Microallopatry Caused Strong Diversification in Buthus scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa)
Identifieur interne : 000180 ( Pmc/Corpus ); précédent : 000179; suivant : 000181Microallopatry Caused Strong Diversification in Buthus scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa)
Auteurs : Jan C. Habel ; Martin Husemann ; Thomas Schmitt ; Frank E. Zachos ; Ann-Christin Honnen ; Britt Petersen ; Aristeidis Parmakelis ; Iasmi StathiSource :
- PLoS ONE [ 1932-6203 ] ; 2012.
Abstract
The immense biodiversity of the Atlas Mountains in North Africa might be the result of high rates of microallopatry caused by mountain barriers surpassing 4000 meters leading to patchy habitat distributions. We test the influence of geographic structures on the phylogenetic patterns among
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DOI: 10.1371/journal.pone.0029403
PubMed: 22383951
PubMed Central: 3287997
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Microallopatry Caused Strong Diversification in <italic>Buthus</italic> scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa)</title><author><name sortKey="Habel, Jan C" sort="Habel, Jan C" uniqKey="Habel J" first="Jan C." last="Habel">Jan C. Habel</name><affiliation><nlm:aff id="aff1"><addr-line>Natural History Museum Luxembourg, Invertebrate Biology, Luxembourg</addr-line></nlm:aff></affiliation></author><author><name sortKey="Husemann, Martin" sort="Husemann, Martin" uniqKey="Husemann M" first="Martin" last="Husemann">Martin Husemann</name><affiliation><nlm:aff id="aff2"><addr-line>Baylor University, Biology Department, Waco, Texas, United States of America</addr-line></nlm:aff></affiliation></author><author><name sortKey="Schmitt, Thomas" sort="Schmitt, Thomas" uniqKey="Schmitt T" first="Thomas" last="Schmitt">Thomas Schmitt</name><affiliation><nlm:aff id="aff3"><addr-line>Trier University, Department of Biogeography, Trier, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Zachos, Frank E" sort="Zachos, Frank E" uniqKey="Zachos F" first="Frank E." last="Zachos">Frank E. Zachos</name><affiliation><nlm:aff id="aff4"><addr-line>Natural History Museum Vienna, Vienna, Austria</addr-line></nlm:aff></affiliation></author><author><name sortKey="Honnen, Ann Christin" sort="Honnen, Ann Christin" uniqKey="Honnen A" first="Ann-Christin" last="Honnen">Ann-Christin Honnen</name><affiliation><nlm:aff id="aff5"><addr-line>Zoological Institute, Christian-Albrechts-University, Kiel, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Petersen, Britt" sort="Petersen, Britt" uniqKey="Petersen B" first="Britt" last="Petersen">Britt Petersen</name><affiliation><nlm:aff id="aff6"><addr-line>Institute of Clinical Molecular Biology, University Hospital SH Campus Kiel, Kiel, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Parmakelis, Aristeidis" sort="Parmakelis, Aristeidis" uniqKey="Parmakelis A" first="Aristeidis" last="Parmakelis">Aristeidis Parmakelis</name><affiliation><nlm:aff id="aff7"><addr-line>National and Kapodistrian University of Athens, Department of Ecology and Taxonomy, Athens, Greece</addr-line></nlm:aff></affiliation></author><author><name sortKey="Stathi, Iasmi" sort="Stathi, Iasmi" uniqKey="Stathi I" first="Iasmi" last="Stathi">Iasmi Stathi</name><affiliation><nlm:aff id="aff8"><addr-line>Natural History Museum of Crete, University of Crete, Heraklion, Crete, Greece</addr-line></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">22383951</idno><idno type="pmc">3287997</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3287997</idno><idno type="RBID">PMC:3287997</idno><idno type="doi">10.1371/journal.pone.0029403</idno><date when="2012">2012</date><idno type="wicri:Area/Pmc/Corpus">000180</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000180</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Microallopatry Caused Strong Diversification in <italic>Buthus</italic> scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa)</title><author><name sortKey="Habel, Jan C" sort="Habel, Jan C" uniqKey="Habel J" first="Jan C." last="Habel">Jan C. Habel</name><affiliation><nlm:aff id="aff1"><addr-line>Natural History Museum Luxembourg, Invertebrate Biology, Luxembourg</addr-line></nlm:aff></affiliation></author><author><name sortKey="Husemann, Martin" sort="Husemann, Martin" uniqKey="Husemann M" first="Martin" last="Husemann">Martin Husemann</name><affiliation><nlm:aff id="aff2"><addr-line>Baylor University, Biology Department, Waco, Texas, United States of America</addr-line></nlm:aff></affiliation></author><author><name sortKey="Schmitt, Thomas" sort="Schmitt, Thomas" uniqKey="Schmitt T" first="Thomas" last="Schmitt">Thomas Schmitt</name><affiliation><nlm:aff id="aff3"><addr-line>Trier University, Department of Biogeography, Trier, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Zachos, Frank E" sort="Zachos, Frank E" uniqKey="Zachos F" first="Frank E." last="Zachos">Frank E. Zachos</name><affiliation><nlm:aff id="aff4"><addr-line>Natural History Museum Vienna, Vienna, Austria</addr-line></nlm:aff></affiliation></author><author><name sortKey="Honnen, Ann Christin" sort="Honnen, Ann Christin" uniqKey="Honnen A" first="Ann-Christin" last="Honnen">Ann-Christin Honnen</name><affiliation><nlm:aff id="aff5"><addr-line>Zoological Institute, Christian-Albrechts-University, Kiel, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Petersen, Britt" sort="Petersen, Britt" uniqKey="Petersen B" first="Britt" last="Petersen">Britt Petersen</name><affiliation><nlm:aff id="aff6"><addr-line>Institute of Clinical Molecular Biology, University Hospital SH Campus Kiel, Kiel, Germany</addr-line></nlm:aff></affiliation></author><author><name sortKey="Parmakelis, Aristeidis" sort="Parmakelis, Aristeidis" uniqKey="Parmakelis A" first="Aristeidis" last="Parmakelis">Aristeidis Parmakelis</name><affiliation><nlm:aff id="aff7"><addr-line>National and Kapodistrian University of Athens, Department of Ecology and Taxonomy, Athens, Greece</addr-line></nlm:aff></affiliation></author><author><name sortKey="Stathi, Iasmi" sort="Stathi, Iasmi" uniqKey="Stathi I" first="Iasmi" last="Stathi">Iasmi Stathi</name><affiliation><nlm:aff id="aff8"><addr-line>Natural History Museum of Crete, University of Crete, Heraklion, Crete, Greece</addr-line></nlm:aff></affiliation></author></analytic><series><title level="j">PLoS ONE</title><idno type="eISSN">1932-6203</idno><imprint><date when="2012">2012</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>The immense biodiversity of the Atlas Mountains in North Africa might be the result of high rates of microallopatry caused by mountain barriers surpassing 4000 meters leading to patchy habitat distributions. We test the influence of geographic structures on the phylogenetic patterns among <italic>Buthus</italic> scorpions using mtDNA sequences. We sampled 91 individuals of the genus <italic>Buthus</italic> from 51 locations scattered around the Atlas Mountains (Antiatlas, High Atlas, Middle Atlas and Jebel Sahro). We sequenced 452 bp of the Cytochrome Oxidase I gene which proved to be highly variable within and among <italic>Buthus</italic> species. Our phylogenetic analysis yielded 12 distinct genetic groups one of which comprised three subgroups mostly in accordance with the orographic structure of the mountain systems. Main clades overlap with each other, while subclades are distributed parapatrically. Geographic structures likely acted as long-term barriers among populations causing restriction of gene flow and allowing for strong genetic differentiation. Thus, genetic structure and geographical distribution of genetic (sub)clusters follow the classical theory of allopatric differentiation where distinct groups evolve without range overlap until reproductive isolation and ecological differentiation has built up. Philopatry and low dispersal ability of <italic>Buthus</italic> scorpions are the likely causes for the observed strong genetic differentiation at this small geographic scale.