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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">The Population Structure of <italic>Borrelia lusitaniae</italic> Is Reflected by a Population Division of Its <italic>Ixodes</italic> Vector</title><author><name sortKey="Norte, Ana Claudia" sort="Norte, Ana Claudia" uniqKey="Norte A" first="Ana Cláudia" last="Norte">Ana Cláudia Norte</name><affiliation><nlm:aff id="af1-microorganisms-09-00933">MARE-Marine and Environmental Sciences Centre, Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal;<email>acgnorte@ci.uc.pt</email></nlm:aff></affiliation><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation></author><author><name sortKey="Boyer, Pierre H" sort="Boyer, Pierre H" uniqKey="Boyer P" first="Pierre H." last="Boyer">Pierre H. Boyer</name><affiliation><nlm:aff id="af3-microorganisms-09-00933">CHRU Strasbourg, UR7290 Lyme Borreliosis Group, ITI InnoVec, Fédération de Médecine Translationnelle de Strasbourg, Institut de Bactériologie, University of Strasbourg, 3 rue Koeberlé, 67000 Strasbourg, France;<email>pierreboyer@unistra.fr</email></nlm:aff></affiliation></author><author><name sortKey="Castillo Ramirez, Santiago" sort="Castillo Ramirez, Santiago" uniqKey="Castillo Ramirez S" first="Santiago" last="Castillo-Ramirez">Santiago Castillo-Ramirez</name><affiliation><nlm:aff id="af4-microorganisms-09-00933">Programa de Genómica Evolutiva, Centro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Apartado Postal 565-A, Cuernavaca, CP 62210, Mexico;<email>iago@ccg.unam.mx</email></nlm:aff></affiliation></author><author><name sortKey="Chvosta, Michal" sort="Chvosta, Michal" uniqKey="Chvosta M" first="Michal" last="Chvostá">Michal Chvostá</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation></author><author><name sortKey="Brahami, Mohand O" sort="Brahami, Mohand O" uniqKey="Brahami M" first="Mohand O." last="Brahami">Mohand O. Brahami</name><affiliation><nlm:aff id="af6-microorganisms-09-00933">Laboratory of Ecology and Biology of Terrestrial Ecosystems, Faculty Biological and Agronomic Sciences, University Mouloud Mammeri, 15000 Tizi-Ouzou, Algeria;<email>brahamimohand@yahoo.fr</email></nlm:aff></affiliation></author><author><name sortKey="Rollins, Robert E" sort="Rollins, Robert E" uniqKey="Rollins R" first="Robert E." last="Rollins">Robert E. Rollins</name><affiliation><nlm:aff id="af7-microorganisms-09-00933">Division of Evolutionary Biology, LMU Munich, Faculty of Biology, Grosshaderner Strasse 2, 82152 Planegg-Martinsried, Germany;<email>rollins@biologie.uni-muenchen.de</email></nlm:aff></affiliation></author><author><name sortKey="Woudenberg, Tom" sort="Woudenberg, Tom" uniqKey="Woudenberg T" first="Tom" last="Woudenberg">Tom Woudenberg</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author><author><name sortKey="Didyk, Yuliya M" sort="Didyk, Yuliya M" uniqKey="Didyk Y" first="Yuliya M." last="Didyk">Yuliya M. Didyk</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation><affiliation><nlm:aff id="af9-microorganisms-09-00933">Department of Acarology, I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Khmelnytskogo 15, 01030 Kyiv, Ukraine</nlm:aff></affiliation></author><author><name sortKey="Derdakova, Marketa" sort="Derdakova, Marketa" uniqKey="Derdakova M" first="Marketa" last="Derdakova">Marketa Derdakova</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation></author><author><name sortKey="Nuncio, Maria Sofia" sort="Nuncio, Maria Sofia" uniqKey="Nuncio M" first="Maria Sofia" last="Núncio">Maria Sofia Núncio</name><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation><affiliation><nlm:aff id="af10-microorganisms-09-00933">Environmental Health Institute, Medicine Faculty, University of Lisbon, 1649-026 Lisbon, Portugal</nlm:aff></affiliation></author><author><name sortKey="De Carvalho, Isabel Lopes" sort="De Carvalho, Isabel Lopes" uniqKey="De Carvalho I" first="Isabel Lopes" last="De Carvalho">Isabel Lopes De Carvalho</name><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation><affiliation><nlm:aff id="af10-microorganisms-09-00933">Environmental Health Institute, Medicine Faculty, University of Lisbon, 1649-026 Lisbon, Portugal</nlm:aff></affiliation></author><author><name sortKey="Margos, Gabriele" sort="Margos, Gabriele" uniqKey="Margos G" first="Gabriele" last="Margos">Gabriele Margos</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author><author><name sortKey="Fingerle, Volker" sort="Fingerle, Volker" uniqKey="Fingerle V" first="Volker" last="Fingerle">Volker Fingerle</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">33925391</idno><idno type="pmc">8145215</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8145215</idno><idno type="RBID">PMC:8145215</idno><idno type="doi">10.3390/microorganisms9050933</idno><date when="2021">2021</date><idno type="wicri:Area/Pmc/Corpus">000065</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000065</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">The Population Structure of <italic>Borrelia lusitaniae</italic> Is Reflected by a Population Division of Its <italic>Ixodes</italic> Vector</title><author><name sortKey="Norte, Ana Claudia" sort="Norte, Ana Claudia" uniqKey="Norte A" first="Ana Cláudia" last="Norte">Ana Cláudia Norte</name><affiliation><nlm:aff id="af1-microorganisms-09-00933">MARE-Marine and Environmental Sciences Centre, Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal;<email>acgnorte@ci.uc.pt</email></nlm:aff></affiliation><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation></author><author><name sortKey="Boyer, Pierre H" sort="Boyer, Pierre H" uniqKey="Boyer P" first="Pierre H." last="Boyer">Pierre H. Boyer</name><affiliation><nlm:aff id="af3-microorganisms-09-00933">CHRU Strasbourg, UR7290 Lyme Borreliosis Group, ITI InnoVec, Fédération de Médecine Translationnelle de Strasbourg, Institut de Bactériologie, University of Strasbourg, 3 rue Koeberlé, 67000 Strasbourg, France;<email>pierreboyer@unistra.fr</email></nlm:aff></affiliation></author><author><name sortKey="Castillo Ramirez, Santiago" sort="Castillo Ramirez, Santiago" uniqKey="Castillo Ramirez S" first="Santiago" last="Castillo-Ramirez">Santiago Castillo-Ramirez</name><affiliation><nlm:aff id="af4-microorganisms-09-00933">Programa de Genómica Evolutiva, Centro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Apartado Postal 565-A, Cuernavaca, CP 62210, Mexico;<email>iago@ccg.unam.mx</email></nlm:aff></affiliation></author><author><name sortKey="Chvosta, Michal" sort="Chvosta, Michal" uniqKey="Chvosta M" first="Michal" last="Chvostá">Michal Chvostá</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation></author><author><name sortKey="Brahami, Mohand O" sort="Brahami, Mohand O" uniqKey="Brahami M" first="Mohand O." last="Brahami">Mohand O. Brahami</name><affiliation><nlm:aff id="af6-microorganisms-09-00933">Laboratory of Ecology and Biology of Terrestrial Ecosystems, Faculty Biological and Agronomic Sciences, University Mouloud Mammeri, 15000 Tizi-Ouzou, Algeria;<email>brahamimohand@yahoo.fr</email></nlm:aff></affiliation></author><author><name sortKey="Rollins, Robert E" sort="Rollins, Robert E" uniqKey="Rollins R" first="Robert E." last="Rollins">Robert E. Rollins</name><affiliation><nlm:aff id="af7-microorganisms-09-00933">Division of Evolutionary Biology, LMU Munich, Faculty of Biology, Grosshaderner Strasse 2, 82152 Planegg-Martinsried, Germany;<email>rollins@biologie.uni-muenchen.de</email></nlm:aff></affiliation></author><author><name sortKey="Woudenberg, Tom" sort="Woudenberg, Tom" uniqKey="Woudenberg T" first="Tom" last="Woudenberg">Tom Woudenberg</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author><author><name sortKey="Didyk, Yuliya M" sort="Didyk, Yuliya M" uniqKey="Didyk Y" first="Yuliya M." last="Didyk">Yuliya M. Didyk</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation><affiliation><nlm:aff id="af9-microorganisms-09-00933">Department of Acarology, I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Khmelnytskogo 15, 01030 Kyiv, Ukraine</nlm:aff></affiliation></author><author><name sortKey="Derdakova, Marketa" sort="Derdakova, Marketa" uniqKey="Derdakova M" first="Marketa" last="Derdakova">Marketa Derdakova</name><affiliation><nlm:aff id="af5-microorganisms-09-00933">Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</nlm:aff></affiliation></author><author><name sortKey="Nuncio, Maria Sofia" sort="Nuncio, Maria Sofia" uniqKey="Nuncio M" first="Maria Sofia" last="Núncio">Maria Sofia Núncio</name><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation><affiliation><nlm:aff id="af10-microorganisms-09-00933">Environmental Health Institute, Medicine Faculty, University of Lisbon, 1649-026 Lisbon, Portugal</nlm:aff></affiliation></author><author><name sortKey="De Carvalho, Isabel Lopes" sort="De Carvalho, Isabel Lopes" uniqKey="De Carvalho I" first="Isabel Lopes" last="De Carvalho">Isabel Lopes De Carvalho</name><affiliation><nlm:aff id="af2-microorganisms-09-00933">Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</nlm:aff></affiliation><affiliation><nlm:aff id="af10-microorganisms-09-00933">Environmental Health Institute, Medicine Faculty, University of Lisbon, 1649-026 Lisbon, Portugal</nlm:aff></affiliation></author><author><name sortKey="Margos, Gabriele" sort="Margos, Gabriele" uniqKey="Margos G" first="Gabriele" last="Margos">Gabriele Margos</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author><author><name sortKey="Fingerle, Volker" sort="Fingerle, Volker" uniqKey="Fingerle V" first="Volker" last="Fingerle">Volker Fingerle</name><affiliation><nlm:aff id="af8-microorganisms-09-00933">National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</nlm:aff></affiliation></author></analytic><series><title level="j">Microorganisms</title><idno type="eISSN">2076-2607</idno><imprint><date when="2021">2021</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Populations of vector-borne pathogens are shaped by the distribution and movement of vector and reservoir hosts. To study what impact host and vector association have on tick-borne pathogens, we investigated the population structure of <italic>Borrelia lusitaniae</italic> using multilocus sequence typing (MLST). Novel sequences were acquired from questing ticks collected in multiple North African and European locations and were supplemented by publicly available sequences at the Borrelia Pubmlst database (accessed on 11 February 2020). Population structure of <italic>B. lusitaniae</italic> was inferred using clustering and network analyses. Maximum likelihood phylogenies for two molecular tick markers (the mitochondrial 16S rRNA locus and a nuclear locus, Tick-receptor of outer surface protein A, <italic>trospA</italic>) were used to confirm the morphological species identification of collected ticks. Our results confirmed that <italic>B. lusitaniae</italic> does indeed form two distinguishable populations: one containing mostly European samples and the other mostly Portuguese and North African samples. Of interest, Portuguese samples clustered largely based on being from north (European) or south (North African) of the river Targus. As two different <italic>Ixodes</italic> species (i.e., <italic>I. ricinus</italic> and <italic>I. inopinatus</italic>) may vector <italic>Borrelia</italic> in these regions, reference samples were included for <italic>I. inopinatus</italic> but did not form monophyletic clades in either tree, suggesting some misidentification. Even so, the <italic>trospA</italic> phylogeny showed a monophyletic clade containing tick samples from Northern Africa and Portugal south of the river Tagus suggesting a population division in <italic>Ixodes</italic> on this locus. The pattern mirrored the clustering of <italic>B. lusitaniae</italic> samples, suggesting a potential co-evolution between tick and <italic>Borrelia</italic> populations that deserve further investigation.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Kurtenbach, K" uniqKey="Kurtenbach K">K. Kurtenbach</name></author><author><name sortKey="Hoen, A G" uniqKey="Hoen A">A.G. Hoen</name></author><author><name sortKey="Bent, S J" uniqKey="Bent S">S.J. Bent</name></author><author><name sortKey="Vollmer, S A" uniqKey="Vollmer S">S.A. Vollmer</name></author><author><name sortKey="Ogden, N H" uniqKey="Ogden N">N.H. Ogden</name></author><author><name sortKey="Margos, G" uniqKey="Margos G">G. Margos</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Margos, G" uniqKey="Margos G">G. Margos</name></author><author><name sortKey="Vollmer, S A" uniqKey="Vollmer S">S.A. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Microorganisms</journal-id><journal-id journal-id-type="iso-abbrev">Microorganisms</journal-id><journal-id journal-id-type="publisher-id">microorganisms</journal-id><journal-title-group><journal-title>Microorganisms</journal-title></journal-title-group><issn pub-type="epub">2076-2607</issn><publisher><publisher-name>MDPI</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">33925391</article-id><article-id pub-id-type="pmc">8145215</article-id><article-id pub-id-type="doi">10.3390/microorganisms9050933</article-id><article-id pub-id-type="publisher-id">microorganisms-09-00933</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>The Population Structure of <italic>Borrelia lusitaniae</italic> Is Reflected by a Population Division of Its <italic>Ixodes</italic> Vector</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Norte</surname><given-names>Ana Cláudia</given-names></name><xref ref-type="aff" rid="af1-microorganisms-09-00933">1</xref><xref ref-type="aff" rid="af2-microorganisms-09-00933">2</xref></contrib><contrib contrib-type="author"><name><surname>Boyer</surname><given-names>Pierre H.