Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters
Identifieur interne : 000177 ( Pmc/Corpus ); précédent : 000176; suivant : 000178Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters
Auteurs : Martin Godefroid ; Astrid Cruaud ; Jean-Pierre Rossi ; Jean-Yves RasplusSource :
- PLoS ONE [ 1932-6203 ] ; 2015.
Abstract
Widely distributed species often show strong phylogeographic structure, with lineages potentially adapted to different biotic and abiotic conditions. The success of an invasion process may thus depend on the intraspecific identity of the introduced propagules. However, pest risk analyses are usually performed without accounting for intraspecific diversity. In this study, we developed bioclimatic models using MaxEnt and boosted regression trees approaches, to predict the potential distribution in Europe of six economically important Tephritid pests (
Url:
DOI: 10.1371/journal.pone.0135209
PubMed: 26274582
PubMed Central: 4537207
Links to Exploration step
PMC:4537207Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters</title><author><name sortKey="Godefroid, Martin" sort="Godefroid, Martin" uniqKey="Godefroid M" first="Martin" last="Godefroid">Martin Godefroid</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Cruaud, Astrid" sort="Cruaud, Astrid" uniqKey="Cruaud A" first="Astrid" last="Cruaud">Astrid Cruaud</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Rossi, Jean Pierre" sort="Rossi, Jean Pierre" uniqKey="Rossi J" first="Jean-Pierre" last="Rossi">Jean-Pierre Rossi</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Rasplus, Jean Yves" sort="Rasplus, Jean Yves" uniqKey="Rasplus J" first="Jean-Yves" last="Rasplus">Jean-Yves Rasplus</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26274582</idno><idno type="pmc">4537207</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4537207</idno><idno type="RBID">PMC:4537207</idno><idno type="doi">10.1371/journal.pone.0135209</idno><date when="2015">2015</date><idno type="wicri:Area/Pmc/Corpus">000177</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters</title><author><name sortKey="Godefroid, Martin" sort="Godefroid, Martin" uniqKey="Godefroid M" first="Martin" last="Godefroid">Martin Godefroid</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Cruaud, Astrid" sort="Cruaud, Astrid" uniqKey="Cruaud A" first="Astrid" last="Cruaud">Astrid Cruaud</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Rossi, Jean Pierre" sort="Rossi, Jean Pierre" uniqKey="Rossi J" first="Jean-Pierre" last="Rossi">Jean-Pierre Rossi</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author><author><name sortKey="Rasplus, Jean Yves" sort="Rasplus, Jean Yves" uniqKey="Rasplus J" first="Jean-Yves" last="Rasplus">Jean-Yves Rasplus</name><affiliation><nlm:aff id="aff001"></nlm:aff></affiliation></author></analytic><series><title level="j">PLoS ONE</title><idno type="eISSN">1932-6203</idno><imprint><date when="2015">2015</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Widely distributed species often show strong phylogeographic structure, with lineages potentially adapted to different biotic and abiotic conditions. The success of an invasion process may thus depend on the intraspecific identity of the introduced propagules. However, pest risk analyses are usually performed without accounting for intraspecific diversity. In this study, we developed bioclimatic models using MaxEnt and boosted regression trees approaches, to predict the potential distribution in Europe of six economically important Tephritid pests (<italic>Ceratitis fasciventris</italic> (Bezzi), <italic>Bactrocera oleae</italic> (Rossi), <italic>Anastrepha obliqua</italic> (Macquart), <italic>Anastrepha fraterculus</italic> (Wiedemann), <italic>Rhagoletis pomonella</italic> (Walsh) and <italic>Bactrocera cucurbitae</italic> (Coquillet)). We considered intraspecific diversity in our risk analyses by independently modeling the distributions of conspecific lineages. The six species displayed different potential distributions in Europe. A strong signal of intraspecific climate envelope divergence was observed in most species. In some cases, conspecific lineages differed strongly in potential distributions suggesting that taxonomic resolution should be accounted for in pest risk analyses. No models (lineage- and species-based approaches) predicted high climatic suitability in the entire invaded range of <italic>B</italic>. <italic>oleae</italic>—the only species whose intraspecific identity of invading populations has been elucidated—in California. Host availability appears to play the most important role in shaping the geographic range of this specialist pest. However, climatic suitability values predicted by species-based models are correlated with population densities of <italic>B</italic>. <italic>oleae</italic> globally reported in California. Our study highlights how classical taxonomic boundaries may lead to under- or overestimation of the potential pest distributions and encourages accounting for intraspecific diversity when assessing the risk of biological invasion.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Pimentel, D" uniqKey="Pimentel D">D Pimentel</name></author><author><name sortKey="Mcnair, S" uniqKey="Mcnair S">S McNair</name></author><author><name sortKey="Janecka, J" uniqKey="Janecka J">J Janecka</name></author><author><name sortKey="Wightman, J" uniqKey="Wightman J">J Wightman</name></author><author><name sortKey="Simmonds, C" uniqKey="Simmonds C">C Simmonds</name></author><author><name sortKey="O Connell, C" uniqKey="O Connell C">C O'Connell</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hulme, Pe" uniqKey="Hulme P">PE Hulme</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kolar, Cs" uniqKey="Kolar C">CS Kolar</name></author><author><name sortKey="Lodge, Dm" uniqKey="Lodge D">DM Lodge</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Vieglais, Da" uniqKey="Vieglais D">DA Vieglais</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rafter, Ma" uniqKey="Rafter M">MA Rafter</name></author><author><name sortKey="Hereward, Jp" uniqKey="Hereward J">JP Hereward</name></author><author><name sortKey="Walter, Gh" uniqKey="Walter G">GH Walter</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Santos, H" uniqKey="Santos H">H Santos</name></author><author><name sortKey="Burban, C" uniqKey="Burban C">C Burban</name></author><author><name sortKey="Rousselet, J" uniqKey="Rousselet J">J Rousselet</name></author><author><name sortKey="Rossi, Jp" uniqKey="Rossi J">JP Rossi</name></author><author><name sortKey="Branco, M" uniqKey="Branco M">M Branco</name></author><author><name sortKey="Kerdelhue, C" uniqKey="Kerdelhue C">C Kerdelhué</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rey, O" uniqKey="Rey O">O Rey</name></author><author><name sortKey="Estoup, A" uniqKey="Estoup A">A Estoup</name></author><author><name sortKey="Vonshak, M" uniqKey="Vonshak M">M Vonshak</name></author><author><name sortKey="Loiseau, A" uniqKey="Loiseau A">A Loiseau</name></author><author><name sortKey="Blanchet, S" uniqKey="Blanchet S">S Blanchet</name></author><author><name sortKey="Calcaterra, L" uniqKey="Calcaterra L">L Calcaterra</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kelly, Dw" uniqKey="Kelly D">DW Kelly</name></author><author><name sortKey="Muirhead, Jr" uniqKey="Muirhead J">JR Muirhead</name></author><author><name sortKey="Heath, Dd" uniqKey="Heath D">DD Heath</name></author><author><name sortKey="Macisaac, Hj" uniqKey="Macisaac H">HJ Macisaac</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lachmuth, S" uniqKey="Lachmuth S">S Lachmuth</name></author><author><name sortKey="Durka, W" uniqKey="Durka W">W Durka</name></author><author><name sortKey="Schurr, Fm" uniqKey="Schurr F">FM Schurr</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hamilton, Ja" uniqKey="Hamilton J">JA Hamilton</name></author><author><name sortKey="Okada, M" uniqKey="Okada M">M Okada</name></author><author><name sortKey="Korves, T" uniqKey="Korves T">T Korves</name></author><author><name sortKey="Schmitt, J" uniqKey="Schmitt J">J Schmitt</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Holt, Rd" uniqKey="Holt R">RD Holt</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="White, Im" uniqKey="White I">IM White</name></author><author><name sortKey="Elson Harris, Mm" uniqKey="Elson Harris M">MM Elson-Harris</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Mun, J" uniqKey="Mun J">J Mun</name></author><author><name sortKey="Bohonak, Aj" uniqKey="Bohonak A">AJ Bohonak</name></author><author><name sortKey="Roderick, Gk" uniqKey="Roderick G">GK Roderick</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rice, R" uniqKey="Rice R">R Rice</name></author><author><name sortKey="Phillips, P" uniqKey="Phillips P">P Phillips</name></author><author><name sortKey="Stewart Leslie, J" uniqKey="Stewart Leslie J">J Stewart-Leslie</name></author><author><name sortKey="Sibbett, G" uniqKey="Sibbett G">G Sibbett</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Koyama, J" uniqKey="Koyama J">J Koyama</name></author><author><name sortKey="Kakinohana, H" uniqKey="Kakinohana H">H Kakinohana</name></author><author><name sortKey="Miyatake, T" uniqKey="Miyatake T">T Miyatake</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Drew, Rai" uniqKey="Drew R">RAI Drew</name></author><author><name sortKey="Tsuruta, K" uniqKey="Tsuruta K">K Tsuruta</name></author><author><name sortKey="White, I" uniqKey="White I">I White</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Malacrida, Ar" uniqKey="Malacrida A">AR Malacrida</name></author><author><name sortKey="Gomulski, Lm" uniqKey="Gomulski L">LM Gomulski</name></author><author><name sortKey="Bonizzoni, M" uniqKey="Bonizzoni M">M Bonizzoni</name></author><author><name sortKey="Bertin, S" uniqKey="Bertin S">S Bertin</name></author><author><name sortKey="Gasperi, G" uniqKey="Gasperi G">G Gasperi</name></author><author><name sortKey="Guglielmino, Cr" uniqKey="Guglielmino C">CR Guglielmino</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author><author><name sortKey="Plant, Re" uniqKey="Plant R">RE Plant</name></author><author><name sortKey="Carey, Jr" uniqKey="Carey J">JR Carey</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Clarke, Ar" uniqKey="Clarke A">AR Clarke</name></author><author><name sortKey="Armstrong, Kf" uniqKey="Armstrong K">KF Armstrong</name></author><author><name sortKey="Carmichael, Ae" uniqKey="Carmichael A">AE Carmichael</name></author><author><name sortKey="Milne, Jr" uniqKey="Milne J">JR Milne</name></author><author><name sortKey="Raghu, S" uniqKey="Raghu S">S Raghu</name></author><author><name sortKey="Roderick, Gk" uniqKey="Roderick G">GK Roderick</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Nardi, F" uniqKey="Nardi F">F Nardi</name></author><author><name sortKey="Carapelli, A" uniqKey="Carapelli A">A Carapelli</name></author><author><name sortKey="Dallai, R" uniqKey="Dallai R">R Dallai</name></author><author><name sortKey="Roderick, Gk" uniqKey="Roderick G">GK Roderick</name></author><author><name sortKey="Frati, F" uniqKey="Frati F">F Frati</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Nardi, F" uniqKey="Nardi F">F Nardi</name></author><author><name sortKey="Carapelli, A" uniqKey="Carapelli A">A Carapelli</name></author><author><name sortKey="Boore, Jl" uniqKey="Boore J">JL Boore</name></author><author><name sortKey="Roderick, Gk" uniqKey="Roderick G">GK Roderick</name></author><author><name sortKey="Dallai, R" uniqKey="Dallai R">R Dallai</name></author><author><name sortKey="Frati, F" uniqKey="Frati F">F Frati</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Ruiz Arce, R" uniqKey="Ruiz Arce R">R Ruiz-Arce</name></author><author><name sortKey="Barr, Nb" uniqKey="Barr N">NB Barr</name></author><author><name sortKey="Owen, Cl" uniqKey="Owen C">CL Owen</name></author><author><name sortKey="Thomas, Db" uniqKey="Thomas D">DB Thomas</name></author><author><name sortKey="Mcpheron, Ba" uniqKey="Mcpheron B">BA McPheron</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Virgilio, M" uniqKey="Virgilio M">M Virgilio</name></author><author><name sortKey="Delatte, H" uniqKey="Delatte H">H Delatte</name></author><author><name sortKey="Quilici, S" uniqKey="Quilici S">S Quilici</name></author><author><name sortKey="Backeljau, T" uniqKey="Backeljau T">T Backeljau</name></author><author><name sortKey="De Meyer, M" uniqKey="De Meyer M">M De Meyer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Grout, Tg" uniqKey="Grout T">TG Grout</name></author><author><name sortKey="Stoltz, Kc" uniqKey="Stoltz K">KC Stoltz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Papanastasiou, Sa" uniqKey="Papanastasiou S">SA Papanastasiou</name></author><author><name sortKey="Nestel, D" uniqKey="Nestel D">D Nestel</name></author><author><name sortKey="Diamantidis, Ad" uniqKey="Diamantidis A">AD Diamantidis</name></author><author><name sortKey="Nakas, Ct" uniqKey="Nakas C">CT Nakas</name></author><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Muller, H G" uniqKey="Muller H">H-G Müller</name></author><author><name sortKey="Wu, S" uniqKey="Wu S">S Wu</name></author><author><name sortKey="Diamantidis, Ad" uniqKey="Diamantidis A">AD Diamantidis</name></author><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author><author><name sortKey="Carey, Jr" uniqKey="Carey J">JR Carey</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Diamantidis, Ad" uniqKey="Diamantidis A">AD Diamantidis</name></author><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author><author><name sortKey="Nakas, Ct" uniqKey="Nakas C">CT Nakas</name></author><author><name sortKey="Wu, S" uniqKey="Wu S">S Wu</name></author><author><name sortKey="Muller, Hg" uniqKey="Muller H">HG Müller</name></author><author><name sortKey="Carey, Jr" uniqKey="Carey J">JR Carey</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Diamantidis, Ad" uniqKey="Diamantidis A">AD Diamantidis</name></author><author><name sortKey="Carey, Jr" uniqKey="Carey J">JR Carey</name></author><author><name sortKey="Nakas, Ct" uniqKey="Nakas C">CT Nakas</name></author><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author><author><name sortKey="Perez Staples, D" uniqKey="Perez Staples D">D Pérez-Staples</name></author><author><name sortKey="Macias Ord Ez, R" uniqKey="Macias Ord Ez R">R Macías-Ordóñez</name></author><author><name sortKey="Pi Ero, J" uniqKey="Pi Ero J">J Piñero</name></author><author><name sortKey="Mcpheron, B" uniqKey="Mcpheron B">B Mcpheron</name></author><author><name sortKey="Hernandez Ortiz, V" uniqKey="Hernandez Ortiz V">V Hernández-Ortiz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Smith Caldas, Mr" uniqKey="Smith Caldas M">MR Smith-Caldas</name></author><author><name sortKey="Mcpheron, Ba" uniqKey="Mcpheron B">BA Mcpheron</name></author><author><name sortKey="Silva, Jg" uniqKey="Silva J">JG Silva</name></author><author><name sortKey="Zucchi, Ra" uniqKey="Zucchi R">RA Zucchi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Feder, Jl" uniqKey="Feder J">JL Feder</name></author><author><name sortKey="Berlocher, Sh" uniqKey="Berlocher S">SH Berlocher</name></author><author><name sortKey="Roethele, Jb" uniqKey="Roethele J">JB Roethele</name></author><author><name sortKey="Dambroski, H" uniqKey="Dambroski H">H Dambroski</name></author><author><name sortKey="Smith, Jj" uniqKey="Smith J">JJ Smith</name></author><author><name sortKey="Perry, Wl" uniqKey="Perry W">WL Perry</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Feder, Jl" uniqKey="Feder J">JL Feder</name></author><author><name sortKey="Xie, X" uniqKey="Xie X">X Xie</name></author><author><name sortKey="Rull, J" uniqKey="Rull J">J Rull</name></author><author><name sortKey="Velez, S" uniqKey="Velez S">S Velez</name></author><author><name sortKey="Forbes, A" uniqKey="Forbes A">A Forbes</name></author><author><name sortKey="Leung, B" uniqKey="Leung B">B Leung</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Virgilio, M" uniqKey="Virgilio M">M Virgilio</name></author><author><name sortKey="Delatte, H" uniqKey="Delatte H">H Delatte</name></author><author><name sortKey="Backeljau, T" uniqKey="Backeljau T">T Backeljau</name></author><author><name sortKey="De Meyer, M" uniqKey="De Meyer M">M De Meyer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Michel, Ap" uniqKey="Michel A">AP Michel</name></author><author><name sortKey="Rull, J" uniqKey="Rull J">J Rull</name></author><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author><author><name sortKey="Feder, Jl" uniqKey="Feder J">JL Feder</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Lude A, B" uniqKey="Lude A B">B Ludeña</name></author><author><name sortKey="Bayas, R" uniqKey="Bayas R">R Bayas</name></author><author><name sortKey="Pintaud, J C" uniqKey="Pintaud J">J-C Pintaud</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hernandez Ortiz, V" uniqKey="Hernandez Ortiz V">V Hernández-Ortiz</name></author><author><name sortKey="Bartolucci, Af" uniqKey="Bartolucci A">AF Bartolucci</name></author><author><name sortKey="Morales Valles, P" uniqKey="Morales Valles P">P Morales-Valles</name></author><author><name sortKey="Frias, D" uniqKey="Frias D">D Frías</name></author><author><name sortKey="Selivon, D" uniqKey="Selivon D">D Selivon</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zygouridis, Ne" uniqKey="Zygouridis N">NE Zygouridis</name></author><author><name sortKey="Augustinos, Aa" uniqKey="Augustinos A">AA Augustinos</name></author><author><name sortKey="Zalom, Fg" uniqKey="Zalom F">FG Zalom</name></author><author><name sortKey="Mathiopoulos, Kd" uniqKey="Mathiopoulos K">KD Mathiopoulos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Van Asch, B" uniqKey="Van Asch B">B van Asch</name></author><author><name sortKey="Pereira Castro, I" uniqKey="Pereira Castro I">I Pereira-Castro</name></author><author><name sortKey="Rei, F" uniqKey="Rei F">F Rei</name></author><author><name sortKey="Costa, Lt" uniqKey="Costa L">LT Costa</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Daane, Km" uniqKey="Daane K">KM Daane</name></author><author><name sortKey="Johnson, Mw" uniqKey="Johnson M">MW Johnson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author><author><name sortKey="Pi Ero, J" uniqKey="Pi Ero J">J Piñero</name></author><author><name sortKey="L Pez, M" uniqKey="L Pez M">M López</name></author><author><name sortKey="Ruiz, C" uniqKey="Ruiz C">C Ruíz</name></author><author><name sortKey="Zu Iga, A" uniqKey="Zu Iga A">A Zúñiga</name></author><author><name sortKey="Piedra, E" uniqKey="Piedra E">E Piedra</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Md, Meyer" uniqKey="Md M">Meyer Md</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Copeland, Rs" uniqKey="Copeland R">RS Copeland</name></author><author><name sortKey="Wharton, Ra" uniqKey="Wharton R">RA Wharton</name></author><author><name sortKey="Luke, Q" uniqKey="Luke Q">Q Luke</name></author><author><name sortKey="De Meyer, M" uniqKey="De Meyer M">M De Meyer</name></author><author><name sortKey="Lux, S" uniqKey="Lux S">S Lux</name></author><author><name sortKey="Zenz, N" uniqKey="Zenz N">N Zenz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Virgilio, M" uniqKey="Virgilio M">M Virgilio</name></author><author><name sortKey="Backeljau, T" uniqKey="Backeljau T">T Backeljau</name></author><author><name sortKey="Barr, N" uniqKey="Barr N">N Barr</name></author><author><name sortKey="Meyer, Md" uniqKey="Meyer M">MD Meyer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hernandez Ortiz, V" uniqKey="Hernandez Ortiz V">V Hernández-Ortiz</name></author><author><name sortKey="G Mez Anaya, J" uniqKey="G Mez Anaya J">J Gómez-Anaya</name></author><author><name sortKey="Sanchez, A" uniqKey="Sanchez A">A Sánchez</name></author><author><name sortKey="Mcpheron, B" uniqKey="Mcpheron B">B McPheron</name></author><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Vera, Mt" uniqKey="Vera M">MT Vera</name></author><author><name sortKey="Caceres, C" uniqKey="Caceres C">C Cáceres</name></author><author><name sortKey="Wornoayporn, V" uniqKey="Wornoayporn V">V Wornoayporn</name></author><author><name sortKey="Islam, A" uniqKey="Islam A">A Islam</name></author><author><name sortKey="Robinson, As" uniqKey="Robinson A">AS Robinson</name></author><author><name sortKey="Marcelo, H" uniqKey="Marcelo H">H Marcelo</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Alberti, Ac" uniqKey="Alberti A">AC Alberti</name></author><author><name sortKey="Rodriguero, Ms" uniqKey="Rodriguero M">MS Rodriguero</name></author><author><name sortKey="Cendra, Pg" uniqKey="Cendra P">PG Cendra</name></author><author><name sortKey="Saidman, Bo" uniqKey="Saidman B">BO Saidman</name></author><author><name sortKey="Vilardi, Jc" uniqKey="Vilardi J">JC Vilardi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Selivon, D" uniqKey="Selivon D">D Selivon</name></author><author><name sortKey="Perondini, A" uniqKey="Perondini A">A Perondini</name></author><author><name sortKey="Morgante, J" uniqKey="Morgante J">J Morgante</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Alberti, Ac" uniqKey="Alberti A">AC Alberti</name></author><author><name sortKey="Confalonieri, Va" uniqKey="Confalonieri V">VA Confalonieri</name></author><author><name sortKey="Zandomeni, Ro" uniqKey="Zandomeni R">RO Zandomeni</name></author><author><name sortKey="Vilardi, Jc" uniqKey="Vilardi J">JC Vilardi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Wisz, Ms" uniqKey="Wisz M">MS Wisz</name></author><author><name sortKey="Hijmans, R" uniqKey="Hijmans R">R Hijmans</name></author><author><name sortKey="Li, J" uniqKey="Li J">J Li</name></author><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Graham, C" uniqKey="Graham C">C Graham</name></author><author><name sortKey="Guisan, A" uniqKey="Guisan A">A Guisan</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hernandez, Pa" uniqKey="Hernandez P">PA Hernandez</name></author><author><name sortKey="Graham, Ch" uniqKey="Graham C">CH Graham</name></author><author><name sortKey="Master, Ll" uniqKey="Master L">LL Master</name></author><author><name sortKey="Albert, Dl" uniqKey="Albert D">DL Albert</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rull, J" uniqKey="Rull J">J Rull</name></author><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author><author><name sortKey="Feder, Jl" uniqKey="Feder J">JL Feder</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Wu, Y" uniqKey="Wu Y">Y Wu</name></author><author><name sortKey="Mcpheron, Ba" uniqKey="Mcpheron B">BA McPheron</name></author><author><name sortKey="Wu, Jj" uniqKey="Wu J">JJ Wu</name></author><author><name sortKey="Li, Zh" uniqKey="Li Z">ZH Li</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hijmans, Rj" uniqKey="Hijmans R">RJ Hijmans</name></author><author><name sortKey="Cameron, Se" uniqKey="Cameron S">SE Cameron</name></author><author><name sortKey="Parra, Jl" uniqKey="Parra J">JL Parra</name></author><author><name sortKey="Jones, Pg" uniqKey="Jones P">PG Jones</name></author><author><name sortKey="Jarvis, A" uniqKey="Jarvis A">A Jarvis</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Nakazawa, Y" uniqKey="Nakazawa Y">Y