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   <front>
      <journal-meta>
         <journal-id journal-id-type="publisher-id">COLLBOT</journal-id>
         <journal-title-group>
            <journal-title specific-use="original">Collectanea Botanica</journal-title>
            <abbrev-journal-title abbrev-type="publisher">Collect. Bot.</abbrev-journal-title>
         </journal-title-group>
         <issn publication-format="electronic">1989-1067</issn>
         <issn-l>0010-0730</issn-l>
         <publisher>
            <publisher-name>Consejo Superior de Investigaciones Cient&#x00ED;ficas</publisher-name>
            <publisher-loc>
               <country>Espa&#x00F1;a</country>
            </publisher-loc>
         </publisher>
      </journal-meta>
      <article-meta>
         <article-id pub-id-type="doi">10.3989/collectbot.2024.v43.0004</article-id>
         <article-id pub-id-type="publisher-id">collectbot.2024.v43.0004</article-id>
         <article-categories>
            <subj-group subj-group-type="heading">
               <subject>Art&#x00ED;culos</subject>
            </subj-group>
         </article-categories>
         <title-group>
            <article-title>
               <italic toggle="yes">Sempervivum druentianum</italic> (Crassulaceae subfam. Sempervivoideae), a new endemic species discovered in the Durance Valley (D&#x00E9;p. Hautes-Alpes, Region Provence-Alpes-C&#x00F4;te d&#x2019;Azur, France), and description of a new nothospecies</article-title>
            <trans-title-group xml:lang="es">
               <trans-title>
                  <italic toggle="yes">Sempervivum druentianum</italic> (Crassulaceae subfam. Sempervivoideae), nueva especie end&#x00E9;mica descubierta en el Valle del Durance (Dpto. Altos Alpes, Regi&#x00F3;n Provenza-Alpes-Costa AzuL, Francia), y descripci&#x00F3;n de una nueva notospecie</trans-title>
            </trans-title-group>
            <alt-title alt-title-type="running-head">
               <italic toggle="yes">Sempervivum druentianum,</italic> a new endemic species in France  </alt-title>
         </title-group>
         <contrib-group>
            <contrib contrib-type="author" corresp="yes">
               <contrib-id contrib-id-type="orcid" authenticated="false">https://orcid.org/0000-0001-6884-0805</contrib-id>
               <name name-style="western">
                  <surname>Gallo</surname>
                  <given-names>Lorenzo</given-names>
               </name>
               <xref ref-type="corresp" rid="corr-1-e004"/>
               <xref ref-type="author-notes" rid="fn-1-e004"/>
            </contrib>
         </contrib-group>
         <author-notes>
            <corresp id="corr-1-e004">Email: L. Gallo <email xlink:href="lorenzogallo1959@hotmail.it">lorenzogallo1959@hotmail.it</email>
            </corresp>
            <fn id="fn-1-e004">
               <p>Strada Val San Martino sup. 194, Torino 10131, Italy</p>
            </fn>
            <fn id="fn-2-e004" fn-type="edited-by">
               <p>Editor: M. Bonifacino</p>
            </fn>
         </author-notes>
         <pub-date date-type="pub"
                   publication-format="electronic"
                   iso-8601-date="2024-12-30">
            <day>30</day>
            <month>12</month>
            <year>2024</year>
         </pub-date>
         <pub-date date-type="collection"
                   publication-format="electronic"
                   iso-8601-date="2024-12-30">
            <day>30</day>
            <month>12</month>
            <year>2024</year>
         </pub-date>
         <volume>43</volume>
         <elocation-id>e004</elocation-id>
         <pub-history>
            <event>
               <event-desc>Received</event-desc>
               <date date-type="received" iso-8601-date="2023-04-05">
                  <day>05</day>
                  <month>04</month>
                  <year>2023</year>
               </date>
            </event>
            <event>
               <event-desc>accepted</event-desc>
               <date date-type="accepted" iso-8601-date="2024-05-14">
                  <day>14</day>
                  <month>05</month>
                  <year>2024</year>
               </date>
            </event>
            <event>
               <event-desc>published on line</event-desc>
               <date date-type="pub" iso-8601-date="2025-01-27">
                  <day>27</day>
                  <month>01</month>
                  <year>2025</year>
               </date>
            </event>
         </pub-history>
         <permissions>
            <copyright-statement>&#x00A9; 2024 CSIC</copyright-statement>
            <copyright-year>2024</copyright-year>
            <copyright-holder>CSIC</copyright-holder>
            <ali:free_to_read/>
            <license license-type="open-access"
                     xlink:href="https://creativecommons.org/licenses/by/4.0/">
               <ali:license_ref>https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
			   <license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.</license-p>
            </license>
         </permissions>
         <self-uri xlink:href="XXXXXXXXXXXXXXXXXXXXXX"/>
         <abstract>
            <title>Abstract</title>