</given-names></name><xref ref-type="aff" rid="af3-microorganisms-09-00933">3</xref></contrib><contrib contrib-type="author"><name><surname>Castillo-Ramirez</surname><given-names>Santiago</given-names></name><xref ref-type="aff" rid="af4-microorganisms-09-00933">4</xref></contrib><contrib contrib-type="author"><name><surname>Chvostáč</surname><given-names>Michal</given-names></name><xref ref-type="aff" rid="af5-microorganisms-09-00933">5</xref></contrib><contrib contrib-type="author"><name><surname>Brahami</surname><given-names>Mohand O.</given-names></name><xref ref-type="aff" rid="af6-microorganisms-09-00933">6</xref></contrib><contrib contrib-type="author"><name><surname>Rollins</surname><given-names>Robert E.</given-names></name><xref ref-type="aff" rid="af7-microorganisms-09-00933">7</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-5675-7470</contrib-id><name><surname>Woudenberg</surname><given-names>Tom</given-names></name><xref ref-type="aff" rid="af8-microorganisms-09-00933">8</xref></contrib><contrib contrib-type="author"><name><surname>Didyk</surname><given-names>Yuliya M.</given-names></name><xref ref-type="aff" rid="af5-microorganisms-09-00933">5</xref><xref ref-type="aff" rid="af9-microorganisms-09-00933">9</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0003-2669-6969</contrib-id><name><surname>Derdakova</surname><given-names>Marketa</given-names></name><xref ref-type="aff" rid="af5-microorganisms-09-00933">5</xref></contrib><contrib contrib-type="author"><name><surname>Núncio</surname><given-names>Maria Sofia</given-names></name><xref ref-type="aff" rid="af2-microorganisms-09-00933">2</xref><xref ref-type="aff" rid="af10-microorganisms-09-00933">10</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-4693-3952</contrib-id><name><surname>de Carvalho</surname><given-names>Isabel Lopes</given-names></name><xref ref-type="aff" rid="af2-microorganisms-09-00933">2</xref><xref ref-type="aff" rid="af10-microorganisms-09-00933">10</xref></contrib><contrib contrib-type="author"><name><surname>Margos</surname><given-names>Gabriele</given-names></name><xref ref-type="aff" rid="af8-microorganisms-09-00933">8</xref><xref rid="c1-microorganisms-09-00933" ref-type="corresp">*</xref></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-3835-5646</contrib-id><name><surname>Fingerle</surname><given-names>Volker</given-names></name><xref ref-type="aff" rid="af8-microorganisms-09-00933">8</xref></contrib></contrib-group><contrib-group><contrib contrib-type="editor"><name><surname>Stefanoff</surname><given-names>Paweł</given-names></name><role>Academic Editor</role></contrib><contrib contrib-type="editor"><name><surname>Fingerle</surname><given-names>Volker</given-names></name><role>Academic Editor</role></contrib><contrib contrib-type="editor"><name><surname>Henningsson</surname><given-names>Anna J</given-names></name><role>Academic Editor</role></contrib><contrib contrib-type="editor"><name><surname>Markowicz</surname><given-names>Mateusz</given-names></name><role>Academic Editor</role></contrib></contrib-group><aff id="af1-microorganisms-09-00933"><label>1</label>MARE-Marine and Environmental Sciences Centre, Department of Life Sciences, University of Coimbra, 3000-456 Coimbra, Portugal;<email>acgnorte@ci.uc.pt</email></aff><aff id="af2-microorganisms-09-00933"><label>2</label>Centre for Vector and Infectious Diseases Research, National Institute of Health Doutor Ricardo Jorge, Águas de Moura, 2965-575 Setúbal, Portugal;<email>sofia.nuncio@insa.min-saude.pt</email> (M.S.N.);<email>isabel.carvalho@insa.min-saude.pt</email> (I.L.d.C.)</aff><aff id="af3-microorganisms-09-00933"><label>3</label>CHRU Strasbourg, UR7290 Lyme Borreliosis Group, ITI InnoVec, Fédération de Médecine Translationnelle de Strasbourg, Institut de Bactériologie, University of Strasbourg, 3 rue Koeberlé, 67000 Strasbourg, France;<email>pierreboyer@unistra.fr</email></aff><aff id="af4-microorganisms-09-00933"><label>4</label>Programa de Genómica Evolutiva, Centro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Apartado Postal 565-A, Cuernavaca, CP 62210, Mexico;<email>iago@ccg.unam.mx</email></aff><aff id="af5-microorganisms-09-00933"><label>5</label>Institute of Zoology, Slovak Academy of Sciences, 84506 Bratislava, Slovakia;<email>michalchvostac@gmail.com</email> (M.C.);<email>yu.m.didyk@gmail.com</email> (Y.M.D.);<email>marketa.derdakova@gmail.com</email> (M.D.)</aff><aff id="af6-microorganisms-09-00933"><label>6</label>Laboratory of Ecology and Biology of Terrestrial Ecosystems, Faculty Biological and Agronomic Sciences, University Mouloud Mammeri, 15000 Tizi-Ouzou, Algeria;<email>brahamimohand@yahoo.fr</email></aff><aff id="af7-microorganisms-09-00933"><label>7</label>Division of Evolutionary Biology, LMU Munich, Faculty of Biology, Grosshaderner Strasse 2, 82152 Planegg-Martinsried, Germany;<email>rollins@biologie.uni-muenchen.de</email></aff><aff id="af8-microorganisms-09-00933"><label>8</label>National Reference Center for Borrelia, Bavarian Health and Food Safety Authority, 85764 Oberschleissheim, Germany;<email>tom.woudenberg@gmail.com</email> (T.W.);<email>volker.fingerle@lgl.bayern.de</email> (V.F.)</aff><aff id="af9-microorganisms-09-00933"><label>9</label>Department of Acarology, I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Khmelnytskogo 15, 01030 Kyiv, Ukraine</aff><aff id="af10-microorganisms-09-00933"><label>10</label>Environmental Health Institute, Medicine Faculty, University of Lisbon, 1649-026 Lisbon, Portugal</aff><author-notes><corresp id="c1-microorganisms-09-00933"><label>*</label>Correspondence: <email>gabriele.margos@lgl.bayern.de</email> or <email>gmargos1@gmail.com</email>; Tel.: +49-9131-6808-5883</corresp></author-notes><pub-date pub-type="epub"><day>27</day><month>4</month><year>2021</year></pub-date><pub-date pub-type="collection"><month>5</month><year>2021</year></pub-date><volume>9</volume><issue>5</issue><elocation-id>933</elocation-id><history><date date-type="received"><day>16</day><month>3</month><year>2021</year></date><date date-type="accepted"><day>21</day><month>4</month><year>2021</year></date></history><permissions><copyright-statement>© 2021 by the authors.</copyright-statement><copyright-year>2021</copyright-year><license><ali:license_ref specific-use="textmining" content-type="ccbylicense">https://creativecommons.org/licenses/by/4.0/</ali:license_ref><license-p>Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license (<ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link>).</license-p></license></permissions><abstract><p>Populations of vector-borne pathogens are shaped by the distribution and movement of vector and reservoir hosts. To study what impact host and vector association have on tick-borne pathogens, we investigated the population structure of <italic>Borrelia lusitaniae</italic> using multilocus sequence typing (MLST). Novel sequences were acquired from questing ticks collected in multiple North African and European locations and were supplemented by publicly available sequences at the Borrelia Pubmlst database (accessed on 11 February 2020). Population structure of <italic>B. lusitaniae</italic> was inferred using clustering and network analyses. Maximum likelihood phylogenies for two molecular tick markers (the mitochondrial 16S rRNA locus and a nuclear locus, Tick-receptor of outer surface protein A, <italic>trospA</italic>) were used to confirm the morphological species identification of collected ticks. Our results confirmed that <italic>B. lusitaniae</italic> does indeed form two distinguishable populations: one containing mostly European samples and the other mostly Portuguese and North African samples. Of interest, Portuguese samples clustered largely based on being from north (European) or south (North African) of the river Targus. As two different <italic>Ixodes</italic> species (i.e., <italic>I. ricinus</italic> and <italic>I. inopinatus</italic>) may vector <italic>Borrelia</italic> in these regions, reference samples were included for <italic>I. inopinatus</italic> but did not form monophyletic clades in either tree, suggesting some misidentification. Even so, the <italic>trospA</italic> phylogeny showed a monophyletic clade containing tick samples from Northern Africa and Portugal south of the river Tagus suggesting a population division in <italic>Ixodes</italic> on this locus. The pattern mirrored the clustering of <italic>B. lusitaniae</italic> samples, suggesting a potential co-evolution between tick and <italic>Borrelia</italic> populations that deserve further investigation.</p></abstract><kwd-group><kwd>population structure</kwd><kwd>vector association</kwd><kwd><italic>Borrelia lusitaniae</italic></kwd><kwd>Ixodes</kwd><kwd>MLST</kwd></kwd-group></article-meta></front><body><sec sec-type="intro" id="sec1-microorganisms-09-00933"><title>1. Introduction</title><p>Populations of vector-borne infectious disease agents are shaped by migration of, and selection pressure exerted by, their hosts and vectors [<xref rid="B1-microorganisms-09-00933" ref-type="bibr">1</xref>,<xref rid="B2-microorganisms-09-00933" ref-type="bibr">2</xref>,<xref rid="B3-microorganisms-09-00933" ref-type="bibr">3</xref>,<xref rid="B4-microorganisms-09-00933" ref-type="bibr">4</xref>] and these processes leave genetic signatures in the genomes of the organisms. Investigation of the geographical distribution of genetic lineages and their diversity permit inference of evolutionary processes such as selection or migration that have shaped populations [<xref rid="B4-microorganisms-09-00933" ref-type="bibr">4</xref>]. Ticks range among the most important vectors for microbial pathogens [<xref rid="B5-microorganisms-09-00933" ref-type="bibr">5</xref>] but many pathogens they transmit are difficult to study [<xref rid="B6-microorganisms-09-00933" ref-type="bibr">6</xref>]. Disease agents that have been associated with <italic>Ixodes</italic> ticks include <italic>Borrelia</italic> spp., <italic>Babesia</italic> spp., <italic>Anaplasma</italic> spp., <italic>Rickettsia</italic> spp., and viruses such as tick-borne encephalitis virus or Powassan virus with species varying in their geographical distribution and pathogenic potential [<xref rid="B5-microorganisms-09-00933" ref-type="bibr">5</xref>], [<xref rid="B7-microorganisms-09-00933" ref-type="bibr">7</xref>]. The abundance of ticks, tick-borne disease agents and resulting host-parasite interactions have been reported to be influenced by environmental changes, among them changes in land use, increases in temperature due to climate change and others [<xref rid="B3-microorganisms-09-00933" ref-type="bibr">3</xref>,<xref rid="B8-microorganisms-09-00933" ref-type="bibr">8</xref>,<xref rid="B9-microorganisms-09-00933" ref-type="bibr">9</xref>]. In this study, as a proxy for tick-transmitted pathogens, we investigated a population of the most frequently found microbes transmitted by ticks in the temperate northern hemisphere, i.e., microbes that belong to the <italic>Borrelia burgdorferi</italic> sensu lato species complex.</p><p>The <italic>Borrelia burgdorferi</italic> sensu lato species complex comprises more than 20 spirochetal bacterial species that use ticks of the genus <italic>Ixodes</italic> as vector and small to medium sized vertebrate species as reservoir hosts. <italic>Borrelia</italic> species vary considerably in the scale of their host and vector specificity [<xref rid="B1-microorganisms-09-00933" ref-type="bibr">1</xref>,<xref rid="B6-microorganisms-09-00933" ref-type="bibr">6</xref>,<xref rid="B10-microorganisms-09-00933" ref-type="bibr">10</xref>,<xref rid="B11-microorganisms-09-00933" ref-type="bibr">11</xref>], and this is reflected in the geographic distribution of their lineages [<xref rid="B11-microorganisms-09-00933" ref-type="bibr">11</xref>,<xref rid="B12-microorganisms-09-00933" ref-type="bibr">12</xref>,<xref rid="B13-microorganisms-09-00933" ref-type="bibr">13</xref>]. Furthermore, host- and vector associations have been identified as drivers for diversification and speciation in <italic>Borrelia</italic> [<xref rid="B6-microorganisms-09-00933" ref-type="bibr">6</xref>,<xref rid="B11-microorganisms-09-00933" ref-type="bibr">11</xref>,<xref rid="B14-microorganisms-09-00933" ref-type="bibr">14</xref>,<xref rid="B15-microorganisms-09-00933" ref-type="bibr">15</xref>]. For example, <italic>B. burgdorferi</italic> sensu stricto (s.s.) Johnson et al. 1984, the main cause of human Lyme borreliosis in North America, is considered a generalist species because it can have different groups of vertebrate species such as rodents or birds as reservoir hosts and several <italic>Ixodes</italic> species as vectors [<xref rid="B16-microorganisms-09-00933" ref-type="bibr">16</xref>,<xref rid="B17-microorganisms-09-00933" ref-type="bibr">17</xref>]. The distribution range in North America includes the Northeast and Midwest of the USA [<xref rid="B18-microorganisms-09-00933" ref-type="bibr">18</xref>] ranging now into Canada [<xref rid="B19-microorganisms-09-00933" ref-type="bibr">19</xref>,<xref rid="B20-microorganisms-09-00933" ref-type="bibr">20</xref>,<xref rid="B21-microorganisms-09-00933" ref-type="bibr">21</xref>] and Western coastal regions [<xref rid="B22-microorganisms-09-00933" ref-type="bibr">22</xref>,<xref rid="B23-microorganisms-09-00933" ref-type="bibr">23</xref>]. Phylogenies based on multilocus sequence typing (MLST) or genomic SNPs have revealed a complex population structure, but strains do not cluster according to geography [<xref rid="B24-microorganisms-09-00933" ref-type="bibr">24</xref>,<xref rid="B25-microorganisms-09-00933" ref-type="bibr">25</xref>,<xref rid="B26-microorganisms-09-00933" ref-type="bibr">26</xref>,<xref rid="B27-microorganisms-09-00933" ref-type="bibr">27</xref>]. Species using only avian reservoirs, such as <italic>Borrelia garinii</italic> Baranton et al. 1982 or <italic>Borrelia valaisiana</italic> Wang et al. 1997, showed very little population structure likely owing to the migration and dispersal pattern of their avian reservoir hosts [<xref rid="B12-microorganisms-09-00933" ref-type="bibr">12</xref>,<xref rid="B13-microorganisms-09-00933" ref-type="bibr">13</xref>,<xref rid="B28-microorganisms-09-00933" ref-type="bibr">28</xref>,<xref rid="B29-microorganisms-09-00933" ref-type="bibr">29</xref>,<xref rid="B30-microorganisms-09-00933" ref-type="bibr">30</xref>,<xref rid="B31-microorganisms-09-00933" ref-type="bibr">31</xref>,<xref rid="B32-microorganisms-09-00933" ref-type="bibr">32</xref>]. In comparison, species that use hosts with smaller migration ranges such as rodents or lizards show more pronounced population structure, supposedly reflecting the migration range of their hosts [<xref rid="B12-microorganisms-09-00933" ref-type="bibr">12</xref>,<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>].