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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">AJBM</journal-id>
<journal-title-group>
<journal-title>Anales del Jard&#x00ED;n Bot&#x00E1;nico de Madrid</journal-title>
</journal-title-group>
<issn pub-type="ppub">0211-1322</issn>
<issn pub-type="epub">1988-3196</issn>
<publisher>
<publisher-name>Consejo Superior de Investigaciones Cientificas</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">AJBM201713-2472</article-id>
<article-id pub-id-type="doi">10.3989/ajbm.2472</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Articles</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>On the identity of <italic>Helianthemum mathezii</italic> and <italic>H. pomeridianum (Cistaceae)</italic></article-title>
<trans-title-group xml:lang="es">
<trans-title>Acerca de la identidad de <italic>Helianthemum mathezii</italic> y <italic>H. pomeridianum (Cistaceae)</italic></trans-title>
</trans-title-group>
<alt-title alt-title-type="running-head">On the identity of <italic>Helianthemum mathezii</italic> and <italic>H. pomeridianum</italic></alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Aparicio</surname>
<given-names>Abelardo</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
<xref ref-type="corresp" rid="cor1">&#x002A;</xref>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Albaladejo</surname>
<given-names>Rafael G.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">2</xref>
</contrib>
</contrib-group>
<aff id="aff0001">
<label>1,2</label>Dept. of Plant Biology and Ecology, Universidad de Sevilla, Spain</aff>
<author-notes>
<corresp id="cor1">
<label>&#x002A;</label>Corresponding author: <email xlink:href="abelardo@us.es">abelardo@us.es</email>, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-7122-4421">http://orcid.org/0000-0001-7122-4421</ext-link></corresp>
<corresp id="cor2">
<label>2</label><email xlink:href="albadalejo@us.es">albadalejo@us.es</email>, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0003-2101-5204">http://orcid.org/0000-0003-2101-5204</ext-link></corresp>
<fn>
<p>Associate Editor: C. Aedo.</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>30</day>
<month>09</month>
<year>2017</year>
</pub-date>
<pub-date pub-type="collection">
<year>2017</year>
</pub-date>
<volume>74</volume>
<issue>2</issue>
<elocation-id content-type="doi">10.3989/ajbm.2472</elocation-id>
<history>
<date date-type="received">
<day>27</day>
<month>02</month>
<year>2017</year>
</date>
<date date-type="accepted">
<day>25</day>
<month>05</month>
<year>2017</year>
</date>
<date date-type="published online">
<day>26</day>
<month>10</month>
<year>2017</year>
</date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2017 CSIC</copyright-statement>
<copyright-year>2017</copyright-year>
<license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc/3.0/">
<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution-Non Commercial (by-nc) Spain 3.0 License.</license-p>
</license>
</permissions>
<abstract>
<title>Abstract</title>
<p>The original description of <italic>Helianthemum mathezii</italic> regarded the species to be a therophyte. However, the detailed observation of the holotype of <italic>H. mathezii,</italic> as well as newly collected specimens from the type locality, does not support its condition of annual plant. Further study has led to the conclusion that all these plants can readily be identified as <italic>H. pomeridianum;</italic> the descriptions of <italic>H. mathezii</italic> and <italic>H. pomeridianum</italic> are equivalent except for the habit, being the former annual and the latter suffruticose. We show in this study that <italic>H. mathezii</italic> is to be considered a synonym of <italic>H. pomeridianum,</italic> and report a new locality of <italic>H. pomeridianum</italic> in Morocco. Also, we provide the first chromosome count for this species (2<italic>n</italic> = 22) which emphasizes the close relationship of <italic>H.</italic> pomeridianum to <italic>H.</italic> subgen. <italic>Plectolobum</italic>.</p>
</abstract>
<trans-abstract xml:lang="es">
<title>Resumen</title>