            <p>A new species of <italic toggle="yes">Sempervivum</italic> was discovered in southern France and is here named <italic toggle="yes">Sempervivum druentianum</italic>. This new species, which has habitus like that of <italic toggle="yes">S. tectorum</italic>, is mainly characterised by leaves that are always covered with glandular trichomes and carpels that are glandular on the back. To date, only one population is known, but true or alleged hybrids with other <italic toggle="yes">Sempervivum</italic> species have been discovered in the surroundings of the type locality of <italic toggle="yes">S. druentianum</italic>; one of these, <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic> (= <italic toggle="yes">S. arachnoideum</italic> &#x00D7; <italic toggle="yes">S. druentianum</italic>), is also present and described here. A key to all taxa belonging to the <italic toggle="yes">S. tectorum</italic> L. group located in southern France is provided.</p>
         </abstract>
         <trans-abstract xml:lang="es">
            <title>Resumen</title>
            <p>Una nueva especie de <italic toggle="yes">Sempervivum</italic> fue descubierta en el sur de Francia y se denomina aqu&#x00ED; <italic toggle="yes">Sempervivum druentianum</italic>. Esta nueva especie, que tiene un h&#x00E1;bito como el de <italic toggle="yes">S. tectorum</italic>, se caracteriza principalmente por hojas que siempre est&#x00E1;n cubiertas de tricomas glandulares y carpelos que son glandulares en el env&#x00E9;s. Hasta la fecha, solo se conoce una poblaci&#x00F3;n, pero se han descubierto h&#x00ED;bridos verdaderos o supuestos con otras especies de <italic toggle="yes">Sempervivum</italic> en los alrededores de la localidad tipo de <italic toggle="yes">S. druentianum</italic>; una de ellas, <italic toggle="yes">S.</italic> &#x00D7;<italic toggle="yes">brigantiacum</italic> (= <italic toggle="yes">S. arachnoideum</italic> &#x00D7; <italic toggle="yes">S. druentianum</italic>), tambi&#x00E9;n est&#x00E1; presente y descrita aqu&#x00ED;. Se proporciona una clave para todos los taxones pertenecientes al grupo <italic toggle="yes">S. tectorum</italic> L. ubicado en el sur de Francia.</p>
         </trans-abstract>
         <kwd-group>
            <kwd>France</kwd>
            <kwd>S. arachnoideum</kwd>
            <kwd>S. druentianum</kwd>
            <kwd>S. &#x00D7;brigantiacum</kwd>
            <kwd>S. gr. tectorum</kwd>
            <kwd>taxonomy</kwd>
         </kwd-group>
         <kwd-group xml:lang="es">
            <kwd>Francia</kwd>
            <kwd>S. arachnoideum</kwd>
            <kwd>S. druentianum</kwd>
            <kwd>S. &#x00D7;brigantiacum</kwd>
            <kwd>S. gr. tectorum</kwd>
            <kwd>taxonom&#x00ED;a</kwd>
         </kwd-group>
         <counts>
            <fig-count count="5"/>
            <table-count count="2"/>
            <equation-count count="0"/>
            <ref-count count="24"/>
            <page-count count="11"/>
         </counts>
      </article-meta>
   </front>
   <body>
      <sec sec-type="intro" id="sec-1-e004">
         <title>INTRODUCTION</title>
         <p>The genus <italic toggle="yes">Sempervivum</italic> L. <italic toggle="yes">s. str.</italic> (excl. <italic toggle="yes">Jovibarba</italic> Opiz) is subendemic to the European continent and was most recently recognised as consisting of 40 to 60 species (Gowler &#x0026; Tebbitt, <xref rid="ref-11-e004" ref-type="bibr">1995</xref>; &#x2018;t Hart <italic toggle="yes">et al.</italic>, <xref rid="ref-22-e004" ref-type="bibr">2003</xref>; Thiede &#x0026; Eggli, <xref rid="ref-23-e004" ref-type="bibr">2007</xref>). The type of this genus is <italic toggle="yes">S. tectorum</italic> L., established by Parnell (<xref rid="ref-16-e004" ref-type="bibr">1993</xref>: 87), with a lectotype designated from the Clifford herbarium (BM). Unfortunately, the proposed lectotype represents <italic toggle="yes">S. &#x00D7;fauconnetii</italic> Reut.; therefore, the conservation of the name with a conserved type was proposed (Gallo &#x0026; Jarvis, <xref rid="ref-9-e004" ref-type="bibr">2016</xref>: 1431) and recommended (Applequist, <xref rid="ref-2-e004" ref-type="bibr">2019</xref>: 849). The conservation of the name has facilitated the clarification of the taxonomic application of the binomial <italic toggle="yes">S. tectorum</italic>, which, in some cases, had been used too frequently to identify different taxa (e.g. <italic toggle="yes">S. alpinum</italic> Griseb. &#x0026; Schenk, <italic toggle="yes">S. riccii</italic> Iberite &#x0026; Anzal. and, perhaps, <italic toggle="yes">S. boutignyanum</italic> Billot &#x0026; Gren. in F. Schultz) while in other cases obvious synonymies were not implemented, as in the case of <italic toggle="yes">S. arvernense</italic> Lecoq &#x0026; Lamotte and <italic toggle="yes">S. andreanum</italic> Wale.</p>
         <p>The <italic toggle="yes">S. tectorum</italic> group represents a morphologically variable taxon, which is distributed over a very wide range, including particularly central-western Europe where it is almost always spontaneous, while in eastern Europe it is frequently cultivated and more or less naturalized (&#x2018;t Hart <italic toggle="yes">et al</italic>., <xref rid="ref-22-e004" ref-type="bibr">2003</xref>). Research carried out to date has already made it possible to understand, at least in part, the taxonomic complexity of this group but much work remains to be done especially in the westernmost part of its range. This study contributes to a better understanding of the populations of <italic toggle="yes">Sempervivum</italic> in southern France and presents the results of a field-based study of a population of <italic toggle="yes">Sempervivum</italic> discovered near Champcella, in the Durance valley (D&#x00E9;p. Hautes-Alpes, Region Provence-Alpes-C&#x00F4;te d&#x2019;Azur, France) that was, morphologically and ecologically attributable, at first sight, to <italic toggle="yes">S. tectorum</italic>. A more in-depth study has instead revealed that this population is characterized by peculiar morphological features and a precise geographical distribution beyond that of the <italic toggle="yes">S. tectorum</italic>. This material is described as a new species, named <italic toggle="yes">S. druentianum</italic> L. Gallo.</p>
