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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Environ. Sci.</journal-id>
<journal-title>Frontiers in Environmental Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Environ. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-665X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">880626</article-id>
<article-id pub-id-type="doi">10.3389/fenvs.2022.880626</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Environmental Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Synergetic use of unmanned aerial vehicle and satellite images for detecting non-native tree species: An insight into <italic>Acacia saligna</italic> invasion in the Mediterranean coast</article-title>
<alt-title alt-title-type="left-running-head">Marzialetti et al.</alt-title>
<alt-title alt-title-type="right-running-head">
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/fenvs.2022.880626">10.3389/fenvs.2022.880626</ext-link>
</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Marzialetti</surname>
<given-names>Flavio</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1554059/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Di Febbraro</surname>
<given-names>Mirko</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1689211/overview"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Frate</surname>
<given-names>Ludovico</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>De Simone</surname>
<given-names>Walter</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1874372/overview"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Acosta</surname>
<given-names>Alicia Teresa Rosario</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Carranza</surname>
<given-names>Maria Laura</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1689341/overview"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Envix-Lab, Department of Biosciences and Territory, Molise University, Isernia, </institution>
<country>Italy</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>LaCEMod, Department of Life, Health and Environmental Sciences, Environmental Sciences Sect., University of L&#x0027;Aquila, L&#x0027;Aquila, </institution>
<country>Italy</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Vegetation Ecology Laboratory, Department of Sciences, Roma Tre University, Roma, </institution>
<country>Italy</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>
<bold>Edited by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1242445/overview">Thomas Campagnaro</ext-link>, University of Padua, Italy</p>
</fn>
<fn fn-type="edited-by">
<p>
<bold>Reviewed by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1803393/overview">M Arasumani</ext-link>, University of Greifswald, Germany</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1829520">Francesco Chianucci</ext-link>, Council for Agricultural and Economics Research (CREA), Italy</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1829774/overview">Joseph J Erinjery</ext-link>, Kannur University, India</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1830062/overview">Vahid Nasiri</ext-link>, University of Tehran, Iran</p>
</fn>
<corresp id="c001">&#x2a;Correspondence: Mirko Di Febbraro, <email>mirko.difebbraro@unimol.it</email>; Alicia Teresa Rosario Acosta, <email>aliciateresarosario.acosta@uniroma3.it</email>
</corresp>
<fn fn-type="other">
<p>This article was submitted to Conservation and Restoration Ecology, a section of the journal Frontiers in Environmental Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>08</day>
<month>08</month>
<year>2022</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>10</volume>
<elocation-id>880626</elocation-id>
<history>
<date date-type="received">
<day>21</day>
<month>02</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>07</day>
<month>07</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2022 Marzialetti, Di Febbraro, Frate, De Simone, Acosta and Carranza.</copyright-statement>
<copyright-year>2022</copyright-year>
<copyright-holder>Marzialetti, Di Febbraro, Frate, De Simone, Acosta and Carranza</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>Invasive alien plants (IAPs) are increasingly threatening biodiversity worldwide; thus, early detection and monitoring tools are needed. Here, we explored the potential of unmanned aerial vehicle (UAV) images in providing intermediate reference data which are able to link IAP field occurrence and satellite information. Specifically, we used very high spatial resolution (VHR) UAV maps of <italic>A. saligna</italic> as calibration data for satellite-based predictions of its spread in the Mediterranean coastal dunes. Based on two satellite platforms (PlanetScope and Sentinel-2), we developed and tested a dedicated procedure to predict <italic>A. saligna</italic> spread organized in four steps: 1) setting of calibration data for satellite-based predictions, by aggregating UAV-based VHR IAP maps to satellite spatial resolution (3 and 10&#xa0;m); 2) selection of monthly multispectral (blue, green, red, and near infra-red bands) cloud-free images for both satellite platforms; 3) calculation of monthly spectral variables depicting leaf and plant characteristics, canopy biomass, soil features, surface water and hue, intensity, and saturation values; 4) prediction of <italic>A. saligna</italic> distribution and identification of the most important spectral variables discriminating IAP occurrence using a fandom forest (RF) model. RF models calibrated for both satellite platforms showed high predictive performances (<italic>R</italic>
<sup>2</sup> &#x3e; 0.6; RMSE &#x3c;0.008), with accurate spatially explicit predictions of the invaded areas. While Sentinel-2 performed slightly better, the PlanetScope-based model effectively delineated invaded area edges and small patches. The summer leaf chlorophyll content followed by soil spectral variables was regarded as the most important variables discriminating <italic>A. saligna</italic> patches from native vegetation. Such variables depicted the characteristic IAP phenology and typically altered leaf litter and soil organic matter of invaded patches. Overall, we presented new evidence of the importance of VHR UAV data to fill the gap between field observation of <italic>A. saligna</italic> and satellite data, offering new tools for detecting and monitoring non-native tree spread in a cost-effective and timely manner.</p>
</abstract>
<kwd-group>
<kwd>upscaling</kwd>
<kwd>invasive alien plants</kwd>
<kwd>Sentinel-2</kwd>
<kwd>PlanetScope</kwd>
<kwd>environmental monitoring</kwd>
<kwd>random forest model</kwd>
<kwd>coastal dune landscapes</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec id="s1">
<title>1 Introduction</title>
<p>Biological invasions are among the major threats impinging on biodiversity across the world (<xref ref-type="bibr" rid="B104">Simberloff et al., 2013</xref>; <xref ref-type="bibr" rid="B114">Vil&#xe0; and Hulme, 2017</xref>; <xref ref-type="bibr" rid="B98">Py&#x161;ek et al., 2020</xref>). Invasive alien species (IAS), i.e., non-native species introduced by humans into a natural system outside of their native range, are causing intense direct impact and indirect impact on invaded ecosystems, compromising their ecological functions and services (<xref ref-type="bibr" rid="B7">Bartz and Kowarik, 2019</xref>). IAS invasions may alter community composition and species assemblage strategies (e.g., photosynthetic rate and standing and dead biomass; <xref ref-type="bibr" rid="B70">Linders et al., 2019</xref>) and degrade soil properties (e.g., nutrient content and water surface; <xref ref-type="bibr" rid="B20">Castro-D&#xed;ez al., 2019</xref>).</p>
