaeidae), as a serious pest of palms such as <italic>Phoenix dactylifera</italic> L. (<xref ref-type="bibr" rid="B48">Kabiru et al., 2014</xref>), <italic>Elaeis guineensis</italic>, Jacq. (<xref ref-type="bibr" rid="B15">Bedford, 2014</xref>), and <italic>Cocos nucifera</italic> L., is responsible for about 40% of coconut losses in tropical Africa (<xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>). Apart from palms, <italic>O. monoceros</italic> attacks non-palm hosts, such as <italic>Saccharum officinarum</italic> L., <italic>Ananas comosus</italic> L., and <italic>Musa paradisiaca</italic> L. <italic>O. monoceros</italic> inflicts damage to palms in both direct and indirect ways. The beetle directly stunts the growth of the palms by boring and laying eggs in the soft growing points (<xref ref-type="bibr" rid="B14">Bedford, 2013</xref>). Indirectly, <italic>O. monoceros</italic> feeding exposes palms to secondary infestation by the African palm weevil, <italic>Rhynchophorus phoenicis</italic> L., which enters the palms via holes or galleries created by <italic>O. monoceros</italic>. At present, <italic>O. monoceros</italic> has been reported primarily in Africa (<xref ref-type="bibr" rid="B67">Philippe and Dery, 2004</xref>; <xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>; <xref ref-type="bibr" rid="B25">Edijala et al., 2009</xref>; <xref ref-type="bibr" rid="B16">Bila et al., 2019</xref>; <xref ref-type="bibr" rid="B28">EPPO, 2021</xref>), and Middle East (<xref ref-type="bibr" rid="B8">Arrow, 1937</xref>; <xref ref-type="bibr" rid="B62">NHM, 1938</xref>; <xref ref-type="bibr" rid="B55">Lepesme, 1947</xref>; <xref ref-type="bibr" rid="B28">EPPO, 2021</xref>). However, predictions of potential suitable climate regions for preventive and quarantine measures are generally lacking for <italic>O. monoceros</italic>.</p>
<p>There are over 40 Oryctes species worldwide (<xref ref-type="bibr" rid="B11">Bedford, 1976</xref>). Despite the species&#x2019; similarities, morphometric features on different life stages, such as adults and larvae, have been critical in distinguishing these species. For example, in <italic>O. monoceros</italic>, the anteroventral border of the prothoracic shield is rectilinear, whereas in <italic>Oryctes rhinoceros</italic> Linnaeus, it is scalloped and more concave (<xref ref-type="bibr" rid="B42">Hurpin and Fresneau, 1970</xref>). Adults of <italic>O. rhinoceros</italic> are gregarious, however just a single <italic>O. monoceros</italic> is seen feeding on the growing point of palms in most occasions (<xref ref-type="bibr" rid="B67">Philippe and Dery, 2004</xref>). <italic>O. monoceros</italic> is found across Africa, whereas <italic>O. rhinoceros</italic> is widely distributed in South and Southeast Asia (<xref ref-type="bibr" rid="B28">EPPO, 2021</xref>). <italic>O. monoceros</italic> differs from other Oryctes species like <italic>Oryctes boas</italic> Fabricius and <italic>Oryctes gigas</italic> Verkr&#x00FC;zen by having a third tooth on the right mandible between the scissorial and molar sections. Furthermore, <italic>O. monoceros</italic> has raster teges with 78&#x2013;120 small sharp setae and a lower anal lip with 60&#x2013;80 similar setae (<xref ref-type="bibr" rid="B12">Bedford, 1979</xref>). On the other hand, each claw of <italic>O. gigas</italic> has three to five long setae, whereas each claw of <italic>O. boas</italic> has two long setae. The diameter of the <italic>O. boas</italic> prothoracic shield is almost equal to its length, according to <xref ref-type="bibr" rid="B41">Hurpin (1969)</xref>, with rectilinear and parallel anterior and posterior edges.</p>
<p>Changes in the earth&#x2019;s temperature due to an increase in greenhouse gases have been shown to alter the distribution of species (<xref ref-type="bibr" rid="B22">Dhaliwal et al., 2010</xref>; <xref ref-type="bibr" rid="B61">Netherer and Schopf, 2010</xref>). Insect life cycles and population dynamics will undoubtedly be affected by global warming, including their growth rate, voltinism, and dispersal patterns (<xref ref-type="bibr" rid="B10">Bale et al., 2002</xref>), which influence the distribution and seasonal activity of species (<xref ref-type="bibr" rid="B37">Halsch et al., 2021</xref>). It can also induce changes in the habitat suitability of agricultural pests, thereby increasing invasion or exacerbating outbreaks and eventually optimizing the likelihood of damage to food production and food security.</p>
<p>Environmental niche models (ENMs) combine species occurrence records and environmental datasets, build a model using a machine learning algorithm, simulate environmental requirements of species, and project the analysis at a different time and space to predict its habitat suitability (<xref ref-type="bibr" rid="B82">Warren and Seifert, 2011</xref>; <xref ref-type="bibr" rid="B80">Valencia-Rodr&#x00ED;guez et al., 2021</xref>). Environmental niche modeling uses correlation and mechanistic or process-based models to estimate species&#x2019; habitat requirements. The mechanistic species distribution model (e.g., CLIMEX) considers how environmental conditions constrain the physiological characteristics of species at a given location (<xref ref-type="bibr" rid="B57">Louvrier et al., 2020</xref>). In contrast, correlative models establish mathematical correlations between observable species distributions (presence or absence) and environmental variables [e.g., Maximum Entropy, Boosted Regression Trees (BRT), and Random Forest (RF)] (<xref ref-type="bibr" rid="B68">Phillips et al., 2006</xref>; <xref ref-type="bibr" rid="B83">Yu et al., 2020</xref>). Each model is theoretically distinct, has distinct data requirements, and employs unique analytic techniques. However, the BRT model is considered as a more robust technique than the other ecological niche models (<xref ref-type="bibr" rid="B83">Yu et al., 2020</xref>). The model works well with large datasets or when there are a lot of environmental factors compared to the number of observations (<xref ref-type="bibr" rid="B27">Elith et al., 2008</xref>). No prior data modification or outlier removal is required to fit the BRT model&#x2019;s complicated non-linear relationships and automatically manage predictor interaction effects (<xref ref-type="bibr" rid="B21">De&#x2019;Ath, 2007</xref>). The main limitation of single tree models is its low predictive powers, which can be improved by using BRT to fit multiple trees (<xref ref-type="bibr" rid="B26">Elith and Leathwick, 2017</xref>). The BRT model has been used to identify habitat suitability for <italic>Aedes aegypti</italic> and <italic>Aedes albopictus</italic> (the arbovirus vectors), based on land-cover and environmental variables (<xref ref-type="bibr" rid="B53">Kraemer et al., 2015</xref>). The model has also been used to model a mountain pine beetle distribution (<xref ref-type="bibr" rid="B70">Ramazi et al., 2021</xref>). The output from such models provides baseline information for monitoring, surveillance, and development of ecologically friendly management strategies (<xref ref-type="bibr" rid="B60">Mu et al., 2013</xref>).</p>
<p>Several studies on <italic>O. monoceros</italic> have focused on its biology and ecology (<xref ref-type="bibr" rid="B79">Ukeh et al., 2003</xref>; <xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>; <xref ref-type="bibr" rid="B25">Edijala et al., 2009</xref>; <xref ref-type="bibr" rid="B48">Kabiru et al., 2014</xref>; <xref ref-type="bibr" rid="B33">Gbangboche et al., 2016</xref>), and cultural management practices, such as farm sanitation by destroying felled wood and palms or covering them with cover crops to minimize larval breeding (<xref ref-type="bibr" rid="B58">Mariau and Calvez, 1973</xref>).</p>
<p>However, due to the high planting density of palms and the expensive expense of these management measures, they are rarely adopted (<xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>). Attempts to manage the pests using biological control have been unsuccessful because none of the parasitoids or pathogens (<italic>Baculovirus oryctes</italic>) have provided a long-lasting solution to the threat posed by <italic>O. monoceros</italic> (<xref ref-type="bibr" rid="B73">Seguni et al., 1999</xref>). In addition, application of synthetic pesticides has not been recommended due to environmental and health risks (<xref ref-type="bibr" rid="B66">Parven et al., 2021</xref>). In Ghana, a new method for controlling <italic>O. monoceros</italic> in coconut plantations is extracting adults using a metal mini-harpoon from their feeding galleries (<xref ref-type="bibr" rid="B67">Philippe and Dery, 2004</xref>). This mechanical control strategy is labor-intensive and costly in high-density palm plantations. Also, since preventive measures are economically cheaper than post entry management (<xref ref-type="bibr" rid="B9">Arthur et al., 2015</xref>), creating risk maps would serve as an early warning against the future spread, especially in areas where the pest is currently absent.</p>
<p>The average global temperatures would rise ranging from 2 and 9.7&#x00B0;F (1.1 to 5.4&#x00B0;C) by 2100 in a variety of greenhouse gas emission scenarios (<xref ref-type="bibr" rid="B39">Herring, 2012</xref>). In recent decades, it is accepted that a rise in global temperature is considered as one of the causes of the drastic change in climate (<xref ref-type="bibr" rid="B78">Soytas and Sari, 2006</xref>; <xref ref-type="bibr" rid="B49">Karki, 2007</xref>). Climate change is an important factor restricting insects&#x2019; survival and population growth (<xref ref-type="bibr" rid="B81">Vanhanen et al., 2007</xref>). However, there is a scarcity of information on the effects of climate change on the global distribution of <italic>O. monoceros</italic> as well as identifying environmental factors driving the geographical distribution for surveillance, monitoring and preventive programs is generally lacking. To address this research gap, we simulated the potential distribution of <italic>O. monoceros</italic> under different climate change scenarios to identify climate suitable areas for biosecurity and risk assessment.</p>
</sec>
<sec id="S2" sec-type="materials|methods">
<title>Materials and Methods</title>
<p>Our research included two key procedures: (i) obtaining occurrence records and climate datasets; and (ii) predicting the pest&#x2019;s existing and future potential environmental niche shifts. The BRT modeling framework is a prominent technique to ecological studies that has been used to accurately forecast the potential geographic range of many species (<xref ref-type="bibr" rid="B46">Jiang et al., 2019</xref>; <xref ref-type="bibr" rid="B52">Kraemer et al., 2019</xref>; <xref ref-type="bibr" rid="B20">Chen et al., 2020</xref>; <xref ref-type="bibr" rid="B23">Ding et al., 2020</xref>). The BRT modeling approach combines the advantages of regression trees (<xref ref-type="bibr" rid="B17">Breiman et al., 1984</xref>) and gradient boosting (<xref ref-type="bibr" rid="B32">Friedman, 2001</xref>), and has a good ability in fitting complex non-linear response functions from outcomes based on given environmental covariates. To assess the effects of climate change on environmental suitability for <italic>O. monoceros</italic>, we used the following datasets as key input data for modeling analysis: (a) the assembled contemporary occurrence records of <italic>O. monoceros</italic>, and a set of pseudo-absence points of pest; (b) a set of environmental covariates reflecting a baseline scenario for the year 2020; and (c) a set of environmental covariates reflecting two representative concentration pathways (RCPs) (R 4.5 and 8.5) scenarios on future periods (the 2030s, 2050s, and 2080s).</p>