</p><p><italic>Borrelia lusitaniae</italic> Le Fleche et al. 1997 was isolated for the first time in Portugal [<xref rid="B34-microorganisms-09-00933" ref-type="bibr">34</xref>] and described as a species in 1997 [<xref rid="B35-microorganisms-09-00933" ref-type="bibr">35</xref>]. The species has been associated with lizards as reservoir hosts [<xref rid="B36-microorganisms-09-00933" ref-type="bibr">36</xref>,<xref rid="B37-microorganisms-09-00933" ref-type="bibr">37</xref>,<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>] and <italic>Ixodes ricinus</italic> Linnaeus, 1758 as its main vector [<xref rid="B39-microorganisms-09-00933" ref-type="bibr">39</xref>,<xref rid="B40-microorganisms-09-00933" ref-type="bibr">40</xref>,<xref rid="B41-microorganisms-09-00933" ref-type="bibr">41</xref>]. Several species of the family Lacertidae, e.g., <italic>Psammodromus algirus</italic> Linnaeus, 1758 [<xref rid="B36-microorganisms-09-00933" ref-type="bibr">36</xref>,<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>,<xref rid="B42-microorganisms-09-00933" ref-type="bibr">42</xref>], <italic>Podarcis</italic> spp. Wagler, 1830 [<xref rid="B37-microorganisms-09-00933" ref-type="bibr">37</xref>,<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>,<xref rid="B43-microorganisms-09-00933" ref-type="bibr">43</xref>,<xref rid="B44-microorganisms-09-00933" ref-type="bibr">44</xref>], <italic>Teira dugesii</italic> Milne-Edwards 1829 [<xref rid="B45-microorganisms-09-00933" ref-type="bibr">45</xref>] as well as <italic>Lacerta</italic> spp. Linnaeus, 1758 [<xref rid="B43-microorganisms-09-00933" ref-type="bibr">43</xref>,<xref rid="B46-microorganisms-09-00933" ref-type="bibr">46</xref>] were incriminated as potential reservoir hosts for <italic>B. lusitaniae</italic>. Some studies reported isolation of <italic>B. lusitaniae</italic> from, or detection of <italic>B. lusitaniae</italic> by PCR in, larvae feeding on the rodent <italic>Apodemus sylvaticus</italic> Linnaeus, 1758 [<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>,<xref rid="B47-microorganisms-09-00933" ref-type="bibr">47</xref>]. Although this is not proof of reservoir competence, it cannot be immediately discounted that hosts other than lizards may serve as reservoirs for <italic>B. lusitaniae</italic>; further studies are needed to clarify this point. Initial reports of <italic>B. lusitaniae</italic> came from around the Mediterranean basin (Tunisia, Morocco, Portugal, Italy) [<xref rid="B34-microorganisms-09-00933" ref-type="bibr">34</xref>,<xref rid="B41-microorganisms-09-00933" ref-type="bibr">41</xref>,<xref rid="B48-microorganisms-09-00933" ref-type="bibr">48</xref>,<xref rid="B49-microorganisms-09-00933" ref-type="bibr">49</xref>,<xref rid="B50-microorganisms-09-00933" ref-type="bibr">50</xref>] but now it has also been reported in countries such as Slovakia, Poland, and Latvia [<xref rid="B46-microorganisms-09-00933" ref-type="bibr">46</xref>,<xref rid="B51-microorganisms-09-00933" ref-type="bibr">51</xref>,<xref rid="B52-microorganisms-09-00933" ref-type="bibr">52</xref>,<xref rid="B53-microorganisms-09-00933" ref-type="bibr">53</xref>]. Remarkably high <italic>Borrelia</italic> infection rates of adult ticks (at the time identified as <italic>I. ricinus</italic>) (75%, 41 out of 55) were found in Portugal near Grândola of which a majority were identified as <italic>B. lusitaniae</italic> [<xref rid="B54-microorganisms-09-00933" ref-type="bibr">54</xref>] suggesting favorable conditions in terms of reservoir hosts and vector ticks for this <italic>Borrelia</italic> species. First studies showed that <italic>B. lusitaniae</italic> induced disease in C3H/HeN mice, causing both nodular interstitial cystitis and vasculitis of the great vessels of the heart [<xref rid="B55-microorganisms-09-00933" ref-type="bibr">55</xref>]. From an epidemiological perspective, further investigations of <italic>B. lusitaniae</italic> populations will be important as to date two isolates have been obtained from human clinical cases [<xref rid="B56-microorganisms-09-00933" ref-type="bibr">56</xref>,<xref rid="B57-microorganisms-09-00933" ref-type="bibr">57</xref>,<xref rid="B58-microorganisms-09-00933" ref-type="bibr">58</xref>].</p><p>The population structure of <italic>B. lusitaniae</italic> was analyzed based on single genes (outer surface protein A; <italic>ospA</italic>) [<xref rid="B59-microorganisms-09-00933" ref-type="bibr">59</xref>] and multilocus sequence typing (MLST) [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>]. The study on <italic>ospA</italic> included samples from Italy, Germany, Morocco and Portugal. Phylogenetic analyses demonstrated that Portuguese tick isolates (PoTiB1, PoTiB2 and PoTiB3, all isolated from ticks from around Águas de Moura, south of the river Tagus) formed a clade together with North African isolates while isolates from Germany, Italy and a Portuguese human isolate (from Lisbon region) formed a second clade [<xref rid="B59-microorganisms-09-00933" ref-type="bibr">59</xref>]. The MLST study included samples from Portugal collected in Mafra (north of the river Tagus) and Grândola (south of the river Tagus). Phylogenetic analyses of MLST sequences showed that isolates from north of the Tagus river mostly clustered together while samples from Grândola formed a second cluster suggesting two populations in Portugal–one from the north of the Tagus river, the other one from south of the Tagus river [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>]. To confirm the population structure of <italic>B. lusitaniae</italic>, we analyzed, using MLST, samples from various European countries (Austria, Croatia, Latvia, Serbia, Slovakia, Ukraine and Portugal) and from North Africa (Algeria). The Portuguese samples included specimens from north (Mafra and Coimbra) and south (Santiago do Cacém) of the river Tagus and from Madeira island. In addition, we tested the hypothesis that the population structure of <italic>B. lusitaniae</italic> is shaped by association with different vector species. In 2014, a new <italic>Ixodes</italic> species was described and named <italic>Ixodes inopinatus</italic> Estrada-Peña, Nava et Petney 2014 [<xref rid="B60-microorganisms-09-00933" ref-type="bibr">60</xref>]. The authors describe that <italic>I. inopinatus</italic> can be found on lizards and suggest that it replaces <italic>I. ricinus</italic> in dry Mediterranean regions of Algeria, Morocco, Portugal, Spain and Tunisia. A previous study on the population structure of <italic>I. ricinus</italic> had shown that there was a divergent clade in North Africa [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]. The latter study used concatenated sequences of four mitochondrial and nuclear genes including the 16S rRNA locus and Tick-receptor of outer surface protein A (<italic>trospA</italic>). In the present study, we used these two loci to establish whether there are different populations of <italic>Ixodes</italic> in Portugal and whether the ticks collected in Algeria may be identified as <italic>I. inopinatus</italic>.</p></sec><sec id="sec2-microorganisms-09-00933"><title>2. Materials and Methods</title><sec id="sec2dot1-microorganisms-09-00933"><title>2.1. Tick Collection and Processing</title><p>Questing ticks were collected by standardized flagging method [<xref rid="B62-microorganisms-09-00933" ref-type="bibr">62</xref>]. Ticks from Algeria were removed from cattle, one specimen was collected in Portugal feeding on a <italic>P. algirus</italic> (PoTiB11/T3087), three specimens were collected in 2018 in Germany from great tits <italic>Parus major</italic> Linnaeus, 1758 (11-E12, 7-F5, 8-C12) [<xref rid="B63-microorganisms-09-00933" ref-type="bibr">63</xref>], and three specimens feeding on humans (13310PT18, 13360PT18 and 14401PT18). Collected ticks were morphologically determined into stages and species [<xref rid="B60-microorganisms-09-00933" ref-type="bibr">60</xref>,<xref rid="B64-microorganisms-09-00933" ref-type="bibr">64</xref>] and stored in 70% ethanol until further analyses, except for one sample that was inoculated in BSK medium (see <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S1</xref>).</p><p>In Slovakia, ticks were collected in years 2013–2017 in an ecotone between the forest and a small mountain stream on Martinské hole mountains (49.085464, 18.863042; 630–660 m ASL). The vegetation was composed mainly of beech, spruce, hazel and elderberry with herbal undergrowth.</p><p>In Croatia, ticks were collected in 2011 in Grabovac (44.971317, 15.640267; 420 m ASL). The study area consisted of hornbeam forest, shrubs and pastures.</p><p>In Ukraine, ticks were collected during 2015–2016 in urban green areas Kyiv city: (50.446639, 30.570000; 179 m ASL). The vegetation cover at the sampling sites consisted of deciduous forests, shrubs and inhomogeneous meadows.</p><p>In Algeria, ticks were collected during 2015–2016 from cattle, in the mountainous Akfadou forest. Three collection sites were investigated during the collection campaign, which are places where cattle rest and can drink: site A (36.6289361, 4.57211111; 1567 m ASL); site B (36.690625, 4.57448611; 1196 m ASL) and site C (36.6951028, 4.55724694; 1201 m ASL).</p><p>Questing ticks from Portugal were collected in Tapada de Mafra in 2003 and 2013, and in 2009 in the Madeira Archipelago, Parque Natural Peneda-Gerês, and in Santiago do Cacém. Details on vegetation and fauna are given in <xref ref-type="app" rid="app1-microorganisms-09-00933">Supplementary Material</xref>.</p><p>Countries where ticks were collected and <italic>Borrelia</italic> samples characterized are shown in <xref ref-type="fig" rid="microorganisms-09-00933-f001">Figure 1</xref>. Details on geographic origin, species and number of ticks included in the current analysis are given in <xref rid="microorganisms-09-00933-t001" ref-type="table">Table 1</xref> and <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S1</xref>.</p></sec><sec id="sec2dot2-microorganisms-09-00933"><title>2.2. DNA Extraction</title><p>For DNA isolation from ticks, different methods were used. In Slovakia and Ukraine, alkaline-hydrolysis was used as described [<xref rid="B65-microorganisms-09-00933" ref-type="bibr">65</xref>] on individual ticks. In Portugal, a DNAeasy kit (Qiagen, Hilden, Germany) was used. Ticks sampled in Algeria were individually extracted using the MagNA Pure 96 system using the MagNA Pure96 DNA and Viral NA Small Volume Kit (Roche Diagnostics, Meylan, France) according to the manufacturer’s instruction and for ticks from Germany the Qiagen BioSprint 96 DNA Blood kit (Qiagen) was used. DNA samples were stored frozen (at −20 °C and −80 °C, respectively), until further processing. In samples from Croatia, Slovakia, and Ukraine the presence of tick DNA was confirmed by amplification of a 620 bp fragment of the tick mitochondrial gene cytochrome B (Black & Roehrdanz 1998). Only PCR positive samples were further analyzed.</p></sec><sec id="sec2dot3-microorganisms-09-00933"><title>2.3. PCR Borrelia Genes</title><p><italic>Borrelia burgdorferi</italic> s.l. was detected through PCR amplification of a 222–255 bp fragment of the <italic>rrfA</italic>-<italic>rrlB</italic> intergenic spacer (IGS) in samples from Croatia, Slovakia, and Ukraine [<xref rid="B66-microorganisms-09-00933" ref-type="bibr">66</xref>] and Portugal [<xref rid="B67-microorganisms-09-00933" ref-type="bibr">67</xref>]. In Portuguese and Algerian samples <italic>B. burgdorferi</italic> s.l. was confirmed through amplification of the flagellin B (<italic>flaB</italic>) gene [<xref rid="B68-microorganisms-09-00933" ref-type="bibr">68</xref>,<xref rid="B69-microorganisms-09-00933" ref-type="bibr">69</xref>] (see <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S2</xref>).</p><p>MLST on <italic>Borrelia</italic> positive samples was done primarily using a nested or semi-nested PCR (HotStar Taq Master Mix Kit or HotStarTaq Plus DNA Polymerase kit, Qiagen, Hilden, Germany), with primers and PCR conditions as previously described [<xref rid="B70-microorganisms-09-00933" ref-type="bibr">70</xref>,<xref rid="B71-microorganisms-09-00933" ref-type="bibr">71</xref>] [see also <uri xlink:href="https://pubmlst.org/borrelia/">https://pubmlst.org/borrelia/</uri> (11 February 2020)]. As the <italic>uvrA</italic> gene did not amplify in all cases, fragments of the seven housekeeping genes <italic>clpA</italic>, <italic>clpX</italic>, <italic>nifS</italic>, <italic>pepX</italic>, <italic>pyrG</italic>, <italic>recG</italic>, and <italic>rplB</italic> were amplified and sequenced. As positive control DNA of <italic>B. japonica</italic> or <italic>B. turdi</italic> was used while double distilled water was used as negative control. <xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref> and <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S2</xref> give an overview of <italic>B. lusitaniae</italic> positive samples that were typed by MLST and for which at least seven MLST housekeeping genes gave good sequences without ambiguities. MLST sequences were submitted and are available via <uri>pubmlst.org/borrelia/</uri> (11 May 2020) at the University of Oxford.