<p>Among the most invasive plants impinging on coastal ecosystems worldwide, <italic>Acacia saligna</italic> (Labill.) H. Wendl (<xref ref-type="bibr" rid="B107">Starfinger and Schrader, 2021</xref>) is one of the most aggressive plants, and it was recently included in the European Regulation on Invasive Alien Species (EU1143/2014; hereafter, IAS Regulation). <italic>A. saligna</italic> is a small evergreen tree, native of Western Australia (<xref ref-type="bibr" rid="B79">Maslin, 1974</xref>), with fast-growing and intense vegetative and sexual propagation (Witkowski, 1991). Being introduced as fodder (<xref ref-type="bibr" rid="B5">Asefa and Tamir, 2006</xref>; <xref ref-type="bibr" rid="B43">George et al., 2007</xref>), windbreak and dune stabilization (<xref ref-type="bibr" rid="B6">Bar et al., 2004</xref>), and for ornamental purposes (<xref ref-type="bibr" rid="B30">Donaldson et al., 2014</xref>), it has become invasive in coastal areas across the world (e.g., South Africa, North Africa, Horn of Africa, Chile, and the Mediterranean; <xref ref-type="bibr" rid="B108">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B73">Lozano et al., 2020</xref>). Several negative effects of <italic>A. saligna</italic> invasion on natural ecosystems were reported, for e.g., the alteration of biodiversity values, the decline of focal species, and the drastic change of vegetation structure toward dense monospecific <italic>A. saligna</italic> woodlands (<xref ref-type="bibr" rid="B67">Le Maitre et al., 2011</xref>; <xref ref-type="bibr" rid="B27">Del Vecchio et al., 2013</xref>; <xref ref-type="bibr" rid="B109">Tozzi et al., 2021</xref>). <italic>A. saligna</italic> also modifies soil nitrogen and organic matter content (<xref ref-type="bibr" rid="B118">Yelenik et al., 2004</xref>; <xref ref-type="bibr" rid="B35">El-Gawad and El-Amier, 2015</xref>), altering soil microbial communities (<xref ref-type="bibr" rid="B23">Cris&#xf3;stomo et al., 2013</xref>). Furthermore, invaded areas tend to present poorer aesthetic and recreational landscape values than natural ones (<xref ref-type="bibr" rid="B68">Lehrer et al., 2011</xref>).</p>
<p>The growing pressure exerted by IAS across different ecosystems worldwide urges the scientific community and civil society to identify adequate monitoring and management strategies (<xref ref-type="bibr" rid="B16">Brundu et al., 2018</xref>). As the Convention on Biological Diversity (CBD; <ext-link ext-link-type="uri" xlink:href="https://www.cbd.int/">https://www.cbd.int/</ext-link>) claims for a global strategy against IAS by 2030, the European Regulation on Invasive Alien Species (EU1143/2014) provides clear guidelines to prevent, minimize, and mitigate the occurrence and effects of alien species on natural ecosystems (<xref ref-type="bibr" rid="B42">Genovesi et al., 2015</xref>; <xref ref-type="bibr" rid="B14">Branquart et al., 2016</xref>).</p>
<p>The analysis of invasion processes was traditionally based on field campaigns, often combined with visual interpretation of aerial photos; both approaches were widely recognized as expensive, time-consuming, and limited to depicting IAS occurrences on remote and inaccessible areas (<xref ref-type="bibr" rid="B27">Del Vecchio et al., 2013</xref>; <xref ref-type="bibr" rid="B106">Stanisci et al., 2014</xref>; <xref ref-type="bibr" rid="B102">Royimani et al., 2019</xref>). Indeed, field campaigns for IAS spread analysis require an accurate work plan to be carried out in a specific sampling period, often in limited areas (<xref ref-type="bibr" rid="B11">Bolch et al., 2020</xref>). In addition, the visual interpretation of aerial photos is very time-consuming, even for skilled photo-interpreters (<xref ref-type="bibr" rid="B11">Bolch et al., 2020</xref>). During the last decade, there has been increasing evidence of the potential of remote sensing (RS) for IAS early detection, monitoring, and mapping in a cost-effective, spatially contiguous, and timely manner (<xref ref-type="bibr" rid="B57">Huang and Asner, 2009</xref>; <xref ref-type="bibr" rid="B102">Royimani et al., 2019</xref>). Indeed, the distribution of different IAS can be detected by using expensive very high-resolution (spatial and spectral) RS images (e.g., airborne and satellite platforms with multispectral or hyperspectral sensors; <xref ref-type="bibr" rid="B113">Underwood et al., 2003</xref>; <xref ref-type="bibr" rid="B92">Paz-Kagan et al., 2017</xref>; <xref ref-type="bibr" rid="B90">Niphadkar et al., 2017</xref>) or combining free coarser RS images and field-collected occurrences as calibration data (<xref ref-type="bibr" rid="B120">Zhou et al., 2018</xref>; <xref ref-type="bibr" rid="B61">Kattenborn et al., 2019</xref>).</p>
<p>However, the combination of standard satellite RS images and field-collected occurrences for IAS detection has different shortcomings that should be overcome (<xref ref-type="bibr" rid="B13">Bradley 2014</xref>; <xref ref-type="bibr" rid="B95">Pettorelli et al., 2014</xref>) as: 1) satellite images regularly covering the overall Earth surface using standardized multispectral sensors (fixed spatial resolution and zenith angle) could weakly describe some important ecological parameters (e.g., IAS flower color and the blooming period or altered soils on invaded areas) which are essential for IAS detection (<xref ref-type="bibr" rid="B86">M&#xfc;llerov&#xe1; et al., 2017</xref>); 2) the acquisition of high quality IAS occurrence data in the field may be hampered by local conditions, such as a dense canopy cover, trouble in reaching remote invaded areas (<xref ref-type="bibr" rid="B60">Kaartinen et al., 2015</xref>; <xref ref-type="bibr" rid="B82">McGaughey et al., 2017</xref>), or the complexity of the invaded ecosystem mosaic (<xref ref-type="bibr" rid="B12">Bradley 2009</xref>; <xref ref-type="bibr" rid="B52">Hawthorne et al., 2017</xref>); 3) building a reliable database of field IAS occurrences may require a great effort to collect spatially accurate records using Real-Time Kinematics Global Navigation Satellite System (RTK GNSS) devices (<xref ref-type="bibr" rid="B96">Piiroinen et al., 2018</xref>; <xref ref-type="bibr" rid="B25">Dao et al., 2021</xref>).</p>
<p>A valid alternative to increase the spectral information and the amount of accurate IAS occurrence data at a local scale is offered by unmanned aerial vehicles (UAVs). Indeed, the use of UAVs allows the collection of highly customized data as the operator can easily set several parameters that are useful to detect alien species (e.g., type of sensor, angle of view, spatial resolution, time, and frequency of acquisition), which are commonly standardized on most satellite platforms (<xref ref-type="bibr" rid="B61">Kattenborn et al., 2019</xref>; <xref ref-type="bibr" rid="B3">Alvarez-Vanhard et al., 2021</xref>). In addition, since UAV images register environmental complexity as a continuous surface (<xref ref-type="bibr" rid="B61">Kattenborn et al., 2019</xref>; <xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>) at very high spatial resolution (VHR), variations of spectral values caused by IAS can be detected in a smoothed way (<xref ref-type="bibr" rid="B4">Anderson 2018</xref>; <xref ref-type="bibr" rid="B69">Leit&#xe3;o et al., 2018</xref>), helping to fill the gap between field observations and satellite data. On the other side, technical constraints of UAVs (e.g., battery capability and surveying restrictions) often prevent the use of this technology to describe ecosystem patterns on a large scale. However, the ability to acquire highly customized data makes UAVs an interesting option to provide accurate maps of local IAS occurrence (e.g., <xref ref-type="bibr" rid="B116">Wijesingha et al., 2020</xref>; <xref ref-type="bibr" rid="B77">Marzialetti et al., 2021</xref>), supporting satellite detection at wider scales. The contribution of VHR images for aiding IAS satellite detection deserves to be further explored (<xref ref-type="bibr" rid="B38">Elkind et al., 2019</xref>; <xref ref-type="bibr" rid="B3">Alvarez-Vanhard et al., 2021</xref>).</p>
<p>The present work sets out to investigate the potential of UAV data depicting the smooth occurrence of <italic>A. saligna</italic> at a local scale to support its satellite-based detection at a wide scale in complex and dynamic environments such as Mediterranean coastal dunes. Specifically, we used VHR (0.05&#xa0;m) multispectral (blue, green, red, and near infrared) UAV images collected during the <italic>A. saligna</italic> blooming period to provide occurrence data needed to predict the spread of this IAS using two satellite platforms (free for research purposes) with different spatial resolutions (e.g., PlanetScope: 3&#xa0;m and Sentinel-2: 10&#xa0;m). Moreover, we also compared IAS detection performance achieved by the two satellite platforms, highlighting their differences in mapping invaded coastal dune environments.</p>
</sec>
<sec id="s2">
<title>2 Materials and methods</title>
<sec id="s2-1">
<title>2.1 Study area and target species</title>