<sec id="S2.SS1">
<title>Occurrence Records and Pseudo-Absence Points</title>
<p>A 3-year nationwide survey for <italic>O. monoceros</italic> was conducted in palm plantations in Ghana to obtain the pest&#x2019;s occurrence records. The sampling consisted of visual observation for the presence of any developmental stage (eggs, larvae, pupae, and adults), as well as symptomatic fronds, feeding galleries, and debris. A handheld GPS device (Garmin eTrex 22x), was used to obtain GPS coordinates of locations where <italic>O. monoceros</italic> was collected in the field. Having enough data points for accurate modeling is critical for optimal model performance, therefore the field data were supplemented by a comprehensive scientific literature search using internet sources, such as Web of Science, Science Direct, Google Scholar, and PubMed (<xref ref-type="bibr" rid="B8">Arrow, 1937</xref>; <xref ref-type="bibr" rid="B62">NHM, 1938</xref>; <xref ref-type="bibr" rid="B55">Lepesme, 1947</xref>; <xref ref-type="bibr" rid="B47">Julia and Mariau, 1976</xref>; <xref ref-type="bibr" rid="B56">Lomer, 1986</xref>; <xref ref-type="bibr" rid="B35">Gries et al., 1994</xref>; <xref ref-type="bibr" rid="B73">Seguni et al., 1999</xref>; <xref ref-type="bibr" rid="B79">Ukeh et al., 2003</xref>; <xref ref-type="bibr" rid="B67">Philippe and Dery, 2004</xref>; <xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>, <xref ref-type="bibr" rid="B7">2008</xref>, <xref ref-type="bibr" rid="B5">2012</xref>; <xref ref-type="bibr" rid="B25">Edijala et al., 2009</xref>; <xref ref-type="bibr" rid="B14">Bedford, 2013</xref>; <xref ref-type="bibr" rid="B48">Kabiru et al., 2014</xref>; <xref ref-type="bibr" rid="B33">Gbangboche et al., 2016</xref>; <xref ref-type="bibr" rid="B75">S&#x00E9;r&#x00E9; et al., 2018</xref>; <xref ref-type="bibr" rid="B16">Bila et al., 2019</xref>; <xref ref-type="bibr" rid="B43">Idowu et al., 2019</xref>). Google Earth Pro version 7.3 was used to extract the coordinates (i.e., latitudes and longitudes) and elevation information if only locations were provided. In addition, we manually deleted redundant and ambiguous datasets, and those with evident geocoding issues. Finally, 323 <italic>O. monoceros</italic> records were discovered worldwide (<xref ref-type="supplementary-material" rid="TS1">Supplementary Table 1</xref>). An equivalent number of pseudo-absence points were randomly selected from a previously described approach (<xref ref-type="bibr" rid="B74">Senay et al., 2013</xref>). The following steps were used to obtain the pseudo-absence records; to begin with, the pseudo-absence locations were randomly selected from the background points. In addition, the pseudo-absence locations were created within a delimited geographical distance from recorded occurrence points. Finally, we selected pseudo-absence locations with environmentally dissimilar conditions from the occurrence points.</p>
</sec>
<sec id="S2.SS2">
<title>Environmental Variables</title>
<p>The environmental factors chosen as the starting variables to be used in the modeling in this study are shown in <xref ref-type="table" rid="T1">Table 1</xref>, for the various reasons described above. The environmental variables, which included 19 bioclimatic variables, were downloaded with a spatial resolution of 5 km from the WorldClim Global environmental database.<sup><xref ref-type="fn" rid="footnote1">1</xref></sup> In its AR5 assessment, the Intergovernmental Panel on Climate Change (IPCC) chose four RCPs to depict the future climate scenario, namely RCPs 2.6, 4.5, 6.0, and RCP8.5 (<xref ref-type="bibr" rid="B71">Remya et al., 2015</xref>). The future environmental variables generated by the MIROC-ESM-CHEM model for two climatic scenarios (RCPs 4.5 and 8.5) were taken from the Climatic Change, Agriculture and Food Security website.<sup><xref ref-type="fn" rid="footnote2">2</xref></sup></p>
<table-wrap position="float" id="T1">
<label>TABLE 1</label>
<caption><p>The environmental variables with code and units used for the present study.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Code</td>
<td valign="top" align="left">Environmental variable</td>
<td valign="top" align="center">Unit</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Bio1</td>
<td valign="top" align="left">Annual average temperature</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio2</td>
<td valign="top" align="left">Average variation of day time temperature</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio3</td>
<td valign="top" align="left">Isothermality</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio4</td>
<td valign="top" align="left">Seasonality of temperature (SD &#x00D7; 100)</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio5</td>