         <p>Several of the morphological traits defining <italic toggle="yes">S. druentianum</italic> are unique and differentiate this new taxon from what is currently interpreted as <italic toggle="yes">S. tectorum</italic>. Especially the glandular pubescence on both the surfaces of the rosette leaves has been observed to be consistently present in the case of <italic toggle="yes">S. druentianum</italic>. The trait is reminiscent of the leaf vestiture found in <italic toggle="yes">S. riccii</italic>, a species endemic of central Italy, from which <italic toggle="yes">S. druentianum</italic> can be distinguished by the glandular nature of carpels on abaxial surface vs (as in <italic toggle="yes">S. riccii</italic>) (<xref rid="taw-1-e004" ref-type="table">Table 1</xref>).</p>
         <p>The hybrid between <italic toggle="yes">S. arachnoideum</italic> L and <italic toggle="yes">S. druentianum</italic>, which is present at the type locality of <italic toggle="yes">S. druentianum</italic>, is here described as <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic> L. Gallo. An identification key is presented for morphologically similar taxa in southern of France.</p>
         <table-wrap id="taw-1-e004" position="float" orientation="portrait">
            <label>Table 1</label>
            <caption>
               <title>Comparative morphology of <italic toggle="yes">Sempervirum tectorum</italic> L. group.</title>
            </caption>
            <table id="tab-1-e004"
                   frame="border"
                   rules="all"
                   width="90&#x0025;">
               <thead>
                  <tr>
                     <th style="width:76.6pt;text-align:center;vertical-align:top;"
                         rowspan="1"
                         colspan="1">
                        <bold>Taxa/Traits</bold>
                     </th>
                     <th style="width:80.3pt;text-align:center;" rowspan="1" colspan="1">
                        <bold>
                           <italic toggle="yes">Sempervirum tectorum</italic> L. (incl. <italic toggle="yes">Sempervirum andreanum</italic> Wale, <italic toggle="yes">Sempervirum arvernense</italic> Lecoq &#x0026; Lamotte)</bold>
                     </th>
                     <th style="width:82pt;text-align:center;vertical-align:top;"
                         rowspan="1"
                         colspan="1">
                        <bold>
                           <italic toggle="yes">Sempervirum boutignyanum</italic> Billot &#x0026; Gren.</bold>
                     </th>
                     <th style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;text-align:center;vertical-align:top;"
                         rowspan="1"
                         colspan="1">
                        <bold>
                           <italic toggle="yes">Sempervirum druentianum</italic> L. Gallo</bold>
                     </th>
                     <th style="width:74.1pt;text-align:center;vertical-align:top;"
                         rowspan="1"
                         colspan="1">
                        <bold>
                           <italic toggle="yes">Sempervirum riccii</italic> Iberite &#x0026; Anzal.</bold>
                     </th>
                     <th style="width:72.95pt;text-align:center;vertical-align:top;"
                         rowspan="1"
                         colspan="1">
                        <bold>
                           <italic toggle="yes">Sempervirum alpinum</italic> Griseb. &#x0026; Schenk</bold>
                     </th>
                  </tr>
               </thead>
               <tbody>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Leaf induments</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">Glabrous</td>
                     <td style="width:82pt;" rowspan="1" colspan="1">Glabrous</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">Glandular</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1">Glandular</td>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Glabrous</td>
                  </tr>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Leaf base color</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">White</td>
                     <td style="width:82pt;" rowspan="1" colspan="1">White</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">White</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1">White</td>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Violet / Purple</td>
                  </tr>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Inflorescence shape</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">Scorpioid</td>
                     <td style="width:82pt;" rowspan="1" colspan="1">Condensed</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">Branched but not scorpioid</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1"/>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Condensed</td>
                  </tr>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Petals color</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">White with stripe pink</td>
                     <td style="width:82pt;" rowspan="1" colspan="1">Green or white with strip pink</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">White with central green strip, base pink</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1">Pink with white margins</td>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Pink with sometimes stripe white</td>