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="De Lattin, G" uniqKey="De Lattin G">G de Lattin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Hewitt, Gm" uniqKey="Hewitt G">GM Hewitt</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="G Mez, A" uniqKey="G Mez A">A Gómez</name></author><author><name sortKey="Lunt, Dh" uniqKey="Lunt D">DH Lunt</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Joger, U" uniqKey="Joger U">U Joger</name></author><author><name sortKey="Fritz, U" uniqKey="Fritz U">U Fritz</name></author><author><name sortKey="Guicking, D" uniqKey="Guicking D">D Guicking</name></author><author><name sortKey="Tripepi, S" uniqKey="Tripepi S">S 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<front><journal-meta><journal-id journal-id-type="nlm-ta">PLoS One</journal-id><journal-id journal-id-type="publisher-id">plos</journal-id><journal-id journal-id-type="pmc">plosone</journal-id><journal-title-group><journal-title>PLoS ONE</journal-title></journal-title-group><issn pub-type="epub">1932-6203</issn><publisher><publisher-name>Public Library of Science</publisher-name><publisher-loc>San Francisco, USA</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">22383951</article-id><article-id pub-id-type="pmc">3287997</article-id><article-id pub-id-type="publisher-id">PONE-D-11-17600</article-id><article-id pub-id-type="doi">10.1371/journal.pone.0029403</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="Discipline-v2"><subject>Biology</subject><subj-group><subject>Ecology</subject></subj-group><subj-group><subject>Evolutionary Biology</subject></subj-group><subj-group><subject>Population Biology</subject></subj-group><subj-group><subject>Zoology</subject></subj-group></subj-group></article-categories><title-group><article-title>Microallopatry Caused Strong Diversification in <italic>Buthus</italic> scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa)</article-title><alt-title alt-title-type="running-head">Biogeography of Buthus Scorpions in N-Africa</alt-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Habel</surname><given-names>Jan C.</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="corresp" rid="cor1"><sup>*</sup></xref></contrib><contrib contrib-type="author"><name><surname>Husemann</surname><given-names>Martin</given-names></name><xref ref-type="aff" rid="aff2"><sup>2</sup></xref></contrib><contrib contrib-type="author"><name><surname>Schmitt</surname><given-names>Thomas</given-names></name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref></contrib><contrib contrib-type="author"><name><surname>Zachos</surname><given-names>Frank E.</given-names></name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref></contrib><contrib contrib-type="author"><name><surname>Honnen</surname><given-names>Ann-Christin</given-names></name><xref ref-type="aff" rid="aff5"><sup>5</sup></xref></contrib><contrib contrib-type="author"><name><surname>Petersen</surname><given-names>Britt</given-names></name><xref ref-type="aff" rid="aff6"><sup>6</sup></xref></contrib><contrib contrib-type="author"><name><surname>Parmakelis</surname><given-names>Aristeidis</given-names></name><xref ref-type="aff" rid="aff7"><sup>7</sup></xref></contrib><contrib contrib-type="author"><name><surname>Stathi</surname><given-names>Iasmi</given-names></name><xref ref-type="aff" rid="aff8"><sup>8</sup></xref></contrib></contrib-group><aff id="aff1"><label>1</label><addr-line>Natural History Museum Luxembourg, Invertebrate Biology, Luxembourg</addr-line></aff><aff id="aff2"><label>2</label><addr-line>Baylor University, Biology Department, Waco, Texas, United States of America</addr-line></aff><aff id="aff3"><label>3</label><addr-line>Trier University, Department of Biogeography, Trier, Germany</addr-line></aff><aff id="aff4"><label>4</label><addr-line>Natural History Museum Vienna, Vienna, Austria</addr-line></aff><aff id="aff5"><label>5</label><addr-line>Zoological Institute, Christian-Albrechts-University, Kiel, Germany</addr-line></aff><aff id="aff6"><label>6</label><addr-line>Institute of Clinical Molecular Biology, University Hospital SH Campus Kiel, Kiel, Germany</addr-line></aff><aff id="aff7"><label>7</label><addr-line>National and Kapodistrian University of Athens, Department of Ecology and Taxonomy, Athens, Greece</addr-line></aff><aff id="aff8"><label>8</label><addr-line>Natural History Museum of Crete, University of Crete, Heraklion, Crete, Greece</addr-line></aff><contrib-group><contrib contrib-type="editor"><name><surname>Hofreiter</surname><given-names>Michael</given-names></name><role>Editor</role><xref ref-type="aff" rid="edit1"></xref></contrib></contrib-group><aff id="edit1">University of York, United Kingdom</aff><author-notes><corresp id="cor1">* E-mail: <email>Janchristianhabel@gmx.de</email></corresp><fn fn-type="con"><p>Conceived and designed the experiments: JCH MH TS. Performed the experiments: JCH MH. Analyzed the data: JCH MH IS BP AH. Contributed reagents/materials/analysis tools: JCH MH FZ. Wrote the paper: JCH MH TS FZ IS AP.</p></fn></author-notes><pub-date pub-type="collection"><year>2012</year></pub-date><pub-date pub-type="epub"><day>27</day><month>2</month><year>2012</year></pub-date><volume>7</volume><issue>2</issue><elocation-id>e29403</elocation-id><history><date date-type="received"><day>8</day><month>9</month><year>2011</year></date><date date-type="accepted"><day>28</day><month>11</month><year>2011</year></date></history><permissions><copyright-statement>Habel et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</copyright-statement><copyright-year>2012</copyright-year></permissions><abstract><p>The immense biodiversity of the Atlas Mountains in North Africa might be the result of high rates of microallopatry caused by mountain barriers surpassing 4000 meters leading to patchy habitat distributions. We test the influence of geographic structures on the phylogenetic patterns among <italic>Buthus</italic> scorpions using mtDNA sequences. We sampled 91 individuals of the genus <italic>Buthus</italic> from 51 locations scattered around the Atlas Mountains (Antiatlas, High Atlas, Middle Atlas and Jebel Sahro). We sequenced 452 bp of the Cytochrome Oxidase I gene which proved to be highly variable within and among <italic>Buthus</italic> species. Our phylogenetic analysis yielded 12 distinct genetic groups one of which comprised three subgroups mostly in accordance with the orographic structure of the mountain systems. Main clades overlap with each other, while subclades are distributed parapatrically. Geographic structures likely acted as long-term barriers among populations causing restriction of gene flow and allowing for strong genetic differentiation. Thus, genetic structure and geographical distribution of genetic (sub)clusters follow the classical theory of allopatric differentiation where distinct groups evolve without range overlap until reproductive isolation and ecological differentiation has built up. Philopatry and low dispersal ability of <italic>Buthus</italic> scorpions are the likely causes for the observed strong genetic differentiation at this small geographic scale.</p></abstract><counts><page-count count="8"></page-count></counts></article-meta></front><body><sec id="s1"><title>Introduction</title><p>The Mediterranean region comprises a highly diverse geographic area characterized by a variety of peninsulas and islands. This underlying geographic structure is reflected in biogeographical patterns of the region already described by de Lattin (1949) <xref ref-type="bibr" rid="pone.0029403-deLattin1">[1]</xref> who distinguished nine Mediterranean sub-centres. These early hypotheses have largely been confirmed by genetic analyses <xref ref-type="bibr" rid="pone.0029403-Hewitt1">[2]</xref>. However, recent molecular data support even more fine-scaled patterns within these sub-centres; such patterns are especially known from the Iberian Peninsula e.g. <xref ref-type="bibr" rid="pone.0029403-Gmez1">[3]</xref>, peninsular Italy e.g. <xref ref-type="bibr" rid="pone.0029403-Joger1">[4]</xref> and the Balkans <xref ref-type="bibr" rid="pone.0029403-Schmitt1">[5]</xref>. While the European part of the Mediterranean is biogeographically relatively well understood, the Maghreb still remains largely unstudied <xref ref-type="bibr" rid="pone.0029403-Hewitt2">[6]</xref>. Therefore, the question emerges whether similar genetic substructures also exist in NW Africa.</p><p>De Lattin (1949) <xref ref-type="bibr" rid="pone.0029403-deLattin1">[1]</xref> suggested that the faunas of the Maghreb differ north and south of the High Atlas; this pattern is well reflected in the geographic distributions of many species e.g. <xref ref-type="bibr" rid="pone.0029403-Fritz1">[7]</xref>. On the contrary, other genetic studies revealed an east-west split along the Maghreb coast <xref ref-type="bibr" rid="pone.0029403-Cosson1">[8]</xref>. Furthermore, no genetic differentiation from Morocco to Tunisia was observed in the allozyme patterns of two butterfly species <xref ref-type="bibr" rid="pone.0029403-Habel1">[9]</xref>. Nevertheless, all these genetic patterns are much simpler compared to findings in southern Europe. This is particularly astonishing considering the high orographic diversity of the Maghreb in general and the Atlas Mountains in particular. Yet, only few studies have addressed the phylo- or biogeographic patterns within the Atlas Mountains, although this area comprises an excellent region to study allopatric differentiation.