<td valign="top" align="left">Highest temperature of the hottest month</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio6</td>
<td valign="top" align="left">Lowest temperature of the coldest month</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio7</td>
<td valign="top" align="left">Annual temperature variation</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio8</td>
<td valign="top" align="left">Average temperature of the rainy quarter months</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio9</td>
<td valign="top" align="left">Average temperature of the driest quarter months</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio10</td>
<td valign="top" align="left">Average temperature of the hottest quarter months</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio11</td>
<td valign="top" align="left">Average temperature of the coldest quarter months</td>
<td valign="top" align="center">&#x00B0;C</td>
</tr>
<tr>
<td valign="top" align="left">Bio12</td>
<td valign="top" align="left">Annual precipitation</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio13</td>
<td valign="top" align="left">Precipitation of the rainiest month</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio14</td>
<td valign="top" align="left">Precipitation of the driest month</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio15</td>
<td valign="top" align="left">Precipitation seasonality</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio16</td>
<td valign="top" align="left">Precipitation of the rainiest quarter months</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bi017</td>
<td valign="top" align="left">Precipitation of the driest quarter months</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio18</td>
<td valign="top" align="left">Precipitation of the hottest quarter months</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left">Bio19</td>
<td valign="top" align="left">Precipitation of the coldest quarter months</td>
<td valign="top" align="center">mm</td>
</tr>
<tr>
<td valign="top" align="left"/>
<td valign="top" align="left">Elevation</td>
<td valign="top" align="center">m</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="S2.SS3">
<title>Modeling Analysis</title>
<p>The &#x201C;dismo&#x201D; package was used to run the modeling analysis process in the R 3.3.3 statistical programming environment. Also, recommendations from scientific literature were consulted for the BRT model projections (<xref ref-type="bibr" rid="B46">Jiang et al., 2019</xref>, <xref ref-type="bibr" rid="B45">2021</xref>; <xref ref-type="bibr" rid="B84">Zheng et al., 2019</xref>). This allowed us to specify key BRT model parameter values. As an initial step, we used <italic>O. monoceros</italic> records and pseudo-absence points as well as environmental variables to create a baseline scenario for 2020. Pseudo-absence points were minimized by constructing an ensemble of 30 BRT models, and this helped make our modeling technique more resilient. Then, the BRT models were merged with a set of future environmental covariates to generate <italic>O. monoceros</italic> environmental suitability maps for the future. By averaging 30 BRT models, the maps of the world&#x2019;s future geographic distribution were created. In order to avoid overfitting, a tenfold cross-validation area under the receiver operating characteristic curve (ROC-AUC) was chosen. The ROC curve and the area under the ROC curve were used to evaluate the model&#x2019;s accuracy. AUC values range between 0 and 1. As a general rule, an AUC less than 0.9 implies that a model&#x2019;s prediction accuracy is low, while an AUC more than 0.9 suggests that the model&#x2019;s accuracy is outstanding. As this value approaches 1, the model&#x2019;s performance improves (<xref ref-type="bibr" rid="B68">Phillips et al., 2006</xref>). We utilized the ArcGIS software&#x2019;s inbuilt Tabulate Area tool (version 10.1) to calculate the habitat suitability for <italic>O. monoceros</italic>.</p>
</sec>
</sec>
<sec id="S3" sec-type="results">
<title>Results</title>
<sec id="S3.SS1">
<title>Performance of the Model and the Significance of Environmental Variables</title>
<p>The <italic>O. monoceros</italic> model prediction accuracy was shown to be &#x201C;excellent&#x201D; for the current period (AUC mean = 0.992). The results showed that the variables chosen correctly predicted the current <italic>O. monoceros</italic> distribution. <xref ref-type="table" rid="T1">Table 1</xref> listed the relative contribution of the environmental variables, which showed that the annual temperature variation contributed most to the model (39.45 &#x00B1; 11.40%), followed by the seasonality of temperature (23.00% &#x00B1; 8.5%), the isothermality (18.76% &#x00B1; 12%), precipitation of the hottest quarter months (6.07% &#x00B1; 3.46%), and average variation of day time temperature (3.27% &#x00B1; 1.95%) (<xref ref-type="table" rid="T2">Table 2</xref>). The other variables accounted for less than 1% of the total contribution.</p>
<table-wrap position="float" id="T2">
<label>TABLE 2</label>
<caption><p>Environmental variables&#x2019; proportional contributions to predicting global habitat suitability for <italic>O. monoceros</italic> occurrence.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Environmental variables</td>
<td valign="top" align="center">Mean (%)</td>
<td valign="top" align="center">SD</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Bio7</td>
<td valign="top" align="center">39.445</td>