Nakazawa</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jimenez Valverde, A" uniqKey="Jimenez Valverde A">A Jiménez-Valverde</name></author><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Sober N, J" uniqKey="Sober N J">J Soberón</name></author><author><name sortKey="Overton, Jm" uniqKey="Overton J">JM Overton</name></author><author><name sortKey="Arag N, P" uniqKey="Arag N P">P Aragón</name></author><author><name sortKey="Lobo, Jm" uniqKey="Lobo J">JM Lobo</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Phillips, Sj" uniqKey="Phillips S">SJ Phillips</name></author><author><name sortKey="Anderson, Rp" uniqKey="Anderson R">RP Anderson</name></author><author><name sortKey="Schapire, Re" uniqKey="Schapire R">RE Schapire</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Friedman, J" uniqKey="Friedman J">J Friedman</name></author><author><name sortKey="Hastie, T" uniqKey="Hastie T">T Hastie</name></author><author><name sortKey="Tibshirani, R" uniqKey="Tibshirani R">R Tibshirani</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Elith, J" uniqKey="Elith J">J Elith</name></author><author><name sortKey="Leathwick, Jr" uniqKey="Leathwick J">JR Leathwick</name></author><author><name sortKey="Hastie, T" uniqKey="Hastie T">T Hastie</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Ridgeway, G" uniqKey="Ridgeway G">G Ridgeway</name></author><author><name sortKey="Ridgeway, Mg" uniqKey="Ridgeway M">MG Ridgeway</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Elith, J" uniqKey="Elith J">J Elith</name></author><author><name sortKey="H Graham, C" uniqKey="H Graham C">C H. Graham</name></author><author><name sortKey="P Anderson, R" uniqKey="P Anderson R">R P. Anderson</name></author><author><name sortKey="Dudik, M" uniqKey="Dudik M">M Dudík</name></author><author><name sortKey="Ferrier, S" uniqKey="Ferrier S">S Ferrier</name></author><author><name sortKey="Guisan, A" uniqKey="Guisan A">A Guisan</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Vanderwal, J" uniqKey="Vanderwal J">J VanDerWal</name></author><author><name sortKey="Shoo, Lp" uniqKey="Shoo L">LP Shoo</name></author><author><name sortKey="Graham, C" uniqKey="Graham C">C Graham</name></author><author><name sortKey="Williams, Se" uniqKey="Williams S">SE Williams</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Anderson, Rp" uniqKey="Anderson R">RP Anderson</name></author><author><name sortKey="Raza, A" uniqKey="Raza A">A Raza</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Elith, J" uniqKey="Elith J">J Elith</name></author><author><name sortKey="Kearney, M" uniqKey="Kearney M">M Kearney</name></author><author><name sortKey="Phillips, S" uniqKey="Phillips S">S Phillips</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Fielding, Ah" uniqKey="Fielding A">AH Fielding</name></author><author><name sortKey="Bell, Jf" uniqKey="Bell J">JF Bell</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Legendre, L" uniqKey="Legendre L">L Legendre</name></author><author><name sortKey="Legrendre, L" uniqKey="Legrendre L">L Legrendre</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rossi, J P" uniqKey="Rossi J">J-P Rossi</name></author><author><name sortKey="Blanchart, E" uniqKey="Blanchart E">E Blanchart</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Doledec, S" uniqKey="Doledec S">S Dolédec</name></author><author><name sortKey="Chessel, D" uniqKey="Chessel D">D Chessel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Manly, Bf" uniqKey="Manly B">BF Manly</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schoener, Tw" uniqKey="Schoener T">TW Schoener</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Warren, Dl" uniqKey="Warren D">DL Warren</name></author><author><name sortKey="Glor, Re" uniqKey="Glor R">RE Glor</name></author><author><name sortKey="Turelli, M" uniqKey="Turelli M">M Turelli</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mccormack, Je" uniqKey="Mccormack J">JE McCormack</name></author><author><name sortKey="Zellmer, Aj" uniqKey="Zellmer A">AJ Zellmer</name></author><author><name sortKey="Knowles, Ll" uniqKey="Knowles L">LL Knowles</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Broennimann, O" uniqKey="Broennimann O">O Broennimann</name></author><author><name sortKey="Fitzpatrick, Mc" uniqKey="Fitzpatrick M">MC Fitzpatrick</name></author><author><name sortKey="Pearman, Pb" uniqKey="Pearman P">PB Pearman</name></author><author><name sortKey="Petitpierre, B" uniqKey="Petitpierre B">B Petitpierre</name></author><author><name sortKey="Pellissier, L" uniqKey="Pellissier L">L Pellissier</name></author><author><name sortKey="Yoccoz, Ng" uniqKey="Yoccoz N">NG Yoccoz</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Guisan, A" uniqKey="Guisan A">A Guisan</name></author><author><name sortKey="Petitpierre, B" uniqKey="Petitpierre B">B Petitpierre</name></author><author><name sortKey="Broennimann, O" uniqKey="Broennimann O">O Broennimann</name></author><author><name sortKey="Daehler, C" uniqKey="Daehler C">C Daehler</name></author><author><name sortKey="Kueffer, C" uniqKey="Kueffer C">C Kueffer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Godsoe, W" uniqKey="Godsoe W">W Godsoe</name></author><author><name sortKey="Case, Bs" uniqKey="Case B">BS Case</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Barve, N" uniqKey="Barve N">N Barve</name></author><author><name sortKey="Barve, V" uniqKey="Barve V">V Barve</name></author><author><name sortKey="Jimenez Valverde, A" uniqKey="Jimenez Valverde A">A Jiménez-Valverde</name></author><author><name sortKey="Lira Noriega, A" uniqKey="Lira Noriega A">A Lira-Noriega</name></author><author><name sortKey="Maher, Sp" uniqKey="Maher S">SP Maher</name></author><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Webber, Bl" uniqKey="Webber B">BL Webber</name></author><author><name sortKey="Yates, Cj" uniqKey="Yates C">CJ Yates</name></author><author><name sortKey="Le Maitre, Dc" uniqKey="Le Maitre D">DC Le Maitre</name></author><author><name sortKey="Scott, Jk" uniqKey="Scott J">JK Scott</name></author><author><name sortKey="Kriticos, Dj" uniqKey="Kriticos D">DJ Kriticos</name></author><author><name sortKey="Ota, N" uniqKey="Ota N">N Ota</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Duyck, P F" uniqKey="Duyck P">P-F Duyck</name></author><author><name sortKey="David, P" uniqKey="David P">P David</name></author><author><name sortKey="Quilici, S" uniqKey="Quilici S">S Quilici</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Katiyar, Kp" uniqKey="Katiyar K">KP Katiyar</name></author><author><name sortKey="Molina, Jc" uniqKey="Molina J">JC Molina</name></author><author><name sortKey="Matheus, R" uniqKey="Matheus R">R Matheus</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sivinski, J" uniqKey="Sivinski J">J Sivinski</name></author><author><name sortKey="Aluja, M" uniqKey="Aluja M">M Aluja</name></author><author><name sortKey="Pi Ero, J" uniqKey="Pi Ero J">J Piñero</name></author><author><name sortKey="Ojeda, M" uniqKey="Ojeda M">M Ojeda</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="De Villiers, M" uniqKey="De Villiers M">M De Villiers</name></author><author><name sortKey="Hattingh, V" uniqKey="Hattingh V">V Hattingh</name></author><author><name sortKey="Kriticos, D" uniqKey="Kriticos D">D Kriticos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Papadopoulos, Nt" uniqKey="Papadopoulos N">NT Papadopoulos</name></author><author><name sortKey="Carey, Jr" uniqKey="Carey J">JR Carey</name></author><author><name sortKey="Katsoyannos, Bi" uniqKey="Katsoyannos B">BI Katsoyannos</name></author><author><name sortKey="Kouloussis, Na" uniqKey="Kouloussis N">NA Kouloussis</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Vera, Mt" uniqKey="Vera M">MT Vera</name></author><author><name sortKey="Rodriguez, R" uniqKey="Rodriguez R">R Rodriguez</name></author><author><name sortKey="Segura, Df" uniqKey="Segura D">DF Segura</name></author><author><name sortKey="Cladera, Jl" uniqKey="Cladera J">JL Cladera</name></author><author><name sortKey="Sutherst, Rw" uniqKey="Sutherst R">RW Sutherst</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="De Meyer, M" uniqKey="De Meyer M">M De Meyer</name></author><author><name sortKey="Robertson, Mp" uniqKey="Robertson M">MP Robertson</name></author><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author><author><name sortKey="Mansell, Mw" uniqKey="Mansell M">MW Mansell</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Gutierrez, Ap" uniqKey="Gutierrez A">AP Gutierrez</name></author><author><name sortKey="Ponti, L" uniqKey="Ponti L">L Ponti</name></author><author><name sortKey="Cossu, Q" uniqKey="Cossu Q">Q Cossu</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Brooks, Cp" uniqKey="Brooks C">CP Brooks</name></author><author><name sortKey="Ervin, Gn" uniqKey="Ervin G">GN Ervin</name></author><author><name sortKey="Varone, L" uniqKey="Varone L">L Varone</name></author><author><name sortKey="Logarzo, Ga" uniqKey="Logarzo G">GA Logarzo</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Yokoyama, Vy" uniqKey="Yokoyama V">VY Yokoyama</name></author><author><name sortKey="Rend N, Pa" uniqKey="Rend N P">PA Rendón</name></author><author><name sortKey="Sivinski, J" uniqKey="Sivinski J">J Sivinski</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Daane, K" uniqKey="Daane K">K Daane</name></author><author><name sortKey="Johnson, M" uniqKey="Johnson M">M Johnson</name></author><author><name sortKey="Lynn Patterson, K" uniqKey="Lynn Patterson K">K Lynn-Patterson</name></author><author><name sortKey="Nadel, H" uniqKey="Nadel H">H Nadel</name></author><author><name sortKey="Opp, S" uniqKey="Opp S">S Opp</name></author><author><name sortKey="Stewart Leslie, J" uniqKey="Stewart Leslie J">J Stewart-Leslie</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Pearman, Pb" uniqKey="Pearman P">PB Pearman</name></author><author><name sortKey="Guisan, A" uniqKey="Guisan A">A Guisan</name></author><author><name sortKey="Broennimann, O" uniqKey="Broennimann O">O Broennimann</name></author><author><name sortKey="Randin, Cf" uniqKey="Randin C">CF Randin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Thompson, Gd" uniqKey="Thompson G">GD Thompson</name></author><author><name sortKey="Robertson, Mp" uniqKey="Robertson M">MP Robertson</name></author><author><name sortKey="Webber, Bl" uniqKey="Webber B">BL Webber</name></author><author><name sortKey="Richardson, Dm" uniqKey="Richardson D">DM Richardson</name></author><author><name sortKey="Le Roux, Jj" uniqKey="Le Roux J">JJ Le Roux</name></author><author><name sortKey="Wilson, Jru" uniqKey="Wilson J">JRU Wilson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schulte, U" uniqKey="Schulte U">U Schulte</name></author><author><name sortKey="Hochkirch, A" uniqKey="Hochkirch A">A Hochkirch</name></author><author><name sortKey="Lotters, S" uniqKey="Lotters S">S Lötters</name></author><author><name sortKey="Rodder, D" uniqKey="Rodder D">D Rödder</name></author><author><name sortKey="Schweiger, S" uniqKey="Schweiger S">S Schweiger</name></author><author><name sortKey="Weimann, T" uniqKey="Weimann T">T Weimann</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Beaumont, Lj" uniqKey="Beaumont L">LJ Beaumont</name></author><author><name sortKey="Gallagher, Rv" uniqKey="Gallagher R">RV Gallagher</name></author><author><name sortKey="Leishman, Mr" uniqKey="Leishman M">MR Leishman</name></author><author><name sortKey="Hughes, L" uniqKey="Hughes L">L Hughes</name></author><author><name sortKey="Downey, Po" uniqKey="Downey P">PO Downey</name></author><author><name sortKey="Wilson, J" uniqKey="Wilson J">J Wilson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sober N, J" uniqKey="Sober N J">J Soberón</name></author><author><name sortKey="Peterson, At" uniqKey="Peterson A">AT Peterson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Larson, Er" uniqKey="Larson E">ER Larson</name></author><author><name sortKey="Olden, Jd" uniqKey="Olden J">JD Olden</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Larson, Er" uniqKey="Larson E">ER Larson</name></author><author><name sortKey="Gallagher, Rv" uniqKey="Gallagher R">RV Gallagher</name></author><author><name sortKey="Beaumont, Lj" uniqKey="Beaumont L">LJ Beaumont</name></author><author><name sortKey="Olden, Jd" uniqKey="Olden J">JD Olden</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Michalak, P" uniqKey="Michalak P">P Michalak</name></author><author><name sortKey="Minkov, I" uniqKey="Minkov I">I Minkov</name></author><author><name sortKey="Helin, A" uniqKey="Helin A">A Helin</name></author><author><name sortKey="Lerman, Dn" uniqKey="Lerman D">DN Lerman</name></author><author><name sortKey="Bettencourt, Br" uniqKey="Bettencourt B">BR Bettencourt</name></author><author><name sortKey="Feder, Me" uniqKey="Feder M">ME Feder</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Rehfeldt, Ge" uniqKey="Rehfeldt G">GE Rehfeldt</name></author><author><name sortKey="Ying, Cc" uniqKey="Ying C">CC Ying</name></author><author><name sortKey="Spittlehouse, Dl" uniqKey="Spittlehouse D">DL Spittlehouse</name></author><author><name sortKey="Hamilton, Da" uniqKey="Hamilton D">DA Hamilton</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Keller, Rp" uniqKey="Keller R">RP Keller</name></author><author><name sortKey="Lodge, Dm" uniqKey="Lodge D">DM Lodge</name></author><author><name sortKey="Finnoff, Dc" uniqKey="Finnoff D">DC Finnoff</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hebert, Pd" uniqKey="Hebert P">PD Hebert</name></author><author><name sortKey="Cywinska, A" uniqKey="Cywinska A">A Cywinska</name></author><author><name sortKey="Ball, Sl" uniqKey="Ball S">SL Ball</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Armstrong, K" uniqKey="Armstrong K">K Armstrong</name></author><author><name sortKey="Ball, S" uniqKey="Ball S">S Ball</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Coeur D Cier, A" uniqKey="Coeur D Cier A">A Coeur d’acier</name></author><author><name sortKey="Cruaud, A" uniqKey="Cruaud A">A Cruaud</name></author><author><name sortKey="Artige, E" uniqKey="Artige E">E Artige</name></author><author><name sortKey="Genson, G" uniqKey="Genson G">G Genson</name></author><author><name sortKey="Clamens, A L" uniqKey="Clamens A">A-L Clamens</name></author><author><name sortKey="Pierre, E" uniqKey="Pierre E">E Pierre</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Van Houdt, Jkj" uniqKey="Van Houdt J">JKJ Van Houdt</name></author><author><name sortKey="Breman, Fc" uniqKey="Breman F">FC Breman</name></author><author><name sortKey="Virgilio, M" uniqKey="Virgilio M">M Virgilio</name></author><author><name sortKey="De Meyer, M" uniqKey="De Meyer M">M De Meyer</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">PLoS One</journal-id><journal-id journal-id-type="iso-abbrev">PLoS ONE</journal-id><journal-id journal-id-type="publisher-id">plos</journal-id><journal-id journal-id-type="pmc">plosone</journal-id><journal-title-group><journal-title>PLoS ONE</journal-title></journal-title-group><issn pub-type="epub">1932-6203</issn><publisher><publisher-name>Public Library of Science</publisher-name><publisher-loc>San Francisco, CA USA</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26274582</article-id><article-id pub-id-type="pmc">4537207</article-id><article-id pub-id-type="doi">10.1371/journal.pone.0135209</article-id><article-id pub-id-type="publisher-id">PONE-D-15-12728</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters</article-title><alt-title alt-title-type="running-head">Assessing the Risk of Invasion by Tephritid Fruit Flies</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Godefroid</surname><given-names>Martin</given-names></name><xref rid="cor001" ref-type="corresp">*</xref><xref ref-type="aff" rid="aff001"></xref></contrib><contrib contrib-type="author"><name><surname>Cruaud</surname><given-names>Astrid</given-names></name><xref ref-type="aff" rid="aff001"></xref></contrib><contrib contrib-type="author"><name><surname>Rossi</surname><given-names>Jean-Pierre</given-names></name><xref ref-type="aff" rid="aff001"></xref></contrib><contrib contrib-type="author"><name><surname>Rasplus</surname><given-names>Jean-Yves</given-names></name><xref ref-type="aff" rid="aff001"></xref></contrib></contrib-group><aff id="aff001"><addr-line>Centre de Biologie pour la Gestion des Populations, Institut National de la Recherche Agronomique, Montferriez-sur-Lez, France</addr-line></aff><contrib-group><contrib contrib-type="editor"><name><surname>Papadopoulos</surname><given-names>Nikos T</given-names></name><role>Editor</role><xref ref-type="aff" rid="edit1"></xref></contrib></contrib-group><aff id="edit1"><addr-line>University of Thessaly, GREECE</addr-line></aff><author-notes><fn fn-type="conflict" id="coi001"><p><bold>Competing Interests: </bold>The authors have declared that no competing interests exist.</p></fn><fn fn-type="con" id="contrib001"><p>Conceived and designed the experiments: MG AC JYR JPR. Analyzed the data: MG AC JYR JPR. Wrote the paper: MG AC JYR JPR.</p></fn><corresp id="cor001">* E-mail: <email>martin.godefroid@gmail.com</email></corresp></author-notes><pub-date pub-type="epub"><day>14</day><month>8</month><year>2015</year></pub-date><pub-date pub-type="collection"><year>2015</year></pub-date><volume>10</volume><issue>8</issue><elocation-id>e0135209</elocation-id><history><date date-type="received"><day>24</day><month>3</month><year>2015</year></date><date date-type="accepted"><day>19</day><month>7</month><year>2015</year></date></history><permissions><copyright-year>2015</copyright-year><copyright-holder>Godefroid et al</copyright-holder><license xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>This is an open access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited</license-p></license></permissions><self-uri content-type="pdf" xlink:type="simple" xlink:href="pone.0135209.pdf"></self-uri><abstract><p>Widely distributed species often show strong phylogeographic structure, with lineages potentially adapted to different biotic and abiotic conditions. The success of an invasion process may thus depend on the intraspecific identity of the introduced propagules. However, pest risk analyses are usually performed without accounting for intraspecific diversity. In this study, we developed bioclimatic models using MaxEnt and boosted regression trees approaches, to predict the potential distribution in Europe of six economically important Tephritid pests (<italic>Ceratitis fasciventris</italic> (Bezzi), <italic>Bactrocera oleae</italic> (Rossi), <italic>Anastrepha obliqua</italic> (Macquart), <italic>Anastrepha fraterculus</italic> (Wiedemann), <italic>Rhagoletis pomonella</italic> (Walsh) and <italic>Bactrocera cucurbitae</italic> (Coquillet)). We considered intraspecific diversity in our risk analyses by independently modeling the distributions of conspecific lineages. The six species displayed different potential distributions in Europe. A strong signal of intraspecific climate envelope divergence was observed in most species. In some cases, conspecific lineages differed strongly in potential distributions suggesting that taxonomic resolution should be accounted for in pest risk analyses. No models (lineage- and species-based approaches) predicted high climatic suitability in the entire invaded range of <italic>B</italic>. <italic>oleae</italic>—the only species whose intraspecific identity of invading populations has been elucidated—in California. Host availability appears to play the most important role in shaping the geographic range of this specialist pest. However, climatic suitability values predicted by species-based models are correlated with population densities of <italic>B</italic>. <italic>oleae</italic> globally reported in California. Our study highlights how classical taxonomic boundaries may lead to under- or overestimation of the potential pest distributions and encourages accounting for intraspecific diversity when assessing the risk of biological invasion.</p></abstract><funding-group><funding-statement>This work was financially supported by the Metaprogram SMACH (Sustainable Management of Crop Health; <ext-link ext-link-type="uri" xlink:href="http://www.smach.inra.fr/">http://www.smach.inra.fr/</ext-link>) of the French National Agronomic Institute (INRA). This work was also based on financial support received from the BIOFIS (Bioagressors and invasive species: From individual to population to species) project (number 1001-001) allocated by Agropolis Fondation to JYR (Montpellier, France; <ext-link ext-link-type="uri" xlink:href="http://www.agropolis-fondation.fr/fr/accueil.html">http://www.agropolis-fondation.fr/fr/accueil.html</ext-link>).</funding-statement></funding-group><counts><fig-count count="4"></fig-count><table-count count="3"></table-count><page-count count="19"></page-count></counts><custom-meta-group><custom-meta id="data-availability"><meta-name>Data Availability</meta-name><meta-value>All relevant data are within the paper and its Supporting Information files.</meta-value></custom-meta></custom-meta-group></article-meta><notes><title>Data Availability</title><p>All relevant data are within the paper and its Supporting Information files.</p></notes></front><body><sec sec-type="intro" id="sec001"><title>Introduction</title><p>Controlling invasive species has become a task of utmost importance as the rate of biological invasions increases, leading to a global economic cost estimated at US$ 1.4 trillion per year, representing nearly 5% of the worldwide economy [<xref rid="pone.0135209.ref001" ref-type="bibr">1</xref>]. Because commercial exchanges are continuously increasing and since human activities lead to important environmental changes, numerous pest species are—and will be—susceptible to geographical expansion, thereby causing new problems for agriculture [<xref rid="pone.0135209.ref002" ref-type="bibr">2</xref>]. An important element of control lies in the development of powerful and accurate predictive tools, allowing a proper risk assessment [<xref rid="pone.0135209.ref003" ref-type="bibr">3</xref>], which, in turn, will be helpful to prevent the introduction of alien species and design cost-effective management strategies. Pest risk analyses (PRAs) address organisms' biology and ecology to discriminate potentially innocuous exotic species from harmful ones. Besides integrating the multiplicity of ecological and anthropological factors susceptible to enhance invasion risk, estimating climatic tolerances of exotic species constitutes a crucial point in PRAs. Pest risk analyses often rely on species distribution models (SDMs) when estimating the potential distribution of target species outside their current geographical range [<xref rid="pone.0135209.ref004" ref-type="bibr">4</xref>]. The existence of different intraspecific lineages potentially adapted to different biotic (e.g. host-plant [<xref rid="pone.0135209.ref005" ref-type="bibr">5</xref>]) and abiotic conditions (e.g. climatic tolerances [<xref rid="pone.0135209.ref006" ref-type="bibr">6</xref>, <xref rid="pone.0135209.ref007" ref-type="bibr">7</xref>]) is generally ignored. However, there is increasing evidence that the origin of introduced population(s) might determine the success of an invasion [<xref rid="pone.0135209.ref007" ref-type="bibr">7</xref>–<xref rid="pone.0135209.ref010" ref-type="bibr">10</xref>]. Thus, as suggested by Peterson & Holt [<xref rid="pone.0135209.ref011" ref-type="bibr">11</xref>], in cases where models have been developed without taking into account the phylogeographic structure of the potential invader, invasion risk could be under- or overestimated, leading to inappropriate phytosanitary measures.