</p><p>In order to test for the influence of orographic structures on differentiation and speciation processes, we selected the scorpion genus <italic>Buthus</italic>, which has undergone extensive speciation within NW Africa <xref ref-type="bibr" rid="pone.0029403-Loureno1">[10]</xref>, as a model system. <italic>Buthus</italic> scorpions are widely distributed over large parts of Africa and show a circum-Mediterranean stronghold including the Mediterranean islands, with extraordinarily high diversity in North Africa <xref ref-type="bibr" rid="pone.0029403-Loureno1">[10]</xref>–<xref ref-type="bibr" rid="pone.0029403-Gantenbein1">[14]</xref>. Yet, molecular studies showed that diversity may still be underestimated and additional cryptic species may be abundant <xref ref-type="bibr" rid="pone.0029403-Gantenbein1">[14]</xref>. Most buthid scorpions are strictly territorial and philopatric and have low dispersal abilities. Consequently, individuals have limited home ranges which generally are restricted to small areas around their burrows <xref ref-type="bibr" rid="pone.0029403-Polis1">[15]</xref>. Strong genetic differentiation has previously been observed for scorpion species and was probably triggered by barriers such as high elevations, steep valleys or sea barriers <xref ref-type="bibr" rid="pone.0029403-Froufe1">[16]</xref>–<xref ref-type="bibr" rid="pone.0029403-Sousa2">[19]</xref> in combination with low dispersal ability of the organisms <xref ref-type="bibr" rid="pone.0029403-Polis2">[20]</xref>. In addition to being extremely limited, dispersal is also highly asymmetrical in these organisms, with male scorpions dispersing more widely. This apparently is a function of mate, prey and shelter search, where males exhibit a much more opportunistic locomotive behaviour, whereas females stay in their shelter and forage mainly close to the entrance of their burrows (“doorkeeping” strategy) <xref ref-type="bibr" rid="pone.0029403-Kaltsas1">[21]</xref>, <xref ref-type="bibr" rid="pone.0029403-Kaltsas2">[22]</xref>.</p><p>For this study we collected a total of 91 individuals belonging to 4 described <italic>Buthus</italic> species from 51 locations over major parts of the Antiatlas, High Atlas, Middle Atlas and Jebel Sahro during spring 2008 and 2009. Three specimens of the related genus <italic>Androctonus</italic> from two locations served as outgroup. We analysed a fragment of the mitochondrial Cytochrome Oxidase I (<italic>COI</italic>) gene, which has been shown to be highly informative at species level and consequently is one of the most commonly used markers for phylogenetic and phylogeographic studies. Due to its high potential for species identification it is often chosen as the barcoding gene in zoological studies. Here we use its properties to address the following questions:</p><list list-type="roman-lower"><list-item><p>Does genetic variation within <italic>Buthus</italic> exhibit geographic structuring?</p></list-item><list-item><p>Do genetic structures follow any geographical and/or evolutionary pattern?</p></list-item><list-item><p>Are genetic lineages in accordance with current taxonomy?</p></list-item></list></sec><sec id="s2"><title>Results</title><p>We generated 94 sequences (including three outgroup specimens) with a total length of 452 base pairs. One sequence from <italic>Androctonus mauritanicus</italic> (JF820097) was obtained from genbank as an additional outgroup. Details about analyzed individuals, including genbank accession numbers, are given in <xref ref-type="table" rid="pone-0029403-t001">Table 1</xref>. Out of the 452 sites analysed, 135 were polymorphic (29.9%) of which 127 were parsimony informative sites (4 sites had missing data). Within the ingroup we recovered 69 haplotype with a haplotypes diversity of Hd = 0.993 (SD = 0.003) and a nucleotide diversity of Pi = 0.0916 (SD = 0.0018). The average number of nucleotide differences was k = 41.138. Fst estimates among groups obtained from phylogenetic analyses were overall high and mostly significant (<xref ref-type="table" rid="pone-0029403-t002">Table 2</xref>; non significant values are due to low sample sizes in some groups).</p><table-wrap id="pone-0029403-t001" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0029403.t001</object-id><label>Table 1</label><caption><title>Sampling localities of all studied <italic>Buthus</italic> populations.</title></caption><alternatives><graphic id="pone-0029403-t001-1" xlink:href="pone.0029403.t001"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1">Group</td><td align="left" rowspan="1" colspan="1">Locality</td><td align="left" rowspan="1" colspan="1">Site</td><td align="left" rowspan="1" colspan="1">Coordinates(N; W)</td><td align="left" rowspan="1" colspan="1">Species</td><td align="left" rowspan="1" colspan="1">Date of sampling</td><td align="left" rowspan="1" colspan="1">Specimen ID/##</td><td align="left" rowspan="1" colspan="1">Genbank accession#</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><bold>A</bold></td><td align="left" rowspan="1" colspan="1">Tamensourt</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">31.43; 8.01</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">13-III-2009</td><td align="left" rowspan="1" colspan="1">9-80-5</td><td align="left" rowspan="1" colspan="1">JN885948</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>B</bold></td><td align="left" rowspan="1" colspan="1">Chichoaua</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">31.27; 8.47</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">13-III-2009</td><td align="left" rowspan="1" colspan="1">9-79-1</td><td align="left" rowspan="1" colspan="1">JN832012</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-79-2</td><td align="left" rowspan="1" colspan="1">JN832013</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>C</bold></td><td align="left" rowspan="1" colspan="1">Argane</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">30.55; 9.03</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">12-III-2009</td><td align="left" rowspan="1" colspan="1">9-78-2</td><td align="left" rowspan="1" colspan="1">JN832008</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-78-3</td><td align="left" rowspan="1" colspan="1">JN832009</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>D</bold></td><td align="left" rowspan="1" colspan="1">Ameskrout</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">30.37; 9.20</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">12-III-2009</td><td align="left" rowspan="1" colspan="1">9-77-1</td><td align="left" rowspan="1" colspan="1">JN832003</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-77-2</td><td align="left" rowspan="1" colspan="1">JN832004</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Agadir</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">30.28; 9.18</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">12-III-2009</td><td align="left" rowspan="1" colspan="1">9-74-4</td><td align="left" rowspan="1" colspan="1">JN831999</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-74-6</td><td align="left" rowspan="1" colspan="1">JN832000</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>E</bold></td><td align="left" rowspan="1" colspan="1">Tiznit</td><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">29.18; 9.45</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">24-II-2009</td><td align="left" rowspan="1" colspan="1">9-13-1</td><td align="left" rowspan="1" colspan="1">JN831992</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-13-2</td><td align="left" rowspan="1" colspan="1">JN831993</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Et Tnine</td><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">29.42; 9.16</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">11-V-2008</td><td align="left" rowspan="1" colspan="1">8-6-1</td><td align="left" rowspan="1" colspan="1">JN831971</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">8-6-2</td><td align="left" rowspan="1" colspan="1">JN831972</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tighermi</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">29.32; 9.20</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">24-II-2009</td><td align="left" rowspan="1" colspan="1">9-10-3</td><td align="left" rowspan="1" colspan="1">JN831983</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-10-5</td><td align="left" rowspan="1" colspan="1">JN831984</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Onafka</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">29.24; 9.15</td><td align="left" rowspan="1" colspan="1"><italic>B.</italic> spec.</td><td align="left" rowspan="1" colspan="1">24-II-2009</td><td align="left" rowspan="1" colspan="1">9-12-1</td><td align="left" rowspan="1" colspan="1">JN831989</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-12-2</td><td align="left" rowspan="1" colspan="1">JN831990</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Izerbi</td><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1">29.20; 9.04</td><td align="left" rowspan="1" colspan="1"><italic>B.