<td valign="top" align="center">11.401</td>
</tr>
<tr>
<td valign="top" align="left">Bio4</td>
<td valign="top" align="center">23.008</td>
<td valign="top" align="center">8.501</td>
</tr>
<tr>
<td valign="top" align="left">Bio6</td>
<td valign="top" align="center">18.756</td>
<td valign="top" align="center">11.998</td>
</tr>
<tr>
<td valign="top" align="left">Bio3</td>
<td valign="top" align="center">6.071</td>
<td valign="top" align="center">3.458</td>
</tr>
<tr>
<td valign="top" align="left">Bio18</td>
<td valign="top" align="center">3.271</td>
<td valign="top" align="center">1.950</td>
</tr>
<tr>
<td valign="top" align="left">Bio2</td>
<td valign="top" align="center">1.489</td>
<td valign="top" align="center">0.617</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="S3.SS2">
<title>Global Potential Distribution <italic>Oryctes monoceros</italic></title>
<p>Under the current time, the prediction of habitat suitability for <italic>O. monoceros</italic> is illustrated in <xref ref-type="fig" rid="F1">Figure 1</xref>. The model results are in line with the present-day known distribution of the pest. According to the model, suitable areas for <italic>O. monoceros</italic> are largely found in Africa&#x2019;s east, south, and west coastlines, South America&#x2019;s north and east coasts, Asia&#x2019;s south and east coasts, and Oceania&#x2019;s north coast. The prediction demonstrates that the habitat suitability for the pest spans all continents, although the high suitability areas are found in Asia, Africa, Europe, and the Americas. The major oil palm producing countries, such as Thailand, Indonesia, Malaysia, Nigeria, and Colombia, are predicted to be favorable for <italic>O. monoceros</italic>. Similarly, the model forecasts habitat suitability in Indonesia, the Philippines, India, Brazil, and Sri Lanka, all of which produce considerable quantities of coconut. Finally, the predictions show that generally Egypt, Algeria, Iran, and Saudi Arabia, the world&#x2019;s leading producers of date palms are not suitable for <italic>O. monoceros</italic>.</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption><p>Suitability of the environment for the occurrence of <italic>O. monoceros</italic>. Data for <bold>(A)</bold> global probability <bold>(B)</bold> South America and sections of Central America, <bold>(C)</bold> Africa, and <bold>(D)</bold> Asia and Oceania.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-895906-g001.tif"/>
</fig>
<p>In the RCP 4.5 scenario, the BRT model predicted that high suitability areas are mainly distributed in northern parts of southern America, central Africa, southern Asia, and southeastern Asia. In addition, most contraction of suitability was focused on parts of South America, and scattered throughout Africa. While suitability in parts of Africa and Asia shows an expanded trend. The potential geographical distribution of <italic>O. monoceros</italic> was quantified and the results presented in <xref ref-type="table" rid="T3">Table 3</xref>. The potential global suitable areas of <italic>O. monoceros</italic> are shown in <xref ref-type="fig" rid="F2">Figures 2A&#x2013;C</xref> and presented in <xref ref-type="table" rid="T3">Table 3</xref>. The model predicts changes in zones of areas at risk of <italic>O. monoceros</italic> invasion (<xref ref-type="fig" rid="F2">Figures 2D&#x2013;F</xref>). Also, the model predicts more suitable areas in future (7,236,100 km<sup>2</sup>) than the current time (6,525,300 million km<sup>2</sup>).</p>
<table-wrap position="float" id="T3">
<label>TABLE 3</label>
<caption><p>The suitable area (10,000 km<sup>2</sup>) of <italic>O. monoceros</italic> by continent.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Continent</td>
<td valign="top" align="center" colspan="4">Suitable area (10,000 km<sup>2</sup>)<hr/></td>
</tr>
<tr>
<td valign="top" align="left"/>
<td valign="top" align="center">Current</td>
<td valign="top" align="center">2030</td>
<td valign="top" align="center">2050</td>
<td valign="top" align="center">2080</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left"><bold>RCP 4.5 scenario</bold></td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="left">Africa</td>
<td valign="top" align="center">324.91</td>
<td valign="top" align="center">278.94</td>
<td valign="top" align="center">286.97</td>
<td valign="top" align="center">266.62</td>
</tr>
<tr>
<td valign="top" align="left">South America</td>
<td valign="top" align="center">170.01</td>
<td valign="top" align="center">261.17</td>
<td valign="top" align="center">284.53</td>
<td valign="top" align="center">302.17</td>
</tr>
<tr>
<td valign="top" align="left">Asia</td>
<td valign="top" align="center">101.78</td>
<td valign="top" align="center">76.37</td>
<td valign="top" align="center">84.79</td>
<td valign="top" align="center">89.56</td>
</tr>
<tr>
<td valign="top" align="left">North America</td>
<td valign="top" align="center">38.88</td>
<td valign="top" align="center">46.10</td>
<td valign="top" align="center">46.75</td>
<td valign="top" align="center">44.79</td>
</tr>
<tr>
<td valign="top" align="left">Australia</td>
<td valign="top" align="center">10.56</td>
<td valign="top" align="center">8.35</td>
<td valign="top" align="center">6.59</td>
<td valign="top" align="center">15.13</td>
</tr>
<tr>
<td valign="top" align="left">Oceania</td>
<td valign="top" align="center">5.11</td>
<td valign="top" align="center">4.21</td>
<td valign="top" align="center">4.23</td>
<td valign="top" align="center">4.35</td>
</tr>
<tr>