<p>En la descripci&#x00F3;n original de <italic>Helianthemum mathezii</italic> se considera esta especie como un terofito. Sin embargo, la observaci&#x00F3;n minuciosa de su holotipo, as&#x00ED; como de otros espec&#x00ED;menes recientemente recolectados en la misma localidad del tipo, sugiere que no se trata de una planta anual. Concluimos que estas plantas pueden ser bien identificadas como <italic>H. pomeridianum;</italic> las descripciones de <italic>H. mathezii</italic> y <italic>H. pomeridianum</italic> son similares excepto en lo referido a su h&#x00E1;bito, la primera supuestamente es una planta anual y la segunda sufruticosa. As&#x00ED; pues, aqu&#x00ED; recomendamos considerar <italic>H. mathezii</italic> como sin&#x00F3;nimo de <italic>H. pomeridianum,</italic> y citamos una localidad nueva de esta especie en Marruecos. Adem&#x00E1;s, damos cuenta del primer recuento de cromosomas efectuado para la misma (2<italic>n</italic> = 22), que demuestra la estrecha afinidad de <italic>H. pomeridianum</italic> y <italic>H.</italic> subgen. <italic>Plectolobum</italic>.</p>
</trans-abstract>
<kwd-group xml:lang="en">
<title>Keywords</title>
<kwd>Chromosome count</kwd>
<kwd><italic>Cistaceae</italic></kwd>
<kwd><italic>Helianthemum mathezii</italic></kwd>
<kwd><italic>Helianthemum pomeridianum</italic></kwd>
<kwd>taxonomy</kwd>
</kwd-group>
<kwd-group xml:lang="es">
<title>Palabras clave</title>
<kwd><italic>Cistaceae</italic></kwd>
<kwd><italic>Helianthemum mathezii</italic></kwd>
<kwd><italic>Helianthemum pomeridianum</italic></kwd>
<kwd>recuento de cromosomas</kwd>
<kwd>taxonom&#x00ED;a</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec id="sec1" sec-type="intro">
<title>INTRODUCTION</title>
<p>Dobignard (<xref ref-type="bibr" rid="cit0005">2009</xref>) described <italic>H. mathezii</italic> Dobignard &#x2014;Cistaceae Juss.&#x2014; from a single locality from the Moroccan High Atlas &#x2014;Taroudant province&#x2014; as a 20&#x2013;35 cm tall, much-branched, and glandular-hairy annual plant. This author stated that, despite its apparent morphological resemblance to <italic>H. salicifolium</italic> (L.) Mill. <italic>(H.</italic> subg. <italic>Helianthemum), H. mathezii</italic> should be ascribed to <italic>H.</italic> sect. <italic>Atlanthemum</italic> (Raynaud) G. L&#x00F3;pez &#x0026; al. <italic>(H.</italic> subg. <italic>Plectolobum</italic> Willk.) due to its glandular and prominently 5&#x2013;7-veined inner sepals that resembled those of <italic>H. sanguineum</italic> (Lag.) Lag. ex Dunal. Dobignard (<xref ref-type="bibr" rid="cit0005">2009</xref>) also reported that this species inhabited wet travertine deposits accompanied by <italic>Feeria angustifolia</italic> Buser and <italic>Euphorbia rimarum</italic> Coss. &#x0026; Balansa, among other species. Thus described, <italic>H. mathezii</italic> constituted a very interesting species inasmuch as it was the only <italic>Helianthemum</italic> species inhabiting a wet habitat (cf. Arrington &#x0026; Kubitzki <xref ref-type="bibr" rid="cit0002">2003</xref>), and the only annual species in the genus with such a restricted distribution (cf. Proctor &#x0026; Heywood <xref ref-type="bibr" rid="cit0015">1968</xref>; Greuter &#x0026; al. <xref ref-type="bibr" rid="cit0011">1984</xref>), further included in the checklist of North African endemic plant species (El Oualidi &#x0026; al. <xref ref-type="bibr" rid="cit0007">2012</xref>). Consequently, as part of a comprehensive phylogenetic study of the genus <italic>Helianthemum</italic> (Aparicio &#x0026; al. <xref ref-type="bibr" rid="cit0001">2017</xref>), we considered of interest to perform a new collection and a detailed study of <italic>H. mathezii</italic>.</p>
</sec>
<sec id="sec2" sec-type="material|methods">
<title>MATERIAL AND METHODS</title>
<p>To do so, we visited the type locality of <italic>H. mathezii,</italic> at &#x201C;piste d&#x2019;acc&#x00E8;s au plateau du J. Tichka alt. c. 1100 m, 30&#x00B0;47&#x2019;18 N, &#x2013;8&#x00B0;40&#x2019;73 W&#x201D; (Dobignard <xref ref-type="bibr" rid="cit0005">2009</xref>), to survey the species. For the chromosome number determination we collected matured seeds that were disposed for germination in Petri dishes. Then, root tips were immersed in chilling water at 0 &#x00BA;C for 24 h, then fixed in 1:3 glacial acetic acid and absolute ethanol for 3 h. They were finally stained in alcoholic hydrochloric acid-carmine for 48 h and squashed in 45% acetic acid.</p>
<p>No type specimen of <italic>H. pomeridianum</italic> Dunal has been observed. Nevertheless, <italic>H. pomeridianum</italic> has been many times collected by different authors in its Algerian locus classicus &#x2014;circa Oran, Djebel Santo&#x2014;, and vouchers are available on-line at the Mus&#x00E9;um Nationale D&#x2019;Historie Naturelle &#x2014;MNHN&#x2014;. We list the studied ones in <xref ref-type="app" rid="app01">appendix 1</xref>, all identified as <italic>H. pomeridianum</italic>.</p>
</sec>
<sec id="sec3" sec-type="results|discussion">