</italic> spec.</td><td align="left" rowspan="1" colspan="1">24-II-2009</td><td align="left" rowspan="1" colspan="1">9-11-1</td><td align="left" rowspan="1" colspan="1">JN831986</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-11-2</td><td align="left" rowspan="1" colspan="1">JN831987</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>F</bold></td><td align="left" rowspan="1" colspan="1">Ait Saha</td><td align="left" rowspan="1" colspan="1">11</td><td align="left" rowspan="1" colspan="1">30.06; 9.12</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">23-II-2009</td><td align="left" rowspan="1" colspan="1">9-6-1</td><td align="left" rowspan="1" colspan="1">JN831995</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-6-3</td><td align="left" rowspan="1" colspan="1">JN831996</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tiz-n-Test</td><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">30.52; 8.23</td><td align="left" rowspan="1" colspan="1"><italic>B.</italic> spec.</td><td align="left" rowspan="1" colspan="1">12-V-2008</td><td align="left" rowspan="1" colspan="1">8-10-1</td><td align="left" rowspan="1" colspan="1">JN831974</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">8-10-2</td><td align="left" rowspan="1" colspan="1">JN831975</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">W Tassoumate</td><td align="left" rowspan="1" colspan="1">13</td><td align="left" rowspan="1" colspan="1">30.30; 8.34</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">11-III-2009</td><td align="left" rowspan="1" colspan="1">9-73-2</td><td align="left" rowspan="1" colspan="1">JN832029</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-73-3</td><td align="left" rowspan="1" colspan="1">JN832030</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ait Aissa</td><td align="left" rowspan="1" colspan="1">14</td><td align="left" rowspan="1" colspan="1">30.18; 8.31</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">22-II-2009</td><td align="left" rowspan="1" colspan="1">9-3-1</td><td align="left" rowspan="1" colspan="1">JN832016</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-3-2</td><td align="left" rowspan="1" colspan="1">JN832017</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tassga</td><td align="left" rowspan="1" colspan="1">15</td><td align="left" rowspan="1" colspan="1">30.10; 8.28</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">22-II-2009</td><td align="left" rowspan="1" colspan="1">9-5-1</td><td align="left" rowspan="1" colspan="1">JN831978</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-5-2</td><td align="left" rowspan="1" colspan="1">JN831981</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">S Tassoumate</td><td align="left" rowspan="1" colspan="1">16</td><td align="left" rowspan="1" colspan="1">30.33; 8.15</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">11-III-2009</td><td align="left" rowspan="1" colspan="1">9-72-1</td><td align="left" rowspan="1" colspan="1">JN832026</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-72-3</td><td align="left" rowspan="1" colspan="1">JN832027</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tassoumate</td><td align="left" rowspan="1" colspan="1">17</td><td align="left" rowspan="1" colspan="1">30.35; 8.15</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">11-III-2009</td><td align="left" rowspan="1" colspan="1">9-71-1</td><td align="left" rowspan="1" colspan="1">JN832023</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-71-2</td><td align="left" rowspan="1" colspan="1">JN832024</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">SW Taliouine</td><td align="left" rowspan="1" colspan="1">18</td><td align="left" rowspan="1" colspan="1">30.22; 8.09</td><td align="left" rowspan="1" colspan="1"><italic>B. elmoutaouakili</italic></td><td align="left" rowspan="1" colspan="1">11-III-2009</td><td align="left" rowspan="1" colspan="1">9-70-1</td><td align="left" rowspan="1" colspan="1">JN831997</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-70-3</td><td align="left" rowspan="1" colspan="1">JN831998</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Igherm</td><td align="left" rowspan="1" colspan="1">19</td><td align="left" rowspan="1" colspan="1">30.06; 8.70</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">22-II-2009</td><td align="left" rowspan="1" colspan="1">9-4-1</td><td align="left" rowspan="1" colspan="1">JN832019</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>G</bold></td><td align="left" rowspan="1" colspan="1">75 km W Tazenakht</td><td align="left" rowspan="1" colspan="1">20</td><td align="left" rowspan="1" colspan="1">30.30; 7.52</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">10-III-2009</td><td align="left" rowspan="1" colspan="1">9-67-1</td><td align="left" rowspan="1" colspan="1">JN885945</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-67-4</td><td align="left" rowspan="1" colspan="1">JN885946</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-67-5</td><td align="left" rowspan="1" colspan="1">JN885947</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">W Tazenakht</td><td align="left" rowspan="1" colspan="1">21</td><td align="left" rowspan="1" colspan="1">30.27; 7.39</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">10-III-2009</td><td align="left" rowspan="1" colspan="1">9-66-2</td><td align="left" rowspan="1" colspan="1">JN885943</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-66-3</td><td align="left" rowspan="1" colspan="1">JN885944</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tazenakht</td><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">30.28; 7.22</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">10-III-2009</td><td align="left" rowspan="1" colspan="1">9-65-2</td><td align="left" rowspan="1" colspan="1">JN885941</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-65-5</td><td align="left" rowspan="1" colspan="1">JN885942</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tazenakht</td><td align="left" rowspan="1" colspan="1">23</td><td align="left" rowspan="1" colspan="1">30.36; 7.16</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">28-II-2009</td><td align="left" rowspan="1" colspan="1">9-22-1</td><td align="left" rowspan="1" colspan="1">JN885906</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-22-2</td><td align="left" rowspan="1" colspan="1">JN885907</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Agouine</td><td align="left" rowspan="1" colspan="1">24</td><td align="left" rowspan="1" colspan="1">31.05; 7.17</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">3-III-2009</td><td align="left" rowspan="1" colspan="1">9-30-1</td><td align="left" rowspan="1" colspan="1">JN885912</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-30-2</td><td align="left" rowspan="1" colspan="1">JN885913</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Iriri</td><td align="left" rowspan="1" colspan="1">25</td><td align="left" rowspan="1" colspan="1">30.56; 7.13</td><td align="left" rowspan="1" colspan="1"><italic>B. albengai</italic></td><td align="left" rowspan="1" colspan="1">28-II-2009</td><td align="left" rowspan="1" colspan="1">9-24-1</td><td align="left" rowspan="1" colspan="1">JN885908</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-24-2</td><td align="left" rowspan="1" colspan="1">JN885909</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>H</bold></td><td align="left" rowspan="1" colspan="1">Aid Ben Haddou</td><td align="left" rowspan="1" colspan="1">26</td><td align="left" rowspan="1" colspan="1">31.05; 7.08</td><td align="left" rowspan="1" colspan="1"><italic>B. albengai</italic></td><td align="left" rowspan="1" colspan="1">1-III-2009</td><td align="left" rowspan="1" colspan="1">9-25-1</td><td align="left" rowspan="1" colspan="1">JN885910</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-25-2</td><td align="left" rowspan="1" colspan="1">JN885911</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ouarzazate</td><td align="left" rowspan="1" colspan="1">27</td><td align="left" rowspan="1" colspan="1">30.55; 6.51</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">9-III-2009</td><td align="left" rowspan="1" colspan="1">9-62-2</td><td align="left" rowspan="1" colspan="1">JN885939</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-62-3</td><td align="left" rowspan="1" colspan="1">JN885940</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>I</bold></td><td align="left" rowspan="1" colspan="1">Tata</td><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">29.41; 8.09</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">25-II-2009</td><td align="left" rowspan="1" colspan="1">9-15-1</td><td