</p><p>The ‘true’ fruit flies (Diptera, Tephritidae) are important agricultural pests worldwide [<xref rid="pone.0135209.ref012" ref-type="bibr">12</xref>]. The larvae of most species are phytophagous and attack tissues of a wide spectrum of plants, especially fruits and flowers, significantly reducing crop yields. As a result, many species are of high economic importance (especially in the genera <italic>Bactrocera</italic>, <italic>Anastrepha</italic>, <italic>Rhagoletis</italic> and <italic>Ceratitis)</italic>. In addition, several species (e.g. <italic>Bactrocera dorsalis</italic> (Hendel, 1912), <italic>Bactrocera cucurbitae</italic> (Coquillet, 1899), <italic>Bactrocera oleae</italic> (Rossi, 1790), <italic>Bactrocera depressa</italic> (Shiraki, 1933), <italic>Ceratitis capitata</italic> (Wiedemann, 1824), and <italic>Ceratitis rosa</italic> Karsch, 1887) have been successful invasive species over the past decades [<xref rid="pone.0135209.ref013" ref-type="bibr">13</xref>–<xref rid="pone.0135209.ref019" ref-type="bibr">19</xref>]. For these reasons, tephritid flies are indiscriminately listed as quarantine organisms for Europe. Several tephritid species show strong genetic structure and some are considered species complexes (e.g. the <italic>Anastrepha fraterculus</italic> group, the <italic>Bactrocera dorsalis</italic> group and the <italic>Ceratitis fasciventris</italic>, <italic>C</italic>. <italic>rosa</italic>, <italic>C</italic>. <italic>anonae</italic> complexes) [<xref rid="pone.0135209.ref020" ref-type="bibr">20</xref>–<xref rid="pone.0135209.ref024" ref-type="bibr">24</xref>]. Lineages can display different life history traits related to adaptations to local climate [<xref rid="pone.0135209.ref025" ref-type="bibr">25</xref>–<xref rid="pone.0135209.ref029" ref-type="bibr">29</xref>] or different host preferences [<xref rid="pone.0135209.ref030" ref-type="bibr">30</xref>], which make control strategies more difficult and lead to the validity of species-based PRAs to be questioned. Tephritid flies are thus good candidates to test whether the phylogeographic structure of a potential invasive pest should be considered when modeling species distribution in the context of invasion risk assessment.</p><p>In the present paper, we first used the classical species-based approach to assess the potential distribution in Europe of these economically important tephritid species. In a second step, we developed a set of models accounting the phylogeographic structure of the species.</p></sec><sec id="sec002"><title>Material & Methods</title><sec id="sec003"><title>Species data and phylogeographic patterns</title><p>We studied six, widely distributed and economically important tephritid species, namely <italic>Bactrocera oleae</italic>, <italic>Bactrocera cucurbitae</italic>, <italic>Anastrepha obliqua</italic> (Macquart, 1835), <italic>Anastrepha fraterculus</italic> (Wiedemann, 1830), <italic>Rhagoletis pomonella</italic> (Walsh, 1867) and <italic>Ceratitis fasciventris</italic> (Bezzi, 1920). We chose these species because their phylogeographic structure has previously been described in the literature [<xref rid="pone.0135209.ref021" ref-type="bibr">21</xref>, <xref rid="pone.0135209.ref023" ref-type="bibr">23</xref>, <xref rid="pone.0135209.ref031" ref-type="bibr">31</xref>–<xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>]. Most occurrences were collected from the scientific literature (more than 95 publications; <xref rid="pone.0135209.s001" ref-type="supplementary-material">S1 File</xref>) and online databases (e.g. Global Biodiversity Information Facility (GBIF), National Agricultural Pest Information System (NAPIS) and the BioSystematics Database of World Diptera). Doubtful or imprecise records were removed from our datasets. We assigned each occurrence to a genetic lineage according to the phylogeographic pattern described in the relevant publication (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). Several records were assigned to more than one lineage, since several lineages occurred in the same geographical area. Some records were used in species-based models only when assignation to one lineage was challenging according to phylogeographic studies (<xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>).</p><table-wrap id="pone.0135209.t001" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.t001</object-id><label>Table 1</label><caption><title>Tephritid fruit flies selected to assess invasion risk and integrate intraspecific diversity in species distribution models.</title></caption><alternatives><graphic id="pone.0135209.t001g" xlink:href="pone.0135209.t001"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col></colgroup><thead><tr><th align="left" rowspan="1" colspan="1">Species</th><th align="left" rowspan="1" colspan="1">Lineages</th><th align="left" rowspan="1" colspan="1">Symbol</th><th align="left" rowspan="1" colspan="1">Geographic distribution</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>Bactrocera oleae</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Bo_all</td><td align="left" rowspan="1" colspan="1">Africa, Asia, Europe, Americas</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Middle East lineage</td><td align="left" rowspan="1" colspan="1">Bo_me</td><td align="left" rowspan="1" colspan="1">Israel, Turkey, Cyprus, Syria</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Western Europe lineage</td><td align="left" rowspan="1" colspan="1">Bo_we</td><td align="left" rowspan="1" colspan="1">Spain, Italy, Greece, France, Morocco, Tunisia, Turkey, Algeria</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Africa lineage</td><td align="left" rowspan="1" colspan="1">Bo_af</td><td align="left" rowspan="1" colspan="1">Africa</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Asia lineage</td><td align="left" rowspan="1" colspan="1">bo_as</td><td align="left" rowspan="1" colspan="1">Pakistan</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Ceratitis fasciventris</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Cf_all</td><td align="left" rowspan="1" colspan="1">Africa</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Western Africa lineage</td><td align="left" rowspan="1" colspan="1">Cf_we</td><td align="left" rowspan="1" colspan="1">Western Africa & Eastern Tanzania</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Eastern Africa lineage</td><td align="left" rowspan="1" colspan="1">Cf_ke</td><td align="left" rowspan="1" colspan="1">Eastern & Central Africa</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Anastrepha obliqua</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Ao_all</td><td align="left" rowspan="1" colspan="1">Mexico, South and Central America, Caribbean</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Northern lineage</td><td align="left" rowspan="1" colspan="1">Ao_01</td><td align="left" rowspan="1" colspan="1">Central America, Caribbean, Northern Andean</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Western Mexico lineage</td><td align="left" rowspan="1" colspan="1">Ao_wm</td><td align="left" rowspan="1" colspan="1">Western & Southeastern Mexico</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">South America lineage</td><td align="left" rowspan="1" colspan="1">Ao_03</td><td align="left" rowspan="1" colspan="1">South America, Northern Andean, Panama</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Anastrepha fraterculus</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Af_all</td><td align="left" rowspan="1" colspan="1">USA, Mexico, Central America, South America</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Mexican lineage</td><td align="left" rowspan="1" colspan="1">Af_mex</td><td align="left" rowspan="1" colspan="1">Mexico & Central America</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Brazilian lineage</td><td align="left" rowspan="1" colspan="1">Af_bra</td><td align="left" rowspan="1" colspan="1">South America</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Andean lineage</td><td align="left" rowspan="1" colspan="1">Af_and</td><td align="left" rowspan="1" colspan="1">Northern Andean (high altitude)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Rhagoletis pomonella</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Rp_all</td><td align="left" rowspan="1" colspan="1">USA, Canada & Mexico</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">USA lineage</td><td align="left" rowspan="1" colspan="1">Rp_usa</td><td align="left" rowspan="1" colspan="1">USA & Canada</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Mexican lineages</td><td align="left" rowspan="1" colspan="1">Rp_mex</td><td align="left" rowspan="1" colspan="1">Mexico</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Bactrocera cucurbitae</italic></td><td align="left" rowspan="1" colspan="1">All lineages</td><td align="left" rowspan="1" colspan="1">Bc_all</td><td align="left" rowspan="1" colspan="1">Asia, Africa, USA, La Réunion</td></tr></tbody></table></alternatives></table-wrap><p>The olive fruit fly <italic>B</italic>. <italic>oleae</italic> occurs in Africa, Europe and Asia and has recently invaded California and Mexico [<xref rid="pone.0135209.ref038" ref-type="bibr">38</xref>]. Genetic analyses suggest the Middle East region as the most likely origin of invading populations [<xref rid="pone.0135209.ref021" ref-type="bibr">21</xref>, <xref rid="pone.0135209.ref022" ref-type="bibr">22</xref>, <xref rid="pone.0135209.ref038" ref-type="bibr">38</xref>]. A deep genetic differentiation exists between Asian (var. <italic>asiatica</italic>), Mediterranean and African populations and closely concurs with the pattern of Quaternary differentiation of its hosts <italic>Olea</italic> spp. [<xref rid="pone.0135209.ref022" ref-type="bibr">22</xref>]. Within the Mediterranean region, microsatellite and mitochondrial markers suggested divergence between populations from the Near East (eastern Turkey, Israel, Cyprus) and populations from Western Europe and Northern Africa (Italy, Spain, Greece, France, Morocco, Algeria) [<xref rid="pone.0135209.ref021" ref-type="bibr">21</xref>, <xref rid="pone.0135209.ref022" ref-type="bibr">22</xref>, <xref rid="pone.0135209.ref038" ref-type="bibr">38</xref>, <xref rid="pone.0135209.ref039" ref-type="bibr">39</xref>]. Recently, <italic>B</italic>. <italic>oleae</italic> was also detected on olive in China, but whether these populations are native or recent invaders is still a matter of debate [<xref rid="pone.0135209.ref040" ref-type="bibr">40</xref>]; for that reason, we did not consider the Chinese populations in our study. Given the phylogeographic pattern of <italic>B</italic>. <italic>oleae</italic>, we considered four native lineages as relevant in our study (Africa, Western Europe, Middle East and Pakistan) and one invasive population (America) (Fig A in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). As the reasons for the existence of shared haplotypes among phylogeographic regions is still debated (e.g. cryptic invasions due to global olive trade or incomplete lineage sorting [<xref rid="pone.0135209.ref021" ref-type="bibr">21</xref>]), we deliberately ignored potential geographic overlapping among conspecific lineages.</p><p>The West Indian fruit fly <italic>Anastrepha obliqua</italic> occurs in the Neotropics from northern Mexico to southern Brazil as well as in the Caribbean islands. This polyphagous pest species mainly feeds on <italic>Mangifera indica</italic> and <italic>Spondias</italic> species (Anacardiaceae) [<xref rid="pone.0135209.ref041" ref-type="bibr">41</xref>]. Three well-supported infraspecific lineages were identified using mitochondrial markers [<xref rid="pone.0135209.ref023" ref-type="bibr">23</xref>]. One lineage encompasses specimens sampled across Central America, Mexico, Caribbean islands and northern Andean highlands and all records from this area were assigned to this lineage (Fig C in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref> and <xref rid="pone.0135209.t001" ref-type="table">Table 1</xref>). A second lineage encompasses specimens sampled mostly across Mexico west to the Sierra Madre Occidental (SMOC). However, one haplotype associated with this lineage was recovered in Southeastern Mexico (locality of Agua Blanca in Tabasco province). The occurrence of this lineage in this region is being debated, so we did not consider this record, which could probably have resulted from a recent invasion process [<xref rid="pone.0135209.ref023" ref-type="bibr">23</xref>]. All occurrences of <italic>A</italic>. <italic>obliqua</italic> in Western Mexico were assigned to this lineage (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). Finally, a third lineage encompasses haplotypes recovered from South America, including the northern Andean highlands and Panama. All documented records from these regions were assigned to this lineage (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and Fig C in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>).</p><p>The Afrotropical <italic>Ceratitis fasciventris</italic> is a polyphagous fruit fly [<xref rid="pone.0135209.ref042" ref-type="bibr">42</xref>, <xref rid="pone.0135209.ref043" ref-type="bibr">43</xref>]. Genetic structure was detected within <italic>C</italic>. <italic>fasciventris</italic> by both microsatellite data and mitochondrial markers [<xref rid="pone.0135209.ref024" ref-type="bibr">24</xref>, <xref rid="pone.0135209.ref044" ref-type="bibr">44</xref>]. Molecular markers suggest that one infraspecific lineage occurs in western Africa and in coastal Tanzania, while a second lineage occurs in the highlands of eastern Africa (Kenya, Uganda, Ethiopia, Zambia) [<xref rid="pone.0135209.ref024" ref-type="bibr">24</xref>, <xref rid="pone.0135209.ref044" ref-type="bibr">44</xref>]. Additionally, two clades composed of specimens from Benin and Mali/Côte d’Ivoire were highlighted by the analysis of mitochondrial sequences of specimens from Western Africa [<xref rid="pone.0135209.ref044" ref-type="bibr">44</xref>]. As the preliminary results showed that climatic envelopes of these two clades were broadly similar, we considered these lineages as a single set. We removed from our dataset the records of <italic>C</italic>. <italic>fasciventris</italic> in Democratic Republic of Congo, Rwanda, Namibia, Angola and Congo, since these geographic zones were not sampled in the phylogeographic studies [<xref rid="pone.0135209.ref024" ref-type="bibr">24</xref>, <xref rid="pone.0135209.ref044" ref-type="bibr">44</xref>]. Finally, we assigned all records from Western Africa and the lowlands of Tanzania to one lineage (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and Fig B in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). The records from the highlands of Eastern Africa were assigned to a second lineage (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and Fig B in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>).</p><p><italic>Anastrepha fraterculus</italic> is considered a species complex (<italic>AF</italic> complex) ranging from South America to the USA [<xref rid="pone.0135209.ref045" ref-type="bibr">45</xref>]. Cryptic species of <italic>A</italic>. <italic>fraterculus</italic> occur in different geographic areas, displaying different host preferences and showing signals of reproductive isolation [<xref rid="pone.0135209.ref030" ref-type="bibr">30</xref>, <xref rid="pone.0135209.ref046" ref-type="bibr">46</xref>]. Although the taxonomic status of several entities within the complex is still debated, some of them are differentiated based on genetic and morphological evidence [<xref rid="pone.0135209.ref031" ref-type="bibr">31</xref>, <xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>, <xref rid="pone.0135209.ref047" ref-type="bibr">47</xref>–<xref rid="pone.0135209.ref049" ref-type="bibr">49</xref>]. In this study, we considered independently a "Mexican' lineage occurring across Central America from Mexico to Panama [<xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>], an 'Andean' lineage [<xref rid="pone.0135209.ref031" ref-type="bibr">31</xref>, <xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>] occurring in the highlands of the Andean region (approximately in regions located at altitudes ranging from 1200 to 2500 meters (see [<xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>]) and a 'Brazilian' lineage occurring in Brazil and Argentina [<xref rid="pone.0135209.ref037" ref-type="bibr">37</xref>, <xref rid="pone.0135209.ref047" ref-type="bibr">47</xref>, <xref rid="pone.0135209.ref049" ref-type="bibr">49</xref>] (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and Fig D in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). We did not consider types occurring in the lowlands of Peru, Venezuela and Colombia because we lacked occurrences in these regions. As occurrences were also lacking to model the distribution of the Andean lineage, we artificially generated 1,000 additional presences at altitudes ranging between 1,200 and 2,500m to perform our SDMs. We estimated this amount of occurrences to be large enough to accurately model the distribution of species [<xref rid="pone.0135209.ref050" ref-type="bibr">50</xref>, <xref rid="pone.0135209.ref051" ref-type="bibr">51</xref>].</p><p>The apple maggot fly, <italic>R</italic>. <italic>pomonella</italic>, is widely distributed in the USA, Canada and mountainous ranges of Mexico. This species has deep economic implications since <italic>R</italic>. <italic>pomonella</italic> recently shifted to the introduced cultivated apple (<italic>Malus pumila</italic> Mill.). This species encompasses at least four genetically distinct populations [<xref rid="pone.0135209.ref032" ref-type="bibr">32</xref>, <xref rid="pone.0135209.ref033" ref-type="bibr">33</xref>, <xref rid="pone.0135209.ref035" ref-type="bibr">35</xref>] displaying mating incompatibilities [<xref rid="pone.0135209.ref052" ref-type="bibr">52</xref>]. These populations occur respectively in the USA, the Eje Volcanico Trans Mexicano (EVTM), the Sierra Madre Oriental (SMO) and the elevated regions of Chiapas in Mexico. Because we lacked occurrences of <italic>R</italic>. <italic>pomonella</italic> in Mexico, and the preliminary results showed high climatic envelope similarity among Mexican populations, we considered all Mexican records as belonging to a 'Mexican' lineage (<xref rid="pone.0135209.t001" ref-type="table">Table 1</xref> and Fig E in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>).</p><p>When assessing the invasion risk of <italic>B</italic>. <italic>cucurbitae</italic>, we only considered the species level (Fig F in <xref rid="pone.0135209.s002" ref-type="supplementary-material">S2 File</xref>). Although genetic structure exists in <italic>B</italic>. <italic>cucurbitae</italic> [<xref rid="pone.0135209.ref034" ref-type="bibr">34</xref>], differentiation among geographically distant populations appears mostly as a result of human-mediated range expansions rather than ancient geographic and climatic events [<xref rid="pone.0135209.ref053" ref-type="bibr">53</xref>].</p></sec><sec id="sec004"><title>Species distribution modeling</title><p>We used climatic data available from the WorldClim database version 1.4 [<xref rid="pone.0135209.ref054" ref-type="bibr">54</xref>] to investigate niche divergence between lineages and to model their distributions (<xref rid="pone.0135209.t002" ref-type="table">Table 2</xref>). These climatic data are derived from temperature and rainfall annual trends between 1950 and 2000. The choice of environmental data set is crucial when modeling potential distribution in a new area [<xref rid="pone.0135209.ref055" ref-type="bibr">55</xref>, <xref rid="pone.0135209.ref056" ref-type="bibr">56</xref>]. We consequently decided to model the distribution of fruit flies species with a strongly restricted climatic data set of moderately correlated (Pearson correlation index r < 0.7) and biologically meaningful variables that were susceptible to inducing physiological stress for these organisms. We did not include climatic descriptors depicting the annual range of temperatures and the seasonal structure of the climate because these variables probably influence the phenology of the insects rather than inducing physiological stress. We thus selected the following climatic variables: the mean temperature of the warmest month (bio5), the mean temperature of the coldest month (bio6), the precipitation of the wettest quarter (bio16) and the precipitation of the driest quarter (bio17).</p><table-wrap id="pone.0135209.t002" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.t002</object-id><label>Table 2</label><caption><title>Bioclimatic variables used to investigate the climatic niche of tephritid fruit flies species and lineages.