<p>The study area includes a representative tract of recent (i.e., Holocenic) dunes along the Adriatic coast of Central Italy, characterized by a Mediterranean climate (<xref ref-type="fig" rid="F1">Figure 1A</xref>; <xref ref-type="bibr" rid="B2">Acosta et al., 2009</xref>; <xref ref-type="bibr" rid="B17">Carranza et al., 2008</xref>). We selected two areas: 1) an invaded one of approximately 11&#xa0;ha as the UAV flight calibration area (<xref ref-type="fig" rid="F1">Figure 1B</xref>, red polygon, Marzialetti et al., 2021) and 2) a wider one at high invasion risk (Marzialetti et al., 2019) of approximately 70&#xa0;ha as the prediction area (<xref ref-type="fig" rid="F1">Figure 1B</xref>, green polygon). As in other coastal areas of the Adriatic coast, the analyzed dunes are low (less than 8&#x2013;10&#xa0;m) and occupy a narrow strip parallel to the seashore. Under natural conditions, the psammophilous vegetation mosaic follows a sea&#x2013;inland gradient ranging from the annual pioneer communities on the upper beach to the Mediterranean maquis and <italic>Pinus</italic> spp. woods in the inner fore dune sectors (<xref ref-type="bibr" rid="B1">Acosta et al., 2003</xref>; <xref ref-type="bibr" rid="B17">Carranza et al., 2008</xref>; <xref ref-type="bibr" rid="B8">Bazzichetto et al., 2016</xref>). This zonation promotes the development of highly specialized biodiversity, which shares few species with other terrestrial communities (<xref ref-type="bibr" rid="B32">Drius et al., 2016</xref>; <xref ref-type="bibr" rid="B76">Marzialetti et al., 2020</xref>), and its integrity assures manifold ecosystem services (<xref ref-type="bibr" rid="B33">Drius et al., 2013</xref>). This area is impinged by several human-related disturbances as most of the Mediterranean coastal landscapes: agricultural pressure (<xref ref-type="bibr" rid="B74">Malavasi et al., 2013</xref>), tourism and urban expansion (<xref ref-type="bibr" rid="B18">Carranza et al., 2018</xref>), beach pollution (<xref ref-type="bibr" rid="B29">Di Febbraro et al., 2021</xref>), and alien species invasions (<xref ref-type="bibr" rid="B27">Del Vecchio et al., 2013</xref>; <xref ref-type="bibr" rid="B75">Marzialetti et al., 2019</xref>). The analyzed area is inside a special area of conservation (SAC, Habitat Directive 92/43/EEC; Foce Trigno&#x2014;Marina di Petacciato IT7228221) and is a node of the Long-Term Ecological Research Network (LTER, <ext-link ext-link-type="uri" xlink:href="http://www.lter-europe.net/">http://www.lter-europe.net/</ext-link>; <xref ref-type="bibr" rid="B106">Stanisci et al., 2014</xref>; <xref ref-type="bibr" rid="B33">Drius et al., 2013</xref>), which makes it an excellent testing ground to develop methodologies which are able to evaluate and monitor invasion processes.</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption>
<p>
<bold>(A)</bold> Study area reporting the UAV flight calibration area in red and the prediction area in green (coordinate system in WGS 84 epsg: 4326). SAC and LTER site shape is reported in black (Foce Trigno&#x2013;Marina di Petacciato&#x2014;IT7228221, <ext-link ext-link-type="uri" xlink:href="https://deims.org/1835cda2-b56d-400a-b413-ab5c74086dc5">https://deims.org/1835cda2-b56d-400a-b413-ab5c74086dc5</ext-link>). <bold>(B)</bold> Enlargement of the study area projected on the PlanetScope image of 20 July 2021. Red polygon represents UAV flight coverage used for preparing calibration data, and the green polygon represents the satellite prediction area.</p>
</caption>
<graphic xlink:href="fenvs-10-880626-g001.tif"/>
</fig>
</sec>
<sec id="s2-2">
<title>2.2 Data collection and analysis</title>
<p>The proposed framework for detecting and predicting <italic>A. saligna</italic> distribution was structured following four steps that are schematically reported in <xref ref-type="fig" rid="F2">Figure 2</xref>: (A) UAV-based <italic>A. saligna</italic> maps and calibration data, (B) satellite imagery selection, (C) remote sensing variable calculation, and (D) <italic>A. saligna</italic> satellite-based predictions.</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption>
<p>Workflow synthesizing the procedure for the satellite-based prediction of <italic>A. saligna</italic> distribution using the UAV-based IAS VHR information as calibration data. The proposal framework: <bold>(A)</bold> UAV-based <italic>A. saligna</italic> maps and calibration data, <bold>(B)</bold> Satellite imagery selection, <bold>(C)</bold> Remote sensing variables calculation, <bold>(D)</bold> <italic>A. saligna</italic> satellite-based predictions.</p>
</caption>
<graphic xlink:href="fenvs-10-880626-g002.tif"/>
</fig>
<sec id="s2-2-1">
<title>2.2.1 Unmanned aerial vehicle-based <italic>Acacia saligna</italic> calibration data</title>
<p>As calibration data for satellite-based predictions, we used a VHR (0.05&#xa0;m) presence/absence map of <italic>A. saligna</italic> derived from the combination of a set of highly accurate UAV-based maps produced for the study area (on <xref ref-type="fig" rid="F1">Figure 1</xref>, red polygon) and recently published by Marzialetti et al. (2021; Supplementary material 1 <xref ref-type="sec" rid="s11">Supplementary Figure S1</xref>). In that research, UAV images were collected in pre-flowering and flowering periods using a multirotor quadcopter (DJI Phantom 4 Pro V2.0) equipped with two sensors: the CMOS (complementary metal oxide semiconductor) RGB (red&#x2013;green&#x2013;blue) camera with 20&#xa0;Mpx and the Parrot Sequoia multispectral sensor G, R, REdge, and NIR bands (Green: 550 &#xb1; 40&#xa0;nm; Red: 660 &#xb1; 40&#xa0;nm; Red Edge: 735 &#xb1; 10&#xa0;nm; and Near Infrared: 790 &#xb1; 40&#xa0;nm) with 1.2&#xa0;Mpx for each band. This study (<xref ref-type="bibr" rid="B77">Marzialetti et al., 2021</xref>) indicated a very high predictive performance (overall accuracy &#x3e;95%, Kappa statistic &#x3e;0.75) of four VHR maps derived from images registered during the IAP flowering period. So, in order to define robust calibration data (<xref ref-type="fig" rid="F2">Figure 2A</xref>, calibration area) and reduce possible errors, we stacked the flowering period maps reporting as occurrences only those pixels in which <italic>A. saligna</italic> was predicted in at least three of the four maps. Then, based on these VHR calibration data (0.05&#xa0;m), we calculated, either for PlanetScope or for Sentinel-2 platform, the fractional cover maps of <italic>A. saligna</italic> reporting the percent of VHR-invaded pixels in 3&#xa0;m (FCoverPS) and 10&#xa0;m (FcoverS2) grid cells, respectively.</p>
</sec>
<sec id="s2-2-2">
<title>2.2.2 Satellite imagery selection</title>
<p>As Mediterranean coastal areas are characterized by a highly dynamic landscape undergoing substantial seasonal changes, we relied on RS multi-temporal stacks for gathering monthly images. In order to capture the annual phenology both for <italic>A. saligna</italic> and for native coastal dune vegetation, we used monthly multispectral images recorded by two satellite platforms, which are accessible free of charge for research purposes (<xref ref-type="fig" rid="F2">Figure 2B</xref>). PlanetScope (PS) satellite constellation consists of multiple DOVE CubeSat acquiring four bands: blue (B<sub>PS</sub>, 455&#x2013;515&#xa0;nm), green (G<sub>PS</sub>, 500&#x2013;590&#xa0;nm), red (R<sub>PS</sub>, 590&#x2013;670&#xa0;nm), and near infrared (NIR<sub>PS</sub>, 780&#x2013;860&#xa0;nm), with 3&#xa0;m of spatial resolution (PlanetLabs Inc 2021; <xref ref-type="bibr" rid="B22">Cheng et al., 2020</xref>). We downloaded 12 cloud-free PlanetScope images (November 2020&#x2013;October 2021) with a zenith view angle lower than 5&#xb0; (i.e., nadir viewing; <ext-link ext-link-type="uri" xlink:href="https://www.planet.com/explorer/">https://www.planet.com/explorer/</ext-link>). We specifically used the surface reflectance products (i.e., processing level 3B) with atmospherical correction already performed by PlanetLabs (Supplementary Material 2 <xref ref-type="sec" rid="s11">Supplementary Table S1</xref>, <xref ref-type="bibr" rid="B62">Kotchenova et al., 2006</xref>; <xref ref-type="bibr" rid="B63">Kotchenova and Vermote, 2007</xref>).</p>
<p>The Sentinel-2 satellite (S2) is equipped with a multispectral instrument (MSI) sensor including 13 bands in the range of visible, near-infrared (10&#xa0;m), and short-wave infrared (20&#xa0;m). We used the 10&#xa0;m bands: blue (B<sub>S2</sub>, 459&#x2013;525&#xa0;nm), green (G<sub>S2</sub>, 541&#x2013;577&#xa0;nm), red (R<sub>S2</sub>, 649&#x2013;680&#xa0;nm), and near infrared (NIR<sub>S2</sub>, 780&#x2013;886&#xa0;nm) bands (<xref ref-type="bibr" rid="B34">Drusch et al., 2012</xref>). We downloaded 12 cloud-free Sentinel-2 images (November 2020&#x2013;October 2021; Copernicus Open Access Hub: <ext-link ext-link-type="uri" xlink:href="https://scihub.copernicus.eu/">https://scihub.copernicus.eu/</ext-link>). As the image of January 2021 showed high cloud coverage, we replaced it with a cloud-free image registered in January 2020 (Supplementary Material 2 <xref ref-type="sec" rid="s11">Supplementary Table S2</xref>). We used atmospherically corrected images at the surface reflectance (processing level 2A; European Space Agency using a Sen2Cor processor, <xref ref-type="bibr" rid="B72">Louis et al., 2019</xref>).</p>