                  </tr>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Filaments indument</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">Glandular </td>
                     <td style="width:82pt;" rowspan="1" colspan="1">Glabrous or with some papillae</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">Glabrous or glandular</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1">Glandular</td>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Always glandular</td>
                  </tr>
                  <tr>
                     <td style="width:76.6pt;" rowspan="1" colspan="1">
                        <bold>Carpels induments</bold>
                     </td>
                     <td style="width:80.3pt;" rowspan="1" colspan="1">Glandular but naked on the back</td>
                     <td style="width:82pt;" rowspan="1" colspan="1">Glandular but naked on the back</td>
                     <td style="width:67.05pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                         rowspan="1"
                         colspan="1">Glandular throughout</td>
                     <td style="width:74.1pt;" rowspan="1" colspan="1">Glandular</td>
                     <td style="width:72.95pt;" rowspan="1" colspan="1">Glandular but naked on the back</td>
                  </tr>
               </tbody>
            </table>
         </table-wrap>
      </sec>
      <sec sec-type="materials&#x007C;methods" id="sec-2-e004">
         <title>MATERIALS AND METHODS</title>
         <p>Comparative morphological, ecological and biological (protandry and protogyny) studies that compare <italic toggle="yes">S. druentianum</italic> with related taxa present in southern France (<italic toggle="yes">S. alpinum</italic>, <italic toggle="yes">S. calcareum</italic> Jord. and <italic toggle="yes">S. tectorum</italic>) were carried out in the field and in the private collection of the author for three years. In addition to the specialised literature on the genus <italic toggle="yes">Sempervivum</italic> as well as French floristic bibliography pertaining to the study area, the more than 200 names that have been included in the synonymys of <italic toggle="yes">S. tectorum s. l.</italic> (L. Gallo, ined.) were considered to rule out the possibility that <italic toggle="yes">S. druentianum</italic> had already been described earlier.</p>
      </sec>
      <sec sec-type="results" id="sec-3-e004">
         <title>RESULTS (TAXONOMY)</title>
         <p>Neither the literature survey not the study of the more than 200 synonyms attributable to <italic toggle="yes">S. tectorum s. l.</italic> indicated any data associated with &#x00D7;<italic toggle="yes">S. druentianum.</italic> The population of <italic toggle="yes">Sempervivum</italic> of Champcella is therefore not yet known and is here described.</p>
         <p>
            <bold>
               <italic toggle="yes">Sempervivum druentianum</italic>
            </bold> L. Gallo <bold>spec. nov.</bold> (<xref rid="fig-1-e004" ref-type="fig">Fig. 1</xref>)</p>
         <fig id="fig-1-e004" position="float" orientation="portrait">
            <label>Figure 1</label>
            <caption>
               <title>
                  <italic toggle="yes">Sempervivum druentianum</italic> L. Gallo sp. nov.: (A), plants in habitat; (B), habitat at the type-locality of Champcella (D&#x00E9;p. Hautes-Alpes. Region Provence-Alpes-C&#x00F4;te d&#x2019;Azur. France); (C), close-up of the rosettes; (D), putative hybrid <italic toggle="yes">S. druentianum</italic> &#x00D7; <italic toggle="yes">S. tectorum</italic> L. (not in the type-locality); (E), close-up of an inflorescence.</title>
            </caption>
            <graphic xlink:href="e004_001.jpeg"
                     position="anchor"
                     orientation="portrait"
                     id="gra-1-e004"/>
         </fig>
         <p>
            <italic toggle="yes">Type</italic>: France, Provence-Alpes-C&#x00F4;te d&#x2019;Azur, D&#x00E9;p. Hautes-Alpes, between Le Chambon and Chanteloube, neighborhood of Champcella, before the place named &#x201C;La Bourgea&#x201D;, on the side of the D38, 986 m a.s.l., rocks covered with xerophilous vegetation, 44.711056 N, 06.587453 E, 22-30.VI.2019, <italic toggle="yes">L. Gallo</italic> &#x0026; <italic toggle="yes">M. Gai</italic> (holotype: MRSN-12989) (<xref rid="fig-2-e004" ref-type="fig">Fig. 2</xref>).</p>
         <fig id="fig-2-e004" position="float" orientation="portrait">
            <label>Figure 2</label>
            <caption>
               <title>Holotype of <italic toggle="yes">Sempervivum druentianum</italic> L. Gallo (MRSN-12989) (photograph: F. Longo).</title>
            </caption>
            <graphic xlink:href="e004_002.jpeg"
                     position="anchor"
                     orientation="portrait"
                     id="gra-2-e004"/>
         </fig>
         <p>
            <italic toggle="yes">Diagnosis</italic>: <italic toggle="yes">Sempervivum druentianum</italic> is very similar to <italic toggle="yes">S. tectorum</italic> but differs in its leaves being glandular on both surfaces (vs. glabrous in <italic toggle="yes">S. tectorum</italic>), and oblanceolate to sometimes rhombeus (<italic toggle="yes">cf</italic>. Stearn <xref rid="ref-21-e004" ref-type="bibr">1987</xref>: 320, 493) (vs. oblanceolate to linear in <italic toggle="yes">S. tectorum</italic>) and the carpels more or less glandular on the back (vs. glabrous in <italic toggle="yes">S. tectorum</italic>). Other peculiar traits are inconsistently present, including that the filaments are sometimes glabrous at the base and not hairy as in <italic toggle="yes">S. tectorum</italic> and <italic toggle="yes">S. alpinum</italic>, and the inflorescence branches of <italic toggle="yes">S. druentianum</italic> are not scorpioid &#x005B;these are morphological characters that are present in natural populations and cannot be overlooked. We must be honest and say that there are objective difficulties in determining them&#x005D;.</p>