align="left" rowspan="1" colspan="1">JN885900</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-15-2</td><td align="left" rowspan="1" colspan="1">JN885901</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ouarzazate</td><td align="left" rowspan="1" colspan="1">29</td><td align="left" rowspan="1" colspan="1">30.33; 7.09</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">28-II-2009</td><td align="left" rowspan="1" colspan="1">9-21-1</td><td align="left" rowspan="1" colspan="1">JN885904</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-21-2</td><td align="left" rowspan="1" colspan="1">JN885905</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ouarzazate</td><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">30.51; 6.51</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">4-III-2009</td><td align="left" rowspan="1" colspan="1">9-31-4</td><td align="left" rowspan="1" colspan="1">JN885914</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">SE Ouarzazate</td><td align="left" rowspan="1" colspan="1">31</td><td align="left" rowspan="1" colspan="1">30.48; 6.44</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">4-III-2009</td><td align="left" rowspan="1" colspan="1">9-32-4</td><td align="left" rowspan="1" colspan="1">JN885915</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ait Sarin</td><td align="left" rowspan="1" colspan="1">32</td><td align="left" rowspan="1" colspan="1">30.44; 6.38</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">4-III-2009</td><td align="left" rowspan="1" colspan="1">9-33-1</td><td align="left" rowspan="1" colspan="1">JN885916</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-33-2</td><td align="left" rowspan="1" colspan="1">JN885917</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Rebat</td><td align="left" rowspan="1" colspan="1">33</td><td align="left" rowspan="1" colspan="1">30.44; 6.26</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">5-III-2009</td><td align="left" rowspan="1" colspan="1">9-34-3</td><td align="left" rowspan="1" colspan="1">JN885918</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-34-4</td><td align="left" rowspan="1" colspan="1">JN885919</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Agdz</td><td align="left" rowspan="1" colspan="1">34</td><td align="left" rowspan="1" colspan="1">30.41; 6.26</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">5-III-2009</td><td align="left" rowspan="1" colspan="1">9-Agdz-1</td><td align="left" rowspan="1" colspan="1">JN885956</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-Agdz-2</td><td align="left" rowspan="1" colspan="1">JN885957</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>J</bold></td><td align="left" rowspan="1" colspan="1">Fumzguid</td><td align="left" rowspan="1" colspan="1">35</td><td align="left" rowspan="1" colspan="1">30.04; 6.52</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">27-II-2009</td><td align="left" rowspan="1" colspan="1">9-20-1</td><td align="left" rowspan="1" colspan="1">JN885902</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-20-2</td><td align="left" rowspan="1" colspan="1">JN885903</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>K_a</bold></td><td align="left" rowspan="1" colspan="1">Sidi-Flah</td><td align="left" rowspan="1" colspan="1">36</td><td align="left" rowspan="1" colspan="1">31.00; 6.27</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">9-III-2009</td><td align="left" rowspan="1" colspan="1">9-61-1</td><td align="left" rowspan="1" colspan="1">JN885937</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-61-2</td><td align="left" rowspan="1" colspan="1">JN885938</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Skoura</td><td align="left" rowspan="1" colspan="1">37</td><td align="left" rowspan="1" colspan="1">31.01; 6.29</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">9-III-2009</td><td align="left" rowspan="1" colspan="1">9-60-1</td><td align="left" rowspan="1" colspan="1">JN885936</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Skoura</td><td align="left" rowspan="1" colspan="1">38</td><td align="left" rowspan="1" colspan="1">31.08; 6.21</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">9-III-2009</td><td align="left" rowspan="1" colspan="1">9-59-5</td><td align="left" rowspan="1" colspan="1">JN885935</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">M'Gouna</td><td align="left" rowspan="1" colspan="1">39</td><td align="left" rowspan="1" colspan="1">31.14; 6.06</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">9-III-2009</td><td align="left" rowspan="1" colspan="1">9-58-1</td><td align="left" rowspan="1" colspan="1">JN885934</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">El-Kelaa</td><td align="left" rowspan="1" colspan="1">40</td><td align="left" rowspan="1" colspan="1">31.17; 6.08</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">8-III-2009</td><td align="left" rowspan="1" colspan="1">9-57-5</td><td align="left" rowspan="1" colspan="1">JN885933</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Boumalne de Dades</td><td align="left" rowspan="1" colspan="1">41</td><td align="left" rowspan="1" colspan="1">31.22; 5.57</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">8-III-2009</td><td align="left" rowspan="1" colspan="1">9-53-1</td><td align="left" rowspan="1" colspan="1">JN885929</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-53-2</td><td align="left" rowspan="1" colspan="1">JN885930</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Juniter</td><td align="left" rowspan="1" colspan="1">42</td><td align="left" rowspan="1" colspan="1">31.22; 5.47</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">7-III-2009</td><td align="left" rowspan="1" colspan="1">9-52-1</td><td align="left" rowspan="1" colspan="1">JN885928</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Emsoudar Aitsdrat</td><td align="left" rowspan="1" colspan="1">43</td><td align="left" rowspan="1" colspan="1">31.27; 5.58</td><td align="left" rowspan="1" colspan="1"><italic>B. malhommei</italic></td><td align="left" rowspan="1" colspan="1">8-III-2009</td><td align="left" rowspan="1" colspan="1">9-54-1</td><td align="left" rowspan="1" colspan="1">JN885931</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-54-3</td><td align="left" rowspan="1" colspan="1">JN885932</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>K_b</bold></td><td align="left" rowspan="1" colspan="1">N'Kob</td><td align="left" rowspan="1" colspan="1">44</td><td align="left" rowspan="1" colspan="1">30.51; 5.51</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">4-III-2009</td><td align="left" rowspan="1" colspan="1">9-35-1</td><td align="left" rowspan="1" colspan="1">JN885920</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Tazzarine</td><td align="left" rowspan="1" colspan="1">45</td><td align="left" rowspan="1" colspan="1">30.50; 5.30</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">5-III-2009</td><td align="left" rowspan="1" colspan="1">9-38-2</td><td align="left" rowspan="1" colspan="1">JN885921</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">N Alnif</td><td align="left" rowspan="1" colspan="1">46</td><td align="left" rowspan="1" colspan="1">31.10; 5.13</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">5-III-2009</td><td align="left" rowspan="1" colspan="1">9-40-2</td><td align="left" rowspan="1" colspan="1">JN885922</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>K_c</bold></td><td align="left" rowspan="1" colspan="1">Tinerhir</td><td align="left" rowspan="1" colspan="1">47</td><td align="left" rowspan="1" colspan="1">31.27; 5.36</td><td align="left" rowspan="1" colspan="1"><italic>B. albengai</italic></td><td align="left" rowspan="1" colspan="1">7-III-2009</td><td align="left" rowspan="1" colspan="1">9-51-2</td><td align="left" rowspan="1" colspan="1">JN885927</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Asselab</td><td align="left" rowspan="1" colspan="1">48</td><td align="left" rowspan="1" colspan="1">31.32; 4.40</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">6-III-2009</td><td align="left" rowspan="1" colspan="1">9-44-2</td><td align="left" rowspan="1" colspan="1">JN885923</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-44-3</td><td align="left" rowspan="1" colspan="1">JN885924</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Ait El Farsi</td><td align="left" rowspan="1" colspan="1">49</td><td align="left" rowspan="1" colspan="1">31.22; 5.17</td><td align="left" rowspan="1" colspan="1"><italic>B. draa</italic></td><td align="left" rowspan="1" colspan="1">6-III-2009</td><td align="left" rowspan="1" colspan="1">9-47-1</td><td align="left" rowspan="1" colspan="1">JN885925</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-47-5</td><td align="left" rowspan="1" colspan="1">JN885926</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Oulad Driss</td><td align="left" rowspan="1" colspan="1">50</td><td align="left" rowspan="1" colspan="1">29.49; 5.38</td><td align="left" rowspan="1" colspan="1"><italic>Buthus</italic> spec.