</sec>
<sec id="s2-2-3">
<title>2.2.3 Remote sensing variable calculation</title>
<p>As IAS spread could alter environmental features such as biomass, chlorophyll content, and soil features in invaded native ecosystems (<xref ref-type="bibr" rid="B70">Linders et al., 2019</xref>; Castro-D&#xed;ez al., 2019), we calculated monthly RS variables related to these main biophysical parameters (<xref ref-type="fig" rid="F2">Figure 2C</xref>; <xref ref-type="table" rid="T1">Table 1</xref>, <xref ref-type="sec" rid="s11">Supplementary Material S3</xref>). In particular, we calculated a set of spectral variables using the R environmental ( R: <ext-link ext-link-type="uri" xlink:href="https://www.r-project.org/">https://www.r-project.org/</ext-link>) that could help in distinguishing <italic>A. saligna</italic> invaded areas from native vegetation, depicting leaf and plant characteristics (e.g., leaf content of chlorophyll and carotenoids) and canopy biomass, as well as soil features (e.g., bare surfaces and organic content), and the presence of surface water (<xref ref-type="bibr" rid="B78">Masemola et al., 2020</xref>; <xref ref-type="bibr" rid="B103">Shoko et al., 2020</xref>). For leaf and plant characteristics, we estimated leaf pigments using six spectral indices: the chlorophyll vegetation index (CVI) and the green leaf index (GLI) for chlorophyll content; the chlorophyll index green (CIgreen), which quantifies the photosynthetic activity; the carotenoid reflectance index 550 (CRI550) and the structure intensive pigment index 3 (SIPI3), which quantify the carotenoids and photosynthetic pigments; and the green difference vegetation index (GDVI), which estimates the content of nitrogen (<xref ref-type="bibr" rid="B111">Tucker et al., 1979</xref>; <xref ref-type="bibr" rid="B94">Pe&#xf1;uelas et al., 1995</xref>; <xref ref-type="bibr" rid="B48">Gobron et al., 2000</xref>; <xref ref-type="bibr" rid="B47">Gitelson et al., 2005</xref>; <xref ref-type="bibr" rid="B115">Vincini et al., 2008</xref>). Canopy biomass characteristics were described by computing five spectral indices: the normalized difference vegetation index (NDVI) and simple ratio (SR), which are canonical biomass indices; the enhanced vegetation index (EVI), which improves the biomass estimate compared to two previous indices, reducing the atmospheric influence; the transformed vegetation index (TVI), which is sensitive to the photosynthetically active biomass; and the visible atmospherically resistant index green (VARIgreen), able to estimate the canopy biomass using visible bands (B, G, and R, <xref ref-type="bibr" rid="B58">Huete et al., 2002</xref>; <xref ref-type="bibr" rid="B112">Tucker, 1979</xref>; <xref ref-type="bibr" rid="B10">Birth and McVey, 1968</xref>; <xref ref-type="bibr" rid="B100">Rouse et al., 1973</xref>; <xref ref-type="bibr" rid="B46">Gitelson et al., 2002</xref>). As for soil features, we calculated the brightness index (BI) and brightness index 2 (BI2) as proxies of the bare surface, while the coloration index (CI) was used for organic content (<xref ref-type="bibr" rid="B39">Escadafal et al., 1989</xref>; <xref ref-type="bibr" rid="B80">Mathieu et al., 1998</xref>). We also computed the normalized difference water index (NDWI) to estimate seasonal water surface fluctuations along coastal dunes (<xref ref-type="bibr" rid="B81">McFeeters, 1996</xref>).</p>
<table-wrap id="T1" position="float">
<label>TABLE 1</label>
<caption>
<p>Computed spectral indices and metrics: acronyms, full names, formulas, and references. Concerning the spectral indices are grouped by the main biophysical parameters related to leaf and plant characteristics, canopy biomass, soil features, and surface water (see for details <xref ref-type="sec" rid="s11">Supplementary Material S2</xref>).</p>
</caption>
<table>
<thead valign="top">
<tr>
<th align="left">Acronym</th>
<th align="left">Name</th>
<th align="left">Formula</th>
<th align="left">Reference</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td colspan="4" align="left">
<italic>Leaf and plant characteristics</italic>
</td>
</tr>
<tr>
<td align="left">&#x2003;CIgreen</td>
<td align="left">Chlorophyll index green</td>
<td align="left">
<inline-formula id="inf1">
<mml:math id="m1">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mi>G</mml:mi>
</mml:mfrac>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B47">Gitelson et al. (2005)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;CVI</td>
<td align="left">Chlorophyll vegetation index</td>
<td align="left">
<inline-formula id="inf2">
<mml:math id="m2">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mi>E</mml:mi>
<mml:mi>D</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:msup>
<mml:mi>G</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B115">Vincini et al. (2008)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;CRI550</td>
<td align="left">Carotenoid reflectance index 550</td>
<td align="left">
<inline-formula id="inf3">
<mml:math id="m3">
<mml:mrow>
<mml:mfrac>
<mml:mn>1</mml:mn>
<mml:mi>B</mml:mi>
</mml:mfrac>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mfrac>
<mml:mn>1</mml:mn>
<mml:mi>G</mml:mi>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B48">Gobron et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;GDVI</td>
<td align="left">Green difference vegetation index</td>
<td align="left">NIR &#x2013; G</td>
<td align="left">
<xref ref-type="bibr" rid="B111">Tucker et al. (1979)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;GLI</td>
<td align="left">Green leaf index</td>
<td align="left">
<inline-formula id="inf4">
<mml:math id="m4">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mn>2</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>G</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>B</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mn>2</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>G</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>B</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B48">Gobron et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;SIPI3</td>
<td align="left">Structure intensive pigment index 3</td>
<td align="left">
<inline-formula id="inf5">
<mml:math id="m5">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>B</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>R</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B94">Pe&#xf1;uelas et al. (1995)</xref>
</td>
</tr>
<tr>
<td colspan="4" align="left">
<italic>Canopy biomass</italic>
</td>
</tr>
<tr>
<td align="left">&#x2003;EVI</td>
<td align="left">Enhanced vegetation index</td>
<td align="left">
<inline-formula id="inf6">
<mml:math id="m6">
<mml:mrow>
<mml:mn>2.5</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mrow>
<mml:mo>(</mml:mo>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#x2b;</mml:mo>
<mml:mn>6</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>7.5</mml:mn>
<mml:mo>&#x2217;</mml:mo>
<mml:mi>B</mml:mi>
</mml:mrow>
<mml:mo>)</mml:mo>
</mml:mrow>
<mml:mo>&#x2b;</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B58">Huete et al. (2002)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;NDVI</td>
<td align="left">Normalized difference vegetation index</td>
<td align="left">
<inline-formula id="inf7">
<mml:math id="m7">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>R</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B112">Tucker (1979)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;SR</td>
<td align="left">Simple ratio</td>
<td align="left">
<inline-formula id="inf8">
<mml:math id="m8">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mi>R</mml:mi>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B10">Birth and McVey (1968)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;TVI</td>
<td align="left">Transformed vegetation index</td>
<td align="left">
<inline-formula id="inf9">
<mml:math id="m9">
<mml:mrow>
<mml:msqrt>
<mml:mrow>
<mml:mn>0.5</mml:mn>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mfrac>
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>G</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mo>&#x2b;</mml:mo>
<mml:mi>G</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:msqrt>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B100">Rouse et al. (1973)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;VARIgreen</td>