</title></caption><alternatives><graphic id="pone.0135209.t002g" xlink:href="pone.0135209.t002"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col></colgroup><thead><tr><th align="center" rowspan="1" colspan="1">Bioclimatic variables</th><th align="center" rowspan="1" colspan="1">symbol</th></tr></thead><tbody><tr><td align="center" rowspan="1" colspan="1">Annual Mean Temperature</td><td align="center" rowspan="1" colspan="1">BIO1</td></tr><tr><td align="center" rowspan="1" colspan="1">Mean Diurnal Range</td><td align="center" rowspan="1" colspan="1">BIO2</td></tr><tr><td align="center" rowspan="1" colspan="1">Isothermality</td><td align="center" rowspan="1" colspan="1">BIO3</td></tr><tr><td align="center" rowspan="1" colspan="1">Temperature Seasonality</td><td align="center" rowspan="1" colspan="1">BIO4</td></tr><tr><td align="center" rowspan="1" colspan="1">Maximum Temperature of Warmest Month</td><td align="center" rowspan="1" colspan="1">BIO5</td></tr><tr><td align="center" rowspan="1" colspan="1">Minimum Temperature of Coldest Month</td><td align="center" rowspan="1" colspan="1">BIO6</td></tr><tr><td align="center" rowspan="1" colspan="1">Temperature Annual Range</td><td align="center" rowspan="1" colspan="1">BIO7</td></tr><tr><td align="center" rowspan="1" colspan="1">Mean Temperature of Wettest Quarter</td><td align="center" rowspan="1" colspan="1">BIO8</td></tr><tr><td align="center" rowspan="1" colspan="1">Mean Temperature of Driest Quarter</td><td align="center" rowspan="1" colspan="1">BIO9</td></tr><tr><td align="center" rowspan="1" colspan="1">Mean Temperature of Warmest Quarter</td><td align="center" rowspan="1" colspan="1">BIO10</td></tr><tr><td align="center" rowspan="1" colspan="1">Mean Temperature of Coldest Quarter</td><td align="center" rowspan="1" colspan="1">BIO11</td></tr><tr><td align="center" rowspan="1" colspan="1">Annual Precipitation</td><td align="center" rowspan="1" colspan="1">BIO12</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Wettest Month</td><td align="center" rowspan="1" colspan="1">BIO13</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Driest Month</td><td align="center" rowspan="1" colspan="1">BIO14</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation Seasonality</td><td align="center" rowspan="1" colspan="1">BIO15</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Wettest Quarter</td><td align="center" rowspan="1" colspan="1">BIO16</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Driest Quarter</td><td align="center" rowspan="1" colspan="1">BIO17</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Warmest Quarter</td><td align="center" rowspan="1" colspan="1">BIO18</td></tr><tr><td align="center" rowspan="1" colspan="1">Precipitation of Coldest Quarter</td><td align="center" rowspan="1" colspan="1">BIO19</td></tr></tbody></table></alternatives></table-wrap><p>We modeled the distribution of species and intraspecific entities using MaxEnt version 3.3.3 with default settings [<xref rid="pone.0135209.ref057" ref-type="bibr">57</xref>] available in the 'dismo' R package [<xref rid="pone.0135209.ref058" ref-type="bibr">58</xref>] and the boosted regression trees (BRT) [<xref rid="pone.0135209.ref059" ref-type="bibr">59</xref>, <xref rid="pone.0135209.ref060" ref-type="bibr">60</xref>] available in the ‘gbm’ R package [<xref rid="pone.0135209.ref061" ref-type="bibr">61</xref>]. These two techniques were selected because they are among the best-performing approaches to model species distributions [<xref rid="pone.0135209.ref050" ref-type="bibr">50</xref>, <xref rid="pone.0135209.ref062" ref-type="bibr">62</xref>]. The MaxEnt approach is a ‘presence-only’ method that performs well, even with a small occurrence dataset [<xref rid="pone.0135209.ref050" ref-type="bibr">50</xref>, <xref rid="pone.0135209.ref051" ref-type="bibr">51</xref>]. This algorithm contrasts the climatic envelope experienced by the species with a set of localities randomly chosen within the background environment, where the presence of the species is unknown. The choice of background area is crucial when modeling distribution with MaxEnt and depends on the purpose of the study [<xref rid="pone.0135209.ref063" ref-type="bibr">63</xref>–<xref rid="pone.0135209.ref065" ref-type="bibr">65</xref>]. The BRT modeling technique is a machine-learning approach relying on presences and absences that can fit complex non-linear relationships and handle correlation effects between predictors [<xref rid="pone.0135209.ref060" ref-type="bibr">60</xref>]. All information regarding the positioning of absences, pseudo-absences and background data used to model the distribution of each species and lineage are available in Supplementary information (<xref rid="pone.0135209.s008" ref-type="supplementary-material">S1 Text</xref>).</p><p>We modeled the distribution of all Species (‘S-models’) and IntraSpecific lineages (‘IS-models’). Each model was fitted using a random subset of 80% of the relevant occurrences (training dataset) and tested with the remaining records. The predictive power of each model was assessed by measuring the area under the receiver operating characteristics (ROC) curve (AUC) [<xref rid="pone.0135209.ref066" ref-type="bibr">66</xref>]. The AUC is widely used in species distribution modeling because it is a threshold-independent evaluation approach that gets around the problem of subjectivity of threshold selection when assessing the accuracy of bioclimatic models. Values of AUC ranging from 0.5 (random prediction) to 1 (perfect prediction) indicate model performance better than random. The AUC of each model for <italic>B</italic>. <italic>oleae</italic> in the invaded range was also assessed with observed presences in Western America and 10000 pseudo-absences generated in Western America (i.e. in a square with longitude ranging from 122.3 to -115.4 and latitude ranging from 31.86 to 39.51).</p><p>All models were projected onto both world and Europe maps to assess the risk of biological invasion at worldwide and European scales. Since projections onto new areas may be particularly uncertain, we computed multivariate environmental similarity surface maps (MESS) [<xref rid="pone.0135209.ref065" ref-type="bibr">65</xref>] that describe the similarity between the climatic spaces of the training and the projected areas. Negative values of MESS indicate that the model is projected onto the area outside the climatic space where the model was fitted (model extrapolation), whilst positive values of MESS indicate model interpolation. We constructed MESS maps using occurrence and background data.</p></sec><sec id="sec005"><title>Niche comparison</title><p>We carried out direct climate comparisons by means of Principal Component Analysis (PCA). This multivariate technique allows climate typologies to be drawn and groups of occurrences associated to similar climate descriptors to be identified [<xref rid="pone.0135209.ref067" ref-type="bibr">67</xref>, <xref rid="pone.0135209.ref068" ref-type="bibr">68</xref>]. The differences between lineages of a species were assessed using a between-class inertia test [<xref rid="pone.0135209.ref069" ref-type="bibr">69</xref>] based on 999 permutations [<xref rid="pone.0135209.ref070" ref-type="bibr">70</xref>]. The higher the between-class inertia, the greater the difference between classes in the multivariate-space under study. The analyses were performed using the 19 bioclimatic variables from the WorldClim database (<xref rid="pone.0135209.t002" ref-type="table">Table 2</xref>).</p><p>We investigated the degree of niche similarity between conspecific lineages by calculating Schoener’ D index [<xref rid="pone.0135209.ref071" ref-type="bibr">71</xref>]. This metric measures the overlap of the SDM projections, which varies from 0 for non-overlapping model predictions to 1 for complete overlap. We deliberately did not address intraspecific niche divergence using recently developed statistical approaches [<xref rid="pone.0135209.ref072" ref-type="bibr">72</xref>–<xref rid="pone.0135209.ref076" ref-type="bibr">76</xref>] because of the difficulty in delimiting different and non-overlapping background regions (i.e. pseudoabsences) in SDMs among conspecific lineages. The uncertainty related to the positioning of pseudoabsences may, indeed, have large effects on SDM predictions and consequently alter the reliability of tests of climatic niche divergence [<xref rid="pone.0135209.ref072" ref-type="bibr">72</xref>, <xref rid="pone.0135209.ref077" ref-type="bibr">77</xref>].</p></sec></sec><sec sec-type="results" id="sec006"><title>Results</title><sec id="sec007"><title>Realized climatic niche similarity between conspecific lineages</title><p>Between-class PCA analyses showed significant climatic envelope divergence for all of the species considered (values of inertia are available in <xref rid="pone.0135209.s006" ref-type="supplementary-material">S1 Table</xref>). High divergence was observed among conspecific lineages of <italic>C</italic>. <italic>fasciventris</italic> (p<0.001) and <italic>A</italic>. <italic>fraterculus</italic> (p<0.001) according to temperature variables such as the mean annual temperature (bio1), the mean temperature of the warmest months (bio5, bio10), and the mean temperature of the coldest months (bio6, bio11) (<xref rid="pone.0135209.g001" ref-type="fig">Fig 1</xref> and Fig B in <xref rid="pone.0135209.s003" ref-type="supplementary-material">S3 File</xref>). The 'Andean' lineage of <italic>A</italic>. <italic>fraterculus</italic> occurred in regions characterized by high precipitations (bio14, bio17) and fresh maximal temperatures (bio5, bio10).</p><fig id="pone.0135209.g001" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.g001</object-id><label>Fig 1</label><caption><title>Principal component analysis (PCA) performed on 19 bioclimatic variables extracted from the Worldclim database [<xref rid="pone.0135209.ref054" ref-type="bibr">54</xref>] for <italic>Ceratitis fasciventris</italic> occurrences (see <xref rid="pone.0135209.t002" ref-type="table">Table 2</xref> for symbol meanings).</title><p>These multivariate analyses draw the bioclimatic envelopes of the different phylogeographic lineages belonging to <italic>C</italic>. <italic>fasciventris</italic>. Circles of correlation (a) and factorial scores of records (b) are shown. Percentages of variance explained by each PCA axis are indicated in correlation circle.</p></caption><graphic xlink:href="pone.0135209.g001"></graphic></fig><p>Conspecific lineages of <italic>R</italic>. <italic>pomonella</italic> mainly differed (p<0.001) according to minimal temperatures (bio6, bio11), temperature annual range (bio2, bio3, bio4, bio7) and maximal precipitations (bio13, bio16) (Fig C in <xref rid="pone.0135209.s003" ref-type="supplementary-material">S3 File</xref>). However, conspecific lineages of <italic>R</italic>. <italic>pomonella</italic> occur in areas with similar maximal temperatures (bio5, bio10).</p><p>No strong differences were observed between the Ao_01 and Ao_03 lineages of <italic>A</italic>. <italic>obliqua</italic> (p<0.001) along the two first PCA axes performed on the climatic data corresponding to occurrence points (Fig A in <xref rid="pone.0135209.s003" ref-type="supplementary-material">S3 File</xref>). However, the climatic envelope of population Ao_02 was different from that of populations Ao_01 and Ao_03 according to precipitations (bio12, bio13, bio15) and maximal temperatures (bio5, bio10).</p><p>Lineages of <italic>B</italic>. <italic>oleae</italic> (p<0.001) were opposed on the first PCA axis by warmest temperatures (bio5, bio10; <xref rid="pone.0135209.g002" ref-type="fig">Fig 2</xref>) and by precipitation descriptors such as the mean annual temperature (bio12), the precipitation of the driest month (bio14) and the precipitations of the warmest quarter (bio 18). On the second PCA axis, populations were opposed by the temperature annual range (bio7, bio4) and by the minimum temperature (bio6, bio11; <xref rid="pone.0135209.g002" ref-type="fig">Fig 2</xref>). The bioclimatic envelope of <italic>B</italic>. <italic>oleae</italic> in the invaded range was different from the other lineages with respect to maximal temperatures (bio5, bio10) and minimal temperatures (bio6, bio11).</p><fig id="pone.0135209.g002" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.g002</object-id><label>Fig 2</label><caption><title>Principal component analysis (PCA) performed on 19 bioclimatic variables extracted from the Worldclim database [<xref rid="pone.0135209.ref054" ref-type="bibr">54</xref>] for <italic>Bactrocera oleae</italic> occurrences (see <xref rid="pone.0135209.t002" ref-type="table">Table 2</xref> for symbol meanings).</title><p>These multivariate analyses draw the bioclimatic envelopes of the different phylogeographic lineages belonging to <italic>B</italic>. <italic>oleae</italic> as well as the invading populations in Americas. Circles of correlation (a) and factorial scores of records (b) are shown. Percentages of variance explained by each PCA axis are indicated in correlation circle.</p></caption><graphic xlink:href="pone.0135209.g002"></graphic></fig></sec><sec id="sec008"><title>Potential distributions of tephritid species</title><p>Our SDM models displayed relatively high values of cross-validated AUC ranging from 0.76 to 0.98 (<xref rid="pone.0135209.s007" ref-type="supplementary-material">S2 Table</xref>). The BRT models globally displayed higher values of cross-validated AUC than MaxEnt models. The distribution of suitable climatic conditions for <italic>C</italic>. <italic>fasciventris</italic>, <italic>A</italic>. <italic>fraterculus</italic>, <italic>B</italic>. <italic>cucurbitae</italic> and <italic>A</italic>. <italic>obliqua</italic> predicted by SDM models encompassed most of the tropical and subtropical areas of the world and a few regions with Mediterranean or temperate climate (<xref rid="pone.0135209.s004" ref-type="supplementary-material">S4</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5</xref> Files). However, the distribution of suitable conditions for <italic>B</italic>. <italic>oleae</italic> and <italic>R</italic>. <italic>pomonella</italic> mainly encompassed temperate regions of the world (Figs A and E in <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5</xref> Files). The potential distributions of all tephritid species encompassed areas in Europe (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Both <italic>R</italic>. <italic>pomonella</italic> and <italic>B</italic>. <italic>oleae</italic> displayed the larger extents of suitable climatic conditions in Europe (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs A and E in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Both <italic>C</italic>. <italic>fasciventris</italic> and <italic>A</italic>. <italic>obliqua</italic> showed a small potential distribution range in Europe (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs B and C in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Maxent maps of <italic>C</italic>. <italic>fasciventris</italic> predicted large potential areas in Europe (e.g. in the United Kingdom and Scandinavia; Fig B in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). However, MESS maps indicated that large model extrapolation occurs when predicting the potential distribution of <italic>C</italic>. <italic>fasciventris</italic> in Northern Europe. We argue that the BRT prediction is the most reliable for <italic>C</italic>. <italic>fasciventris</italic> since the MaxEnt model predicts climatic suitability in implausible areas in northern Europe (such inconsistency appears common with the MaxEnt algorithm, see [<xref rid="pone.0135209.ref078" ref-type="bibr">78</xref>]). Both <italic>A</italic>. <italic>fraterculus</italic> and <italic>B</italic>. <italic>cucurbitae</italic> displayed a larger extent of suitable climatic conditions with respect to <italic>A</italic>. <italic>obliqua</italic> and <italic>C</italic>. <italic>fasciventris</italic> (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs D and F in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>).</p><fig id="pone.0135209.g003" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.g003</object-id><label>Fig 3</label><caption><title>Projections of climatic suitability for six tephritid species and intraspecific lineages in Europe ((a-d) <italic>Bactrocera oleae</italic>, (e-h) <italic>Anastrepha obliqua</italic>, (i-l) <italic>Anastrepha fraterculus</italic>, (m-o) <italic>Ceratitis fasciventris</italic>, (p-r) <italic>Rhagoletis pomonella</italic> and (s) <italic>Bactrocera cucurbitae</italic>) as predicted by species distribution models (SDMs).</title><p>Species- and lineage-based SDMs were performed using the Boosted Regression Trees (BRT) [<xref rid="pone.0135209.ref060" ref-type="bibr">60</xref>]. Climatic suitability is shown by a color gradient, which goes from green (high probability) to light orange (low probability).</p></caption><graphic xlink:href="pone.0135209.g003"></graphic></fig><p>We observed strong differences in the modeled distributions among conspecific lineages of <italic>C</italic>. <italic>fasciventris</italic>, <italic>A</italic>. <italic>fraterculus</italic> and <italic>R</italic>. <italic>pomonella</italic> (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref>, <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5</xref> Files; <xref rid="pone.0135209.t003" ref-type="table">Table 3</xref>). However, conspecific lineages of <italic>B</italic>. <italic>oleae</italic> and <italic>A</italic>. <italic>obliqua</italic> showed similar but not identical modeled distributions (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs A and C in <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5</xref> Files; <xref rid="pone.0135209.t003" ref-type="table">Table 3</xref>).</p><table-wrap id="pone.0135209.t003" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.t003</object-id><label>Table 3</label><caption><title>Measures of species distribution models overlapping among phylogeographic lineages of tephritid fruit flies.</title><p>We calculated projection overlapping by calculating the Schoener' D index, which range from 0 (no overlapping) to 1 (perfect overlapping).</p></caption><alternatives><graphic id="pone.0135209.t003g" xlink:href="pone.0135209.t003"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col><col align="left" valign="middle" span="1"></col></colgroup><thead><tr><th align="left" rowspan="1" colspan="1">Species</th><th align="center" rowspan="1" colspan="1">Lineage 1</th><th align="center" rowspan="1" colspan="1">Lineage 2</th><th colspan="2" align="center" rowspan="1">Schoener' Index</th></tr><tr><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="center" rowspan="1" colspan="1">BRT</th><th align="center" rowspan="1" colspan="1">MaxEnt</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>Ceratitis fasciventris</italic></td><td align="center" rowspan="1" colspan="1">lineage Eastern Africa</td><td align="center" rowspan="1" colspan="1">lineage Western Africa</td><td align="char" char="." rowspan="1" colspan="1">0.46</td><td align="char" char="." rowspan="1" colspan="1">0.23</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Rhagoletis pomonella</italic></td><td align="center" rowspan="1" colspan="1">lineage USA</td><td align="center" rowspan="1" colspan="1">lineage Mexico</td><td align="char" char="." rowspan="1" colspan="1">0.1</td><td align="char" char="." rowspan="1" colspan="1">0.13</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Anastrepha fraterculus</italic></td><td align="center" rowspan="1" colspan="1">lineage Mexico</td><td align="center" rowspan="1" colspan="1">lineage Brazil</td><td align="char" char="." rowspan="1" colspan="1">0.8</td><td align="char" char="." rowspan="1" colspan="1">0.64</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage Mexico</td><td align="center" rowspan="1" colspan="1">lineage Andean</td><td align="char" char="." rowspan="1" colspan="1">0.13</td><td align="char" char="." rowspan="1" colspan="1">0.12</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage Brazil</td><td align="center" rowspan="1" colspan="1">lineage Andean</td><td align="char" char="." rowspan="1" colspan="1">0.09</td><td align="char" char="." rowspan="1" colspan="1">0.11</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Anastrepha obliqua</italic></td><td align="center" rowspan="1" colspan="1">lineage Central America</td><td align="center" rowspan="1" colspan="1">lineage Western Mexico</td><td align="char" char="." rowspan="1" colspan="1">0.91</td><td align="char" char="." rowspan="1" colspan="1">0.55</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage Central America</td><td align="center" rowspan="1" colspan="1">lineage South America</td><td align="char" char="." rowspan="1" colspan="1">0.86</td><td align="char" char="." rowspan="1" colspan="1">0.59</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage South America</td><td align="center" rowspan="1" colspan="1">lineage Western Mexico</td><td align="char" char="." rowspan="1" colspan="1">0.87</td><td align="char" char="." rowspan="1" colspan="1">0.29</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Bactrocera oleae</italic></td><td align="center" rowspan="1" colspan="1">lineage Africa</td><td align="center" rowspan="1" colspan="1">lineage Western Europe</td><td align="char" char="." rowspan="1" colspan="1">0.32</td><td align="char" char="." rowspan="1" colspan="1">0.55</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage Africa</td><td align="center" rowspan="1" colspan="1">lineage Middle East</td><td align="char" char="." rowspan="1" colspan="1">0.25</td><td align="char" char="." rowspan="1" colspan="1">0.42</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1">lineage Western Europe</td><td align="center" rowspan="1" colspan="1">lineage Middle East</td><td align="char" char="." rowspan="1" colspan="1">0.29</td><td align="char" char="." rowspan="1" colspan="1">0.48</td></tr></tbody></table></alternatives></table-wrap><p>We suggest that the MaxEnt algorithm provides the most reliable potential distribution of the Ao_03 lineage since the BRT model predicts climatic suitability in implausible areas (cold mountainous regions of Europe and Greenland). The distributions of suitable climatic conditions predicted by the BRT and MaxEnt S-models for <italic>B</italic>. <italic>oleae</italic> were broadly congruent. They matched the geographic distribution of mild temperate climates and Mediterranean-climate regions where native and cultivated olives occur (e.g. California, Australia, Southeast China, Argentina, Chile, Mexico, Mediterranean area), while also encompassing colder temperate regions. Our SDMs also predicted climatic suitability for <italic>B</italic>. <italic>oleae</italic> in temperate regions (the United Kingdom, Northern France, Belgium, south-eastern United states), where olive trees are not currently cultivated. The records of the genetically distinct lineage occurring in Pakistan were well predicted by S-models. Conspecific lineages of <italic>B</italic>. <italic>oleae</italic> showed slightly different modeled distributions in the invaded range. The models predicted higher climatic suitability in coastal region of California for the African lineage while inland regions were predicted as more suitable for Western European and Middle Eastern lineages (<xref rid="pone.0135209.g004" ref-type="fig">Fig 4</xref>). The AUC values displayed by MaxEnt and BRT S-models for <italic>B</italic>. <italic>oleae</italic> in the invaded range were 0.65 and 0.68 respectively indicating model performance better than random. The AUC values in the invaded range displayed by the species-based BRT model was higher than those displayed by lineages-based BRT models (0.55, 0.46 and 0.57 for African, Western European and Middle Eastern lineages respectively). The AUC value in the invaded range displayed by species-based MaxEnt model was higher than those displayed by western European and Middle Eastern lineages-based BRT models (0.51 and 0.60 respectively) but slightly inferior to AUC displayed by African lineage-based MaxEnt model (0.73).</p><fig id="pone.0135209.g004" orientation="portrait" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0135209.g004</object-id><label>Fig 4</label><caption><title>Projections of species- and lineage-based models (MaxEnt and Boosted Regression Trees (BRT) for <italic>Bactrocera oleae</italic> in the invaded range in Americas.</title><p>Black crosses represent the occurrence of the olive fruit fly. Models were calibrated independently for the species and the different conspecific lineages. Climatic suitability is shown by a color gradient, which goes from green (high probability) to light orange (low probability).</p></caption><graphic xlink:href="pone.0135209.g004"></graphic></fig></sec></sec><sec sec-type="conclusions" id="sec009"><title>Discussion</title><sec id="sec010"><title>Tephritid fruit flies: a current major threat with potential for future expansion</title><p>Tephritid fruit flies are among the most destructive fruit pests and constitute a major threat to worldwide agriculture. Because many species have invaded new regions where they cause severe damage to fruit and other crops [<xref rid="pone.0135209.ref079" ref-type="bibr">79</xref>], the group is under surveillance by biosecurity agencies to design cost-effective management strategies. In addition, investigating the climatic suitability of Europe for mainly tropical tephritid should help to uncover why paradoxically few species have invaded areas in Europe and in Northern Africa compared to other geographic regions such as California [<xref rid="pone.0135209.ref019" ref-type="bibr">19</xref>]. Our results show that all target tephritid species under investigation could potentially find suitable climate conditions in Europe, albeit to different degrees (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Three groups of species may be distinguished according to their potential distribution range in Europe:
<list list-type="order"><list-item><p><italic>C</italic>. <italic>fasciventris</italic> and <italic>A</italic>. <italic>obliqua</italic>, which are restricted to low and mid-elevated areas of tropical regions and consequently display a low probability of being established in European countries (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs B and C in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Only areas with mild winters (e.g. the Atlantic coasts of Portugal and Spain, southern Greece, southern Turkey and Near East region) appeared to be suitable for these species (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref>). Cold stress would likely be the most limiting climatic variable for the long-term establishment of these flies in Europe. Indeed, <italic>A</italic>. <italic>obliqua</italic> has been reported to be less abundant at high altitudes compared to its close relatives (e.g. <italic>A</italic>. <italic>fraterculus</italic>, <italic>A</italic>. <italic>ludens</italic>) and <italic>Ceratitis</italic> species (e.g. <italic>C</italic>. <italic>capitata</italic>) [<xref rid="pone.0135209.ref080" ref-type="bibr">80</xref>, <xref rid="pone.0135209.ref081" ref-type="bibr">81</xref>], while <italic>C</italic>. <italic>fasciventris</italic> is absent from colder regions in South Africa where a congeneric species <italic>C</italic>. <italic>rosa</italic> occurs [<xref rid="pone.0135209.ref082" ref-type="bibr">82</xref>]. However, we cannot exclude that invasive fruit flies might overwinter as larvae within infested host fruit as reported for <italic>C</italic>. <italic>capitata</italic> [<xref rid="pone.0135209.ref083" ref-type="bibr">83</xref>], and consequently could become established in colder areas.</p></list-item><list-item><p><italic>Anastrepha fraterculus</italic> and <italic>B</italic>. <italic>cucurbitae</italic> displayed a larger potential distribution in Europe (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs D and F in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). In their native range, these species occur in tropical lowlands and in colder elevated regions, suggesting that low winter temperatures may be less constraining than for <italic>C</italic>. <italic>fasciventris</italic> and <italic>A</italic>. <italic>obliqua</italic>. For example, <italic>A</italic>. <italic>fraterculus</italic> is sympatric with <italic>C</italic>. <italic>capitata</italic> in South America and is thus expected to display climatic tolerances allowing it to become established in parts of temperate Europe, which is also the case for <italic>C</italic>. <italic>capitata</italic> [<xref rid="pone.0135209.ref084" ref-type="bibr">84</xref>, <xref rid="pone.0135209.ref085" ref-type="bibr">85</xref>]. In the case of <italic>A</italic>. <italic>fraterculus</italic>, we deliberately calibrated SDMs with occurrences located in dry areas where the species is associated with human settlements (e.g. irrigated areas in the 'Chaco' region and in dry highlands of the Argentinean province of Jujuy). We acknowledge that these records might lead to an overestimation of the potential distribution of <italic>A</italic>. <italic>fraterculus</italic> into dry regions, since climatic layers do not reflect microclimates resulting from human activities. We nevertheless added these records to our SDMs to avoid omission error [<xref rid="pone.0135209.ref084" ref-type="bibr">84</xref>]. Indeed, irrigated agriculture is performed in many dry regions of the world and might allow tephritid colonization despite apparently unfavorable macroclimatic conditions. These results also highlighted that <italic>B</italic>. <italic>cucurbitae</italic> might expand its invaded range towards southern parts of Africa where it has not yet been detected (Fig F in <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4</xref> and <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5</xref> Files).</p></list-item><list-item><p><italic>Rhagoletis pomonella</italic> and <italic>B</italic>. <italic>oleae</italic> occur only in temperate regions and in some tropical highlands. Apart from very cold regions, the potential distribution of <italic>R</italic>. <italic>pomonella</italic> covers all of Europe (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Figs A and E in <xref rid="pone.0135209.s005" ref-type="supplementary-material">S5 File</xref>). Since it naturally occurs in the Mediterranean basin, <italic>B</italic>. <italic>oleae</italic> does not constitute a new threat for Europe and we will not discuss its potential distribution in detail.</p></list-item></list></p></sec><sec id="sec011"><title>Niche variation among conspecific lineages</title><p>Although there is increasing evidence that climatic niche variation among conspecific lineages may have strong implications in invasion risk management [<xref rid="pone.0135209.ref007" ref-type="bibr">7</xref>, <xref rid="pone.0135209.ref009" ref-type="bibr">9</xref>, <xref rid="pone.0135209.ref010" ref-type="bibr">10</xref>], such intraspecific variation in tephritid fruit flies has been poorly investigated so far (but see [<xref rid="pone.0135209.ref025" ref-type="bibr">25</xref>]). We illustrate here how intraspecific structure of widely distributed pests is, in some cases, associated with strong climate envelope divergence. Three fruit flies (<italic>C</italic>. <italic>fasciventris</italic>, <italic>A</italic>. <italic>fraterculus</italic>, <italic>R</italic>. <italic>pomonella</italic>) highly differ in realized climatic niche and consequently display different potential distributions (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref>). Interestingly, models predict one lineage of <italic>C</italic>. <italic>fasciventris</italic> to be more susceptible to expanding its geographic range into Europe. Similarly, parts of the Mediterranean Basin are predicted to be suitable for the Mexican and Brazilian lineages of <italic>A</italic>. <italic>fraterculus</italic> but not for the Andean lineage (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Fig D in <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4 File</xref>). However, we detected no marked climate divergence among lineages of <italic>A</italic>. <italic>obliqua</italic> (Fig A in <xref rid="pone.0135209.s003" ref-type="supplementary-material">S3 File</xref>). The distributions of conspecific lineages of <italic>A</italic>. <italic>obliqua</italic> appear to be constrained by cold temperatures in Mexico as well as in South America, meaning that conspecific lineages display similar potential distributions (<xref rid="pone.0135209.g003" ref-type="fig">Fig 3</xref> and Fig C in <xref rid="pone.0135209.s004" ref-type="supplementary-material">S4 File</xref>). Similarly, relatively high overlapping of SDM projections was observed among lineages of <italic>B</italic>. <italic>oleae</italic>.</p><p>Among these species, we only addressed the invasion of <italic>B</italic>. <italic>oleae</italic> since the origin of invading populations of the other target fruit flies under study is still unknown. Our species-based climatic suitability maps indicate low climatic suitability in mountainous ranges and in southern and central California, as previously reported [<xref rid="pone.0135209.ref086" ref-type="bibr">86</xref>]. Species-based and African lineage-based models best explain the invaded range of <italic>B</italic>. <italic>oleae</italic>, suggesting that no marked ecological divergence is associated with intraspecific genetic structure of this pest and invasive populations might have potentially originated from every location in its entire native range. In addition, no SDMs (species- and lineage-based approaches) predict the entire invasive range as climatically suitable for <italic>B</italic>. <italic>oleae</italic>, suggesting that, given the monophagy in this fly, host distribution mainly governs its distribution, as reported for other invasive specialist phytophagous insects [<xref rid="pone.0135209.ref087" ref-type="bibr">87</xref>]. Interestingly, our species-based models predict high climatic suitability in coastal region of California and in Sacramento valley where high <italic>B</italic>. <italic>oleae</italic> population densities are reported [<xref rid="pone.0135209.ref015" ref-type="bibr">15</xref>, <xref rid="pone.0135209.ref088" ref-type="bibr">88</xref>]. However, SDMs predict low climatic suitability in San Joaquin Valley and in southeastern counties where lower population densities of <italic>B</italic>. <italic>oleae</italic> are reported probably because of the extremely hot summer temperatures that occur in these regions [<xref rid="pone.0135209.ref089" ref-type="bibr">89</xref>]. This case highlights how lowering the taxonomic resolution may lead, in some cases, to an increase of omission error in PRA and underlines why such approaches still need to be interpreted with caution by risk analysts.</p><sec id="sec012"><title>Perspectives and pitfalls for plant biosecurity purposes</title><p>Niche divergence exists among conspecific lineages in several of these pests (e.g. host preferences in <italic>AF</italic> complex, see [<xref rid="pone.0135209.ref030" ref-type="bibr">30</xref>]). However, addressing climatic niche divergence among organisms is complex [<xref rid="pone.0135209.ref090" ref-type="bibr">90</xref>] and we acknowledge that additional data are required to claim differences in invasion risk among conspecific lineages with respect to climate. Other studies showed the poor performance of lineage-based models to predict the infraspecific identity of invasive populations [<xref rid="pone.0135209.ref091" ref-type="bibr">91</xref>–<xref rid="pone.0135209.ref093" ref-type="bibr">93</xref>] and suggested the existence of a cryptic niche conservatism that may explain, in some cases, the poor predictive power of lineage-based models. In other words, these studies highlighted that differences in the <italic>realized</italic> climatic niche [<xref rid="pone.0135209.ref094" ref-type="bibr">94</xref>] do not always reflect differences in physiological tolerances (the <italic>fundamental</italic> climatic niche [<xref rid="pone.0135209.ref094" ref-type="bibr">94</xref>]). Realized climatic niche divergence may reflect non-evolutionary mechanisms, since the geographic range of populations/lineages can also be shaped by various biotic and historical factors in addition to their unique physiological tolerances [<xref rid="pone.0135209.ref094" ref-type="bibr">94</xref>]. As most species, to our knowledge, do not show intraspecific divergence in their diet (with the exception of the lineages of <italic>A</italic>. <italic>fraterculus</italic> occurring in Mesoamerica and in Brazil and Argentina), we suggest that hosts do not artificially inflate climatic niche divergence signals among the lineages under study. In this context, several recent studies underlined the need to increase the taxonomic resolution in PRA to avoid underestimation of invasion risk and capture the risks of niche shifts during the invasion process [<xref rid="pone.0135209.ref095" ref-type="bibr">95</xref>, <xref rid="pone.0135209.ref096" ref-type="bibr">96</xref>]. Moreover, as phylogeographic and taxonomic studies are usually based on neutral genetic markers, we lack evidence for genetic adaptations of lineages to particular climatic conditions and the role of phenotypic plasticity in shaping geographic distributions still needs to be addressed. Testing the predictive power of such SDMs usually involves <italic>a posteriori</italic> analyses of successful invasions [<xref rid="pone.0135209.ref004" ref-type="bibr">4</xref>]; however, data were unfortunately lacking to test the validity of our approaches with most of the species under scrutiny.</p><p>Such approaches open up new perspectives for the control of exotic species. There is now clear evidence that a climatic niche evolves over moderate evolutionary times, as for those involved in phylogeographic divergence and speciation [<xref rid="pone.0135209.ref097" ref-type="bibr">97</xref>, <xref rid="pone.0135209.ref098" ref-type="bibr">98</xref>] with crucial consequences in invasion context [<xref rid="pone.0135209.ref007" ref-type="bibr">7</xref>, <xref rid="pone.0135209.ref009" ref-type="bibr">9</xref>]. Our study illustrates that the intraspecific structure of widely distributed pests played an important role in pest risk analysis and should be considered when assessing quarantine status. Nowadays, the European list of quarantined organisms encompasses all non-European tephritids and does not consider taxonomic entities below the species level. Given the costs of invasive species management and quarantine measures, predicting with more accuracy the potential distribution of intercepted propagules might generate substantial economic benefits [<xref rid="pone.0135209.ref099" ref-type="bibr">99</xref>]. In addition, rapid and accurate molecular tools (e.g. barcoding) are now available for biosecurity agencies to identify intercepted propagules to species and even lineage levels [<xref rid="pone.0135209.ref100" ref-type="bibr">100</xref>–<xref rid="pone.0135209.ref103" ref-type="bibr">103</xref>]. The availability of such tools consequently holds great promise for the design of cost-effective control strategies against exotic species.</p></sec></sec></sec><sec sec-type="supplementary-material" id="sec013"><title>Supporting Information</title><supplementary-material content-type="local-data" id="pone.0135209.s001"><label>S1 File</label><caption><title>Occurrence data used in this study and list of scientific publications where fruit flies occurrences were retrieved.</title><p>(XLS)</p></caption><media xlink:href="pone.0135209.s001.xls"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s002"><label>S2 File</label><caption><title>Distribution of phylogeographic lineages of six tephritid fruit flies.</title><p>(A) <italic>Bactrocera oleae</italic>, (B) <italic>Ceratitis fasciventris</italic>, (C) <italic>Anastrepha obliqua</italic>, (D) <italic>Anastrepha fraterculus</italic>, (E) <italic>Rhagoletis pomonella</italic>, (F) <italic>Bacrocera cucurbitae</italic>.</p><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s002.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s003"><label>S3 File</label><caption><title>Principal component analysis on climatic data extracted from lineage occurrences.</title><p>(A) <italic>Anastrepha obliqua</italic>, (B) <italic>Anastrepha fraterculus</italic>, (C) <italic>Rhagoletis pomonella</italic>.</p><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s003.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s004"><label>S4 File</label><caption><title>Worldwide projections of MaxEnt predictions for six tephritid fruit flies.</title><p>(A) <italic>Bactrocera oleae</italic>, (B) <italic>Ceratitis fasciventris</italic>, (C) <italic>Anastrepha obliqua</italic>, (D) <italic>Anastrepha fraterculus</italic>, (E) <italic>Rhagoletis pomonella</italic>, (F) <italic>Bacrocera cucurbitae</italic>.</p><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s004.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s005"><label>S5 File</label><caption><title>Worldwide projections of BRT predictions for six tephritid fruit flies.</title><p>(A) <italic>Bactrocera oleae</italic>, (B) <italic>Ceratitis fasciventris</italic>, (C) <italic>Anastrepha obliqua</italic>, (D) <italic>Anastrepha fraterculus</italic>, (E) <italic>Rhagoletis pomonella</italic>, (F) <italic>Bacrocera cucurbitae</italic>.</p><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s005.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s006"><label>S1 Table</label><caption><title>Inertia values displayed by between-class analyses.</title><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s006.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s007"><label>S2 Table</label><caption><title>Area under the receiver operating characteristics (ROC) curve (AUC) for each model.</title><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s007.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="pone.0135209.s008"><label>S1 Text</label><caption><title>Positioning of absences and background data in SDM of six tephritid fruit flies.</title><p>(DOCX)</p></caption><media xlink:href="pone.0135209.s008.docx"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material></sec></body><back><ack><p>We thank Raul Ruiz-Arce (USDA-APHIS) and Marshall W. Johnson (University of California, Riverside) for their help in the data collection. We thank Robert S. Copeland for providing pictures of fruit flies. We thank Andrea Sánchez Meseguer for constructive discussions and comments on the manuscript.</p></ack><ref-list><title>References</title><ref id="pone.0135209.ref001"><label>1</label><mixed-citation publication-type="journal"><name><surname>Pimentel</surname><given-names>D</given-names></name>, <name><surname>McNair</surname><given-names>S</given-names></name>, <name><surname>Janecka</surname><given-names>J</given-names></name>, <name><surname>Wightman</surname><given-names>J</given-names></name>, <name><surname>Simmonds</surname><given-names>C</given-names></name>, <name><surname>O'Connell</surname><given-names>C</given-names></name>, <etal>et al</etal><article-title>Economic and environmental threats of alien plant, animal, and microbe invasions</article-title>. <source>Agric Ecosyst Environ</source>. <year>2001</year>;<volume>84</volume>(<issue>1</issue>):<fpage>1</fpage>–<lpage>20</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/S0167-8809(00)00178-X">10.1016/S0167-8809(00)00178-X</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref002"><label>2</label><mixed-citation publication-type="journal"><name><surname>Hulme</surname><given-names>PE</given-names></name>. <article-title>Trade, transport and trouble: managing invasive species pathways in an era of globalization</article-title>. <source>J Appl Ecol</source>. <year>2009</year>;<volume>46</volume>(<issue>1</issue>):<fpage>10</fpage>–<lpage>8</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref003"><label>3</label><mixed-citation publication-type="journal"><name><surname>Kolar</surname><given-names>CS</given-names></name>, <name><surname>Lodge</surname><given-names>DM</given-names></name>. <article-title>Progress in invasion biology: predicting invaders</article-title>. <source>Trends Ecol Evol</source>. <year>2001</year>;<volume>16</volume>(<issue>4</issue>):<fpage>199</fpage>–<lpage>204</lpage>. <pub-id pub-id-type="pmid">11245943</pub-id></mixed-citation></ref><ref id="pone.0135209.ref004"><label>4</label><mixed-citation publication-type="journal"><name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Vieglais</surname><given-names>DA</given-names></name>. <article-title>Predicting species invasions using ecological niche modeling: new approaches from bioinformatics attack a pressing problem</article-title>. <source>Bioscience</source>. <year>2001</year>;<volume>51</volume>(<issue>5</issue>):<fpage>363</fpage>–<lpage>71</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1641/0006-3568(2001)051[0363:psiuen]2.0.co;2">10.1641/0006-3568(2001)051[0363:psiuen]2.0.co;2</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref005"><label>5</label><mixed-citation publication-type="journal"><name><surname>Rafter</surname><given-names>MA</given-names></name>, <name><surname>Hereward</surname><given-names>JP</given-names></name>, <name><surname>Walter</surname><given-names>GH</given-names></name>. <article-title>Species limits, quarantine risk and the intrigue of a polyphagous invasive pest with highly restricted host relationships in its area of invasion</article-title>. <source>Evol App</source>. <year>2013</year>;<volume>6</volume>(<issue>8</issue>):<fpage>1195</fpage>–<lpage>207</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref006"><label>6</label><mixed-citation publication-type="journal"><name><surname>Santos</surname><given-names>H</given-names></name>, <name><surname>Burban</surname><given-names>C</given-names></name>, <name><surname>Rousselet</surname><given-names>J</given-names></name>, <name><surname>Rossi</surname><given-names>JP</given-names></name>, <name><surname>Branco</surname><given-names>M</given-names></name>, <name><surname>Kerdelhué</surname><given-names>C</given-names></name>. <article-title>Incipient allochronic speciation in the pine processionary moth (<italic>Thaumetopoea pityocampa</italic>, Lepidoptera, Notodontidae)</article-title>. <source>J Evol Biol</source>. <year>2011</year>;<volume>24</volume>(<issue>1</issue>):<fpage>146</fpage>–<lpage>58</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1420-9101.2010.02147.x">10.1111/j.1420-9101.2010.02147.x</ext-link></comment><pub-id pub-id-type="pmid">20964783</pub-id></mixed-citation></ref><ref id="pone.0135209.ref007"><label>7</label><mixed-citation publication-type="journal"><name><surname>Rey</surname><given-names>O</given-names></name>, <name><surname>Estoup</surname><given-names>A</given-names></name>, <name><surname>Vonshak</surname><given-names>M</given-names></name>, <name><surname>Loiseau</surname><given-names>A</given-names></name>, <name><surname>Blanchet</surname><given-names>S</given-names></name>, <name><surname>Calcaterra</surname><given-names>L</given-names></name>, <etal>et al</etal><article-title>Where do adaptive shifts occur during invasion? A multidisciplinary approach to unravelling cold adaptation in a tropical ant species invading the Mediterranean area</article-title>. <source>Ecol Lett</source>. <year>2012</year>;<volume>15</volume>(<issue>11</issue>):<fpage>1266</fpage>–<lpage>75</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1461-0248.2012.01849.x">10.1111/j.1461-0248.2012.01849.x</ext-link></comment><pub-id pub-id-type="pmid">22906215</pub-id></mixed-citation></ref><ref id="pone.0135209.ref008"><label>8</label><mixed-citation publication-type="journal"><name><surname>Kelly</surname><given-names>DW</given-names></name>, <name><surname>Muirhead</surname><given-names>JR</given-names></name>, <name><surname>Heath</surname><given-names>DD</given-names></name>, <name><surname>Macisaac</surname><given-names>HJ</given-names></name>. <article-title>Contrasting patterns in genetic diversity following multiple invasions of fresh and brackish waters</article-title>. <source>Mol Ecol</source>. <year>2006</year>;<volume>15</volume>(<issue>12</issue>):<fpage>3641</fpage>–<lpage>53</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-294X.2006.03012.x">10.1111/j.1365-294X.2006.03012.x</ext-link></comment><pub-id pub-id-type="pmid">17032263</pub-id></mixed-citation></ref><ref id="pone.0135209.ref009"><label>9</label><mixed-citation publication-type="journal"><name><surname>Lachmuth</surname><given-names>S</given-names></name>, <name><surname>Durka</surname><given-names>W</given-names></name>, <name><surname>Schurr</surname><given-names>FM</given-names></name>. <article-title>The making of a rapid plant invader: genetic diversity and differentiation in the native and invaded range of <italic>Senecio inaequidens</italic></article-title>. <source>Mol Ecol</source>. <year>2010</year>;<volume>19</volume>(<issue>18</issue>):<fpage>3952</fpage>–<lpage>67</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-294X.2010.04797.x">10.1111/j.1365-294X.2010.04797.x</ext-link></comment><pub-id pub-id-type="pmid">20854275</pub-id></mixed-citation></ref><ref id="pone.0135209.ref010"><label>10</label><mixed-citation publication-type="journal"><name><surname>Hamilton</surname><given-names>JA</given-names></name>, <name><surname>Okada</surname><given-names>M</given-names></name>, <name><surname>Korves</surname><given-names>T</given-names></name>, <name><surname>Schmitt</surname><given-names>J</given-names></name>. <article-title>The role of climate adaptation in colonization success in <italic>Arabidopsis thaliana</italic></article-title>. <source>Mol Ecol</source>. <year>2015</year><comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/mec.13099">10.1111/mec.13099</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref011"><label>11</label><mixed-citation publication-type="journal"><name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Holt</surname><given-names>RD</given-names></name>. <article-title>Niche differentiation in Mexican birds: using point occurrences to detect ecological innovation</article-title>. <source>Ecol Lett</source>. <year>2003</year>;<volume>6</volume>(<issue>8</issue>):<fpage>774</fpage>–<lpage>82</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1046/j.1461-0248.2003.00502.x">10.1046/j.1461-0248.2003.00502.