</td><td align="left" rowspan="1" colspan="1">16-III-2009</td><td align="left" rowspan="1" colspan="1">9-91-3</td><td align="left" rowspan="1" colspan="1">JN885952</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-91-6</td><td align="left" rowspan="1" colspan="1">JN885953</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>L</bold></td><td align="left" rowspan="1" colspan="1">Draa Valley</td><td align="left" rowspan="1" colspan="1">51</td><td align="left" rowspan="1" colspan="1">30.00; 5.32</td><td align="left" rowspan="1" colspan="1"><italic>Buthus draa</italic></td><td align="left" rowspan="1" colspan="1">16-III-2009</td><td align="left" rowspan="1" colspan="1">9-A14-1</td><td align="left" rowspan="1" colspan="1">JN885954</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-A14-4</td><td align="left" rowspan="1" colspan="1">JN885955</td></tr><tr><td align="left" rowspan="1" colspan="1"><bold>Out</bold></td><td align="left" rowspan="1" colspan="1">Zagora</td><td align="left" rowspan="1" colspan="1">52</td><td align="left" rowspan="1" colspan="1">33.85; 6.29</td><td align="left" rowspan="1" colspan="1"><italic>Androctonus spec.</italic></td><td align="left" rowspan="1" colspan="1">17-III-2009</td><td align="left" rowspan="1" colspan="1">9-88-1</td><td align="left" rowspan="1" colspan="1">JN885949</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">Mesrate</td><td align="left" rowspan="1" colspan="1">53</td><td align="left" rowspan="1" colspan="1">30.03; 5.36</td><td align="left" rowspan="1" colspan="1"><italic>Androctonus</italic> spec.</td><td align="left" rowspan="1" colspan="1">16-III-2009</td><td align="left" rowspan="1" colspan="1">9-89-1</td><td align="left" rowspan="1" colspan="1">JN885950</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">9-89-2</td><td align="left" rowspan="1" colspan="1">JN885951</td></tr></tbody></table></alternatives><table-wrap-foot><fn id="nt101"><label></label><p>Given are the respective genetic groups, the locality names, a running number for each site (coinciding with <xref ref-type="fig" rid="pone-0029403-g001">figure 1</xref>), GPS coordinates of localities, morphological species assignment, date of captures and Genbank accession numbers.</p></fn></table-wrap-foot></table-wrap><table-wrap id="pone-0029403-t002" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0029403.t002</object-id><label>Table 2</label><caption><title>Fst values between populations estimated under the K2P substitution model; significance values (bold, α = 0.05) were assigned using 1000 permutations.</title></caption><alternatives><graphic id="pone-0029403-t002-2" xlink:href="pone.0029403.t002"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col align="left" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col><col align="center" span="1"></col></colgroup><thead><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1">B</td><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">D</td><td align="left" rowspan="1" colspan="1">E</td><td align="left" rowspan="1" colspan="1">F</td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">H</td><td align="left" rowspan="1" colspan="1">I</td><td align="left" rowspan="1" colspan="1">J</td><td align="left" rowspan="1" colspan="1">K</td><td align="left" rowspan="1" colspan="1">L</td></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">A</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">B</td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">C</td><td align="left" rowspan="1" colspan="1">0.975</td><td align="left" rowspan="1" colspan="1">0.984</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">D</td><td align="left" rowspan="1" colspan="1">0.973</td><td align="left" rowspan="1" colspan="1">0.978</td><td align="left" rowspan="1" colspan="1">0.976</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">E</td><td align="left" rowspan="1" colspan="1">0.881</td><td align="left" rowspan="1" colspan="1"><bold>0.903</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.901</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.884</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">F</td><td align="left" rowspan="1" colspan="1">0.680</td><td align="left" rowspan="1" colspan="1"><bold>0.768</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.801</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.706</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.827</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">0.754</td><td align="left" rowspan="1" colspan="1"><bold>0.772</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.768</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.801</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.832</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.722</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">H</td><td align="left" rowspan="1" colspan="1">0.957</td><td align="left" rowspan="1" colspan="1">0.971</td><td align="left" rowspan="1" colspan="1">0.962</td><td align="left" rowspan="1" colspan="1"><bold>0.963</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.887</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.825</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.819</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">I</td><td align="left" rowspan="1" colspan="1">0.813</td><td align="left" rowspan="1" colspan="1"><bold>0.842</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.832</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.821</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.750</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.789</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.801</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.841</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">J</td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1">0.991</td><td align="left" rowspan="1" colspan="1">0.982</td><td align="left" rowspan="1" colspan="1"><bold>0.902</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.834</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.823</bold></td><td align="left" rowspan="1" colspan="1">0.962</td><td align="left" rowspan="1" colspan="1"><bold>0.852</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">K</td><td align="left" rowspan="1" colspan="1">0.797</td><td align="left" rowspan="1" colspan="1"><bold>0.832</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.815</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.821</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.809</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.809</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.804</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.712</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.768</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.797</bold></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">L</td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1">0.992</td><td align="left" rowspan="1" colspan="1">0.982</td><td align="left" rowspan="1" colspan="1"><bold>0.906</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.837</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.814</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.972</bold></td><td align="left" rowspan="1" colspan="1"><bold>0.851</bold></td><td align="left" rowspan="1" colspan="1">1.000</td><td align="left" rowspan="1" colspan="1"><bold>0.840</bold></td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table></alternatives></table-wrap><p>The second dataset included 84 additional sequences obtained from genbank (<xref ref-type="supplementary-material" rid="pone.0029403.s001">appendix SI</xref>). In total this more comprehensive dataset consisted of 179 sequences with 383 bp each. This included 148 variable sites (38.6%), 99 of which were parsimony informative. Many of these sites were uninformative due to a relatively large amount (60 sites) of missing data in genbank sequences.</p><p>Reconstructed trees indicate the presence of 12 distinct <italic>Buthus</italic> lineages, which were recovered in all analyses (<xref ref-type="fig" rid="pone-0029403-g001">Fig. 1a–c</xref>). However, the relationships among groups differed depending on the tree reconstruction method. This is also reflected in the low statistical support for among group relationships. The support for most groups is high. The genetic patterns strongly coincide with the geographical distribution of the respective sampling locations. Lineages that occur sympatrically (K; most probably H/G/I, <xref ref-type="fig" rid="pone-0029403-g001">Figure 1</xref>) are genetically well separated in the trees, while genetically closer groups are all strictly allopatric or parapatric (e.g. I/E; H/K). The distributional patterns of most of the genetic clusters conform to the orographic pattern of the region (e.g. along valleys, southern or northern slopes of mountain systems).