<td align="left">Visible atmospherically resistant index green</td>
<td align="left">
<inline-formula id="inf10">
<mml:math id="m10">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>G</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>G</mml:mi>
<mml:mo>&#x2b;</mml:mo>
<mml:mi>R</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>B</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B46">Gitelson et al. (2002)</xref>
</td>
</tr>
<tr>
<td colspan="4" align="left">
<italic>Soil features</italic>
</td>
</tr>
<tr>
<td align="left">&#x2003;BI</td>
<td align="left">Brightness index</td>
<td align="left">
<inline-formula id="inf11">
<mml:math id="m11">
<mml:mrow>
<mml:msqrt>
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:msup>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mn>2</mml:mn>
<mml:mo>&#xa0;</mml:mo>
</mml:mrow>
</mml:msup>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:msup>
<mml:mi>G</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
<mml:mn>2</mml:mn>
</mml:mfrac>
</mml:mrow>
</mml:msqrt>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B39">Escadafal et al. (1989)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;BI2</td>
<td align="left">Brightness index 2</td>
<td align="left">
<inline-formula id="inf12">
<mml:math id="m12">
<mml:mrow>
<mml:msqrt>
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:msup>
<mml:mi>R</mml:mi>
<mml:mrow>
<mml:mn>2</mml:mn>
<mml:mo>&#xa0;</mml:mo>
</mml:mrow>
</mml:msup>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:msup>
<mml:mi>G</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:msup>
<mml:mi>R</mml:mi>
<mml:mn>2</mml:mn>
</mml:msup>
</mml:mrow>
<mml:mn>3</mml:mn>
</mml:mfrac>
</mml:mrow>
</mml:msqrt>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B39">Escadafal et al. (1989)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;CI</td>
<td align="left">Coloration index</td>
<td align="left">
<inline-formula id="inf13">
<mml:math id="m13">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>G</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>G</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B80">Mathieu et al. (1998)</xref>
</td>
</tr>
<tr>
<td colspan="4" align="left">
<italic>Surface water</italic>
</td>
</tr>
<tr>
<td align="left">&#x2003;NDWI</td>
<td align="left">Normalized difference water index</td>
<td align="left">
<inline-formula id="inf14">
<mml:math id="m14">
<mml:mrow>
<mml:mfrac>
<mml:mrow>
<mml:mi>G</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
</mml:mrow>
<mml:mrow>
<mml:mi>G</mml:mi>
<mml:mo>&#xa0;</mml:mo>
<mml:mo>&#x2b;</mml:mo>
<mml:mo>&#xa0;</mml:mo>
<mml:mi>N</mml:mi>
<mml:mi>I</mml:mi>
<mml:mi>R</mml:mi>
</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</inline-formula>
</td>
<td align="left">
<xref ref-type="bibr" rid="B81">McFeeters (1996)</xref>
</td>
</tr>
<tr>
<td colspan="4" align="left">
<italic>Hue, Saturation</italic>, <italic>and Intensity metrics</italic>
</td>
</tr>
<tr>
<td align="left">&#x2003;H</td>
<td align="left">Hue</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B64">Koutsias et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;S</td>
<td align="left">Saturation</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B64">Koutsias et al. (2000)</xref>
</td>
</tr>
<tr>
<td align="left">&#x2003;I</td>
<td align="left">Intensity</td>
<td align="left"/>
<td align="left">
<xref ref-type="bibr" rid="B64">Koutsias et al. (2000)</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>In addition, we transformed the surface reflectance values of RGB bands for all the images into digital numbers and converted them into hue (H), intensity (I), and saturation (S) metrics (<xref ref-type="bibr" rid="B89">Neteler and Mitasova, 2004</xref>; <xref ref-type="bibr" rid="B119">Zhang, 2004</xref>, <xref ref-type="fig" rid="F2">Figure 2C</xref>; <italic>i.rgb.his</italic> tool in GRASS GIS 7.8; <xref ref-type="bibr" rid="B50">GRASS Development Team</xref>, 2020, <xref ref-type="table" rid="T1">Table 1</xref>), so as to catch color differences among <italic>A. saligna</italic> leaves and inflorescences and native vegetation. HIS variables are particularly effective in discriminating <italic>A. saligna</italic> from native vegetation at a local scale (<xref ref-type="bibr" rid="B117">Yang et al., 2020</xref>; <xref ref-type="bibr" rid="B77">Marzialetti et al., 2021</xref>). For each pixel, the H metric depicts the dominant wavelength, the I metric depicts the brightness of a color (e.g., the relative degree of black or white), and the S metric depicts the purity of the color, defined as the absence of mixture with other wavelengths (<xref ref-type="bibr" rid="B64">Koutsias et al., 2000</xref>; <xref ref-type="bibr" rid="B110">Tu et al., 2005</xref>).</p>
<p>For each satellite (e.g., PlanetScope and Sentinel-2 images) and area (e.g., calibration and prediction), we built a multi-temporal stack including monthly bands (RGB and NIR), HIS values, and spectral indices (i.e., 264 layers; <xref ref-type="fig" rid="F2">Figure 2C</xref> and <xref ref-type="table" rid="T1">Table 1</xref>).</p>
</sec>
<sec id="s2-2-4">
<title>2.2.4 <italic>A. saligna</italic> satellite-based predictions</title>
<p>To upscale <italic>A. saligna</italic> FCover derived from UAV to PlanetScope (predicted-FCover<sub>PS</sub>) and Sentinel-2 (predicted-FCover<sub>S2</sub>) resolutions, we modelled this variable against the spectral indices using random forest (RF; <xref ref-type="fig" rid="F2">Figure 2D</xref>, <xref ref-type="bibr" rid="B15">Breiman, 2001</xref>). RF is a machine learning algorithm that operates with a large combination of decision trees, reducing the error in classification and regression by using bootstrap in the considered explanatory variables (<xref ref-type="bibr" rid="B24">Cutler et al., 2007</xref>; <xref ref-type="bibr" rid="B21">Chan and Paelinckx, 2008</xref>). RF is commonly applied to spatial regression and classification of remote sensing data (<xref ref-type="bibr" rid="B9">Belgiu and Dr&#x103;gu&#x163;, 2016</xref>; <xref ref-type="bibr" rid="B59">Izquierdo-Verdiguier and Zurita-Milla, 2020</xref>), also accommodating well with highly correlated explanatory variables (<xref ref-type="bibr" rid="B83">Meyer et al., 2017</xref>).</p>
<p>Given the high dimensionality of our data, we reduced the multitemporal spectral variables using the recursive feature elimination algorithm (RFE, <xref ref-type="fig" rid="F2">Figure 2D</xref>, <xref ref-type="bibr" rid="B65">Kuhn and Johnson, 2013</xref>). RFE is a feature selection algorithm that estimates the lowest possible number of features without reducing the final performance metrics of the RF model (<xref ref-type="bibr" rid="B28">Demarchi et al., 2020</xref>). RFE iteratively computes RF models with all the explanatory variables in the calibration data and then drops in each cycle the lowest important variable (i.e., scoring the lowest value of the mean decrease impurity (MDI) index). At the end of all cycles, RFE identifies the best number of explanatory variables by comparing the predictive performance metrics of all the RF models produced (<xref ref-type="bibr" rid="B83">Meyer et al., 2017</xref>; <xref ref-type="bibr" rid="B71">Lou et al., 2020</xref>). We performed the RFE algorithm through the &#x201c;caret&#x201d; R package (function <italic>rfe</italic>, <xref ref-type="bibr" rid="B66">Kuhn et al., 2021</xref>) applying a 10-fold cross-validation and calculating the root mean square error (RMSE, <xref ref-type="bibr" rid="B19">Castillo-Riffart et al., 2017</xref>) to assess the predictive performance of each subset of variables (i.e., we set a maximum of 80 variables). Once we identified the best number of variables, we fine-tuned the RF model according to three parameters: the number of uncorrelated decision trees (<italic>Ntree</italic>),the number of variables randomly selected at each node of decision trees (<italic>Mtry</italic>), and the minimum number of observations in a terminal node (minimal node size, <xref ref-type="bibr" rid="B9">Belgiu and Dr&#x103;gu&#x163;, 2016</xref>; <xref ref-type="bibr" rid="B97">Probst et al., 2018</xref>). We set a high number of uncorrelated decision trees (<italic>Ntree</italic> &#x3d; 1,000), tested different <italic>Mtry</italic> values ranging from 2 to the number of variables as indicated by RFE results, and checked a range of minimal node size from 1 to 5 (<xref ref-type="fig" rid="F2">Figure 2D</xref>). Furthermore, we used the Extra-Trees algorithm as a splitting procedure to apply the regression on independent tree nodes (<xref ref-type="bibr" rid="B44">Geurts et al., 2006</xref>).</p>