</p><p>Purification of PCR products was done using a NucleoSpin Gel and PCR Clean-up according to the manufacturer’s recommendation (MACHEREY-NAGEL, Düren, Germany). For Slovakian and Algerian samples Sanger sequencing of all the genes was performed by Eurofins Genomics (Eurofins Genomics Germany GmbH, Ebersberg, Germany and Basel, Switzerland).</p></sec><sec id="sec2dot4-microorganisms-09-00933"><title>2.4. PCR on Tick Genes</title><p>To confirm the morphological identification of ticks and to differentiate <italic>I. ricinus</italic> and <italic>I. inopinatus</italic>, two previously used loci, the mitochondrial 16S rRNA locus and the nuclear gene <italic>trospA</italic> were amplified [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>,<xref rid="B72-microorganisms-09-00933" ref-type="bibr">72</xref>] and sequenced. We included tick samples that were characterized for <italic>Borrelia</italic> or originated from the same regions as the <italic>Borrelia</italic> positive tick samples, but they were not necessarily identical. Three morphologically identified samples that matched the 16S rRNA of <italic>I. inopinatus</italic> were available, designated as such, and were included as controls (for details see <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S1</xref>). In tick samples from Algeria, Slovakia and Portugal, the 16S rRNA gene was amplified according to [<xref rid="B72-microorganisms-09-00933" ref-type="bibr">72</xref>] and the <italic>trospA</italic> locus was amplified using primers and conditions with slight modification according to [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]. The implemented modification was a touch-down protocol for the initial six rounds of amplification to minimize non-specific background; the annealing temperatures started at 61 °C and decreased 1 °C per cycle until a temperature of 56 °C was reached. Further amplification rounds were performed at 56 °C. Further details on PCR conditions are given in <xref ref-type="app" rid="app1-microorganisms-09-00933">supplementary information</xref>.</p></sec><sec id="sec2dot5-microorganisms-09-00933"><title>2.5. Bioinformatic Analysis Recombination and Network Analysis on MLST Genes of Borrelia Samples</title><p>Recombination signals were analyzed in the MLST sequence alignment and in both <italic>trospA</italic> and 16S rRNA sequences employing the PhiTest implemented in SplitTree4 [<xref rid="B73-microorganisms-09-00933" ref-type="bibr">73</xref>]. Because the MLST alignment had recombination signals NeighborNet was run instead of a regular phylogeny; we used SplitTree4 to construct the net and selected the uncorrected P distance and equal angle display.</p><p>To identify the potential recombination events in the MLST alignment we used Gubbins with default parameters [<xref rid="B74-microorganisms-09-00933" ref-type="bibr">74</xref>]. We also conduct individual recombination analysis via PhiPack [<xref rid="B75-microorganisms-09-00933" ref-type="bibr">75</xref>], following the method described for <italic>A. baumannii</italic> [<xref rid="B76-microorganisms-09-00933" ref-type="bibr">76</xref>], on each the fragments of the seven housekeeping genes employed for the MLST. Of note, out of the seven housekeeping genes in <italic>clpA</italic> we found signals of recombination (<italic>p</italic>-value = 3.66 × 10<sup>−2</sup>).</p></sec><sec id="sec2dot6-microorganisms-09-00933"><title>2.6. Hierarchical Population Structure Analysis (HPSA) of Borrelia</title><p>For a hierarchical population structure analysis using hierBAPS [<xref rid="B77-microorganisms-09-00933" ref-type="bibr">77</xref>], we used the alignment of concatenated MLST sequences. The analysis was run at four levels of clustering and the number of initial clusters was set to 20. We also computed the coefficient of differentiation via MEGA X [<xref rid="B78-microorganisms-09-00933" ref-type="bibr">78</xref>] for the two major groups found at the first level of HPSA clustering.</p></sec><sec id="sec2dot7-microorganisms-09-00933"><title>2.7. Hierarchical Clustering of MLST Genes of Borrelia Samples</title><p>To assess the hypothesis of having two genetic clades in Europe, we applied a hierarchical clustering algorithm [<xref rid="B79-microorganisms-09-00933" ref-type="bibr">79</xref>]. This algorithm aggregated the isolates towards one group using the allelic profile data on the seven genes (<italic>clpA</italic>, <italic>clpX</italic>, <italic>nifS</italic>, <italic>pepX</italic>, <italic>pyrG</italic>, <italic>recG</italic> and <italic>rplB</italic>). The hierarchical clustering algorithm treats every isolate as its own cluster and will then look for the closest related isolate based on a similarity measure (allelic profile) and fuses these into a cluster. This step is repeated until all isolates belong to a cluster which can be visualized in a dendrogram. The similarity measure was Gower’s similarity coefficient [<xref rid="B80-microorganisms-09-00933" ref-type="bibr">80</xref>]. This coefficient can range from 0 to 1, where a score of 1 indicates complete similarity. Identical isolates fuse directly in the dendrogram, whereas dissimilar isolates only fuse at a more upward branch. R (version 4.0.2) [<xref rid="B81-microorganisms-09-00933" ref-type="bibr">81</xref>] was used to map the data with package “rnaturalearth” and “ggplot2”. The dendrogram was build using the R package “dendextend” [<xref rid="B82-microorganisms-09-00933" ref-type="bibr">82</xref>].</p></sec><sec id="sec2dot8-microorganisms-09-00933"><title>2.8. GoeBURST Analysis of Borrelia</title><p>We used PHYLOVIZ 1.1 [<xref rid="B83-microorganisms-09-00933" ref-type="bibr">83</xref>] to conduct goeBURST analyses using the triple locus variant setting [<xref rid="B84-microorganisms-09-00933" ref-type="bibr">84</xref>] using ST numbers, allelic profiles and meta information. Previous data available for <italic>B. lusitaniae</italic> isolates were downloaded from the <italic>Borrelia</italic> MLST database [<uri xlink:href="www.pubmlst.org/borrelia/">www.pubmlst.org/borrelia/</uri> (11 February 2020)] and included in our network and goeBURST analyses.</p></sec><sec id="sec2dot9-microorganisms-09-00933"><title>2.9. Maximum Likelihood Phylogenies of Tick Samples</title><p>As 16S rRNA and <italic>trospA</italic> did not show signals of recombination regular phylogenies were constructed. For both loci, the best model for the phylogeny was selected via jModelTest [<xref rid="B85-microorganisms-09-00933" ref-type="bibr">85</xref>]. For each locus, a maximum likelihood (ML) phylogeny was run using PhyML [<xref rid="B86-microorganisms-09-00933" ref-type="bibr">86</xref>] setting the best model, which was HK+I for 16S and K80+I for <italic>trospA</italic>.</p></sec></sec><sec sec-type="results" id="sec3-microorganisms-09-00933"><title>3. Results</title><sec id="sec3dot1-microorganisms-09-00933"><title>3.1. Identification of Borrelia lusitaniae in Ticks</title><p>Ticks collected in Europe (Croatia, Portugal, Slovakia, Ukraine,) and North Africa (Algeria) that tested positive for <italic>B. burgdorferi</italic> in screening PCRs were further analyzed by MLST to identify <italic>B. lusitaniae</italic>. Details of <italic>B. lusitaniae</italic> samples (country of origin, region, MLST alleles and STs) included in the present study are given in <xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref> and <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S2</xref>. An overview of the geographical distribution can be found in <xref ref-type="fig" rid="microorganisms-09-00933-f001">Figure 1</xref>. </p></sec><sec id="sec3dot2-microorganisms-09-00933"><title>3.2. MLST, goeBURST and Phylogenetic Analyses of Borrelia Samples</title><sec id="sec3dot2dot1-microorganisms-09-00933"><title>3.2.1. Multilocus Sequence Typing (MLST)</title><p>For 15 <italic>B. lusitaniae</italic> positive samples at least seven MLST genes provided good sequences and these samples were included in the MLST analysis conducted here (<xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref>). These samples were complemented with 40 <italic>B. lusitaniae</italic> samples available via the <italic>Borrelia</italic> MLST database [(pubmlst.org/borrelia/ (11 February 2020)]. For several of the samples, <italic>uvrA</italic> did not yield a PCR product and, thus, we describe the isolates by seven genes, namely: <italic>clpA, clpX, nifS</italic>, <italic>pepX, pyrG, recG</italic> and <italic>rplB</italic>. Most samples of <italic>B. lusitaniae</italic> were from Portugal (<italic>n</italic> = 24), followed by Serbia (<italic>n</italic> = 15) and Algeria (<italic>n</italic> = 6) (<xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref>). Among the 55 isolates there are 19 differing alleles of <italic>clpA</italic>, 19 different alleles of gene <italic>clpX</italic>, nine different alleles of gene <italic>nifS</italic>, 12 alleles of <italic>pepX</italic>, 14 alleles of <italic>pyrG</italic>, 10 alleles of <italic>recG</italic> genes and 17 <italic>rplB</italic> alleles.</p></sec><sec id="sec3dot2dot2-microorganisms-09-00933"><title>3.2.2. Clustering Analysis of <italic>B. lusitaniae</italic></title><p>Based on previous results for <italic>B. lusitaniae</italic> [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>,<xref rid="B59-microorganisms-09-00933" ref-type="bibr">59</xref>], we investigated the hypothesis that isolates may fall into two groups, one cluster of isolates from Portugal and Algeria and one cluster of isolates from Central/Eastern Europe. We assessed this hypothesis by using hierarchical clustering, numbering the isolates 1–55 (<xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref>). In the dendogram in <xref ref-type="fig" rid="microorganisms-09-00933-f002">Figure 2</xref>A two different clusters are obvious (depicted in red and green branches). Within these groups identical isolates were identified: e.g., isolates 18, 8, 9, and 1, 2, 30 were identical.</p><p>The isolates (numbered 1–55) were plotted on a map (<xref ref-type="fig" rid="microorganisms-09-00933-f002">Figure 2</xref>B); numbers and color coding for cluster 1 (red) and cluster 2 (green) correspond in <xref ref-type="fig" rid="microorganisms-09-00933-f002">Figure 2</xref>A,B. Samples from Central/Eastern Europe all fell into cluster 1 while all samples from Algeria fell into cluster 2. Interestingly, isolates from Portugal were assigned to different clusters, a subset fell into cluster 1 and a subset into cluster 2 (inlet in <xref ref-type="fig" rid="microorganisms-09-00933-f002">Figure 2</xref>B). Six isolates from Mafra and Lisbon seem to be different from other Portuguese and North African isolates.</p></sec><sec id="sec3dot2dot3-microorganisms-09-00933"><title>3.2.3. goeBURST Analysis</title><p>Since the cluster analysis was in line with previous results suggesting a population division of <italic>B. lusitaniae</italic>, we used goeBURST to obtain an additional view of the population using a different method. The goeBURST diagram also indicated a population division (<xref ref-type="fig" rid="microorganisms-09-00933-f003">Figure 3</xref>). Although there was one link based on tiebreak rule 1 between ST7-007* from Algeria and ST7-004* from Mafra in Portugal, several connections (one link without choice of tiebreak rules and several lower level edges; see figure legend <xref ref-type="fig" rid="microorganisms-09-00933-f003">Figure 3</xref> for further information) were found between STs from Algeria and STs originating from south of the Tagus river in Portugal (Grândola and Águas de Moura), resulting in a clonal complex (CC) that included 17 STs (CC7-003). A bushy clonal complex (CC918) was also formed consisting of STs from Slovakia, Serbia, Austria, Latvia, Ukraine and one ST (ST7-005) from Mafra, Portugal.</p><p>When we used goeBURST on higher level settings (level 4, using four alleles to infer inter-isolate relationships; level 5, using 5 alleles and level 6, six alleles), the picture of the relationship within populations became more obvious (see <xref ref-type="app" rid="app1-microorganisms-09-00933">Supplementary Figures S1–S3</xref>). Most of the Portuguese isolates clustered with North African isolates, except for some isolates from Lisbon (human), Mafra and Madeira island which clustered with those from Austria, Croatia, Latvia, Serbia and Slovakia.