         <p>
            <italic toggle="yes">Description</italic>: rosette (20)30-60 mm diameter. Leaf erect &#x00B1; oblanceolate-pyriform (pear-shaped), light green with white base, apex brown-violet apex, 25-35 &#x00D7; 10-14 mm, glandular on both the faces and on the margins. Inflorescence (8)15-25 cm height, branched but not scorpioid, often reddish, with rare bracts; the cauline leaves are similar to rosette leaves, green often reddish, glandular, with long cilia and apex violet. Flowers round in bud, sessile or with very short pedicel (&#x003C;1 mm), 20 mm of diameter. Petals 13-14, patent, linear, 6-9 &#x00D7; 1.0-1.5 mm, white with central green strip, base pink, sometimes pink-spotted adaxially. Filaments violet, glabrous or glandular. Anthers oval, not mucronate, bronze or violet. Carpels green, glandular throughout. Stylus erect, green, sometimes spotted of violet toward the apex.</p>
         <p>
            <italic toggle="yes">Area and habitat</italic>: to date, only a single population of <italic toggle="yes">S. druentianum</italic> is known in the D&#x00E9;partement Hautes-Alpes (Region Provence-Alpes-C&#x00F4;te d&#x2019;Azur), on the hills near the Durance River at Champcella, between Le Chambon and Chanteloube (St. Cr&#x00E9;pin) (<xref rid="fig-3-e004" ref-type="fig">Fig. 3</xref>) at about 1000 m a.s.l., on huge rounded limestone rocks. This locality was strongly affected by the periods of glaciations/deglaciations in the Alps, with a permanence of ice up to 20,000 BP (Jorda <italic toggle="yes">et al.</italic>, <xref rid="ref-13-e004" ref-type="bibr">2000</xref>; Buoncristiani &#x0026; Campy, <xref rid="ref-6-e004" ref-type="bibr">2004</xref>; Rosique, <xref rid="ref-19-e004" ref-type="bibr">2004</xref>). In the last glacial maximum (LGM), on the French side, the alpine glacier occupied an elevation between 1200 and 400 m a.s.l. with widespread diffusion in the Durance valley and therefore also in the Champcella area, which was covered by the glacier (Buoncristiani &#x0026; Campy, <xref rid="ref-6-e004" ref-type="bibr">2004</xref>). Climatically, the area is characterised by strong thermal fluctuations with many cold days and many very hot days. Precipitation is highest in autumn and lowest in summer, a Mediterranean type rainfall pattern, with consequent atmospheric aridity in the vegetative period, which is increased by the very high insolation rate (Bardin, <xref rid="ref-3-e004" ref-type="bibr">1967</xref>; Guiter, <xref rid="ref-12-e004" ref-type="bibr">1975</xref>). P&#x00E9;guy (<xref rid="ref-17-e004" ref-type="bibr">1947</xref>) exhaustively compared environmental parameters between different localities of the Durance Valley. In the neighbourhood of Guillestre at 980 m a.s.l., which is close to Champcella (type locality of <italic toggle="yes">S. druentianum</italic>), annual rainfall of 690 to 763 mm was recorded. Widmann (<xref rid="ref-24-e004" ref-type="bibr">1950</xref>) reported rainfall at the well-known location of <italic toggle="yes">Juniperus thurifera</italic> L. at St. Cr&#x00E9;pin, which is opposite that of <italic toggle="yes">S. druentianum</italic>, on the left of the valley, of 744 mm. Rain is mainly concentrated between September and October while the winter appears rather dry (see also Braun-Blanquet, <xref rid="ref-4-e004" ref-type="bibr">1922</xref>); moderate rainfall from April to June is of little use for the vegetation due to the wind and the strong insolation (Widmann, <xref rid="ref-24-e004" ref-type="bibr">1950</xref>). The area comprising Gap and Sisteron is located on the boundary between the Mediterranean region that is characterised, at least potentially, by <italic toggle="yes">Quercus pubescens</italic> Willd. and an alpine forest with prevailing <italic toggle="yes">Pinus sylvestris</italic> L. (Braun-Blanquet, <xref rid="ref-4-e004" ref-type="bibr">1922</xref>; Breistroffer, <xref rid="ref-5-e004" ref-type="bibr">1946</xref>; Widmann, <xref rid="ref-24-e004" ref-type="bibr">1950</xref>; Ozenda, <xref rid="ref-15-e004" ref-type="bibr">1966</xref>; Richard &#x0026; Tonnel, <xref rid="ref-18-e004" ref-type="bibr">1987</xref>; Garraud, <xref rid="ref-10-e004" ref-type="bibr">1998</xref>) on dry, calcareous soils. The type locality of <italic toggle="yes">S. druentianum</italic> is located near 1000 m, but it is possible to hypothesize that the range of its elevation may be between 750 and 1500 m in the area near the Mediterranean zone, where <italic toggle="yes">Sedetum brigantiacum</italic> Br.-Bl. is a pioneer on rocks overlain with little calcareous soil (Ozenda, <xref rid="ref-15-e004" ref-type="bibr">1966</xref>). These plants along with several other representatives of the Crassulaceae are exposed to drought conditions and frost (Bardin, <xref rid="ref-3-e004" ref-type="bibr">1967</xref>). <italic toggle="yes">Sempervivum druentianum</italic> has very limited natural geographical distribution range and its remarkable morphological similarity to <italic toggle="yes">S. tectorum</italic>, together with taxonomic confusion in the genus <italic toggle="yes">Sempervivum</italic> as a whole, have delayed its recognition as a new species.</p>