<p>The RF model reporting the lowest RMSE under a 10-fold cross-validation (<xref ref-type="fig" rid="F2">Figure 2D</xref>, <xref ref-type="bibr" rid="B101">Routh et al., 2018</xref>) was selected as the optimal one and used to predict <italic>A. saligna</italic> FCover. Along with RMSE, RF predictive performance was assessed by calculating the coefficient of determination <italic>R</italic>
<sup>2</sup> between observed and predicted values under cross-validation. RMSE and <italic>R</italic>
<sup>2</sup> values by RF models obtained from PlanetScope and Sentinel-2 images were compared through the Mann&#x2013;Whitney test to assess which of the two satellites performed best in predicting <italic>A. saligna</italic> FCover.</p>
<p>The relative importance of spectral variables in RF models was estimated through the MDI index (i.e., the Gini index with the sum of squares as an impurity measure, <xref ref-type="bibr" rid="B88">Nembrini et al., 2018</xref>; <xref ref-type="fig" rid="F2">Figure 2D</xref>). Then, we explored the shape of the relationships between predicted-FCover of <italic>A. saligna</italic> and the most important variables in both RF models by using partial dependence (PD, <xref ref-type="bibr" rid="B41">Friedman, 2001</xref>) plots. PD plots measure the marginal effect of a given explanatory variable on the predicted values of RF model, f constant, and the other variables&#x2019; constant (e.g., to their median value; <xref ref-type="bibr" rid="B37">Elith et al., 2008</xref>).</p>
<p>Lastly, we evaluated the degree of extrapolation on values of spectral variables outside the UAV flight area by computing the multivariate environmental similarity surface (MESS) in both RF models (<xref ref-type="fig" rid="F2">Figure 2D</xref>, <xref ref-type="bibr" rid="B36">Elith et al., 2010</xref>). MESS measures the similarity between values of spectral variables inside the calibration area and values in the prediction area. Negative values (MESS &#x3c;0) indicate pixels with values of spectral variables dissimilar from the calibration area (<xref ref-type="bibr" rid="B36">Elith et al., 2010</xref>). We computed MESS using the R package &#x201c;dismo&#x201d; (function <italic>mess</italic>, <xref ref-type="bibr" rid="B55">Hijmans and Elith 2021</xref>) and reported the percentage of negative MESS values in both RF models.</p>
</sec>
</sec>
</sec>
<sec id="s3">
<title>3 Results</title>
<p>The RF model on Sentinel-2 data retained 25 spectral variables after RFE with <italic>Mtry</italic> and minimal node size equal to 24 and 1, respectively, performing slightly better (<italic>R</italic>
<sup>2</sup> &#x3d; 0.707 &#xb1; 0.088) than the model on PlanetScope data, which retained 75 variables and obtained <italic>Mtry</italic> and minimal node size equal to 66 and 1, respectively (<italic>R</italic>
<sup>2</sup> &#x3d; 0.628 &#xb1; 0.028, <xref ref-type="fig" rid="F3">Figure 3</xref>, see also <xref ref-type="sec" rid="s11">Supplementary Material S4</xref>). The RMSE of the Sentinel-2 model was significantly lower than that of PlanetScope (Mann&#x2013;Whitney U &#x3d; 0, <italic>p</italic> &#x3c; 0.001; <xref ref-type="sec" rid="s11">Supplementary Material S5</xref>).</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption>
<p>Relationship between calibration <italic>A. saligna</italic> fractional cover (e.g., UAV calibration-FCover) and predicted fractional cover obtained by <bold>(A)</bold> PlanetScope (predicted-FCoverPS) and <bold>(B)</bold> Sentinel-2 (predicted-FCoverS2) classification models. Blue depicts the linear regression between predicted FCover and calibration FCover. Red dashed lines depict ideal regressions of perfect match among predicted-FCover and calibration-FCover values. RF predictive performance (<italic>R</italic>
<sup>2</sup>) and root mean square error (RMSE) for each regression are also reported.</p>
</caption>
<graphic xlink:href="fenvs-10-880626-g003.tif"/>
</fig>
<p>The relationship between <italic>A. saligna</italic> observed (i.e., derived from UAV) and predicted (i.e., satellite-based) FCover values varied between the two satellite platforms, with <italic>R</italic>
<sup>2</sup> PlanetScope &#x3d; 0.628 &#xb1; 0.028 and <italic>R</italic>
<sup>2</sup> Sentinel-2 &#x3d; 0.707 &#xb1; 0.088 (<xref ref-type="fig" rid="F3">Figure 3</xref>). We observed a moderate variability as well as a wider range (e.g., from 0 to 95%) in FCover values predicted by the PlanetScope RF model. Predicted values of the Sentinel-2 model presented a lower variability and a reduced range (from 0 to 75%). In both RF models, predicted FCover values are slightly over-estimated compared with the lowest observed values and under-estimated compared with the highest observed values (<xref ref-type="fig" rid="F3">Figure 3</xref>).</p>
<p>Spatially explicit predictions of <italic>A. saligna</italic> FCover evidenced differences in the ability of the two satellite platforms to depict the smoothed spatial variations in <italic>A. saligna</italic> observed cover as well as the invaded patch edges, which varied markedly with the spatial resolution of the considered satellite images (<xref ref-type="fig" rid="F4">Figure 4</xref>). The RF model based on PlanetScope, with a finer spatial resolution (3&#xa0;m), was able to delineate invaded patches and accurately describe the smooth FCover gradient between the invaded core areas and edges (<xref ref-type="fig" rid="F4">Figure 4C</xref>), predicting even small patches correctly. On the contrary, the model based on Sentinel-2 images, with a coarser spatial resolution (10&#xa0;m), delineated <italic>A. saligna</italic> edges quite poorly, as well as the fuzzy variability of the observed FCover values in invaded areas. In addition, it was unable to map small invaded patches (<xref ref-type="fig" rid="F4">Figure 4D</xref>).</p>
<fig id="F4" position="float">
<label>FIGURE 4</label>
<caption>
<p>Visual example of <bold>(A)</bold> UAV orthophotographs captured during the <italic>A. saligna</italic> flowering period, <bold>(B)</bold> calibration data derived by the UAV-based VHR <italic>A. saligna</italic> map 0.05&#xa0;m, <bold>(C)</bold> <italic>A. saligna</italic> fractional cover on the PlanetScope model (predicted-FCoverPS), and <bold>(D)</bold> <italic>A. saligna</italic> fractional cover on the Sentinel-2 model (predicted-FCoverS2). Black edges reported on boxes C and D depict the shape of UAV-based <italic>A. saligna</italic> distribution used as calibration data.</p>
</caption>
<graphic xlink:href="fenvs-10-880626-g004.tif"/>
</fig>
<p>The relative importance of spectral variables measured by the MDI index differed between Sentinel-2 and PlanetScope models (<xref ref-type="fig" rid="F5">Figure 5</xref>). In the PlanetScope model, the first 60% of the cumulated MDI percentage was achieved with 18 spectral variables (i.e., from CVI.07 to NDWI.07; <xref ref-type="fig" rid="F5">Figure 5A</xref>), while in the Sentinel-2 model the same percentage was achieved with six spectral variables (from H.07 to CVI.08). In the PlanetScope RF model, the three most important variables were chlorophyll vegetation index of July (CVI.07; 7%), hue of July (H.07; 6,9%), and hue of May (H.05; 5,8%; <xref ref-type="fig" rid="F5">Figure 5A</xref>). According to PD plots, high CVI.07 and the extreme values (low and high) of H.07 and H.05 were consistently associated with an increase in <italic>A. saligna</italic> FCover (<xref ref-type="sec" rid="s11">Supplementary Material S6</xref>). Other important variables in the PlanetScope model were related to summer (e.g. BI2.06, BI2.07, GLI.07, CI.07, VARIgreen.07, CIgreen.07, TVI.07, NDWI.07, NIR.07, and GDVI.08; <xref ref-type="fig" rid="F5">Figure 5A</xref>) and autumn (e.g. G.09, CVI.09, EVI.10, SR.10, SIPI3.10, and NDVI.10; <xref ref-type="fig" rid="F5">Figure 5A</xref>).</p>
<fig id="F5" position="float">
<label>FIGURE 5</label>
<caption>
<p>Relative importance of RS spectral variables in percentage (%) calculated using the mean decrease impurity for <bold>(A)</bold> PlanetScope (75 variables) and <bold>(B)</bold> Sentinel-2 (25 variables) RF models. Spectral variables are reported in <xref ref-type="table" rid="T1">Table 1</xref>. Red dashed lines indicate the 60% of cumulated MDI percentage.</p>
</caption>
<graphic xlink:href="fenvs-10-880626-g005.tif"/>
</fig>