</p></sec><sec id="sec3dot2dot4-microorganisms-09-00933"><title>3.2.4. Sequence Analysis</title><p>All analyses were conducted with concatenated sequences of seven MLST genes. As previously reported [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>] we found evidence for recombination (<italic>p</italic> = 0.00125) in the <italic>B. lusitaniae</italic> dataset. Therefore, a NeighborNet [<xref rid="B73-microorganisms-09-00933" ref-type="bibr">73</xref>] was constructed instead of a phylogeny. The net shows that there are two very well differentiated groups (see <xref ref-type="fig" rid="microorganisms-09-00933-f004">Figure 4</xref>A,B), which was confirmed by the first level of the HPSA (<xref ref-type="app" rid="app1-microorganisms-09-00933">Table S3</xref>). One group consisted of Portuguese and Algerian isolates (<xref ref-type="fig" rid="microorganisms-09-00933-f004">Figure 4</xref>A); the other group combined isolates from Eastern Europe, Austria and Portugal (<xref ref-type="fig" rid="microorganisms-09-00933-f004">Figure 4</xref>B). <xref ref-type="app" rid="app1-microorganisms-09-00933">Supplementary Figure S4</xref> provides the whole network, where two major groups are well differentiated. Notably, we did not find any recombination events between these two groups using Gubbins; further suggesting that these are well differentiated groups that may not exchange DNA. We also computed the coefficient of differentiation (N<sub>ST</sub>) using MEGA X; the value of the coefficient was 0.71 (SE 0.04), which implies considerable levels of population differentiation between the two groups.</p><p>The second level of clustering using HPSA found clearly defined sub-clusters (see <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S3</xref>) within the two major groups, which suggests further geographic structuring. For instance, in major group 2 (<xref ref-type="fig" rid="microorganisms-09-00933-f004">Figure 4</xref>B) there is a cluster of Serbian isolates (blue label) and there is also a tight cluster of three Portuguese isolates (green label).</p></sec></sec><sec id="sec3dot3-microorganisms-09-00933"><title>3.3. Analysis of Tick Samples</title><p>One hypothesis for the strong population division of <italic>B. lusitaniae</italic> was that this was due to vector association. To test this hypothesis, we investigated tick samples that originated from the same geographic region as the <italic>B. lusitaniae</italic> samples as well as from other regions. Ticks were identified to species status using morphological criteria [<xref rid="B60-microorganisms-09-00933" ref-type="bibr">60</xref>,<xref rid="B64-microorganisms-09-00933" ref-type="bibr">64</xref>,<xref rid="B87-microorganisms-09-00933" ref-type="bibr">87</xref>]. Sequence data for the mitochondrial 16S rRNA gene and a nuclear gene, <italic>trospA</italic>, were used for corroboration. These loci were chosen because they showed in previous studies that there was a population division of <italic>I. ricinus</italic> between Europe and North Africa [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]. Because a new <italic>Ixodes</italic> species, <italic>I. inopinatus</italic>, was described originally in the Mediterranean region [<xref rid="B60-microorganisms-09-00933" ref-type="bibr">60</xref>], samples from Germany that were identified as <italic>I. inopinatus</italic> morphologically and using molecular data (16S rRNA) were used as a control for <italic>I. inopinatus</italic>.</p><p>Given that neither of the loci, 16S rRNA and <italic>trospA</italic>, showed recombination signals (16S rRNA <italic>p</italic> = 0.82 and <italic>trospA p</italic> = 0.13), we employed ML phylogenies to depict the evolutionary relationships for each locus. All ticks tested in Slovakia were determined as <italic>I. ricinus</italic> by sequencing both 16S rRNA and <italic>trospA</italic> genes and comparing the sequences via BLASTn [<xref rid="B88-microorganisms-09-00933" ref-type="bibr">88</xref>] searches to sequences available in GenBank. In the 16S rRNA ML phylogeny and as far as <italic>I. ricinus</italic> samples are concerned, the Portuguese and North African samples were dispersed in several groups across the tree (<xref ref-type="fig" rid="microorganisms-09-00933-f005">Figure 5</xref>A, green color coding). Furthermore, the <italic>I. inopinatus</italic> samples (both from GenBank and our “reference samples”) grouped together with samples that were classified as <italic>I. ricinus</italic> morphologically. <xref ref-type="fig" rid="microorganisms-09-00933-f005">Figure 5</xref>B shows the phylogeny of <italic>trospA</italic>, which seems to agree with the MLST NeighborNet in <xref ref-type="fig" rid="microorganisms-09-00933-f004">Figure 4</xref>: Two clusters emerged in the <italic>trospA</italic> phylogeny according to the geographical distribution (whatever the tick species) as in the <italic>B. lusitaniae</italic> MLST NeighborNet. Interestingly, in the <italic>trospA</italic> phylogeny, with only one exception all the Portuguese samples cluster with the North African samples (<xref ref-type="fig" rid="microorganisms-09-00933-f005">Figure 5</xref>B, green color coding). The only exception being the Portuguese sample R1756PT13 collected in Mafra (north of the river Tagus), which clusters with other non-Portuguese samples from Europe (red color coding). Of note, the three <italic>I. inopinatus</italic> control (“reference”) samples from Germany (blue color coding) did not form a monophyletic group, as they were located on different branches far apart in the tree. Taken together these results suggest that unlike the 16S rRNA-based phylogeny, the <italic>trospA</italic> phylogeny is consistent with the MLST NeighborNet in that samples from North Africa and Portugal form a separate clade. Notably, neither in the <italic>trospA</italic> phylogeny nor in the 16S RNA phylogeny did the <italic>I. inopinatus</italic> samples form a monophyletic group.</p></sec></sec><sec sec-type="discussion" id="sec4-microorganisms-09-00933"><title>4. Discussion</title><sec id="sec4dot1-microorganisms-09-00933"><title>4.1. Population Structure of Borrelia lusitaniae</title><p>In this study, we have further evaluated the population structure of <italic>B. lusitaniae</italic> including additional samples from Croatia, Portugal, Slovakia, Ukraine as well as from Algeria using MLST. Consistent with previously reported data [<xref rid="B59-microorganisms-09-00933" ref-type="bibr">59</xref>], our data clearly show a subdivision between North African isolates and isolates from Latvia, Serbia, Slovakia, and Austria. The population division in Portugal [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>] was also corroborated. In the cluster analysis isolates from north of the Tagus river clustered together (cluster 1) and only two isolates fell into cluster 2 together with isolates from south of the Tagus river which is consistent with previously reported data by [<xref rid="B33-microorganisms-09-00933" ref-type="bibr">33</xref>]. Southern Portuguese isolates showed a higher degree of genetic relatedness to isolates from Algeria using goeBURST than to isolates from Mafra. However, two Portuguese STs from south of the Tagus river clustered as singletons in the goeBURST diagram with TLV settings. When higher level settings were considered, two singletons from south of the Tagus river (ST67, ST64) clustered with Algerian samples while singletons from Mafra (ST69, ST7-006) and Madeira island (ST925) clustered with samples from Europe (<xref ref-type="app" rid="app1-microorganisms-09-00933">Supplementary Information Figures S1–S3</xref>). Thus, of the two major clonal complexes, one (CC918) consisted of STs from Austria, Croatia, Slovakia, Serbia, Latvia, Ukraine and one Portuguese ST from Mafra, the other (CC7-007*) consisted primarily of STs from Algeria, southern Portugal and two isolates from Mafra and Coimbra. The division was strongly supported through the sequenced-based network analysis where most STs from Mafra (except one) clustered with STs from other European countries and all STs from Grândola and Algeria fell into one cluster together with PoTiB2 and PoTiB3 from Águas de Moura (south of river Tagus), corroborating data published using <italic>ospA</italic> as molecular marker [<xref rid="B47-microorganisms-09-00933" ref-type="bibr">47</xref>,<xref rid="B59-microorganisms-09-00933" ref-type="bibr">59</xref>]. In summary, our data support, and agree with, previously published data demonstrating a population division of <italic>B. lusitaniae</italic>.</p><p>Recombination analysis and an analysis of the degree of differentiation between the two <italic>B. lusitaniae</italic> clusters suggested considerable differentiation. Previous work on <italic>Borrelia</italic> has shown that intraspecific recombination rates are 50 times higher than interspecific recombination rates [<xref rid="B89-microorganisms-09-00933" ref-type="bibr">89</xref>]. Furthermore, population genomics analyses of ocean bacteria investigating early events in population differentiation suggest that gradual separation of genes pools is accompanied by population-specific recombination [<xref rid="B90-microorganisms-09-00933" ref-type="bibr">90</xref>,<xref rid="B91-microorganisms-09-00933" ref-type="bibr">91</xref>]. It appears that the two major groups (populations) maybe in the early stages of speciation; each one on its own trajectory. First, both the population structure analysis and Neighbor net analysis clearly unveiled two very well-defined groups. Secondly, the absence of recombination events between these two groups may suggest that there has been no recent gene flow between the two groups. Clearly further investigations considering whole genome sequences are needed to establish if there is gene flow between the two major populations.</p><p>The observed population division of <italic>B. lusitaniae</italic> did not seem to follow climate or vegetation zones known for Europe [<xref rid="B92-microorganisms-09-00933" ref-type="bibr">92</xref>]. North Portugal may be influence by the Atlantic region while countries such as Italy, Spain and Algeria belong to the sub-Mediterranean or Mediterranean region. Although one could speculate that the division of Portuguese <italic>B. lusitaniae</italic> populations may follow the pattern of floristic regions or may be influenced by a geographic barrier (river Tagus), it would not explain the observed clustering of <italic>B. lusitaniae</italic> STs from North Portugal with STs from other European regions.</p></sec><sec id="sec4dot2-microorganisms-09-00933"><title>4.2. Host and Vector Associations</title><p>Previous work has suggested that host association is a key factor for driving diversification and speciation in <italic>Borrelia</italic> [<xref rid="B6-microorganisms-09-00933" ref-type="bibr">6</xref>,<xref rid="B14-microorganisms-09-00933" ref-type="bibr">14</xref>]. Subsequently it was suggested that vector associations may also drive diversification and ultimately speciation in <italic>Borrelia</italic> [<xref rid="B11-microorganisms-09-00933" ref-type="bibr">11</xref>]. In view of the latest described <italic>Ixodes</italic> species (<italic>I. inopinatus</italic>) and its geographic distribution in North Africa, southern Spain and Portugal [<xref rid="B60-microorganisms-09-00933" ref-type="bibr">60</xref>], it was a very attractive hypothesis to investigate whether the association with different vector species may be responsible for the observed population division. However, our data do not unambiguously support the hypothesis that an association with <italic>I. inopinatus</italic> is responsible for the observed population division of <italic>B. lusitaniae</italic>. We added three sample of <italic>I. inopinatus</italic> as reference because these were morphologically identified as <italic>I. inopinatus</italic> and their 16S rRNA sequences were identical to samples designated <italic>I. inopinatus</italic> in GenBank. Surprisingly, the <italic>I. inopinatus</italic> sequences from GenBank (GenBank accession numbers for 16S rRNA/<italic>trospA</italic>, respectively: GU074596TN/GU074839TN; GU074598DZ/GU074841DZ; GU074602MA/GU074845MA) and our own <italic>I. inopinatus</italic> reference samples did not form a monophyletic group. These data call for good reference sequences for this tick species and currently we cannot confirm the hypothesis of <italic>I. inopinatus</italic> association being the reason for the population division of <italic>B. lusitaniae</italic>. Intriguingly, in the <italic>trospA</italic> tree sequences clustered according to geographical origin: a divergent clade contained (with one exception) all the Portuguese and North African samples, samples designated as <italic>I. inopinatus</italic> in GenBank and one sample that the 16S rRNA BLAST search identified as <italic>I. inopinatus</italic> (3117DZ16). A separation between North African and Eurasian <italic>I. ricinus</italic> was also found by [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>] and [<xref rid="B93-microorganisms-09-00933" ref-type="bibr">93</xref>] using concatenated sequences of four mitochondrial and nuclear genes or 125 single nucleotide polymorphisms, respectively, although none of the studies included tick samples from Portugal. Thus, the <italic>B. lusitaniae</italic> population structure obtained by MLST seems to match the population structure of <italic>Ixodes</italic> using the <italic>trospA</italic> gene. Although these data are suggestive of co-evolution between <italic>B. lusitaniae</italic> and its vector, the drivers for this process remain to be elucidated.