         <fig id="fig-3-e004" position="float" orientation="portrait">
            <label>Figure 3</label>
            <caption>
               <title>Natural populations of <italic toggle="yes">Sempervivum druentianum</italic> L. Gallo (red circle) and <italic toggle="yes">S.</italic> &#x00D7; <italic toggle="yes">brigantiacum</italic> L. Gallo (blue circle).</title>
            </caption>
            <graphic xlink:href="e004_003.jpeg"
                     position="anchor"
                     orientation="portrait"
                     id="gra-3-e004"/>
         </fig>
         <p>
            <italic toggle="yes">Phenology</italic>: <italic toggle="yes">Sempervivum druentianum</italic> predominantly flowers in July. The first flowers opened on 24 June (during the 2019 season) and the last flower opened on the 3 August (during the 2014 season).</p>
         <p>
            <italic toggle="yes">Etymology</italic>: from Druentia, old classic name of the Durance River where, in its surroundings, <italic toggle="yes">S. druentianum</italic> has been found. This name was cited by some classic authors and sometimes reported as Durentia, Drancia, Drance, etc. (Alessio, <xref rid="ref-1-e004" ref-type="bibr">1949</xref>).</p>
         <p>
            <italic toggle="yes">Remarks</italic>: despite repeated field expeditions in the vicinity of Champcella, no other populations of <italic toggle="yes">S. druentianum</italic> have been found. However, traces of its former presence, in the form of hybrids with other species, such as <italic toggle="yes">S. arachnoideum</italic>, have been identified in two locations not far from Champcella: at the Plateau d&#x0027;Herbonne (Ch&#x00E2;teauroux-les-Alpes) between 900 and 930 m a.s.l. and at the Lac de Siguret (Embrun) between 1060 and 1100 m. No other species belonging to the <italic toggle="yes">S. tectorum</italic> group were found at the type locality of <italic toggle="yes">S. druentianum</italic>, although near La Roche-de-Rame, a population of <italic toggle="yes">S. alpinum</italic> was located by the author (GL-9007).</p>
         <p>Different distributions of the <italic toggle="yes">S. tectorum</italic> group were reported in Chas (<xref rid="ref-7-e004" ref-type="bibr">1994</xref>: 203) and <italic toggle="yes">Atlas en ligne de la flore du d&#x00E9;partement des Hautes-Alpes</italic> (SAPN-GF, <xref rid="ref-20-e004" ref-type="bibr">2022</xref>), where <italic toggle="yes">S. alpinum</italic>, <italic toggle="yes">S. tectorum</italic> and perhaps hybrids between them, all been brought together under the same name (<italic toggle="yes">S. tectorum</italic>), which have not yet been separated. Another species of the <italic toggle="yes">S. tectorum</italic> group found in this area is <italic toggle="yes">S. calcareum</italic>, which reaches the northern limit of its distribution range at Lac de Serre-Pon&#x00E7;on (Embrun) (see SAPN-GF, <xref rid="ref-20-e004" ref-type="bibr">2022</xref>), about 30 km north from the type locality of <italic toggle="yes">S. druentianum.</italic>
         </p>
         <sec id="sec-4-e004">
            <title>Key for the species of <italic toggle="yes">S. tectorum</italic> group in southern France:</title>
            <p>
               <bold>1.</bold> Rosette leaves glandulose ..................................<bold>2</bold>
            </p>
            <p>
               <bold>-.</bold> Rosette leaves glabrous ....................................<bold>3</bold>
            </p>
            <p>
               <bold>2.</bold> Carpels glabrous on the back, petals white-green ..................................<bold>
                  <italic toggle="yes">S. calcareum</italic>
               </bold>
            </p>
            <p>
               <bold>-.</bold> Carpels glandular throughout, petals white-pinkish, filament glabrous or papillate ...<bold>
                  <italic toggle="yes">S. druentianum</italic>
               </bold>
            </p>
            <p>
               <bold>3.</bold> Leaves glaucous with apical rhombic red or red/brown dot, filament glabrous ...<bold>
                  <italic toggle="yes">S. calcareum</italic>
               </bold> (excl. ..........................type locality with glandular leaves)</p>
            <p>
               <bold>-.</bold> Leaves green or glaucous with or without apical rhombic red dot, filament glandulose .......................<bold>4</bold>
            </p>
            <p>
               <bold>4.</bold> Leaves green with red-brown apical red-brown dot and base white ..................................<bold>
                  <italic toggle="yes">S. tectorum</italic>
               </bold>
            </p>
            <p>
               <bold>-.</bold> Leaves green or glaucous without apical red dot and with base violet ................................<bold>
                  <italic toggle="yes">S. alpinum</italic>
               </bold>
            </p>
            <p>Many clumps of <italic toggle="yes">S. arachnoideum</italic> were found around <italic toggle="yes">S. druentianum</italic> occurs, and also material identified as a natural hybrid between both species, which is described here:</p>
            <p>
               <bold>
                  <italic toggle="yes">Sempervivum</italic> &#x00D7;<italic toggle="yes">brigantiacum</italic>
               </bold> L. Gallo <bold>hyb. nov.</bold> (<xref rid="fig-4-e004" ref-type="fig">Fig. 4</xref>)</p>
            <fig id="fig-4-e004" position="float" orientation="portrait">
               <label>Figure 4</label>
               <caption>