<p>In the Sentinel-2 RF model, the spectral variables included in the first 60% of the cumulated MDI percentage are related to summer (H.07, BI2.06, and CVI.08), spring (H.05 and CVI.05), and winter (S.01) features. Specifically, the hues of July (H.07) and May (H.05), with 21 and 11% MDI, respectively, were the two most important variables (<xref ref-type="fig" rid="F5">Figure 5B</xref>). H.07, H.05, CVI.05, CVI.08, and S.01 showed a positive relationship with predicted FCover values, while BI2.06 showed a slightly decreasing relationship (<xref ref-type="sec" rid="s11">Supplementary Material S6</xref>).</p>
<p>The degree of extrapolation in the predicted area varied among the two RF models. The PlanetScope model showed a low percentage of pixels with negative MESS values (6.043%) in areas where <italic>A. saligna</italic> occurrence is very unlikely (e.g., on the sandy beach close to the sea; <xref ref-type="sec" rid="s11">Supplementary Material S7</xref>). The Sentinel-2 RF model showed a higher degree of extrapolation with 13.792% of pixels with negative MESS values (<xref ref-type="sec" rid="s11">Supplementary Material S7</xref>).</p>
</sec>
<sec id="s4">
<title>4 Discussion</title>
<p>The present work contributed to improving monitoring tools for <italic>A. saligna</italic> detection and spread, extending the utilization of UAV data to support IAS satellite modelling on Mediterranean coastal dunes. This approach was previously tested in a temperate broadleaved forest (<xref ref-type="bibr" rid="B61">Kattenborn et al., 2019</xref>; <xref ref-type="bibr" rid="B49">Gr&#xe4;nzig et al., 2021</xref>; <xref ref-type="bibr" rid="B56">Holden et al., 2021</xref>), tundra coniferous formations (<xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>), and wetlands (<xref ref-type="bibr" rid="B120">Zhou et al., 2018</xref>; <xref ref-type="bibr" rid="B31">Doughty et al., 2021</xref>). It is implemented in this study for the first time in complex coastal landscapes.</p>
<p>Our results evidenced a good potential of UAV-based fine-scale maps as a source of calibration data for satellite-based prediction of a woody IAP in partially invaded Mediterranean coasts.</p>
<p>The RF models for both satellite platforms predicted <italic>A. saligna</italic> FCover properly, although they slightly over-estimated <italic>A. saligna</italic> FCover comparedwith lower values and under-estimated compared with the higher ones. A similar overestimation of cells with low cover values was registered by <xref ref-type="bibr" rid="B61">Kattenborn et al. (2019)</xref> with other IAPs (e.g., <italic>Pinus radiata</italic>, invading forest and <italic>Ulex europaeus</italic> invading scrublands) in South America. These authors underlined the limits for detecting cover fractions below a &#x223c;12% threshold. Differently, the under-estimation of pixels with high Fcover values may be related to the small number of coarse satellite grids dominated by <italic>A. saligna</italic> inside the calibration area.</p>
<p>Concerning the predictive accuracy, as previously observed in tundra and taiga ecosystems (<xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>; <xref ref-type="bibr" rid="B40">Fraser et al., 2021</xref>), and also on coastal dune landscapes, satellite images with a coarse spatial resolution supported distribution models with higher predictive accuracy than those with fine spatial resolution. According to RMSE values, the 10&#xa0;m resolution Sentinel-2 model predicted the VHR UAV calibration data more accurately than the 3&#xa0;m PlanetScope model. As reported in previous studies (<xref ref-type="bibr" rid="B26">Dark and Bram, 2007</xref>; <xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>; <xref ref-type="bibr" rid="B40">Fraser et al., 2021</xref>), such differences may be due to the modifiable area unit problem (MAUP) and the decreased variance of the coarser-resolution data, arising due to the aggregation of finer scale data (e.g., 0.05&#xa0;m VHR IAP map) on coarser grids (e.g., 10&#xa0;m resolution FcoverS2). MAUP affects the statistical analyses because with the aggregation of fine cell information in the larger ones (e.g., 3&#xa0;m or 10&#xa0;m), a decrease of the between-cell variability is verified and an increase of the explanatory power of derived models occurs.</p>
<p>On the other hand, the finer spatial resolution of PlanetScope (3&#xa0;m) allowed the maintenance of a good part of the spatial information derived by UAV data, as verified by the variability of its FCover values ranging from 0 to almost 100 percent. In the Sentinel-2 FCover values, the relation among VHR UAV occurrences is limited to a smaller range.</p>
<p>Other scale issues that might influence the IAS model accuracy are the low number of pixels registering &#x201c;pure&#x201d; <italic>A. saligna</italic> patches in Sentinel-2 images and the relatively limited number of total pixels in the prediction area. In fact, the fuzzy shape of <italic>A. saligna</italic> patches as derived by Sentinel-2 models corresponds to better model performance values (<xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>; <xref ref-type="bibr" rid="B53">He et al., 2021</xref>). On the other hand, the finer spatial resolution of PlanetScope images allowed us to model the variability of FCover values registered by the very high UAV-based map, thus offering a good support for identifying <italic>A. saligna</italic> patch edges. Similarly, this complementarity between coarse and fine-scale images has been observed in previous studies predicting the fractional cover of tundra vegetation and lichen of taiga using UAV and satellite images with different spatial resolutions (e.g., PlanetScope, Sentinel-2, and Landsat; <xref ref-type="bibr" rid="B99">Riihim&#xe4;ki et al., 2019</xref>; <xref ref-type="bibr" rid="B53">He et al., 2021</xref>).</p>
<p>The analysis of multi-temporal spectral variables derived from visible (blue, green, and red) and NIR bands acquired by Sentinel-2 and PlanetScope platforms effectively depicted the phenological behavior of specific IAS. Our results pinpointed the summer biomass production peak (see CVI of July for PlanetScope and CVI of August for Sentinel-2) among the most important parameters for modelling the fractional cover of <italic>A. saligna</italic> along coastal dune systems. Previous research studies based on medium-resolution satellite images (RGB and NIR bands of Sentinel-2) evidenced the importance of spectral indices depicting the characteristic <italic>Acacia</italic> spp. summer productivity peak to detect invaded patches embedded in natural landscapes (e.g., montane forest and alluvial wetlands; see <xref ref-type="bibr" rid="B78">Masemola et al., 2020</xref>; <xref ref-type="bibr" rid="B61">Kattenborn et al., 2019</xref>). As observed for other <italic>Acacia</italic> species (e.g., <italic>A. dealbata, A. mearnsii</italic>, and <italic>A. longifolia</italic>; <xref ref-type="bibr" rid="B78">Masemola et al., 2020</xref>; <xref ref-type="bibr" rid="B51">Gro&#x3b2;e-Stoltenberg et al., 2016</xref>), and also for <italic>A. saligna</italic>, the analyzed RS variables denoted high leaf chlorophyll content during the summer season. Such photosynthetic temporal patterns can be described by several monthly RS spectral indices (e.g., CIgreen, CVI, GLI, and GDVI) and, in our case, by the high July CVI values. This peak of chlorophyll content is consistent with the previous field ecophysiological studies along the Mediterranean coastal environments (<xref ref-type="bibr" rid="B87">Nativ et al., 1999</xref>; <xref ref-type="bibr" rid="B85">Morris et al., 2011</xref>).</p>
<p>Our results also evidenced that soil RS variables (summer BI2 and CI values) are important for modelling the fractional cover of <italic>A. saligna</italic>, which is consistent with the alterations in leaf litter content and soil organic matter in IAS patches usually observed in the field (<xref ref-type="bibr" rid="B27">Del Vecchio et al., 2013</xref>; <xref ref-type="bibr" rid="B91">Nsikani et al., 2017</xref>; <xref ref-type="bibr" rid="B109">Tozzi et al., 2021</xref>). In fact, <italic>Acacia</italic> species, being nitrogen-fixing plants, modify soil features, increasing the organic content and the development of soil microbiomes (<xref ref-type="bibr" rid="B27">Del Vecchio et al., 2013</xref>). This nitrogen-fixing action alters the nature of the unconsolidated coastal dune soil characterized by low organic content (<xref ref-type="bibr" rid="B67">Le Maitre et al., 2011</xref>). Both PlanetScope and Sentinel-2 models highlighted a reduced summer bare surface on <italic>A. saligna</italic> patches compared to the coastal dune native vegetation, indicating a dense canopy of the IAS as well as an enriched soil organic content. It is also noteworthy that the nitrogen content in leaves, as calculated by GDVI, was markedly higher in <italic>A. saligna</italic> patches than native vegetation ones (<xref ref-type="bibr" rid="B118">Yelenik et al., 2004</xref>; <xref ref-type="bibr" rid="B54">Hellmann et al., 2011</xref>).</p>