</p><fig id="pone-0029403-g001" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0029403.g001</object-id><label>Figure 1</label><caption><title>Maximum Likelihood (ML) (a) and Bayesian Inference (BI) tree (b) rooted with <italic>Androctonus mauretanicus</italic> (N = 1) and <italic>Androctonus</italic> spec. (N = 3) as outgroups.</title><p>Numbers at nodes refer to bootstrap support for ML analyses and posterior probabilities for BI. <xref ref-type="fig" rid="pone-0029403-g001">Figure 1c</xref> shows a Bayesian Inference tree including all <italic>Buthus</italic> COI sequences available from genbank (sequences taken from genbank indicated by stars). For detailed information of sequences taken from genbank see <xref ref-type="supplementary-material" rid="pone.0029403.s001">appendix SI</xref>.</p></caption><graphic xlink:href="pone.0029403.g001"></graphic></fig><p>The comprehensive analyses of genbank data together with our own dataset generally yielded geographically meaningful clusters. The European samples represent two monophyletic groups, one in Portugal and one in France also including Spain. These clades likely correspond to the recently described species, <italic>Buthus montanus</italic> Lourenço & Vachon, 2004 and <italic>Buthus ibericus</italic> Lourenço & Vachon, 2004. However, the European <italic>Buthus</italic> are polyphyletic, yet restricted to one branch of the entire tree. They also make the North African <italic>Buthus</italic> paraphyletic (see <xref ref-type="fig" rid="pone-0029403-g001">Figure 1c</xref>).</p><p>The 12 lineages A through L do not match with current taxonomy (as reflected by morphology based identification keys): (i) <italic>B. elmoutaouakili</italic> Lourenço & Qui, 2006 is found in six lineages, <italic>B. malhommei</italic> Vachon, 1949 in four, <italic>B. albengai</italic> Lourenço, 2003 in two and <italic>B. draa</italic> Lourenço & Slimani, 2004 in four. None of the four species is monophyletic for our marker gene. (ii) Several lineages are composed of different species: four of the five lineages with ten or more individuals analysed contain two to three species; even lineage H (four individuals) includes two different morpho-types.</p></sec><sec id="s3"><title>Discussion</title><p>The two tree hypotheses (ML and BI) constructed from our original dataset distinguish 12 genetic lineages. One of these clusters (lineage K) is further structured into three sub-clades. The genetic clusters exhibit a clear geographic pattern with all the lineages being confined to relatively restricted areas. None of the lineages is widely distributed. Most lineages are separated from each other by geographic barriers such as high mountain ranges or steep valleys. Only some basal lineages show range overlap (e.g. K) while all younger groups are strictly allopatric or parapatric (e.g. the three sub-groups of K). Four groups (A–D) are situated around the western and north-western parts of the High Atlas. One group is located in the westernmost Antiatlas (E), three others at the northern foothills of these mountains (F–H) and one at the southern foothills of the Antiatlas (J). The largest cluster (K) is distributed south of the High Atlas, east of the Draa valley, and splits further into three subclades, which form distinct strictly parapatric groups following the orographic structures of this region (e.g. along the Dades valley or restricted to the Jebel Sahro). Another cluster (I) extends from the Draa valley to the south western edge of the Antiatlas, probably including parts of the distribution ranges of clade G and H.</p><sec id="s3a"><title>Barriers and retreats</title><p>The geographic distribution of the obtained genetic clusters strongly supports significant impact of orographic structures as dispersal barriers. The extant distribution of <italic>Buthus</italic> scorpions mostly does not exceed elevations higher than 2500 m asl <xref ref-type="bibr" rid="pone.0029403-Brownell1">[23]</xref>, and individuals show low dispersal ability. Thus, high mountain ranges likely acted (and act) as strong barriers to gene flow resulting in genetically distinct clusters found within the Atlas Mountains. This is reflected in our data by fine structured genetic lineages of the sampled <italic>Buthus</italic> individuals. A similar pattern was observed for the same taxonomic group using various molecular markers such as allozymes, nuclear and mitochondrial DNA all of which showed significant genetic differentiation patterns even among local populations <xref ref-type="bibr" rid="pone.0029403-Yamashita1">[24]</xref>, <xref ref-type="bibr" rid="pone.0029403-Gantenbein3">[25]</xref>. Similar results were found for other organisms with low to moderate mobility <xref ref-type="bibr" rid="pone.0029403-Fritz1">[7]</xref>. These extremely sedentary species apparently diverged into several narrowly distributed genetic clusters, restricted to the slopes of mountains and river valleys (e.g. the Ourika and Draa). These lineages are found at different altitudes and might be adapted to various ecological niches. We assume that all actual distribution ranges of <italic>Buthus</italic> lineages also include their last centre of differentiation. Thus, our study area may comprise 12 allopatric refugia during the last ice age revealing a much more fine grained biogeographical sub-structure than previously observed in the Maghreb e.g. <xref ref-type="bibr" rid="pone.0029403-Fritz1">[7]</xref>–<xref ref-type="bibr" rid="pone.0029403-Habel1">[9]</xref>. Consequently, the biogeographical structuring of <italic>Buthus</italic> in Morocco is much more complex than the structures observed in other Mediterranean sub-centres <xref ref-type="bibr" rid="pone.0029403-Schmitt2">[26]</xref>, most probably as a consequence of strong gene flow barriers represented by high mountain ranges and a more constant climatic situation in North Africa compared to the northern part of the Mediterranean. The combination of our data with previously analysed <italic>Buthus</italic> individuals (<xref ref-type="fig" rid="pone-0029403-g001">Figure 1c</xref>) underlines this superposition of the Maghreb region as a speciation and differentiation centre: The European <italic>Buthus</italic> are polyphyletic, but restricted to one branch of the entire tree and make the North African <italic>Buthus</italic> paraphyletic. This may be indicative of multiple colonisation events from North Africa to Europe.</p></sec><sec id="s3b"><title>The trigger of genetic differentiation</title><p>This distribution of genetic lineages supports an allopatric differentiation scenario as opposed to sympatric speciation processes <xref ref-type="bibr" rid="pone.0029403-Barraclough1">[27]</xref>: The lineages originating from the oldest vicariance events are most strongly differentiated from each other. Perhaps this long period of time elapsed since the onset of differentiation allowed for the evolution of characteristics enabling sympatric occurrence of representatives of different genetic groups. Furthermore, the close geographic proximity among the well differentiated lineages G, H and I suggests some areas of range overlap among them. If reproductive barriers are in effect, species might even occur in sympatry without niche differentiation when resources are abundant. Thus, the time elapsed since the onset of the evolution of these lineages must have been sufficient that neither the inter-lineage competition for space nor the strong stenotopy of <italic>Buthus</italic> scorpions has been able to prevent the geographic intermixing of these lineages. However, differentiation among them still might not be sufficient for syntopic occurrences so that these groups exclude each other at each single locality. Such a scenario of competitive exclusion would reinforce a strictly geography-dependent distribution pattern of lineages.