                  <title>
                     <italic toggle="yes">Sempervivum</italic> &#x00D7; <italic toggle="yes">brigantiacum</italic> L. Gallo nothosp. nov.: (A), clump of flowering rosettes; (B), close-up of a rosette with tufts of trichomes at the tip of the leaves; (C), inflorescence with emphasis on the lanceolate petals; (D), parents of the hybrids (<italic toggle="yes">S. arachnoideum</italic> L. and <italic toggle="yes">S. druentianum</italic> L. Gallo).</title>
               </caption>
               <graphic xlink:href="e004_004.jpeg"
                        position="anchor"
                        orientation="portrait"
                        id="gra-4-e004"/>
            </fig>
            <p>
               <italic toggle="yes">Type</italic>: France, Provence-Alpes-C&#x00F4;te d&#x2019;Azur, D&#x00E9;p. Hautes-Alpes, between Le Chambon and Chanteloube, neighborhood of Champcella, before the place named &#x201C;La Bourgea&#x201D;, on the side of the D38, 991 m a.s.l., rocks covered with xerophilous vegetation, 44.710783 N, 06.586438 E, 22-30.VI.2019, <italic toggle="yes">L. Gallo</italic> &#x0026; <italic toggle="yes">M. Gai</italic> (holotype: MRSN-12990) (<xref rid="fig-5-e004" ref-type="fig">Fig. 5</xref>).</p>
            <fig id="fig-5-e004" position="float" orientation="portrait">
               <label>Figure 5</label>
               <caption>
                  <title>Holotype of <italic toggle="yes">Sempervivum</italic> &#x00D7; <italic toggle="yes">brigantiacum</italic> L. Gallo (MRSN-12990) (photograph: A. Pepe).</title>
               </caption>
               <graphic xlink:href="e004_005.png"
                        position="anchor"
                        orientation="portrait"
                        id="gra-5-e004"/>
            </fig>
            <p>
               <italic toggle="yes">Diagnosis</italic>: hybrid of <italic toggle="yes">S. arachnoideum</italic> and <italic toggle="yes">S. druentianum</italic>, very similar to <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">fauconnetii</italic>, differing (with difficulty) by the darker color and the more leathery texture of the leaves of the latter. The presence of red-headed glands in the inflorescence of <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic> may also help to distinguish it from &#x005B;these are morphological characters that are present in natural populations and cannot be overlooked. We must be honest and say that there are objective difficulties in determining them&#x005D; <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">fauconnetii</italic>. The distinction from <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">luisae</italic> L. Gallo is much easier because of the presence of glandular trichomes on the surfaces of the leaves of the latter, which are absent in <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">druentianum</italic> (<xref rid="taw-2-e004" ref-type="table">Table 2</xref>).</p>
            <table-wrap id="taw-2-e004" position="float" orientation="portrait">
               <label>Table 2</label>
               <caption>
                  <title>Comparison table among <italic toggle="yes">Sempervirum</italic> &#x00D7;<italic toggle="yes">brigantiacum</italic> and the parents <italic toggle="yes">Sempervirum arachnoideum</italic> and <italic toggle="yes">Sempervirum druentianum</italic>.</title>
               </caption>
               <table id="tab-2-e004"
                      frame="border"
                      rules="all"
                      width="90&#x0025;">
                  <thead>
                     <tr>
                        <th style="width:120.25pt;text-align:center;" rowspan="1" colspan="1">
                           <bold>Morphological traits</bold>
                        </th>
                        <th style="width:106.3pt;text-align:center;" rowspan="1" colspan="1">
                           <bold>
                              <italic toggle="yes">Sempervirum arachnoideum</italic> L.</bold>
                        </th>
                        <th style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;text-align:center;"
                            rowspan="1"
                            colspan="1">
                           <bold>
                              <italic toggle="yes">Sempervirum</italic> &#x00D7;<italic toggle="yes">brigantiacum</italic> L. Gallo</bold>
                        </th>
                        <th style="width:112.65pt;text-align:center;" rowspan="1" colspan="1">
                           <bold>
                              <italic toggle="yes">Sempervirum druentianum</italic>  L. Gallo</bold>
                        </th>
                     </tr>
                  </thead>
                  <tbody>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Leaves apex (color)</bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Green or red</td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Green</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Brown-violet</td>
                     </tr>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Leaves apex (induments)</bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Long arachnoideus</td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Long sinuous marginal cilia, little-tufted at apex</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Absent</td>
                     </tr>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Leaves shape</bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Linear</td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Linear</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Obovate with apex cuspidate</td>