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref012"><label>12</label><mixed-citation publication-type="book"><name><surname>White</surname><given-names>IM</given-names></name>, <name><surname>Elson-Harris</surname><given-names>MM</given-names></name>. <source>Fruit flies of economic significance: their identification and bionomics</source>: <publisher-name>CAB International</publisher-name>; <year>1992</year>.</mixed-citation></ref><ref id="pone.0135209.ref013"><label>13</label><mixed-citation publication-type="other">White I, Meyer Md, Stonehouse J, Price N, Seewooruthun I. A review of native and introduced fruit flies (Diptera, Tephritidae) in the Indian Ocean islands of Mauritius, Réunion, Rodrigues and Seychelles. Proceedings of the Indian Ocean Commission, Regional Fruit Fly Symposium, Flic en Flac, Mauritius, 5th-9th June, 2000. Indian Ocean Commission.</mixed-citation></ref><ref id="pone.0135209.ref014"><label>14</label><mixed-citation publication-type="journal"><name><surname>Mun</surname><given-names>J</given-names></name>, <name><surname>Bohonak</surname><given-names>AJ</given-names></name>, <name><surname>Roderick</surname><given-names>GK</given-names></name>. <article-title>Population structure of the pumpkin fruit fly <italic>Bactrocera depressa</italic> (Tephritidae) in Korea and Japan: Pliocene allopatry or recent invasion?</article-title><source>Mol Ecol</source>. <year>2003</year>;<volume>12</volume>(<issue>11</issue>):<fpage>2941</fpage>–<lpage>51</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1046/j.1365-294X.2003.01978.x">10.1046/j.1365-294X.2003.01978.x</ext-link></comment><pub-id pub-id-type="pmid">14629375</pub-id></mixed-citation></ref><ref id="pone.0135209.ref015"><label>15</label><mixed-citation publication-type="journal"><name><surname>Rice</surname><given-names>R</given-names></name>, <name><surname>Phillips</surname><given-names>P</given-names></name>, <name><surname>Stewart-Leslie</surname><given-names>J</given-names></name>, <name><surname>Sibbett</surname><given-names>G</given-names></name>. <article-title>Olive fruit fly populations measured in central and southern California</article-title>. <source>Calif Agric</source>. <year>2003</year>;<volume>57</volume>(<issue>4</issue>):<fpage>122</fpage>–<lpage>7</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref016"><label>16</label><mixed-citation publication-type="journal"><name><surname>Koyama</surname><given-names>J</given-names></name>, <name><surname>Kakinohana</surname><given-names>H</given-names></name>, <name><surname>Miyatake</surname><given-names>T</given-names></name>. <article-title>Eradication of the melon fly, <italic>Bactrocera cucurbitae</italic>, in Japan: importance of behavior, ecology, genetics, and evolution</article-title>. <source>Annu Rev Entomol</source>. <year>2004</year>;<volume>49</volume>(<issue>1</issue>):<fpage>331</fpage>–<lpage>49</lpage>.<pub-id pub-id-type="pmid">14651467</pub-id></mixed-citation></ref><ref id="pone.0135209.ref017"><label>17</label><mixed-citation publication-type="journal"><name><surname>Drew</surname><given-names>RAI</given-names></name>, <name><surname>Tsuruta</surname><given-names>K</given-names></name>, <name><surname>White</surname><given-names>I</given-names></name>. <article-title>A new species of pest fruit fly (Diptera: Tephritidae: Dacinae) from Sri Lanka and Africa</article-title>. <source>Afr Entomol</source>. <year>2005</year>;<volume>13</volume>(<issue>1</issue>):p. <fpage>149</fpage>–<lpage>54</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref018"><label>18</label><mixed-citation publication-type="journal"><name><surname>Malacrida</surname><given-names>AR</given-names></name>, <name><surname>Gomulski</surname><given-names>LM</given-names></name>, <name><surname>Bonizzoni</surname><given-names>M</given-names></name>, <name><surname>Bertin</surname><given-names>S</given-names></name>, <name><surname>Gasperi</surname><given-names>G</given-names></name>, <name><surname>Guglielmino</surname><given-names>CR</given-names></name>. <article-title>Globalization and fruitfly invasion and expansion: the medfly paradigm</article-title>. <source>Genetica</source>. <year>2006</year>;<volume>131</volume>(<issue>1</issue>):<fpage>1</fpage>–<lpage>9</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1007/s10709-006-9117-2">10.1007/s10709-006-9117-2</ext-link></comment><pub-id pub-id-type="pmid">17111234</pub-id></mixed-citation></ref><ref id="pone.0135209.ref019"><label>19</label><mixed-citation publication-type="journal"><name><surname>Papadopoulos</surname><given-names>NT</given-names></name>, <name><surname>Plant</surname><given-names>RE</given-names></name>, <name><surname>Carey</surname><given-names>JR</given-names></name>. <article-title>From trickle to flood: the large-scale, cryptic invasion of California by tropical fruit flies</article-title>. <source>Proc R Soc Lond B Biol Sci</source>. <year>2013</year>;<volume>280</volume>(<issue>1768</issue>):<fpage>20131466</fpage>.</mixed-citation></ref><ref id="pone.0135209.ref020"><label>20</label><mixed-citation publication-type="journal"><name><surname>Clarke</surname><given-names>AR</given-names></name>, <name><surname>Armstrong</surname><given-names>KF</given-names></name>, <name><surname>Carmichael</surname><given-names>AE</given-names></name>, <name><surname>Milne</surname><given-names>JR</given-names></name>, <name><surname>Raghu</surname><given-names>S</given-names></name>, <name><surname>Roderick</surname><given-names>GK</given-names></name>, <etal>et al</etal><article-title>Invasive phytophagous pests arising through a recent tropical evolutionary radiation: the <italic>Bactrocera dorsalis</italic> complex of fruit flies</article-title>. <source>Annu Rev Entomol</source>. <year>2005</year>;<volume>50</volume>:<fpage>293</fpage>–<lpage>319</lpage>. <pub-id pub-id-type="pmid">15355242</pub-id></mixed-citation></ref><ref id="pone.0135209.ref021"><label>21</label><mixed-citation publication-type="journal"><name><surname>Nardi</surname><given-names>F</given-names></name>, <name><surname>Carapelli</surname><given-names>A</given-names></name>, <name><surname>Dallai</surname><given-names>R</given-names></name>, <name><surname>Roderick</surname><given-names>GK</given-names></name>, <name><surname>Frati</surname><given-names>F</given-names></name>. <article-title>Population structure and colonization history of the olive fly, <italic>Bactrocera oleae</italic> (Diptera, Tephritidae)</article-title>. <source>Mol Ecol</source>. <year>2005</year>;<volume>14</volume>(<issue>9</issue>):<fpage>2729</fpage>–<lpage>38</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-294X.2005.02610.x">10.1111/j.1365-294X.2005.02610.x</ext-link></comment><pub-id pub-id-type="pmid">16029474</pub-id></mixed-citation></ref><ref id="pone.0135209.ref022"><label>22</label><mixed-citation publication-type="journal"><name><surname>Nardi</surname><given-names>F</given-names></name>, <name><surname>Carapelli</surname><given-names>A</given-names></name>, <name><surname>Boore</surname><given-names>JL</given-names></name>, <name><surname>Roderick</surname><given-names>GK</given-names></name>, <name><surname>Dallai</surname><given-names>R</given-names></name>, <name><surname>Frati</surname><given-names>F</given-names></name>. <article-title>Domestication of olive fly through a multi-regional host shift to cultivated olives: Comparative dating using complete mitochondrial genomes</article-title>. <source>Mol Phylogenet Evol</source>. <year>2010</year>;<volume>57</volume>(<issue>2</issue>):<fpage>678</fpage>–<lpage>86</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.ympev.2010.08.008">10.1016/j.ympev.2010.08.008</ext-link></comment><pub-id pub-id-type="pmid">20723608</pub-id></mixed-citation></ref><ref id="pone.0135209.ref023"><label>23</label><mixed-citation publication-type="journal"><name><surname>Ruiz-Arce</surname><given-names>R</given-names></name>, <name><surname>Barr</surname><given-names>NB</given-names></name>, <name><surname>Owen</surname><given-names>CL</given-names></name>, <name><surname>Thomas</surname><given-names>DB</given-names></name>, <name><surname>McPheron</surname><given-names>BA</given-names></name>. <article-title>Phylogeography of <italic>Anastrepha obliqua</italic> inferred with mtDNA sequencing</article-title>. <source>J Econ Entomol</source>. <year>2012</year>;<volume>105</volume>(<issue>6</issue>):<fpage>2147</fpage>–<lpage>60</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1603/ec12211">10.1603/ec12211</ext-link></comment><pub-id pub-id-type="pmid">23356081</pub-id></mixed-citation></ref><ref id="pone.0135209.ref024"><label>24</label><mixed-citation publication-type="journal"><name><surname>Virgilio</surname><given-names>M</given-names></name>, <name><surname>Delatte</surname><given-names>H</given-names></name>, <name><surname>Quilici</surname><given-names>S</given-names></name>, <name><surname>Backeljau</surname><given-names>T</given-names></name>, <name><surname>De Meyer</surname><given-names>M</given-names></name>. <article-title>Cryptic diversity and gene flow among three African agricultural pests: <italic>Ceratitis rosa</italic>, <italic>Ceratitis fasciventris</italic> and <italic>Ceratitis anonae</italic> (Diptera, Tephritidae)</article-title>. <source>Mol Ecol</source>. <year>2013</year>;<volume>22</volume>(<issue>9</issue>):<fpage>2526</fpage>–<lpage>39</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/mec.12278">10.1111/mec.12278</ext-link></comment><pub-id pub-id-type="pmid">23506441</pub-id></mixed-citation></ref><ref id="pone.0135209.ref025"><label>25</label><mixed-citation publication-type="journal"><name><surname>Grout</surname><given-names>TG</given-names></name>, <name><surname>Stoltz</surname><given-names>KC</given-names></name>. <article-title>Developmental rates at constant temperatures of three economically important <italic>Ceratitis</italic> spp. (Diptera: Tephritidae) from southern Africa</article-title>. <source>Environ Entomol</source>. <year>2007</year>;<volume>36</volume>(<issue>6</issue>):<fpage>1310</fpage>–<lpage>7</lpage>. <pub-id pub-id-type="pmid">18284758</pub-id></mixed-citation></ref><ref id="pone.0135209.ref026"><label>26</label><mixed-citation publication-type="journal"><name><surname>Papanastasiou</surname><given-names>SA</given-names></name>, <name><surname>Nestel</surname><given-names>D</given-names></name>, <name><surname>Diamantidis</surname><given-names>AD</given-names></name>, <name><surname>Nakas</surname><given-names>CT</given-names></name>, <name><surname>Papadopoulos</surname><given-names>NT</given-names></name>. <article-title>Physiological and biological patterns of a highland and a coastal population of the European cherry fruit fly during diapause</article-title>. <source>J Insect Physiol</source>. <year>2011</year>;<volume>57</volume>(<issue>1</issue>):<fpage>83</fpage>–<lpage>93</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.jinsphys.2010.09.008">10.1016/j.jinsphys.2010.09.008</ext-link></comment><pub-id pub-id-type="pmid">20933516</pub-id></mixed-citation></ref><ref id="pone.0135209.ref027"><label>27</label><mixed-citation publication-type="journal"><name><surname>Müller</surname><given-names>H-G</given-names></name>, <name><surname>Wu</surname><given-names>S</given-names></name>, <name><surname>Diamantidis</surname><given-names>AD</given-names></name>, <name><surname>Papadopoulos</surname><given-names>NT</given-names></name>, <name><surname>Carey</surname><given-names>JR</given-names></name>. <article-title>Reproduction is adapted to survival characteristics across geographically isolated medfly populations</article-title>. <source>Proc R Soc Lond B Biol Sci</source>. <year>2009</year>;<volume>276</volume>(<issue>1677</issue>):<fpage>4409</fpage>–<lpage>16</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref028"><label>28</label><mixed-citation publication-type="journal"><name><surname>Diamantidis</surname><given-names>AD</given-names></name>, <name><surname>Papadopoulos</surname><given-names>NT</given-names></name>, <name><surname>Nakas</surname><given-names>CT</given-names></name>, <name><surname>Wu</surname><given-names>S</given-names></name>, <name><surname>Müller</surname><given-names>HG</given-names></name>, <name><surname>Carey</surname><given-names>JR</given-names></name>. <article-title>Life history evolution in a globally invading tephritid: patterns of survival and reproduction in medflies from six world regions</article-title>. <source>Biol J Linn Soc</source>. <year>2009</year>;<volume>97</volume>(<issue>1</issue>):<fpage>106</fpage>–<lpage>17</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref029"><label>29</label><mixed-citation publication-type="journal"><name><surname>Diamantidis</surname><given-names>AD</given-names></name>, <name><surname>Carey</surname><given-names>JR</given-names></name>, <name><surname>Nakas</surname><given-names>CT</given-names></name>, <name><surname>Papadopoulos</surname><given-names>NT</given-names></name>. <article-title>Ancestral populations perform better in a novel environment: domestication of Mediterranean fruit fly populations from five global regions</article-title>. <source>Biol J Linn Soc</source>. <year>2011</year>;<volume>102</volume>(<issue>2</issue>):<fpage>334</fpage>–<lpage>45</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref030"><label>30</label><mixed-citation publication-type="journal"><name><surname>Aluja</surname><given-names>M</given-names></name>, <name><surname>Pérez-Staples</surname><given-names>D</given-names></name>, <name><surname>Macías-Ordóñez</surname><given-names>R</given-names></name>, <name><surname>Piñero</surname><given-names>J</given-names></name>, <name><surname>Mcpheron</surname><given-names>B</given-names></name>, <name><surname>Hernández-Ortiz</surname><given-names>V</given-names></name>. <article-title>Nonhost status of <italic>Citrus sinensis</italic> cultivar Valencia and <italic>C</italic>. <italic>paradisi</italic> cultivar Ruby Red to Mexican <italic>Anastrepha fraterculus</italic> (Diptera: Tephritidae)</article-title>. <source>J Econ Entomol</source>. <year>2003</year>;<volume>96</volume>(<issue>6</issue>):<fpage>1693</fpage>–<lpage>703</lpage>. <pub-id pub-id-type="pmid">14977105</pub-id></mixed-citation></ref><ref id="pone.0135209.ref031"><label>31</label><mixed-citation publication-type="journal"><name><surname>Smith-Caldas</surname><given-names>MR</given-names></name>, <name><surname>Mcpheron</surname><given-names>BA</given-names></name>, <name><surname>Silva</surname><given-names>JG</given-names></name>, <name><surname>Zucchi</surname><given-names>RA</given-names></name>. <article-title>Phylogenetic relationships among species of the <italic>fraterculus</italic> group (<italic>Anastrepha</italic>: Diptera: Tephritidae) inferred from DNA sequences of mitochondrial cytochrome oxidase I</article-title>. <source>Neotrop Entomol</source>. <year>2001</year>;<volume>30</volume>(<issue>4</issue>):<fpage>565</fpage>–<lpage>73</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref032"><label>32</label><mixed-citation publication-type="journal"><name><surname>Feder</surname><given-names>JL</given-names></name>, <name><surname>Berlocher</surname><given-names>SH</given-names></name>, <name><surname>Roethele</surname><given-names>JB</given-names></name>, <name><surname>Dambroski</surname><given-names>H</given-names></name>, <name><surname>Smith</surname><given-names>JJ</given-names></name>, <name><surname>Perry</surname><given-names>WL</given-names></name>, <etal>et al</etal><article-title>Allopatric genetic origins for sympatric host-plant shifts and race formation in <italic>Rhagoletis</italic></article-title>. <source>Proc Natl Acad Sci U S A</source>. <year>2003</year>;<volume>100</volume>(<issue>18</issue>):<fpage>10314</fpage>–<lpage>9</lpage>. <pub-id pub-id-type="pmid">12928500</pub-id></mixed-citation></ref><ref id="pone.0135209.ref033"><label>33</label><mixed-citation publication-type="journal"><name><surname>Feder</surname><given-names>JL</given-names></name>, <name><surname>Xie</surname><given-names>X</given-names></name>, <name><surname>Rull</surname><given-names>J</given-names></name>, <name><surname>Velez</surname><given-names>S</given-names></name>, <name><surname>Forbes</surname><given-names>A</given-names></name>, <name><surname>Leung</surname><given-names>B</given-names></name>, <etal>et al</etal><article-title>Mayr, Dobzhansky, and Bush and the complexities of sympatric speciation in <italic>Rhagoletis</italic></article-title>. <source>Proc Natl Acad Sci U S A</source>. <year>2005</year>;<volume>102</volume>(<issue>suppl 1</issue>):<fpage>6573</fpage>–<lpage>80</lpage>. <pub-id pub-id-type="pmid">15851672</pub-id></mixed-citation></ref><ref id="pone.0135209.ref034"><label>34</label><mixed-citation publication-type="journal"><name><surname>Virgilio</surname><given-names>M</given-names></name>, <name><surname>Delatte</surname><given-names>H</given-names></name>, <name><surname>Backeljau</surname><given-names>T</given-names></name>, <name><surname>De Meyer</surname><given-names>M</given-names></name>. <article-title>Macrogeographic population structuring in the cosmopolitan agricultural pest <italic>Bactrocera cucurbitae</italic> (Diptera: Tephritidae)</article-title>. <source>Mol Ecol</source>. <year>2010</year>;<volume>19</volume>(<issue>13</issue>):<fpage>2713</fpage>–<lpage>24</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-294X.2010.04662.x">10.1111/j.1365-294X.2010.04662.x</ext-link></comment><pub-id pub-id-type="pmid">20561200</pub-id></mixed-citation></ref><ref id="pone.0135209.ref035"><label>35</label><mixed-citation publication-type="journal"><name><surname>Michel</surname><given-names>AP</given-names></name>, <name><surname>Rull</surname><given-names>J</given-names></name>, <name><surname>Aluja</surname><given-names>M</given-names></name>, <name><surname>Feder</surname><given-names>JL</given-names></name>. <article-title>The genetic structure of hawthornorn-294X.20 <italic>Rhagoletis pomonella</italic> populations in Mexico: implications for sympatric host race formation</article-title>. <source>Mol Ecol</source>. <year>2007</year>;<volume>16</volume>(<issue>14</issue>):<fpage>2867</fpage>–<lpage>78</lpage>.<pub-id pub-id-type="pmid">17614903</pub-id></mixed-citation></ref><ref id="pone.0135209.ref036"><label>36</label><mixed-citation publication-type="journal"><name><surname>Ludeña</surname><given-names>B</given-names></name>, <name><surname>Bayas</surname><given-names>R</given-names></name>, <name><surname>Pintaud</surname><given-names>J-C</given-names></name>. <article-title>Phylogenetic relationships of Andean-Ecuadorian populations of <italic>Anastrepha fraterculus</italic> (Wiedemann 1830)(Diptera: Tephritidae) inferred from COI and COII gene sequences</article-title>. <source>Ann Soc Entomol Fr</source>. <year>2010</year>;<volume>46</volume>(<issue>3</issue>):<fpage>344</fpage>.