</p></sec><sec id="s3c"><title>Discordance between morphospecies and COI phylogeny</title><p>In contrast to the stringent geographic pattern, our findings are not congruent with the current taxonomic classification. The different species identified based on morphological characters often cluster within different genetic lineages which, in turn, often comprise more than one species. In addition, many specimens could not be identified due to incomplete or doubtful identification keys, and a lack of diagnostic traits. Hence, the current taxonomy of North African <italic>Buthus</italic> species is in need of revision. Additional diagnostic characters must be identified and new species keys be developed. However, to actually define the status of the detected genetic lineages, further morphological, behavioural and ecological studies are necessary. Besides, it is well known that gene trees and species trees may differ, e. g. due to shared ancestral polymorphisms, incomplete phylogenetic lineage sorting and introgression <xref ref-type="bibr" rid="pone.0029403-Pollard1">[28]</xref>. Evidence of recombination of mtDNA has been found in <italic>Buthus</italic>, which could also influence our results <xref ref-type="bibr" rid="pone.0029403-Gantenbein4">[29]</xref>. Taking our results as an exclusive basis for taxonomical conclusions would therefore doubtlessly be premature. In future studies, the addition of nuclear markers is indispensable to solve the question how prominent hybridization and introgression are among these <italic>Buthus</italic> lineages and species.</p></sec></sec><sec sec-type="materials|methods" id="s4"><title>Materials and Methods</title><sec id="s4a"><title>Sampling and identification</title><p>Most specimens were collected at daylight under stones. Overnight collections were facilitated by black light as all scorpions fluoresce in UV light due to a specific protein in their exoskeleton <xref ref-type="bibr" rid="pone.0029403-Anglade1">[30]</xref>. Collected specimens were stored in absolute ethanol until DNA extraction. All sampling sites are shown in <xref ref-type="fig" rid="pone-0029403-g002">Figure 2</xref> and listed in <xref ref-type="table" rid="pone-0029403-t001">Table 1</xref> which also gives details on the sampling date and location. Species were identified with keys and additional species descriptions <xref ref-type="bibr" rid="pone.0029403-Loureno1">[10]</xref>, <xref ref-type="bibr" rid="pone.0029403-Stockmann1">[12]</xref>, <xref ref-type="bibr" rid="pone.0029403-Vachon1">[31]</xref>–<xref ref-type="bibr" rid="pone.0029403-Vachon3">[33]</xref> based on typical morphological characters used for scorpion identification <xref ref-type="bibr" rid="pone.0029403-Vachon2">[cf. 32]</xref>–<xref ref-type="bibr" rid="pone.0029403-Vachon3">[33]</xref>. All species identifications were performed by the same author (IS).</p><fig id="pone-0029403-g002" position="float"><object-id pub-id-type="doi">10.1371/journal.pone.0029403.g002</object-id><label>Figure 2</label><caption><title>Geographical distribution of all sampling sites and the 12 genetic groups and three subgroups around the Atlas Mountains in Morocco.</title><p>Numbers of sampling sites coincide with numbers given in <xref ref-type="table" rid="pone-0029403-t001">Table 1</xref>. Letters of the groups correspond with <xref ref-type="fig" rid="pone-0029403-g001">Figure 1</xref>. The geographic coordinates of sampling sites are given in <xref ref-type="table" rid="pone-0029403-t001">Table 1</xref>. Abbreviations: SV: Souss valley, DV: Draa valley, DaV: Dades valley, ZV: Ziz valley. Country border between Morocco and Algeria is indicated by a dotted line.</p></caption><graphic xlink:href="pone.0029403.g002"></graphic></fig></sec><sec id="s4b"><title>PCR and sequencing</title><p>For all individuals, DNA was isolated from leg, caudal segment or telson muscle tissue using the Qiagen DNeasy kit with standard protocols for tissue samples. A 452 bp fragment of the mitochondrial Cytochrome Oxidase I (<italic>COI</italic>) gene was amplified using standard PCR procedures with the following primers: forward <named-content content-type="gene">5′ GGT CAA CAA ATC ATA AAG ATA TTG G 3′</named-content>, reverse <named-content content-type="gene">5′ TAA ACT TCA GGG TGA CCA AAA AAT CA 3′</named-content><xref ref-type="bibr" rid="pone.0029403-Folmer1">[34]</xref>. PCRs were performed in 20 µl volumes: 10 µl Mastermix (Thermozyme), 0.2 µl of each Primer (1 µM), 4.6 µl PCR grade water and 5 µl DNA template. The cycle programme comprised an activation step at 94°C for 4 min, followed by 40 cycles of 30 sec denaturation at 94°C, 30 sec annealing at 45°C and 1 min elongation at 72°C. Cycling was terminated by a final extension step at 72°C for 10 min. Amplicons were subsequently purified and sequenced in both directions on an automated sequencer (3730xl DNA Analyzer; Applied Biosystems, Carlsbad, CA, USA) at the University of Kiel. All sequences are deposited at the NCBI genbank (<xref ref-type="supplementary-material" rid="pone.0029403.s001">appendix SI</xref>).</p></sec><sec id="s4c"><title>Data analysis</title><p>We included 91 sequences of <italic>Buthus</italic>, each 452 bp in length, in our analyses. Four sequences of the Buthid genus <italic>Androctonus</italic> served as outgroup (one sequence from <italic>A. mauretanicus</italic> obtained from genbank: accession JF820097 and three sequences from <italic>Androctonus</italic> spec. (09-88-1; 09-89-1; 09-89-2)). Sequences were inspected and aligned using Geneious 5.0.3 <xref ref-type="bibr" rid="pone.0029403-Drummond1">[35]</xref>.</p><p>General alignment statistics (diversity indices, etc.) were calculated using DNAsp v. 5.10 <xref ref-type="bibr" rid="pone.0029403-Librado1">[36]</xref>. Estimates of population divergence were calculated as pairwise Fst under a Kimura2P substitution model in Arlequin v.3.5.1.2. <xref ref-type="bibr" rid="pone.0029403-Excoffier1">[37]</xref>. Populations were defined a posteriori according to our phylogenetic results. Significance of Fst values was tested using 1000 permutations and a significance level of 0.05.</p><p>The best fitting model of sequence evolution for phylogenetic tree reconstruction was chosen using MrModeltest 2.3 <xref ref-type="bibr" rid="pone.0029403-Nylander1">[38]</xref> in PAUP 4.0 b10 <xref ref-type="bibr" rid="pone.0029403-Swofford1">[39]</xref> and was determined to be the GTR+I+G. Maximum Likelihood (ML) trees were constructed using MEGA version 5.01 <xref ref-type="bibr" rid="pone.0029403-Tamura1">[40]</xref>; 1000 bootstrap replicates were performed to obtain a statistical measure for branch support (<xref ref-type="fig" rid="pone-0029403-g001">Figure 1a</xref>). Additionally, we used the Bayesian algorithm implemented in MrBayes 3.1.2 <xref ref-type="bibr" rid="pone.0029403-Huelsenbeck1">[41]</xref> to obtain a second phylogenetic inference and evaluate the stability of our tree reconstruction (<xref ref-type="fig" rid="pone-0029403-g001">Figure 1b</xref>). Using the obtained substitution model we ran the Markov chain Monte Carlo algorithm with four chains and two independent runs for 20 million generations. Trees were sampled every 2,000 generations; the first 10% of generated trees were discarded as burn-in as recommended by Tracer v1.5 <xref ref-type="bibr" rid="pone.0029403-Rambaut1">[42]</xref>. The remaining trees were used to generate a consensus tree.</p><p>In a second analysis, we included 84 sequences from <italic>Buthus</italic> scorpions obtained from genbank (see <xref ref-type="supplementary-material" rid="pone.0029403.s001">appendix SI</xref>) resulting in a total of 175 ingroup sequences. However, due to only partial overlap the alignment was reduced to 383 bp in length. Bayesian analyses of this dataset were performed as described above. While these additional analyses provide further insights into geographic structuring of genetic divergence, they do not help in evaluating species identity and monophyly (<xref ref-type="fig" rid="pone-0029403-g001">Figure 1c</xref>). When these sequences were submitted to Genbank, the vast majority of Moroccan <italic>Buthus</italic> species known today had not yet been described and most Moroccan <italic>Buthus</italic> were characterized as subspecies of <italic>B. occitanus</italic> (<italic>B. o. occitanus</italic> – Europe, <italic>B. o. mardochei</italic> – Morocco, <italic>B. o. tunetanus</italic> – Tunisia) <xref ref-type="bibr" rid="pone.0029403-Vachon1">[31]</xref>.</p></sec></sec><sec sec-type="supplementary-material" id="s5"><title>Supporting Information</title><supplementary-material content-type="local-data" id="pone.0029403.s001"><label>Appendix SI</label><caption><p><bold>DNA sequences obtained from NCBI genbank.</bold></p><p>(DOC)</p></caption><media xlink:href="pone.0029403.s001.doc" mimetype="application" mime-subtype="msword"><caption><p>Click here for additional data file.</p></caption></media></supplementary-material></sec></body><back><fn-group><fn fn-type="conflict"><p><bold>Competing Interests: </bold>The authors have declared that no competing interests exist.</p></fn><fn fn-type="financial-disclosure"><p><bold>Funding: </bold>Funding was performed by the “Fonds National de la Recherche Luxembourg” and the “German Academic Exchange Service”; furthermore, the Natural History Museum Luxembourg supported field work and laboratory analyses. 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