<td valign="top" align="left">Antarctica</td>
<td valign="top" align="center">0.82</td>
<td valign="top" align="center">0.99</td>
<td valign="top" align="center">0.98</td>
<td valign="top" align="center">0.91</td>
</tr>
<tr>
<td valign="top" align="left">Europe</td>
<td valign="top" align="center">0.47</td>
<td valign="top" align="center">0.25</td>
<td valign="top" align="center">0.16</td>
<td valign="top" align="center">0.08</td>
</tr>
<tr>
<td valign="top" align="left">Total</td>
<td valign="top" align="center">652.53</td>
<td valign="top" align="center">676.36</td>
<td valign="top" align="center">715.01</td>
<td valign="top" align="center">723.61</td>
</tr>
<tr>
<td valign="top" align="left"><bold>RCP 8.5 scenario</bold></td>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
<td valign="top" align="center"/>
</tr>
<tr>
<td valign="top" align="left">Africa</td>
<td valign="top" align="center">324.91</td>
<td valign="top" align="center">279.49</td>
<td valign="top" align="center">260.68</td>
<td valign="top" align="center">266.23</td>
</tr>
<tr>
<td valign="top" align="left">South America</td>
<td valign="top" align="center">170.01</td>
<td valign="top" align="center">252.76</td>
<td valign="top" align="center">310.20</td>
<td valign="top" align="center">316.83</td>
</tr>
<tr>
<td valign="top" align="left">Asia</td>
<td valign="top" align="center">101.78</td>
<td valign="top" align="center">79.68</td>
<td valign="top" align="center">86.08</td>
<td valign="top" align="center">78.20</td>
</tr>
<tr>
<td valign="top" align="left">North America</td>
<td valign="top" align="center">38.88</td>
<td valign="top" align="center">43.31</td>
<td valign="top" align="center">43.29</td>
<td valign="top" align="center">42.56</td>
</tr>
<tr>
<td valign="top" align="left">Australia</td>
<td valign="top" align="center">10.56</td>
<td valign="top" align="center">9.29</td>
<td valign="top" align="center">11.54</td>
<td valign="top" align="center">12.15</td>
</tr>
<tr>
<td valign="top" align="left">Oceania</td>
<td valign="top" align="center">5.11</td>
<td valign="top" align="center">4.33</td>
<td valign="top" align="center">4.38</td>
<td valign="top" align="center">4.49</td>
</tr>
<tr>
<td valign="top" align="left">Antarctica</td>
<td valign="top" align="center">0.82</td>
<td valign="top" align="center">0.98</td>
<td valign="top" align="center">0.98</td>
<td valign="top" align="center">0.90</td>
</tr>
<tr>
<td valign="top" align="left">Europe</td>
<td valign="top" align="center">0.47</td>
<td valign="top" align="center">0.16</td>
<td valign="top" align="center">0.05</td>
<td valign="top" align="center">0.62</td>
</tr>
<tr>
<td valign="top" align="left">Total</td>
<td valign="top" align="center">652.53</td>
<td valign="top" align="center">669.99</td>
<td valign="top" align="center">717.18</td>
<td valign="top" align="center">721.98</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption><p>Geographical distribution of <italic>O. monoceros</italic> predicted by RCP4.5 for the future worldwide [2030 <bold>(A)</bold>, 2050 <bold>(B)</bold>, and 2080 <bold>(C)</bold>]. Changes in zones expected to be at risk <bold>(D&#x2013;F)</bold>.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-895906-g002.tif"/>
</fig>
<p>In the RCP 8.5 scenario (<xref ref-type="fig" rid="F3">Figures 3A&#x2013;C</xref>), the suitable areas will increase from the current time (6,525,300 km<sup>2</sup>) to 2080 (7,236,100 km<sup>2</sup>) (<xref ref-type="table" rid="T3">Table 3</xref>). The suitable areas of <italic>O. monoceros</italic> were found in all continents, with the model predicting either contraction or expansion of suitable areas (<xref ref-type="fig" rid="F3">Figures 3D&#x2013;F</xref>). The future expansion of suitable areas suggests that there is a need for regular monitoring and surveillance, especially where it is currently absent. Although the model predicts changes in suitable areas in the RCP 4.5 and 8.5 scenarios, more areas will continue to be suitable for <italic>O. monoceros</italic> in the future, especially coastal belts (<xref ref-type="fig" rid="F2">Figures 2A&#x2013;F</xref>, <xref ref-type="fig" rid="F3">3A&#x2013;F</xref>).</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption><p>Geographical distribution of <italic>O. monoceros</italic> predicted by RCP8.5 for the future worldwide [2030 <bold>(A)</bold>, 2050 <bold>(B)</bold>, and 2080 <bold>(C)</bold>]. Changes in zones expected to be at risk <bold>(D&#x2013;F)</bold>.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-895906-g003.tif"/>
</fig>
</sec>
</sec>
<sec id="S4" sec-type="discussion">
<title>Discussion</title>
<p>The occurrence datasets used in the modeling were mostly taken from scientific papers. This is consistent with the open science movement&#x2019;s definition, which encourages data reuse for further research and decision-making (<xref ref-type="bibr" rid="B4">Allen and Mehler, 2019</xref>). As a result, some studies have used historical species data to estimate invasion risk areas (<xref ref-type="bibr" rid="B44">Jiang et al., 2018</xref>; <xref ref-type="bibr" rid="B52">Kraemer et al., 2019</xref>; <xref ref-type="bibr" rid="B36">Guo et al., 2021</xref>; <xref ref-type="bibr" rid="B85">Zhou et al., 2021</xref>; <xref ref-type="bibr" rid="B2">Aidoo et al., 2022a</xref>,<xref ref-type="bibr" rid="B3">b</xref>). The model outputs from such studies have been utilized in developing surveillance, monitoring, and preventive measures against invasive pests (<xref ref-type="bibr" rid="B38">Hao et al., 2022</xref>).</p>