</mixed-citation></ref><ref id="pone.0135209.ref037"><label>37</label><mixed-citation publication-type="journal"><name><surname>Hernández-Ortiz</surname><given-names>V</given-names></name>, <name><surname>Bartolucci</surname><given-names>AF</given-names></name>, <name><surname>Morales-Valles</surname><given-names>P</given-names></name>, <name><surname>Frías</surname><given-names>D</given-names></name>, <name><surname>Selivon</surname><given-names>D</given-names></name>. <article-title>Cryptic species of the <italic>Anastrepha fraterculus</italic> complex (Diptera: Tephritidae): a multivariate approach for the recognition of South American morphotypes</article-title>. <source>Ann Entomol Soc Am</source>. <year>2012</year>;<volume>105</volume>(<issue>2</issue>):<fpage>305</fpage>–<lpage>18</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref038"><label>38</label><mixed-citation publication-type="journal"><name><surname>Zygouridis</surname><given-names>NE</given-names></name>, <name><surname>Augustinos</surname><given-names>AA</given-names></name>, <name><surname>Zalom</surname><given-names>FG</given-names></name>, <name><surname>Mathiopoulos</surname><given-names>KD</given-names></name>. <article-title>Analysis of olive fly invasion in California based on microsatellite markers</article-title>. <source>Heredity</source>. <year>2008</year>;<volume>102</volume>(<issue>4</issue>):<fpage>402</fpage>–<lpage>12</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1038/hdy.2008.125">10.1038/hdy.2008.125</ext-link></comment><pub-id pub-id-type="pmid">19107137</pub-id></mixed-citation></ref><ref id="pone.0135209.ref039"><label>39</label><mixed-citation publication-type="journal"><name><surname>van Asch</surname><given-names>B</given-names></name>, <name><surname>Pereira-Castro</surname><given-names>I</given-names></name>, <name><surname>Rei</surname><given-names>F</given-names></name>, <name><surname>Costa</surname><given-names>LT</given-names></name>. <article-title>Mitochondrial haplotypes reveal olive fly (<italic>Bactrocera oleae</italic>) population substructure in the Mediterranean</article-title>. <source>Genetica</source>. <year>2012</year>;<volume>140</volume>(<issue>4–6</issue>):<fpage>181</fpage>–<lpage>7</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1007/s10709-012-9669-2">10.1007/s10709-012-9669-2</ext-link></comment><pub-id pub-id-type="pmid">22825843</pub-id></mixed-citation></ref><ref id="pone.0135209.ref040"><label>40</label><mixed-citation publication-type="journal"><name><surname>Daane</surname><given-names>KM</given-names></name>, <name><surname>Johnson</surname><given-names>MW</given-names></name>. <article-title>Olive fruit fly: managing an ancient pest in modern times</article-title>. <source>Annu Rev Entomol</source>. <year>2010</year>;<volume>55</volume>(<issue>1</issue>):<fpage>151</fpage>–<lpage>69</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1146/annurev.ento.54.110807.090553">10.1146/annurev.ento.54.110807.090553</ext-link></comment><pub-id pub-id-type="pmid">19961328</pub-id></mixed-citation></ref><ref id="pone.0135209.ref041"><label>41</label><mixed-citation publication-type="journal"><name><surname>Aluja</surname><given-names>M</given-names></name>, <name><surname>Piñero</surname><given-names>J</given-names></name>, <name><surname>López</surname><given-names>M</given-names></name>, <name><surname>Ruíz</surname><given-names>C</given-names></name>, <name><surname>Zúñiga</surname><given-names>A</given-names></name>, <name><surname>Piedra</surname><given-names>E</given-names></name>, <etal>et al</etal><article-title>New host plant and distribution records in Mexico for <italic>Anastrepha</italic> spp., <italic>Toxotrypana curvicauda</italic> Gerstacker, <italic>Rhagoletis zoqui</italic> Bush, <italic>Rhagoletis</italic> sp., and <italic>Hexachaeta</italic> sp. (Diptera: Tephritidae)</article-title>. <source>Proc Entomol Soc Wash</source>. <year>2000</year>;<volume>102</volume>(<issue>4</issue>):<fpage>802</fpage>–<lpage>15</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref042"><label>42</label><mixed-citation publication-type="journal"><name><surname>Md</surname><given-names>Meyer</given-names></name>. <article-title>On the identity of the Natal fruit fly <italic>Ceratitis rosa</italic> Karsch (Diptera, Tephritidae)</article-title>. <source>Bull Inst R Sci Nat Belg Entomol</source>. <year>2001</year>;<volume>71</volume>:<fpage>55</fpage>–<lpage>62</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref043"><label>43</label><mixed-citation publication-type="journal"><name><surname>Copeland</surname><given-names>RS</given-names></name>, <name><surname>Wharton</surname><given-names>RA</given-names></name>, <name><surname>Luke</surname><given-names>Q</given-names></name>, <name><surname>De Meyer</surname><given-names>M</given-names></name>, <name><surname>Lux</surname><given-names>S</given-names></name>, <name><surname>Zenz</surname><given-names>N</given-names></name>, <etal>et al</etal><article-title>Geographic distribution, host fruit, and parasitoids of African fruit fly pests <italic>Ceratitis anonae</italic>, <italic>Ceratitis cosyra</italic>, <italic>Ceratitis fasciventris</italic>, and <italic>Ceratitis rosa</italic> (Diptera: Tephritidae) in Kenya</article-title>. <source>Ann Entomol Soc Am</source>. <year>2006</year>;<volume>99</volume>(<issue>2</issue>):<fpage>261</fpage>–<lpage>78</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref044"><label>44</label><mixed-citation publication-type="journal"><name><surname>Virgilio</surname><given-names>M</given-names></name>, <name><surname>Backeljau</surname><given-names>T</given-names></name>, <name><surname>Barr</surname><given-names>N</given-names></name>, <name><surname>Meyer</surname><given-names>MD</given-names></name>. <article-title>Molecular evaluation of nominal species in the <italic>Ceratitis fasciventris</italic>, <italic>C</italic>. <italic>anonae</italic>, <italic>C</italic>. <italic>rosa</italic> complex (Diptera: Tephritidae)</article-title>. <source>Mol Phylogenet Evol</source>. <year>2008</year>;<volume>48</volume>(<issue>1</issue>):<fpage>270</fpage>–<lpage>80</lpage>. <pub-id pub-id-type="pmid">18502154</pub-id></mixed-citation></ref><ref id="pone.0135209.ref045"><label>45</label><mixed-citation publication-type="journal"><name><surname>Hernández-Ortiz</surname><given-names>V</given-names></name>, <name><surname>Gómez-Anaya</surname><given-names>J</given-names></name>, <name><surname>Sánchez</surname><given-names>A</given-names></name>, <name><surname>McPheron</surname><given-names>B</given-names></name>, <name><surname>Aluja</surname><given-names>M</given-names></name>. <article-title>Morphometric analysis of Mexican and South American populations of the <italic>Anastrepha fraterculus</italic> complex (Diptera: Tephritidae) and recognition of a distinct Mexican morphotype</article-title>. <source>Bull Entomol Res</source>. <year>2004</year>;<volume>94</volume>(<issue>06</issue>):<fpage>487</fpage>–<lpage>99</lpage>.<pub-id pub-id-type="pmid">15541188</pub-id></mixed-citation></ref><ref id="pone.0135209.ref046"><label>46</label><mixed-citation publication-type="journal"><name><surname>Vera</surname><given-names>MT</given-names></name>, <name><surname>Cáceres</surname><given-names>C</given-names></name>, <name><surname>Wornoayporn</surname><given-names>V</given-names></name>, <name><surname>Islam</surname><given-names>A</given-names></name>, <name><surname>Robinson</surname><given-names>AS</given-names></name>, <name><surname>Marcelo</surname><given-names>H</given-names></name>, <etal>et al</etal><article-title>Mating incompatibility among populations of the South American fruit fly <italic>Anastrepha fraterculus</italic> (Diptera: Tephritidae)</article-title>. <source>Ann Entomol Soc Am</source>. <year>2006</year>;<volume>99</volume>(<issue>2</issue>):<fpage>387</fpage>–<lpage>97</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref047"><label>47</label><mixed-citation publication-type="journal"><name><surname>Alberti</surname><given-names>AC</given-names></name>, <name><surname>Rodriguero</surname><given-names>MS</given-names></name>, <name><surname>Cendra</surname><given-names>PG</given-names></name>, <name><surname>Saidman</surname><given-names>BO</given-names></name>, <name><surname>Vilardi</surname><given-names>JC</given-names></name>. <article-title>Evidence indicating that Argentine populations of <italic>Anastrepha fraterculus</italic> (Diptera: Tephritidae) belong to a single biological species</article-title>. <source>Ann Entomol Soc Am</source>. <year>2002</year>;<volume>95</volume>(<issue>4</issue>):<fpage>505</fpage>–<lpage>12</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref048"><label>48</label><mixed-citation publication-type="journal"><name><surname>Selivon</surname><given-names>D</given-names></name>, <name><surname>Perondini</surname><given-names>A</given-names></name>, <name><surname>Morgante</surname><given-names>J</given-names></name>. <article-title>A genetic–morphological characterization of two cryptic species of the <italic>Anastrepha fraterculus</italic> complex (Diptera: Tephritidae)</article-title>. <source>Ann Entomol Soc Am</source>. <year>2005</year>;<volume>98</volume>(<issue>3</issue>):<fpage>367</fpage>–<lpage>81</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref049"><label>49</label><mixed-citation publication-type="journal"><name><surname>Alberti</surname><given-names>AC</given-names></name>, <name><surname>Confalonieri</surname><given-names>VA</given-names></name>, <name><surname>Zandomeni</surname><given-names>RO</given-names></name>, <name><surname>Vilardi</surname><given-names>JC</given-names></name>. <article-title>Phylogeographic studies on natural populations of the South American fruit fly, <italic>Anastrepha fraterculus</italic> (Diptera: Tephritidae)</article-title>. <source>Genetica</source>. <year>2008</year>;<volume>132</volume>(<issue>1</issue>):<fpage>1</fpage>–<lpage>8</lpage>. <pub-id pub-id-type="pmid">17318315</pub-id></mixed-citation></ref><ref id="pone.0135209.ref050"><label>50</label><mixed-citation publication-type="journal"><name><surname>Wisz</surname><given-names>MS</given-names></name>, <name><surname>Hijmans</surname><given-names>R</given-names></name>, <name><surname>Li</surname><given-names>J</given-names></name>, <name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Graham</surname><given-names>C</given-names></name>, <name><surname>Guisan</surname><given-names>A</given-names></name>. <article-title>Effects of sample size on the performance of species distribution models</article-title>. <source>Divers Distrib</source>. <year>2008</year>;<volume>14</volume>(<issue>5</issue>):<fpage>763</fpage>–<lpage>73</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref051"><label>51</label><mixed-citation publication-type="journal"><name><surname>Hernandez</surname><given-names>PA</given-names></name>, <name><surname>Graham</surname><given-names>CH</given-names></name>, <name><surname>Master</surname><given-names>LL</given-names></name>, <name><surname>Albert</surname><given-names>DL</given-names></name>. <article-title>The effect of sample size and species characteristics on performance of different species distribution modeling methods</article-title>. <source>Ecography</source>. <year>2006</year>;<volume>29</volume>(<issue>5</issue>):<fpage>773</fpage>–<lpage>85</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref052"><label>52</label><mixed-citation publication-type="journal"><name><surname>Rull</surname><given-names>J</given-names></name>, <name><surname>Aluja</surname><given-names>M</given-names></name>, <name><surname>Feder</surname><given-names>JL</given-names></name>. <article-title>Evolution of intrinsic reproductive isolation among four North American populations of <italic>Rhagoletis pomonella</italic> (Diptera: Tephritidae)</article-title>. <source>Biol J Linn Soc</source>. <year>2010</year>;<volume>100</volume>(<issue>1</issue>):<fpage>213</fpage>–<lpage>23</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref053"><label>53</label><mixed-citation publication-type="journal"><name><surname>Wu</surname><given-names>Y</given-names></name>, <name><surname>McPheron</surname><given-names>BA</given-names></name>, <name><surname>Wu</surname><given-names>JJ</given-names></name>, <name><surname>Li</surname><given-names>ZH</given-names></name>. <article-title>Genetic relationship of the melon fly, <italic>Bactrocera cucurbitae</italic> (Diptera: Tephritidae) inferred from mitochondrial DNA</article-title>. <source>Insect Sci</source>. <year>2012</year>;<volume>19</volume>(<issue>2</issue>):<fpage>195</fpage>–<lpage>204</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref054"><label>54</label><mixed-citation publication-type="journal"><name><surname>Hijmans</surname><given-names>RJ</given-names></name>, <name><surname>Cameron</surname><given-names>SE</given-names></name>, <name><surname>Parra</surname><given-names>JL</given-names></name>, <name><surname>Jones</surname><given-names>PG</given-names></name>, <name><surname>Jarvis</surname><given-names>A</given-names></name>. <article-title>Very high resolution interpolated climate surfaces for global land areas</article-title>. <source>Int J Climatol</source>. <year>2005</year>;<volume>25</volume>(<issue>15</issue>):<fpage>1965</fpage>–<lpage>78</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1002/joc.1276">10.1002/joc.1276</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref055"><label>55</label><mixed-citation publication-type="journal"><name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Nakazawa</surname><given-names>Y</given-names></name>. <article-title>Environmental data sets matter in ecological niche modelling: an example with <italic>Solenopsis invicta</italic> and <italic>Solenopsis richteri</italic></article-title>. <source>Glob Ecol Biogeogr</source>. <year>2008</year>;<volume>17</volume>(<issue>0</issue>):<fpage>135</fpage>–<lpage>44</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1466-8238.2007.00347.x">10.1111/j.1466-8238.2007.00347.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref056"><label>56</label><mixed-citation publication-type="journal"><name><surname>Jiménez-Valverde</surname><given-names>A</given-names></name>, <name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Soberón</surname><given-names>J</given-names></name>, <name><surname>Overton</surname><given-names>JM</given-names></name>, <name><surname>Aragón</surname><given-names>P</given-names></name>, <name><surname>Lobo</surname><given-names>JM</given-names></name>. <article-title>Use of niche models in invasive species risk assessments</article-title>. <source>Biol Invasions</source>. <year>2011</year>;<volume>13</volume>(<issue>12</issue>):<fpage>2785</fpage>–<lpage>97</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1007/s10530-011-9963-4">10.1007/s10530-011-9963-4</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref057"><label>57</label><mixed-citation publication-type="journal"><name><surname>Phillips</surname><given-names>SJ</given-names></name>, <name><surname>Anderson</surname><given-names>RP</given-names></name>, <name><surname>Schapire</surname><given-names>RE</given-names></name>. <article-title>Maximum entropy modeling of species geographic distributions</article-title>. <source>Ecol Model</source>. <year>2006</year>;<volume>190</volume>(<issue>3–4</issue>):<fpage>231</fpage>–<lpage>59</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.ecolmodel.2005.03.026">10.1016/j.ecolmodel.2005.03.026</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref058"><label>58</label><mixed-citation publication-type="other">Hijmans RJ, Phillips S, Leathwick J, Elith J. dismo. Species distribution modeling. R package version 0.9–3. 2013.</mixed-citation></ref><ref id="pone.0135209.ref059"><label>59</label><mixed-citation publication-type="journal"><name><surname>Friedman</surname><given-names>J</given-names></name>, <name><surname>Hastie</surname><given-names>T</given-names></name>, <name><surname>Tibshirani</surname><given-names>R</given-names></name>. <article-title>Additive logistic regression: a statistical view of boosting</article-title>. <source>Ann Stat</source>. <year>2000</year>;<volume>28</volume>(<issue>2</issue>):<fpage>337</fpage>–<lpage>407</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref060"><label>60</label><mixed-citation publication-type="journal"><name><surname>Elith</surname><given-names>J</given-names></name>, <name><surname>Leathwick</surname><given-names>JR</given-names></name>, <name><surname>Hastie</surname><given-names>T</given-names></name>. <article-title>A working guide to boosted regression trees</article-title>. <source>J Anim Ecol</source>. <year>2008</year>;<volume>77</volume>(<issue>4</issue>):<fpage>802</fpage>–<lpage>13</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-2656.2008.01390.x">10.1111/j.1365-2656.2008.01390.x</ext-link></comment><pub-id pub-id-type="pmid">18397250</pub-id></mixed-citation></ref><ref id="pone.0135209.ref061"><label>61</label><mixed-citation publication-type="book"><name><surname>Ridgeway</surname><given-names>G</given-names></name>, <name><surname>Ridgeway</surname><given-names>MG</given-names></name>. <source>The gbm package</source>. <publisher-name>R Foundation for Statistical Computing</publisher-name>, <publisher-loc>Vienna, Austria</publisher-loc><year>2004</year>.</mixed-citation></ref><ref id="pone.0135209.ref062"><label>62</label><mixed-citation publication-type="journal"><name><surname>Elith</surname><given-names>J</given-names></name>, <name><surname>H. Graham</surname><given-names>C</given-names></name>, <name><surname>P. Anderson</surname><given-names>R</given-names></name>, <name><surname>Dudík</surname><given-names>M</given-names></name>, <name><surname>Ferrier</surname><given-names>S</given-names></name>, <name><surname>Guisan</surname><given-names>A</given-names></name>, <etal>et al</etal><article-title>Novel methods improve prediction of species’ distributions from occurrence data</article-title>. <source>Ecography</source>. <year>2006</year>;<volume>29</volume>(<issue>2</issue>):<fpage>129</fpage>–<lpage>51</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.2006.0906-7590.04596.x">10.1111/j.2006.0906-7590.04596.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref063"><label>63</label><mixed-citation publication-type="journal"><name><surname>VanDerWal</surname><given-names>J</given-names></name>, <name><surname>Shoo</surname><given-names>LP</given-names></name>, <name><surname>Graham</surname><given-names>C</given-names></name>, <name><surname>Williams</surname><given-names>SE</given-names></name>. <article-title>Selecting pseudo-absence data for presence-only distribution modeling: How far should you stray from what you know?</article-title><source>Ecol Model</source>. <year>2009</year>;<volume>220</volume>(<issue>4</issue>):<fpage>589</fpage>–<lpage>94</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.ecolmodel.2008.11.010">10.1016/j.ecolmodel.2008.11.010</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref064"><label>64</label><mixed-citation publication-type="journal"><name><surname>Anderson</surname><given-names>RP</given-names></name>, <name><surname>Raza</surname><given-names>A</given-names></name>. <article-title>The effect of the extent of the study region on GIS models of species geographic distributions and estimates of niche evolution: preliminary tests with montane rodents (genus <italic>Nephelomys</italic>) in Venezuela</article-title>. <source>J Biogeogr</source>. <year>2010</year>;<volume>37</volume>(<issue>7</issue>):<fpage>1378</fpage>–<lpage>93</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-2699.2010.02290.x">10.1111/j.1365-2699.2010.02290.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref065"><label>65</label><mixed-citation publication-type="journal"><name><surname>Elith</surname><given-names>J</given-names></name>, <name><surname>Kearney</surname><given-names>M</given-names></name>, <name><surname>Phillips</surname><given-names>S</given-names></name>. <article-title>The art of modelling range-shifting species</article-title>. <source>Methods Ecol Evol</source>. <year>2010</year>;<volume>1</volume>(<issue>4</issue>):<fpage>330</fpage>–<lpage>42</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref066"><label>66</label><mixed-citation publication-type="journal"><name><surname>Fielding</surname><given-names>AH</given-names></name>, <name><surname>Bell</surname><given-names>JF</given-names></name>. <article-title>A review of methods for the assessment of prediction errors in conservation presence/absence models</article-title>. <source>Environ Conserv</source>. <year>1997</year>;<volume>24</volume>(<issue>1</issue>):<fpage>38</fpage>–<lpage>49</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref067"><label>67</label><mixed-citation publication-type="book"><name><surname>Legendre</surname><given-names>L</given-names></name>, <name><surname>Legrendre</surname><given-names>L</given-names></name>. <chapter-title>Numerical ecology</chapter-title><source>Developments in environmental modelling</source>: <publisher-name>Elsevier Science & Technology</publisher-name>; <year>1998</year>.</mixed-citation></ref><ref id="pone.0135209.ref068"><label>68</label><mixed-citation publication-type="journal"><name><surname>Rossi</surname><given-names>J-P</given-names></name>, <name><surname>Blanchart</surname><given-names>E</given-names></name>. <article-title>Seasonal and land-use induced variations of soil macrofauna composition in the Western Ghats, southern India</article-title>. <source>Soil Biol Biochem</source>. <year>2005</year>;<volume>37</volume>(<issue>6</issue>):<fpage>1093</fpage>–<lpage>104</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.soilbio.2004.11.008">10.1016/j.soilbio.2004.11.008</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref069"><label>69</label><mixed-citation publication-type="journal"><name><surname>Dolédec</surname><given-names>S</given-names></name>, <name><surname>Chessel</surname><given-names>D</given-names></name>. <article-title>Rythmes saisonniers et composantes stationnelles en milieu aquatique. I: Description d'un plan d'observation complet par projection de variables</article-title>. <source>Acta Oecol</source>. <year>1987</year>;<volume>8</volume>(<issue>3</issue>):<fpage>403</fpage>–<lpage>26</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref070"><label>70</label><mixed-citation publication-type="book"><name><surname>Manly</surname><given-names>BF</given-names></name>. <source>Randomization, bootstrap and Monte Carlo methods in biology</source>: <publisher-name>CRC Press</publisher-name>; <year>2006</year>.</mixed-citation></ref><ref id="pone.0135209.ref071"><label>71</label><mixed-citation publication-type="journal"><name><surname>Schoener</surname><given-names>TW</given-names></name>. <article-title>Sizes of feeding territories among birds</article-title>. <source>Ecology</source>. <year>1968</year>;<volume>49</volume>:<fpage>123</fpage>–<lpage>41</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref072"><label>72</label><mixed-citation publication-type="journal"><name><surname>Warren</surname><given-names>DL</given-names></name>, <name><surname>Glor</surname><given-names>RE</given-names></name>, <name><surname>Turelli</surname><given-names>M</given-names></name>. <article-title>Environmental niche equivalency versus conservatism: quantitative approaches to niche evolution</article-title>. <source>Evolution</source>. <year>2008</year>;<volume>62</volume>(<issue>11</issue>):<fpage>2868</fpage>–<lpage>83</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1558-5646.2008.00482.x">10.1111/j.1558-5646.2008.00482.x</ext-link></comment><pub-id pub-id-type="pmid">18752605</pub-id></mixed-citation></ref><ref id="pone.0135209.ref073"><label>73</label><mixed-citation publication-type="journal"><name><surname>McCormack</surname><given-names>JE</given-names></name>, <name><surname>Zellmer</surname><given-names>AJ</given-names></name>, <name><surname>Knowles</surname><given-names>LL</given-names></name>. <article-title>Does niche divergence accompany allopatric divergence In <italic>Aphelocoma</italic> jays as predicted under ecological speciation?: Insights from tests with niche models</article-title>. <source>Evolution</source>. <year>2009</year>;<volume>64</volume>(<issue>5</issue>):<fpage>1231</fpage>–<lpage>44</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1558-5646.2009.00900.x">10.1111/j.1558-5646.2009.00900.x</ext-link></comment><pub-id pub-id-type="pmid">19922442</pub-id></mixed-citation></ref><ref id="pone.0135209.ref074"><label>74</label><mixed-citation publication-type="journal"><name><surname>Broennimann</surname><given-names>O</given-names></name>, <name><surname>Fitzpatrick</surname><given-names>MC</given-names></name>, <name><surname>Pearman</surname><given-names>PB</given-names></name>, <name><surname>Petitpierre</surname><given-names>B</given-names></name>, <name><surname>Pellissier</surname><given-names>L</given-names></name>, <name><surname>Yoccoz</surname><given-names>NG</given-names></name>, <etal>et al</etal><article-title>Measuring ecological niche overlap from occurrence and spatial environmental data</article-title>. <source>Glob Ecol Biogeogr</source>. <year>2012</year>;<volume>21</volume>(<issue>4</issue>):<fpage>481</fpage>–<lpage>97</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1466-8238.2011.00698.x">10.1111/j.1466-8238.2011.00698.