</p><p>Regarding host association, in the various countries were <italic>B. lusitaniae</italic> has been described, several lizard species have been suggested as reservoir hosts. In Slovakia, Germany and Italy green lizards (<italic>L</italic>. <italic>viridis</italic>) [<xref rid="B46-microorganisms-09-00933" ref-type="bibr">46</xref>], sand lizards (<italic>L. agilis</italic>) [<xref rid="B43-microorganisms-09-00933" ref-type="bibr">43</xref>] and common wall lizards (<italic>Podarcis muralis</italic>) [<xref rid="B37-microorganisms-09-00933" ref-type="bibr">37</xref>,<xref rid="B43-microorganisms-09-00933" ref-type="bibr">43</xref>,<xref rid="B44-microorganisms-09-00933" ref-type="bibr">44</xref>] were described as reservoir hosts. However, these species do not occur on the Iberian Peninsula or in North Africa (see also <uri xlink:href="https://www.Eurolizards.com">https://www.Eurolizards.com</uri>). In Portugal and Spain, <italic>L. schreiberi</italic> is commonly found but its role as reservoir for <italic>B. lusitaniae</italic> cannot be confirmed with current data, although feeding <italic>I. ricinus</italic> nymphs have been found on this host [<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>]. <italic>Psammodromus algirus</italic>, the Algerian lizard, is widespread in North Africa (Algeria, Morocco, Tunisia) and on the Iberian Peninsula but its distribution in Italy or other countries is very restricted. The species has been shown to be divided into differentiated populations in East and West Iberia and North Africa. Furthermore, this species also shows some lineage differentiation in its western range in Iberia, between northern and southern populations, but miscegenation and relatively small sample size has precluded any taxonomic divisions [<xref rid="B94-microorganisms-09-00933" ref-type="bibr">94</xref>,<xref rid="B95-microorganisms-09-00933" ref-type="bibr">95</xref>]. The species has been identified as reservoir host for <italic>B. lusitaniae</italic> in Tunisia [<xref rid="B36-microorganisms-09-00933" ref-type="bibr">36</xref>,<xref rid="B42-microorganisms-09-00933" ref-type="bibr">42</xref>,<xref rid="B96-microorganisms-09-00933" ref-type="bibr">96</xref>] and Portugal [<xref rid="B38-microorganisms-09-00933" ref-type="bibr">38</xref>]. Given the different geographic distributions of potential reservoir hosts for <italic>B. lusitaniae</italic>, it remains to be investigated how narrow the niche for reservoir hosts of <italic>B. lusitaniae</italic> is and serum sensitivity or transmission experiments could help solving this question.</p></sec></sec><sec sec-type="conclusions" id="sec5-microorganisms-09-00933"><title>5. Conclusions</title><p>In this study, we have expanded investigations on the population structure of <italic>B. lusitaniae</italic> and confirmed a population division separating samples from southern Portugal and Algeria from samples from northern Portugal and other European countries. Molecular analyses of morphologically identified <italic>Ixodes</italic> samples (encompassing <italic>I. ricinus</italic> and <italic>I. inopinatus</italic>) acquired from the same geographical regions as <italic>Borrelia</italic> samples showed that the two loci investigated, i.e., 16S rRNA and <italic>trospA</italic>, were phylogenetically incongruous also with respect to clustering of <italic>Ixodes</italic> species. One clade in the <italic>trospA</italic> phylogeny consisted mostly of tick samples from North Africa and Portugal and mirrored the population division found in <italic>Borrelia</italic>. These data suggest that some co-evolution between <italic>Ixodes</italic> and <italic>B. lusitaniae</italic> populations may have occurred which warrants further investigation.</p></sec></body><back><ack><title>Acknowledgments</title><p>We thank Michal Stanko, Bronislava Vichova and Branislav Peťko for providing DNA from <italic>B. lusitaniae</italic> positive ticks from Grabovac, Croatia. Luís Pascoal da Silva for help with sample collection.</p></ack><fn-group><fn><p><bold>Publisher’s Note:</bold> MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.</p></fn></fn-group><app-group><app id="app1-microorganisms-09-00933"><title>Supplementary Materials</title><p>The following are available online at <uri xlink:href="https://www.mdpi.com/article/10.3390/microorganisms9050933/s1">https://www.mdpi.com/article/10.3390/microorganisms9050933/s1</uri>, Table S1: <italic>Ixodes</italic> tick samples included in this study including information on processing of samples, Table S2: <italic>Borrelia</italic> samples included in this study, sequence type (ST) number (if available), allele numbers of the seven genes included for analyses and details of sample processing, Table S3: Detailed results of Hierarchical BAPS analyses at levels 1 and 2, Supplementary Methods, Figure S1: Phyloviz diagram [<xref rid="B83-microorganisms-09-00933" ref-type="bibr">83</xref>] of <italic>B. lusitaniae</italic> samples based on allele profile for seven MLST genes (level 4), Figure S2: Phyloviz diagram of <italic>B. lusitaniae</italic> samples based on allele profiles for seven MLST genes (level 5), Figure S3: Phyloviz diagram of <italic>B. lusitaniae</italic> samples based on allele profiles for seven MLST genes (level 6), Figure S4: NeighborNet of the MLST alignment generated in Splitstree4.</p><supplementary-material content-type="local-data" id="microorganisms-09-00933-s001"><media xlink:href="microorganisms-09-00933-s001.zip"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material></app></app-group><notes><title>Author Contributions</title><p>Conceptualization, A.C.N., G.M., M.S.N., I.L.d.C. and V.F.; methodology, M.D., R.E.R., P.H.B.; formal analysis, A.C.N., S.C.-R., T.W., P.H.B., M.C., Y.M.D., M.D.; resources, A.C.N., P.H.B., M.C., M.O.B., R.E.R., Y.M.D., M.D.; writing—original draft preparation, A.C.N., G.M., V.F.; writing—review and editing, A.C.N., S.C.-R., G.M., P.H.B., T.W., M.D., R.E.R., I.L.d.C.; visualization, A.C.N., S.C.-R.; funding acquisition, A.C.N., V.F., M.D. All authors have read and agreed to the published version of the manuscript.</p></notes><notes><title>Funding</title><p>This research was financially supported by the Slovak Research and Development Agency (grant number APVV-16-0463), by the Fundação para a Ciência e a Tecnologia by the transitory norm contract DL57/2016/CP1370/CT89 to Ana Cláudia Norte and MARE (MARE-UID/MAR/04292/2020), and by the National Institute of Health Doutor Ricardo Jorge, Lisbon, Portugal. The National Reference Center for Borrelia was supported by the Robert-Koch Institute, Berlin, Germany.</p></notes><notes><title>Institutional Review Board Statement</title><p>Not applicable.</p></notes><notes><title>Informed Consent Statement</title><p>Not applicable.</p></notes><notes notes-type="data-availability"><title>Data Availability Statement</title><p>MLST data: All MLST sequence data are available from the MLST database at <uri xlink:href="https://pubmlst.org/borrelia/">https://pubmlst.org/borrelia/</uri> (accessed on 23 April 2021) via isolate names. GenBank Accession numbers: Tick sequences obtained and used in this study were submitted to GenBank under the accession numbers MW017342-MW017360 and MW287230-MW287232 (mitochondrial 16S rRNA locus) and MW379826-MW379844 and MW344866–MW344868 (nuclear gene <italic>trospA</italic>).</p></notes><notes notes-type="COI-statement"><title>Conflicts of Interest</title><p>The authors declare no conflict of interest. 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For details, see <xref rid="microorganisms-09-00933-t001" ref-type="table">Table 1</xref> and <xref rid="microorganisms-09-00933-t002" ref-type="table">Table 2</xref>, <xref ref-type="app" rid="app1-microorganisms-09-00933">Tables S1 and S2</xref>.</p></caption><graphic xlink:href="microorganisms-09-00933-g001"></graphic></fig><fig id="microorganisms-09-00933-f002" orientation="portrait" position="float"><label>Figure 2</label><caption><p>(<bold>A</bold>) Dendrogram of 55 isolates of <italic>B. lusitaniae</italic>. The dendrogram shows the similarity between isolates. Identical isolates are joined together on the same terminal branch, whereas a connection of isolates in the dendrogram further towards the left indicates less similarity (Gower, 1971, R). Color coding of terminal points is according to country (see legend), colors of the branches in the dendrogram specify the two clusters: cluster 1 = red, cluster 2 = green. (<bold>B</bold>) Geographical distribution of isolates belonging to cluster 1 and cluster 2. Color coding of countries corresponds to terminal points in 2A. Color coding of clusters is identical to the branch color in (<bold>A</bold>), cluster 1 = red, cluster 2 = green. The inlet shows the distribution of samples from mainland Portugal belonging to cluster 1 and cluster 2 in more detail. Most samples collected north of the river Tagus fell into cluster 1 while samples from south of the river Tagus fell into cluster 2.</p></caption><graphic xlink:href="microorganisms-09-00933-g002"></graphic></fig><fig id="microorganisms-09-00933-f003" orientation="portrait" position="float"><label>Figure 3</label><caption><p>goeBURST diagram [<xref rid="B83-microorganisms-09-00933" ref-type="bibr">83</xref>] of <italic>B. lusitaniae</italic> samples based on allele profile for seven MLST genes. The triple locus variant setting was chosen. Samples are color coded according to country of origin. For samples from Portugal, the region is indicated. The link color of edges are as follows: Link colors for goeBURST results: Black-Link drawn without recourse to tiebreak rules; Blue-Link drawn using tiebreak rule 1 (number of SLVs); Gray-Links drawn at DLV (darker gray) or TLV (lighter/dotted gray) if the groups are constructed at DLV/TLV level. * indicate STs derived from seven genes only with no correspondence to MLST database ST numbers. § includes one isolate from a human. Dotted line separates samples from the north and south of river Tagus in Portugal.</p></caption><graphic xlink:href="microorganisms-09-00933-g003"></graphic></fig><fig id="microorganisms-09-00933-f004" orientation="portrait" position="float"><label>Figure 4</label><caption><p>NeighborNet of the MLST alignment generated in Splitstree4 [<xref rid="B73-microorganisms-09-00933" ref-type="bibr">73</xref>]. The net was yielded considering uncorrected P distance and equal angle display. Panel A shows the major group that includes the North African and some Portuguese isolates, whereas panel B shows the other major group containing most of the European isolates. The different color labels indicate the clusters found at the second level of clustering of the hierarchical population structure analysis (HPSA) using hierBAPS [<xref rid="B77-microorganisms-09-00933" ref-type="bibr">77</xref>] (see <xref ref-type="app" rid="app1-microorganisms-09-00933">Table S3</xref>). The scale bar shows the number of substitutions per site.</p></caption><graphic xlink:href="microorganisms-09-00933-g004"></graphic></fig><fig id="microorganisms-09-00933-f005" orientation="portrait" position="float"><label>Figure 5</label><caption><p>16S rRNA (<bold>A</bold>) and <italic>trospA</italic> (<bold>B</bold>) Maximum Likelihood phylogeny [<xref rid="B86-microorganisms-09-00933" ref-type="bibr">86</xref>]. Phylogeny based on the 16S RNA gene (<bold>A</bold>). The model HK+I was used to construct the tree. Red labels mark <italic>I. ricinus</italic> non-Portuguese European sequences; green labels highlight <italic>Ixodes</italic> sequences of Portuguese and North African samples; and the blue labels denote <italic>I. inopinatus</italic> “reference” sequences from Germany. Phylogeny based on the <italic>trospA</italic> gene (<bold>B</bold>). The model K80+I was used to construct the tree. Red labels mark <italic>I. ricinus</italic> non-Portuguese European sequences; green labels highlight Portuguese and North African <italic>Ixodes</italic> sequences; and the blue labels denote <italic>I. inopinatus</italic> “reference” sequences. The scale bars show the number of substitutions per site.</p></caption><graphic xlink:href="microorganisms-09-00933-g005"></graphic></fig><table-wrap id="microorganisms-09-00933-t001" orientation="portrait" position="float"><object-id pub-id-type="pii">microorganisms-09-00933-t001_Table 1</object-id><label>Table 1</label><caption><p>Tick samples included in this study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Tick AN for NCBI_16S rRNA</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Tick AN for NCBI_<italic>trospA</italic></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Tick Species<break></break>(16S rRNA Based ID)</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Year</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Stage/Sex</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"><italic>B. lusitaniae</italic><break></break>Infection Status</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">MLST<break></break><italic>B. lusitaniae</italic></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1"></th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Region Specific †</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Reference</th></tr></thead><tbody><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074596TN</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074839TN</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus #</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">TN, Jbel el Jouza</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074598DZ ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074841DZ</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus #</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DZ, El Tarf</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074602MA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074845MA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus #</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">M</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MA, Taza</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074597FR</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074840FR ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">FR, Forêt de Chizé</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074606ES ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074849ES</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">ES, Otxandio</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074595IE ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074838IE</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">IE, Cork, Killarney National park</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074603GB</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074846GB</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">UK</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074607NL</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074850NL</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">NL</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074605DE ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074848DE</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DE, Munich, English garden</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074599IR</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074842IR</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">IR, Mazendaran province</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074592DK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074835DK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DK, Grib Skov Forest</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074593SE ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074836SE</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SE, Alsike</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074588SK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074831SK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Železná Studnička</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074589SK ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074832SK ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Vel’ký Lom</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074590SK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074833SK</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Malá Lehota</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074594EE ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074837EE ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">EE, Tartumaa</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074600FI</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074843FI</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">FI, Turku archipelago</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074604HU</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074847HU</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">HU, Mátrafüred forest</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074591MD</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074834MD ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MD</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074601BG</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">GU074844BG</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">BG, Sofia</td><td align="center" valign="top" style="border-bottom:solid thin" rowspan="1" colspan="1">[<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>]</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">13310PT18 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">13310PT18</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">south Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Santiago do Cacém</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">13360PT18 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">13360PT18</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">south Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">14401PT18 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">14401PT18</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">south Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Santiago do Cacém</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">G101PT09 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">G101PT09</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">unknown</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">north Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Gerês</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1756PT13 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1756PT13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">north Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1773PT13 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1773PT13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">north Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1812PT13 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1812PT13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">negative</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">north Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1794PT13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">R1794PT13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">negative</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">north Tagus</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PT, Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3117DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3117DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">yes: ID 3117</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DZ, Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">T73T04DZ16 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">T73T04DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DZ, Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3115DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3115DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DZ, Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH152SK17 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH152SK17</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2017</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">M</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH149SK14 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH149SK14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2014</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">M</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH139SK14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH139SK14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2014</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">M</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">yes: ID 2653</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH106SK17 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH106SK17</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2017</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH60SK17 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH60SK17</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2017</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2874SK16 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2874SK16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH126SK14 ‡</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH126SK14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2014</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2652SK16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2652SK16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">F</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">positive</td><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1">yes: ID 2652</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SK, Martinské hole</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">11-E12</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">11-E12</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">negative</td><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DE, Starnberg</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-F5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-F5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">negative</td><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DE, Starnberg</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8-C12</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8-C12</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes inopinatus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2018</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">N</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">negative</td><td colspan="2" align="center" valign="middle" style="border-bottom:solid thin" rowspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">DE, Starnberg</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr></tbody></table><table-wrap-foot><fn><p>AN = accession number, M = male, F = female, N = nymh, † the two-letter country code (ISO 3166) is given, ‡ Sequence detected in more than one country., # designated <italic>I. ricinus</italic> in Noureddine et al. 2011 [<xref rid="B61-microorganisms-09-00933" ref-type="bibr">61</xref>] but now designated <italic>I. inopinatus</italic> in GenBank.</p></fn></table-wrap-foot></table-wrap><table-wrap id="microorganisms-09-00933-t002" orientation="portrait" position="float"><object-id pub-id-type="pii">microorganisms-09-00933-t002_Table 2</object-id><label>Table 2</label><caption><p><italic>Borrelia lusitaniae</italic> samples included in this study for which seven MLST genes were available.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Sample<break></break>No.</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Sample ID</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Strain</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Genospecies</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Country</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Area</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Continent</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">ST</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Year of Collection</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Biological Source</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Tick Code</th><th align="center" valign="middle" style="border-top:solid thin;border-bottom:solid thin" rowspan="1" colspan="1">Reference</th></tr></thead><tbody><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">59</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">162/11b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">628</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2011</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">63</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">167/11b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Kosutnjak</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">628</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2011</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">64</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">167/11c</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Kosutnjak</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">136</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2011</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">4</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">124</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr41</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">60</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2002</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">125</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr82</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">61</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2002</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">6</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">126</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr130</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">62</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">127</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr131</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">63</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">128</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr136</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">64</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">129</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTibGr409</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">64</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">10</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">130</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr143</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">65</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">11</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">131</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr211</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">65</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">12</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">132</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr209</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">66</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">13</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">133</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr213</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">66</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">134</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr288</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">67</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">15</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">135</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBGr293</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grândola</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">68</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">136</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoHL1</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Lisbon</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">69</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2002</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>human</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">17</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">137</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiBL37</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">69</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1999</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">18</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">249</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiB2</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">64</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">19</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">374</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">71412L</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Latvia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">218</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2007</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">20</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1164</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">0911001A</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Austria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">North-Tirol</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">336</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">21</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1181</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">0921035A</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Austria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">North-Tirol</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">345</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">22</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1182</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">0921036A</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Austria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">North-Tirol</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">346</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">23</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1250</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">233/13b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Avala</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">148</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">24</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1252</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">221/10c</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">153</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">25</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1253</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">225/10a</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">630</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">26</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1254</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">228/10a</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">194</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">27</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1255</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">229/10a</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Kosutnjak</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">209</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">28</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1585</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">76/12a</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Eastern Serbia-Dobra (Djerdap Gorge)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">580</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2012</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">29</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1586</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">77/12b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Eastern Serbia-Dobra (Djerdap Gorge)</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">586</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2012</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">30</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1815</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">220/10b</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Kosutnjak</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">628</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">31</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1817</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">222/10d</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">629</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">32</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1819</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">224/10c</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">630</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">33</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1821</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">226/10d</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">631</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">34</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1822</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">227/10c</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Serbia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Belgrade-Titov gaj</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">630</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">35</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2073</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiB3</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">766</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">1993</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">36</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2500</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">82NCHG</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Croatia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Grabovac</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">900</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2011</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">37</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2525</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiBmfP147</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-011</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">38</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2526</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiBmfP220</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-004</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">39</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2527</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiBmfJ2</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-005</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2001</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">40</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2528</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiBmfJ50</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-005</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">41</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2529</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PotiBmfP364</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Mafra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-006</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MLST DB</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">42</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2594</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">658 UA</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Ukraine</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kyiv, M.M. Gryshko Nat. Bot. Garden</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">909</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2015</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">43</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2652</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH2016F8</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Slovakia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Martinské hole mountain</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">918</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2652SK16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">44</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2653</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">SMHM139</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Slovakia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Martinské hole mountain</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">919</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2013</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH139SK14</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">52</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3126</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiB10/M436</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Madeira</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">925</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">53</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3127</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiB11/T3087</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Coimbra</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">926</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2014</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">51</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3125</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">PoTiB9/B88</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Portugal</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Santiago do Cacém</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">924</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Dermacentor</italic><break></break><italic>marginatum</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">45</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3114</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube101-Tick25/run1-2</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-000</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">46</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3117</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube104-Tick14/run1-5</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-001</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3117DZ16</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">47</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3118</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube94-Tick5/run1-6</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-002</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">48</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3120</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube101-Tick24/run1-11</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-003</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">49</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3119</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube 95-Tick5/run1-9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-007</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">50</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">3123</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Tube94-tick6/run2-7</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Algeria</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Kabylie</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Africa</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-008</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">54</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2873</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH2016F1</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Slovakia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Martinské hole mountain</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-009</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr><tr><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">55</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2874</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">MH2016F9</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Borrelia lusitaniae</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Slovakia</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Martinské hole mountain</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">Europe</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">7-010</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">2016</td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"><italic>Ixodes ricinus</italic></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1"></td><td align="center" valign="middle" style="border-bottom:solid thin" rowspan="1" colspan="1">This study</td></tr></tbody></table><table-wrap-foot><fn><p>7-0XX = provisional ST number. MLST DB – Multilocus sequence typing database <uri xlink:href="www.pubmlst.org/borrelia/">www.pubmlst.org/borrelia/</uri> (accessed on 11 February 2020).</p></fn></table-wrap-foot></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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