<p>The African rhinoceros beetle has been observed in Africa and Asia (<xref ref-type="bibr" rid="B79">Ukeh et al., 2003</xref>), showing an expanding trend in recent decades. <italic>O. monoceros</italic>, an important pest of palms in tropical Africa (<xref ref-type="bibr" rid="B6">Allou et al., 2006</xref>), causes server damage to these plants, thus leading to economic losses. Therefore, predicting <italic>O. monoceros</italic> suitable areas can help monitor the pest, and develop early warning, and management programmes wherever it occurs. In this study, we used the BRT model and ArcGIS to understand the potential climatic risk factors and estimate the global distribution of <italic>O. monoceros</italic> under climate change.</p>
<p>The environmental factors that influence the pest distribution were annual temperature variation, followed by the seasonality of temperature, the isothermality, precipitation of the hottest quarter months, and average variation of day temperature. This suggests that temperature factors contribute more to the global geographical distribution of the pest, which may explain why temperatures and relative humidity ranging from 27 to 29&#x00B0;C and 85 to 95%, respectively, are ideal for larval growth (<xref ref-type="bibr" rid="B13">Bedford, 1980</xref>).</p>
<p>The African rhinoceros beetle has been reported in two Asian countries: Yemen and Saudi Arabia. According to our predictions, extensive areas of Asia, though remaining uninfested at present, are highly favorable to the pest. For example, the southeastern parts of India, Vietnam, Cambodia, and Thailand are predicted to be suitable for <italic>O. monoceros</italic>. In the present study, our model predicts high habitat suitability in the world&#x2019;s largest coconut-producing countries, such as Indonesia, Philippines, India, Sri Lanka, Brazil, Vietnam, Papua New Guinea, Mexico, Thailand, and Myanmar (<xref ref-type="bibr" rid="B30">FAOSTAT, 2019</xref>). These largest coconut-producing countries will remain suitable to <italic>O. monoceros</italic> in the future. In addition, Indonesia, Malaysia, Thailand, Colombia, Nigeria, Guatemala, Honduras, Papua New Guinea, Ecuador, and Brazil, as the leading producers of oil palm (<xref ref-type="bibr" rid="B30">FAOSTAT, 2019</xref>), were all predicted to have suitable areas for the pest. Our predictions show habitat suitability for <italic>O. monoceros</italic> in Sudan and Saudi Arabia, which is quite discouraging for the date palm industry in these regions since these countries are among the major date palm producing countries in the world. These regions should be alert to the possibility of <italic>O. monoceros</italic> invasion.</p>
<p>In this study, the BRT model predicted habitat suitability for <italic>O. monoceros</italic> in areas where the Asiatic rhinoceros beetle <italic>O. rhinoceros</italic> Linnaeus (Coleoptera: Scarabaeidae) is currently present. <italic>O. monoceros</italic> and <italic>O. rhinoceros</italic> are native to Africa and Asia, respectively (<xref ref-type="bibr" rid="B40">Hinckley, 1973</xref>). <italic>Oryctes rhinoceros</italic> has spread outside its native range inflicting damage to economically important palms in the Americas, Oceania and Africa (<xref ref-type="bibr" rid="B34">Giblin-Davis, 2001</xref>). The countries affected by <italic>O. rhinoceros</italic> include Mauritius, Nigeria, and Reunion in Africa; India, Indonesia, Philippines, Sri Lanka, and Vietnam in Asia; Texas and Hawaii in the Americas; Papua New Guinea and American Samoa in Oceania (<xref ref-type="bibr" rid="B18">Catley, 1969</xref>; <xref ref-type="bibr" rid="B65">Onyeike et al., 2005</xref>; <xref ref-type="bibr" rid="B63">Okaraonye and Ikewuchi, 2009</xref>; <xref ref-type="bibr" rid="B64">Olowu et al., 2012</xref>; <xref ref-type="bibr" rid="B29">EPPO, 2022</xref>). <italic>O. monoceros</italic> has not been reported from Oceania, Southeast Asia, and the Americas. As predicted in this study, parts of these areas have medium to high habitat suitability for <italic>O. monoceros</italic>. Given the substantial economic impacts of Oryctes species, the invasion of <italic>O. monoceros</italic> in countries where <italic>O. rhinoceros</italic> occurs could pose a severe threat to food security and biodiversity conservation. Therefore, our risk maps would guide regular monitoring and surveillance of <italic>O. monoceros</italic>, thereby controlling the pest in an ecologically friendly.</p>
<p>Global warming would cause changes in species distributions (<xref ref-type="bibr" rid="B51">Kiritani, 2006</xref>; <xref ref-type="bibr" rid="B31">Freeman et al., 2018</xref>; <xref ref-type="bibr" rid="B19">Chen et al., 2019</xref>). Our results indicated that future predicted suitable areas occurred outside the present distribution of <italic>O. monoceros</italic>. The new areas were mainly found in South America, the Caribbean, Asia, and Oceania. On the one hand, our predictions revealed that the distribution of <italic>O. monoceros</italic> would expand under future climate scenarios (present &#x003C; 2030 &#x003C; 2050 &#x003C; 2080), indicating that there would be more suitable habitats for <italic>O. monoceros</italic> distribution in the future. The range shift was more noticeable in Central America than in any other region. The predicted range contraction for <italic>O. monoceros</italic> in this study could be useful for pest control and eradication. The range of habitat suitability will potentially contrast in some parts of the world, making this pest less competitive and possibly contributing to its eradication. On the other hand, a few areas are predicted to remain suitable and may require regular monitory and surveillance for early detection and control, especially in regions such as parts of Australia that currently have no occurrence records of <italic>O. monoceros</italic>. The predicted suitable areas are primarily in line with earlier pest occurrence records and show expansion of the pest&#x2019;s geographical distribution.</p>