<p>Interestingly, during the summer period, the two RS models distinguished the <italic>A. saligna</italic> patches from native vegetation by different hue values of leaves (see H of July). This pattern may be due to the different coloration of leaves between <italic>A. saligna</italic> and dominant native species, particularly the gymnosperm species such as <italic>Pinus</italic> spp., <italic>Juniperus oxicedrus</italic>, and the maquis species such as <italic>Pistacia lentiscus</italic> and <italic>Phyllirea angustifolia</italic>.</p>
<p>
<italic>Acacia saligna</italic> is characterized by an early blooming period compared to the coastal dune vegetation and conforms to pure yellow color patches in April&#x2013;May (<xref ref-type="bibr" rid="B84">Milton and Moll, 1982</xref>; <xref ref-type="bibr" rid="B93">Paz-Kagan et al., 2019</xref>). The spectral variables characterizing the <italic>A. saligna</italic> blooming period are very useful for detecting the IAS using remote sensing images with very fine spatial (e.g., VHR UAV images; <xref ref-type="bibr" rid="B77">Marzialetti et al., 2021</xref>) or spectral (e.g., yellow wavelength bands; <xref ref-type="bibr" rid="B93">Paz-Kagan et al., 2019</xref>) resolution. The models developed in this study using coarse spatial and spectral information, freely available from PlanetScope and Sentinel-2 platforms, partially confirm the usefulness of RS variables registered during the blooming period, as suggested by the distinct behavior of hue and the chlorophyll content of leaves in May. Moreover, the combined use of multi-temporal RS variables and machine learning algorithms allowed to describe different RS ecological conditions over time and seasons, supporting the detection of <italic>A. saligna</italic> and the identification of the most important RS variables to distinguish the invaded patches from native vegetation ones.</p>
<p>Finally, the adopted approach offers economic commitments and reduces the time necessary to evaluate <italic>A. saligna</italic> invasion levels in complex environments such as coastal dunes. In this way, this approach supports the prioritization of monitoring and management actions claimed by the EU IAS Regulation 1,143/2014 (<xref ref-type="bibr" rid="B104">Simberloff et al., 2013</xref>; Rai and Sing, 2020; <xref ref-type="bibr" rid="B105">Souza-Alonso et al., 2017</xref>). In addition, this methodology could easily be applied to other IAS in complex environments. UAV images could be considered appropriate candidates to become a major tool to gather reference data. In this context, we could better outline operational workflows for the early detection and monitoring of IAS invasion status over time, which is essential to define adequate management actions and to tackle these invasive species (<xref ref-type="bibr" rid="B56">Holden et al., 2021</xref>).</p>
</sec>
<sec id="s5">
<title>5 Conclusion</title>
<p>Developing early detection and monitoring tools for IAS that are able to operate at wide scales is an urgent challenge requested by the IAS European Regulation (IAS Regulation 1,143/2014). In this study, we tested a methodological workflow, applied for the first time on coastal dune landscapes, to predict the fractional cover of the invasive alien species <italic>A. saligna</italic> by applying a combination of VHR UAV data and two satellite multi-temporal images (PlanetScope and Sentinel-2) with a different spatial resolution (3 and 10&#xa0;m, respectively). We presented new evidence on the importance of VHR UAV data to fill the gap between field observation of <italic>A. saligna</italic> and satellite data in complex and dynamic environments such as the Mediterranean coastal dunes. The predictions based on PlanetScope and Sentinel-2 multi-temporal data accurately predicted <italic>A. saligna</italic> fractional cover derived by the UAV-based map with high performance metrics in both models (<italic>R</italic>
<sup>2</sup> &#x3e; 0.6 and RMSE &#x3c;0.08). The PlanetScope-based model was able to outline invaded area edges even for small patches. Moreover, the model accurately described the smooth FCover gradient between the invaded core areas and edges, confirming the importance of finer spatial resolutions.</p>
<p>The over-estimation and under-estimation of the lowest and highest fractional cover predicted values in both satellites gave evidence of a reduced capability of the satellite-based model to detect early stages of the invasion process and dense monospecific patches.</p>
<p>Nevertheless, from an applied perspective, our results also allow to identify the set of RS variables depicting effective ecological parameters for predicting <italic>A. saligna</italic> occurrence in coastal areas, and then contribute to improving monitoring activities. Our encouraging results support the usefulness of combining UAV and satellite images to detect and monitor <italic>A. saligna</italic> spread in the coastal areas in a timely and cost-effective manner.</p>
<p>Overall, we could conclude that our approach, based on satellite images available worldwide and free of charge for research purposes, could be potentially applied to a wide variety of landscapes and IAPs. This procedure could be important not only in Europe (IAS regulation 1143/2014) but also around the world (global strategy against IAS by CBC) because IAS management requires a common approach worldwide. With this in mind, further case studies could be implemented in the future to better test and extend the synergetic use of unmanned aerial vehicles and satellite images presented here. Moreover, this approach could provide comparable information for other coastal ecosystems, for other invasive alien species (e.g., <italic>Carpobrotus</italic> spp., <italic>Agave americana</italic>, and <italic>Yucca gloriosa</italic>), and for other biogeographical areas as well.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="s6">
<title>Data availability statement</title>
<p>The raw data supporting the conclusion of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s7">
<title>Author contributions</title>
<p>All authors contributed substantially to the work: FM, MC, and MDF conceived and designed the study; FM, LF, and WDS collected remote sensing data; FM, MDF, and MC analyzed the data; FM, MDF, and MC led the writing of the manuscript; AA and MC supervised the research. All authors contributed critically to the drafts and gave final approval for publication. All authors have read and agreed to the published version of the manuscript.</p>
</sec>
<sec id="s8">
<title>Funding</title>
<p>This study was carried out with a support of the bilateral program Italy&#x2013;Israel DERESEMII (Developing state-of-the-art remote sensing tools for monitoring the impact of invasive plant) and the Interreg Italia&#x2014;Croazia CASCADE (CoAStal and marine waters integrated monitoring systems for ecosystems proteCtion AnD management&#x2014;Project ID 10255941). The Grant of Excellence Departments, MIUR&#x2014;Italy (ARTICOLO 1, COMMI 314&#x2013;337 LEGGE 232/2016) is also gratefully acknowledged.</p>
</sec>
<ack>
<p>The authors are grateful to the Long-Term Ecological Research (LTER) and LTER&#x2014;Italia. The authors are grateful to Planet Labs for free PlanetScope imagery for research purposes. Furthermore, the authors acknowledge Silvia Cascone and Francesco Pio Tozzi, for the help given during the acquisition of UAV images. The authors thank the editor and the reviewers for their time and comments which helped them to improve the original version of the article.</p>
</ack>
<sec sec-type="COI-statement" id="s9">
<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 sec-type="disclaimer" id="s10">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors, and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11">
<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/fenvs.2022.880626/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fenvs.2022.880626/full&#x23;supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Datasheet1.docx" id="SM1" mimetype="application/docx" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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