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref075"><label>75</label><mixed-citation publication-type="journal"><name><surname>Guisan</surname><given-names>A</given-names></name>, <name><surname>Petitpierre</surname><given-names>B</given-names></name>, <name><surname>Broennimann</surname><given-names>O</given-names></name>, <name><surname>Daehler</surname><given-names>C</given-names></name>, <name><surname>Kueffer</surname><given-names>C</given-names></name>. <article-title>Unifying niche shift studies: insights from biological invasions</article-title>. <source>Trends Ecol Evol</source>. <year>2014</year>;<volume>29</volume>(<issue>5</issue>):<fpage>260</fpage>–<lpage>9</lpage>. <pub-id pub-id-type="pmid">24656621</pub-id></mixed-citation></ref><ref id="pone.0135209.ref076"><label>76</label><mixed-citation publication-type="journal"><name><surname>Godsoe</surname><given-names>W</given-names></name>, <name><surname>Case</surname><given-names>BS</given-names></name>. <article-title>Accounting for shifts in the frequency of suitable environments when testing for niche overlap</article-title>. <source>Methods Ecol Evol</source>. <year>2014</year><comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/2041-210X.12307">10.1111/2041-210X.12307</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref077"><label>77</label><mixed-citation publication-type="journal"><name><surname>Barve</surname><given-names>N</given-names></name>, <name><surname>Barve</surname><given-names>V</given-names></name>, <name><surname>Jiménez-Valverde</surname><given-names>A</given-names></name>, <name><surname>Lira-Noriega</surname><given-names>A</given-names></name>, <name><surname>Maher</surname><given-names>SP</given-names></name>, <name><surname>Peterson</surname><given-names>AT</given-names></name>, <etal>et al</etal><article-title>The crucial role of the accessible area in ecological niche modeling and species distribution modeling</article-title>. <source>Ecol Model</source>. <year>2011</year>;<volume>222</volume>(<issue>11</issue>):<fpage>1810</fpage>–<lpage>9</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.ecolmodel.2011.02.011">10.1016/j.ecolmodel.2011.02.011</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref078"><label>78</label><mixed-citation publication-type="journal"><name><surname>Webber</surname><given-names>BL</given-names></name>, <name><surname>Yates</surname><given-names>CJ</given-names></name>, <name><surname>Le Maitre</surname><given-names>DC</given-names></name>, <name><surname>Scott</surname><given-names>JK</given-names></name>, <name><surname>Kriticos</surname><given-names>DJ</given-names></name>, <name><surname>Ota</surname><given-names>N</given-names></name>, <etal>et al</etal><article-title>Modelling horses for novel climate courses: insights from projecting potential distributions of native and alien Australian acacias with correlative and mechanistic models</article-title>. <source>Divers Distrib</source>. <year>2011</year>;<volume>17</volume>(<issue>5</issue>):<fpage>978</fpage>–<lpage>1000</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1472-4642.2011.00811.x">10.1111/j.1472-4642.2011.00811.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref079"><label>79</label><mixed-citation publication-type="journal"><name><surname>Duyck</surname><given-names>P-F</given-names></name>, <name><surname>David</surname><given-names>P</given-names></name>, <name><surname>Quilici</surname><given-names>S</given-names></name>. <article-title>A review of relationships between interspecific competition and invasions in fruit flies (Diptera: Tephritidae)</article-title>. <source>Ecol Entomol</source>. <year>2004</year>;<volume>29</volume>(<issue>5</issue>):<fpage>511</fpage>–<lpage>20</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref080"><label>80</label><mixed-citation publication-type="journal"><name><surname>Katiyar</surname><given-names>KP</given-names></name>, <name><surname>Molina</surname><given-names>JC</given-names></name>, <name><surname>Matheus</surname><given-names>R</given-names></name>. <article-title>Fruit flies (Diptera: Tephritidae) infesting fruits of the genus <italic>Psidium</italic> (Myrtaceae) and their altitudinal distribution in Western Venezuela</article-title>. <source>Fla Entomol</source>. <year>2000</year>;<volume>83</volume>:<fpage>480</fpage>–<lpage>6</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref081"><label>81</label><mixed-citation publication-type="journal"><name><surname>Sivinski</surname><given-names>J</given-names></name>, <name><surname>Aluja</surname><given-names>M</given-names></name>, <name><surname>Piñero</surname><given-names>J</given-names></name>, <name><surname>Ojeda</surname><given-names>M</given-names></name>. <article-title>Novel analysis of spatial and temporal patterns of resource use in a group of tephritid flies of the genus <italic>Anastrepha</italic></article-title>. <source>Ann Entomol Soc Am</source>. <year>2004</year>;<volume>97</volume>(<issue>3</issue>):<fpage>504</fpage>–<lpage>12</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref082"><label>82</label><mixed-citation publication-type="journal"><name><surname>De Villiers</surname><given-names>M</given-names></name>, <name><surname>Hattingh</surname><given-names>V</given-names></name>, <name><surname>Kriticos</surname><given-names>D</given-names></name>. <article-title>Combining field phenological observations with distribution data to model the potential distribution of the fruit fly <italic>Ceratitis rosa</italic> Karsch (Diptera: Tephritidae)</article-title>. <source>Bull Entomol Res</source>. <year>2013</year>;<volume>103</volume>(<issue>1</issue>):<fpage>60</fpage>–<lpage>73</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1017/S0007485312000454">10.1017/S0007485312000454</ext-link></comment><pub-id pub-id-type="pmid">22906299</pub-id></mixed-citation></ref><ref id="pone.0135209.ref083"><label>83</label><mixed-citation publication-type="journal"><name><surname>Papadopoulos</surname><given-names>NT</given-names></name>, <name><surname>Carey</surname><given-names>JR</given-names></name>, <name><surname>Katsoyannos</surname><given-names>BI</given-names></name>, <name><surname>Kouloussis</surname><given-names>NA</given-names></name>. <article-title>Overwintering of the Mediterranean fruit fly (Diptera: Tephritidae) in northern Greece</article-title>. <source>Ann Entomol Soc Am</source>. <year>1996</year>;<volume>89</volume>(<issue>4</issue>):<fpage>526</fpage>–<lpage>34</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref084"><label>84</label><mixed-citation publication-type="journal"><name><surname>Vera</surname><given-names>MT</given-names></name>, <name><surname>Rodriguez</surname><given-names>R</given-names></name>, <name><surname>Segura</surname><given-names>DF</given-names></name>, <name><surname>Cladera</surname><given-names>JL</given-names></name>, <name><surname>Sutherst</surname><given-names>RW</given-names></name>. <article-title>Potential geographical distribution of the Mediterranean fruit fly, <italic>Ceratitis capitata</italic> (Diptera: Tephritidae), with emphasis on Argentina and Australia</article-title>. <source>Environ Entomol</source>. <year>2002</year>;<volume>31</volume>(<issue>6</issue>):<fpage>1009</fpage>–<lpage>22</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref085"><label>85</label><mixed-citation publication-type="journal"><name><surname>De Meyer</surname><given-names>M</given-names></name>, <name><surname>Robertson</surname><given-names>MP</given-names></name>, <name><surname>Peterson</surname><given-names>AT</given-names></name>, <name><surname>Mansell</surname><given-names>MW</given-names></name>. <article-title>Ecological niches and potential geographical distributions of Mediterranean fruit fly (<italic>Ceratitis capitata</italic>) and Natal fruit fly (<italic>Ceratitis rosa</italic>)</article-title>. <source>J Biogeogr</source>. <year>2008</year>;<volume>35</volume>:<fpage>270</fpage>–<lpage>81</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1365-2699.2007.01769.x">10.1111/j.1365-2699.2007.01769.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref086"><label>86</label><mixed-citation publication-type="journal"><name><surname>Gutierrez</surname><given-names>AP</given-names></name>, <name><surname>Ponti</surname><given-names>L</given-names></name>, <name><surname>Cossu</surname><given-names>Q</given-names></name>. <article-title>Effects of climate warming on olive and olive fly (<italic>Bactrocera oleae</italic> (Gmelin)) in California and Italy</article-title>. <source>Clim Change</source>. <year>2009</year>;<volume>95</volume>(<issue>1–2</issue>):<fpage>195</fpage>–<lpage>217</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref087"><label>87</label><mixed-citation publication-type="journal"><name><surname>Brooks</surname><given-names>CP</given-names></name>, <name><surname>Ervin</surname><given-names>GN</given-names></name>, <name><surname>Varone</surname><given-names>L</given-names></name>, <name><surname>Logarzo</surname><given-names>GA</given-names></name>. <article-title>Native ecotypic variation and the role of host identity in the spread of an invasive herbivore, <italic>Cactoblastis cactorum</italic></article-title>. <source>Ecology</source>. <year>2012</year>;<volume>93</volume>(<issue>2</issue>):<fpage>402</fpage>–<lpage>10</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1890/11-0541.1">10.1890/11-0541.1</ext-link></comment><pub-id pub-id-type="pmid">22624321</pub-id></mixed-citation></ref><ref id="pone.0135209.ref088"><label>88</label><mixed-citation publication-type="journal"><name><surname>Yokoyama</surname><given-names>VY</given-names></name>, <name><surname>Rendón</surname><given-names>PA</given-names></name>, <name><surname>Sivinski</surname><given-names>J</given-names></name>. <article-title><italic>Psyttalia</italic> cf. <italic>concolor</italic> (Hymenoptera: Braconidae) for Biological Control of Olive Fruit Fly (Diptera: Tephritidae) in California</article-title>. <source>Environ Entomol</source>. <year>2008</year>;<volume>37</volume>(<issue>3</issue>):<fpage>764</fpage>–<lpage>73</lpage>. <pub-id pub-id-type="pmid">18559183</pub-id></mixed-citation></ref><ref id="pone.0135209.ref089"><label>89</label><mixed-citation publication-type="journal"><name><surname>Daane</surname><given-names>K</given-names></name>, <name><surname>Johnson</surname><given-names>M</given-names></name>, <name><surname>Lynn-Patterson</surname><given-names>K</given-names></name>, <name><surname>Nadel</surname><given-names>H</given-names></name>, <name><surname>Opp</surname><given-names>S</given-names></name>, <name><surname>Stewart-Leslie</surname><given-names>J</given-names></name>, <etal>et al</etal><article-title>High temperature affects olive fruit fly populations in California's Central Valley</article-title>. <source>Calif Agric</source>. <year>2011</year>;<volume>65</volume>(<issue>1</issue>):<fpage>29</fpage>–<lpage>33</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref090"><label>90</label><mixed-citation publication-type="journal"><name><surname>Pearman</surname><given-names>PB</given-names></name>, <name><surname>Guisan</surname><given-names>A</given-names></name>, <name><surname>Broennimann</surname><given-names>O</given-names></name>, <name><surname>Randin</surname><given-names>CF</given-names></name>. <article-title>Niche dynamics in space and time</article-title>. <source>Trends Ecol Evol</source>. <year>2008</year>;<volume>23</volume>(<issue>3</issue>):<fpage>149</fpage>–<lpage>58</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.tree.2007.11.005">10.1016/j.tree.2007.11.005</ext-link></comment><pub-id pub-id-type="pmid">18289716</pub-id></mixed-citation></ref><ref id="pone.0135209.ref091"><label>91</label><mixed-citation publication-type="journal"><name><surname>Thompson</surname><given-names>GD</given-names></name>, <name><surname>Robertson</surname><given-names>MP</given-names></name>, <name><surname>Webber</surname><given-names>BL</given-names></name>, <name><surname>Richardson</surname><given-names>DM</given-names></name>, <name><surname>Le Roux</surname><given-names>JJ</given-names></name>, <name><surname>Wilson</surname><given-names>JRU</given-names></name>. <article-title>Predicting the subspecific identity of invasive species using distribution models: <italic>Acacia saligna</italic> as an example</article-title>. <source>Divers Distrib</source>. <year>2011</year>;<volume>17</volume>(<issue>5</issue>):<fpage>1001</fpage>–<lpage>14</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1472-4642.2011.00820.x">10.1111/j.1472-4642.2011.00820.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref092"><label>92</label><mixed-citation publication-type="journal"><name><surname>Schulte</surname><given-names>U</given-names></name>, <name><surname>Hochkirch</surname><given-names>A</given-names></name>, <name><surname>Lötters</surname><given-names>S</given-names></name>, <name><surname>Rödder</surname><given-names>D</given-names></name>, <name><surname>Schweiger</surname><given-names>S</given-names></name>, <name><surname>Weimann</surname><given-names>T</given-names></name>, <etal>et al</etal><article-title>Cryptic niche conservatism among evolutionary lineages of an invasive lizard</article-title>. <source>Glob Ecol Biogeogr</source>. <year>2012</year>;<volume>21</volume>(<issue>2</issue>):<fpage>198</fpage>–<lpage>211</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1466-8238.2011.00665.x">10.1111/j.1466-8238.2011.00665.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref093"><label>93</label><mixed-citation publication-type="journal"><name><surname>Beaumont</surname><given-names>LJ</given-names></name>, <name><surname>Gallagher</surname><given-names>RV</given-names></name>, <name><surname>Leishman</surname><given-names>MR</given-names></name>, <name><surname>Hughes</surname><given-names>L</given-names></name>, <name><surname>Downey</surname><given-names>PO</given-names></name>, <name><surname>Wilson</surname><given-names>J</given-names></name>. <article-title>How can knowledge of the climate niche inform the weed risk assessment process? A case study of <italic>Chrysanthemoides monilifera</italic> in Australia</article-title>. <source>Divers Distrib</source>. <year>2014</year>;<volume>20</volume>(<issue>6</issue>):<fpage>613</fpage>–<lpage>25</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/ddi.12190">10.1111/ddi.12190</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref094"><label>94</label><mixed-citation publication-type="journal"><name><surname>Soberón</surname><given-names>J</given-names></name>, <name><surname>Peterson</surname><given-names>AT</given-names></name>. <article-title>Interpretation of models of fundamental ecological niches and species' distributional areas</article-title>. <source>Biodiversity Informatics</source>. <year>2005</year>;<volume>2</volume>:<fpage>1</fpage>–<lpage>10</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref095"><label>95</label><mixed-citation publication-type="journal"><name><surname>Larson</surname><given-names>ER</given-names></name>, <name><surname>Olden</surname><given-names>JD</given-names></name>. <article-title>Using avatar species to model the potential distribution of emerging invaders</article-title>. <source>Glob Ecol Biogeogr</source>. <year>2012</year>;<volume>21</volume>(<issue>11</issue>):<fpage>1114</fpage>–<lpage>25</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1466-8238.2012.00758.x">10.1111/j.1466-8238.2012.00758.x</ext-link></comment></mixed-citation></ref><ref id="pone.0135209.ref096"><label>96</label><mixed-citation publication-type="journal"><name><surname>Larson</surname><given-names>ER</given-names></name>, <name><surname>Gallagher</surname><given-names>RV</given-names></name>, <name><surname>Beaumont</surname><given-names>LJ</given-names></name>, <name><surname>Olden</surname><given-names>JD</given-names></name>. <article-title>Generalized “avatar” niche shifts improve distribution models for invasive species</article-title>. <source>Divers Distrib</source>. <year>2014</year>;<volume>20</volume>(<issue>11</issue>):<fpage>1296</fpage>–<lpage>306</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref097"><label>97</label><mixed-citation publication-type="journal"><name><surname>Michalak</surname><given-names>P</given-names></name>, <name><surname>Minkov</surname><given-names>I</given-names></name>, <name><surname>Helin</surname><given-names>A</given-names></name>, <name><surname>Lerman</surname><given-names>DN</given-names></name>, <name><surname>Bettencourt</surname><given-names>BR</given-names></name>, <name><surname>Feder</surname><given-names>ME</given-names></name>, <etal>et al</etal><article-title>Genetic evidence for adaptation-driven incipient speciation of <italic>Drosophila melanogaster</italic> along a microclimatic contrast in "Evolution Canyon", Israel</article-title>. <source>Proc Natl Acad Sci U S A</source>. <year>2001</year>;<volume>98</volume>(<issue>23</issue>):<fpage>13195</fpage>–<lpage>200</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1073/pnas.231478298">10.1073/pnas.231478298</ext-link></comment><pub-id pub-id-type="pmid">11687637</pub-id></mixed-citation></ref><ref id="pone.0135209.ref098"><label>98</label><mixed-citation publication-type="journal"><name><surname>Rehfeldt</surname><given-names>GE</given-names></name>, <name><surname>Ying</surname><given-names>CC</given-names></name>, <name><surname>Spittlehouse</surname><given-names>DL</given-names></name>, <name><surname>Hamilton</surname><given-names>DA</given-names><suffix>Jr</suffix></name>. <article-title>Genetic responses to climate in <italic>Pinus contorta</italic>: niche breadth, climate change, and reforestation</article-title>. <source>Ecol Monogr</source>. <year>1999</year>;<volume>69</volume>(<issue>3</issue>):<fpage>375</fpage>–<lpage>407</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref099"><label>99</label><mixed-citation publication-type="journal"><name><surname>Keller</surname><given-names>RP</given-names></name>, <name><surname>Lodge</surname><given-names>DM</given-names></name>, <name><surname>Finnoff</surname><given-names>DC</given-names></name>. <article-title>Risk assessment for invasive species produces net bioeconomic benefits</article-title>. <source>Proc Natl Acad Sci U S A</source>. <year>2007</year>;<volume>104</volume>(<issue>1</issue>):<fpage>203</fpage>–<lpage>7</lpage>. <pub-id pub-id-type="pmid">17190819</pub-id></mixed-citation></ref><ref id="pone.0135209.ref100"><label>100</label><mixed-citation publication-type="journal"><name><surname>Hebert</surname><given-names>PD</given-names></name>, <name><surname>Cywinska</surname><given-names>A</given-names></name>, <name><surname>Ball</surname><given-names>SL</given-names></name>. <article-title>Biological identifications through DNA barcodes</article-title>. <source>Proc R Soc Lond B Biol Sci</source>. <year>2003</year>;<volume>270</volume>(<issue>1512</issue>):<fpage>313</fpage>–<lpage>21</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref101"><label>101</label><mixed-citation publication-type="journal"><name><surname>Armstrong</surname><given-names>K</given-names></name>, <name><surname>Ball</surname><given-names>S</given-names></name>. <article-title>DNA barcodes for biosecurity: invasive species identification</article-title>. <source>Proc R Soc Lond B Biol Sci</source>. <year>2005</year>;<volume>360</volume>(<issue>1462</issue>):<fpage>1813</fpage>–<lpage>23</lpage>.</mixed-citation></ref><ref id="pone.0135209.ref102"><label>102</label><mixed-citation publication-type="journal"><name><surname>Coeur d’acier</surname><given-names>A</given-names></name>, <name><surname>Cruaud</surname><given-names>A</given-names></name>, <name><surname>Artige</surname><given-names>E</given-names></name>, <name><surname>Genson</surname><given-names>G</given-names></name>, <name><surname>Clamens</surname><given-names>A-L</given-names></name>, <name><surname>Pierre</surname><given-names>E</given-names></name>, <etal>et al</etal><article-title>DNA barcoding and the associated PhylAphidB@se website for the identification of European aphids (Insecta: Hemiptera: Aphididae)</article-title>. <source>PLoS ONE</source>. <year>2014</year>;<volume>9</volume>(<issue>6</issue>):<fpage>e97620</fpage><comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1371/journal.pone.0097620">10.1371/journal.pone.0097620</ext-link></comment><pub-id pub-id-type="pmid">24896814</pub-id></mixed-citation></ref><ref id="pone.0135209.ref103"><label>103</label><mixed-citation publication-type="journal"><name><surname>Van Houdt</surname><given-names>JKJ</given-names></name>, <name><surname>Breman</surname><given-names>FC</given-names></name>, <name><surname>Virgilio</surname><given-names>M</given-names></name>, <name><surname>De Meyer</surname><given-names>M</given-names></name>. <article-title>Recovering full DNA barcodes from natural history collections of Tephritid fruitflies (Tephritidae, Diptera) using mini barcodes</article-title>. <source>Mol Ecol Resour</source>. <year>2010</year>;<volume>10</volume>(<issue>3</issue>):<fpage>459</fpage>–<lpage>65</lpage>. <comment>doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1111/j.1755-0998.2009.02800.x">10.1111/j.1755-0998.2009.02800.x</ext-link></comment><pub-id pub-id-type="pmid">21565045</pub-id></mixed-citation></ref></ref-list></back></pmc></record>Pour manipuler ce document sous Unix (Dilib)
EXPLOR_STEP=$WICRI_ROOT/Wicri/Bois/explor/OrangerV1/Data/Pmc/Corpus
HfdSelect -h $EXPLOR_STEP/biblio.hfd -nk 000177 | SxmlIndent | more
Ou
HfdSelect -h $EXPLOR_AREA/Data/Pmc/Corpus/biblio.hfd -nk 000177 | SxmlIndent | more
Pour mettre un lien sur cette page dans le réseau Wicri
{{Explor lien
|wiki= Wicri/Bois
|area= OrangerV1
|flux= Pmc
|étape= Corpus
|type= RBID
|clé= PMC:4537207
|texte= Assessing the Risk of Invasion by Tephritid Fruit Flies: Intraspecific Divergence Matters
}}
Pour générer des pages wiki
HfdIndexSelect -h $EXPLOR_AREA/Data/Pmc/Corpus/RBID.i -Sk "pubmed:26274582" \
| HfdSelect -Kh $EXPLOR_AREA/Data/Pmc/Corpus/biblio.hfd \
| NlmPubMed2Wicri -a OrangerV1
| This area was generated with Dilib version V0.6.25. |  |