<p>Our study identifies the regions where <italic>O. monoceros</italic> would most likely occur, as well as the relationship between the environmental indicators of this pest and the risk of its invasion, especially in areas where it has not yet been observed. It is important to note that there are some limitations in this study. Firstly, the environmental suitability maps generated from the ensemble BRT models can be interpreter to predict the potential geographical distribution of the <italic>O. monoceros</italic>, rather than where the pest will be spread in the future. Secondly, factors such as the interaction of the invader&#x2019;s characteristics (e.g., high propagule output), the abiotic environment (especially disturbance), and biotic interactions within novel environments (e.g., enemy-release) (<xref ref-type="bibr" rid="B72">Sakai et al., 2001</xref>; <xref ref-type="bibr" rid="B50">Keane and Crawley, 2002</xref>; <xref ref-type="bibr" rid="B76">Shea and Chesson, 2002</xref>) were not included in the BRT model because of unavailability of data. However, these biotic and abiotic factors are involved in the ability of a pest to establish in a new geographical area. Among them, competitors, the ability to survive in the absence of hosts, and phenotypic plasticity are considered important influencers of the establishment of species outside their native range (<xref ref-type="bibr" rid="B54">Lee and Lee, 2006</xref>; <xref ref-type="bibr" rid="B1">Aidoo et al., 2021</xref>). Thirdly, local policies, such as the inspection of plants, plant parts, farm machinery, and plant by-products at entry ports, can affect the establishment of this species outside of its natural range (<xref ref-type="bibr" rid="B59">McNeely, 2000</xref>; <xref ref-type="bibr" rid="B69">Py&#x0161;ek et al., 2020</xref>). It is our recommendation that the implementation of plant protection and regulatory procedures, the development of programs to monitor the quality of plant products and byproducts, as well as an increase in academic awareness and the transfer of technology can help prevent the spread of this pest in areas where it has not yet been detected (<xref ref-type="bibr" rid="B77">Shine, 2007</xref>; <xref ref-type="bibr" rid="B24">Early et al., 2016</xref>).</p>
</sec>
<sec id="S5" sec-type="conclusion">
<title>Conclusion</title>
<p>In this study, we have combined the BRT models with <italic>O. monoceros</italic> occurrence and environmental data to predict the potential geographical distribution of the pest. The AUC value of the model was excellent, suggesting that our model predictions are accurate and can be used to inform policy formulation and preventive measures. From now until 2080, we found that the worldwide habitat suitability for <italic>O. monoceros</italic> increased, as predicted by the BRT model. Therefore, to prevent more <italic>O. monoceros</italic> invasions in the face of present and future climate change scenarios, many of the world&#x2019;s largest oil palm, coconut, and date palm plantations must take preventative steps.</p>
</sec>
<sec id="S6" sec-type="data-availability">
<title>Data Availability Statement</title>
<p>The original contributions presented in this study are included in the article/<xref ref-type="supplementary-material" rid="TS1">Supplementary Material</xref>, further inquiries can be directed to the corresponding authors.</p>
</sec>
<sec id="S7">
<title>Author Contributions</title>
<p>OFA, MH, FD, TM, and YQ wrote the manuscript. FD, OFA, TM, NY, CB, ET, and KD conceived and designed the research. FD, DJ, TM, OFA, DW, MH, and TM analyzed the data. All authors reviewed the manuscript.</p>
</sec>
<sec id="conf1" sec-type="COI-statement">
<title>Conflict of Interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="pudiscl1" 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>
</body>
<back>
<sec id="S8" sec-type="funding-information">
<title>Funding</title>
<p>This research was supported and funded by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDA20010203).</p>
</sec>
<ack>
<p>We thank the University of Environment and Sustainable Development and Council for Scientific and Industrial Research, both in Ghana, and the State Key Laboratory of Resources and Environmental Information Systems, Institute of Geographical Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing, China for supporting this study. We also thank the reviewers for the constructive and valuable comments and the editors for their assistance in refining this article.</p>
</ack>
<sec id="S10" sec-type="supplementary-material">
<title>Supplementary Material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fevo.2022.895906/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fevo.2022.895906/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table_1.XLSX" id="TS1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet" xmlns:xlink="http://www.w3.org/1999/xlink"/>
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