                     </tr>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Leaves induments </bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Glandular </td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Glabrous</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Glandular</td>
                     </tr>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Petals shape</bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Lanceolate</td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Lanceolate/linear</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Linear</td>
                     </tr>
                     <tr>
                        <td style="width:120.25pt;" rowspan="1" colspan="1">
                           <bold>Carpels induments</bold>
                        </td>
                        <td style="width:106.3pt;" rowspan="1" colspan="1">Glandular but not on the back</td>
                        <td style="width:113.8pt;background-color:&#x0023;C3BD96;color:&#x0023;000;"
                            rowspan="1"
                            colspan="1">Glandular but not on the back</td>
                        <td style="width:112.65pt;" rowspan="1" colspan="1">Glandular everywhere</td>
                     </tr>
                  </tbody>
               </table>
            </table-wrap>
            <p>
               <italic toggle="yes">Description</italic>: rosette 25-35 mm diameter. Leaf green, erect and linear, without red or brown spots at apex and with white base; glabrous on surfaces with long sinuous marginal cilia, little-tufted at apex. Inflorescence 18-25 cm tall, bifurcated, robust, green sometimes reddish, with long branches and rare bracts; many (also 20 or more) flowers. Caulinar leaves glandular with cilia on the margin; glands often red and not diaphanous. Flowers round or slightly elongated in bud, 20 mm diameter, with a long pedicel, covered with red glands. Petals nine, patent or somewhat decumbent, linear and not much lanceolate at base, pink with central dark line, sometimes green with white margins. Filaments violet, glandular. Anthers oval, not mucronate, violet. Carpels green, glandular but not on the back. Stylus green, violet at the apex, erect at the anthesis.</p>
            <p>
               <italic toggle="yes">Area and habitat</italic>: the type locality is close to that of <italic toggle="yes">S. druentianum</italic> but <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic> was found in two other places on either side of the Durance: at Lac de Siguret (Embrun) where the presence of <italic toggle="yes">S. druentianum</italic> has not was doubtful (only sterile specimens viewed to date, so the distinction from hybrids with <italic toggle="yes">S. alpinum</italic> or <italic toggle="yes">S. calcareum</italic> is not certain) and Plateau d&#x2019;Herbonne (Ch&#x00E2;teauroux-les-Alpes) where the presence has yet to be confirmed (<xref rid="fig-3-e004" ref-type="fig">Fig. 3</xref>).</p>
            <p>
               <italic toggle="yes">Etymology</italic>: the nothospecific epithet originates from &#x201C;Brigantium&#x201D;, the ancient name of the town of Briancon, which is in the neighborhood was found the type locality of <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic>.</p>
         </sec>
         <sec id="sec-5-e004">
            <title>Living specimens examined</title>
            <p>
               <italic toggle="yes">Sempervivum druentianum</italic> L. Gallo. France, D&#x00E9;p. Hautes-Alpes, Provence-Alpes-C&#x00F4;te d&#x2019;Azur: Tra le Chambon e Chanteloube, prima di La Bourgea a lato della D38, 989 m a.s.l., 44.711131 N, 06.587540 E, 5.VII.2019, <italic toggle="yes">L. Gallo</italic> (GL-8868, GL-8936, GL-8937, GL-8939, GL-8943); between Le Chambon e Chanteloube, neighborhood of Champcella, before the place named &#x201C;La Bourgea&#x201D;, on the side of the D38, 991 m a.s.l., rocks covered with xerophilous vegetation, 44.710783 N, 06.586438 E, 31.VII.2020, <italic toggle="yes">L. Gallo</italic> (GL-9021, GL-9022, GL-9023, GL-9024, GL-9025).</p>
            <p>
               <italic toggle="yes">Sempervivum</italic> &#x00D7;<italic toggle="yes">brigantiacum</italic> L. Gallo. France, D&#x00E9;p. Hautes-Alpes, Provence-Alpes-C&#x00F4;te d&#x2019;Azur : between Le Chambon e Chanteloube, neighborhood of Champcella, before the place named &#x201C;La Bourgea&#x201D;, on the side of the D38, 991 m a.s.l., rocks covered with xerophilous vegetation, 44.710783 N, 06.586438 E, 31.VII.2020, <italic toggle="yes">L. Gallo</italic> (GL-9020).</p>
         </sec>
      </sec>
   </body>
   <back>
      <ack id="ack-1-e004">
         <title>ACKNOWLEDGEMENTS</title>
         <p>Thanks to Urs Eggli (Zurich, Switzerland) and Gideon Smith (Port Elizabeth, South Africa) for their constructive proposals which greatly improved the manuscript. Thanks also to Fabrizio Longo and Alessandra Pepe (Museo Regionale di Scienze Naturali, Turin, Italy) which taken the photos of the holotypes of <italic toggle="yes">S. druentianum</italic> and <italic toggle="yes">S</italic>. &#x00D7;<italic toggle="yes">brigantiacum</italic>.</p>
      </ack>
      <ref-list id="refl-1-e004">
         <title>REFERENCES</title>
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               <year>1949</year>. <article-title>Une nouvelle base hydronymique m&#x00E9;diterran&#x00E9;enne: &#x002A;<italic toggle="yes">drava</italic>
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            </mixed-citation>
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