<title>RESULTS AND DISCUSSION</title>
<p>During our field trip to the type locality, we found a population integrated by about 30 individual plants readily identifiable as <italic>H. mathezii,</italic> other than for the fact that these specimens, albeit having slender branches, were perennial suffruticose plants (<xref ref-type="fig" rid="f0001">fig. 1</xref>). Moreover, they were located in a sunny dry area on the top of a calcareous travertine deposit accompanied by <italic>Micromeria hochreutineri</italic> Maire and <italic>Lavandula multifida</italic> L. rather than in the wet vertical run-off area of the same travertine outcrop, where Feeria angustifolia and species of <italic>Euphorbia</italic> L., <italic>Campanula</italic> L., and <italic>Erodium</italic> L&#x0027;H&#x00E9;r. were common.</p>
<fig id="f0001">
<label>Fig. 1</label>
<caption>
<p>Specimen of <italic>Helianthemum pomeridianum</italic> Dunal (SEV 286762).</p>
</caption>
<graphic xlink:href="AJBM201713-2472-g001.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<p>The detailed observation of the holotype of <italic>H. mathezii</italic> (<xref ref-type="fig" rid="f0002">fig. 2</xref>), as well as the figure no. 7 of the original description of the species (Dobignard <xref ref-type="bibr" rid="cit0005">2009</xref>), does not provide conclusive evidence that <italic>H. mathezii</italic> is a therophyte since the root system cannot be observed, a fact clearly evident in the specimens of <italic>H. sanguineum</italic> and <italic>H. salicifolium</italic> therein included for illustration &#x2014;not shown here&#x2014;. Moreover, the analysis of the Dobignard&#x2019;s material and our newly collected plants, following Qu&#x00E9;zel &#x0026; Santa (<xref ref-type="bibr" rid="cit0016">1962</xref>) and Fennane &#x0026; al. (<xref ref-type="bibr" rid="cit0008">1999</xref>), has resulted in that all these plants can be identified as <italic>H. pomeridianum</italic>. Further study has also shown that the description of <italic>H. mathezii</italic> is equivalent to that of <italic>H. pomeridianum</italic> except for its habit &#x2014;the former annual, the latter suffruticose&#x2014; since both descriptions depict glandular plants which have: 1) slender branches bearing very shortly petiolate, exstipulate, flat, ovate-lanceolate leaves markedly veined beneath &#x2014;falsely stipulate because of the presence of small axillary crowded leaves&#x2014;; 2) very loosely arranged flowers in glandular terminal cymes composed by a few pedicellate and bracteate solitary flowers; 3) inner sepals 4&#x2013;6 &#x00D7; 2 mm, striate with prominent veins; 4) stamens numerous; 5) pubescent ovary bearing a short capitate style; 6) capsule shorter than the inner sepals, glabrous, with ciliate margins at the apex (Dunal <xref ref-type="bibr" rid="cit0006">1847</xref>; Willkomm <xref ref-type="bibr" rid="cit0018">1856</xref>; Dobignard <xref ref-type="bibr" rid="cit0005">2009</xref>). Overall, these arguments show that <italic>H. mathezii</italic> is a synonym of <italic>H. pomeridianum,</italic> and that both Dobignard&#x2019;s specimen &#x2014;G 00441364&#x2014; and our own collection &#x2014;SEV 286762 (Morocco: Taroudant, between Sidi Abdellah Oussaid and Alegjane, 1100 m a.s.l., 30&#x00B0;47&#x0027;10.38&#x02BA; N, 8&#x00B0;40&#x0027;43.74&#x02BA; W, 13 Nov. 2016, <italic>A. Aparicio</italic> &#x0026; <italic>FJ. Aparicio</italic> s.n.&#x2014; are to be regarded as a new locality of <italic>H. pomeridianum</italic> in Morocco.</p>
<fig id="f0002">
<label>Fig. 2</label>
<caption>
<p>Holotype of <italic>Helianthemum mathezii</italic> Dobignard (G 00441364).</p>
</caption>
<graphic xlink:href="AJBM201713-2472-g002.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<p><italic>Helianthemum pomeridianum</italic> is a rather poorly known North African endemic species (El Oualidi &#x0026; al. <xref ref-type="bibr" rid="cit0007">2012</xref>), which was originally described, and many times collected by different authors, from around Oran &#x2014;Djebel Santo, Santa Cruz&#x2014; in Algeria. To date, <italic>H. pomeridianum</italic> has also been recorded in Melilla (Caballero <xref ref-type="bibr" rid="cit0004">1917</xref>), the Trara Mountains, also in the northern Algeria (Medjahdi &#x0026; al. <xref ref-type="bibr" rid="cit0013">2009</xref>), and in a few far away locations in the Moroccan High Atlas area, between Taroudant and Marrakech &#x2014;RAB 078906, MA 472377 (P 04729082 photo&#x0021;), MGC 80897 (MA 321784&#x2013;1&#x0021;)&#x2014; (F&#x00F6;rther &#x0026; Podlech <xref ref-type="bibr" rid="cit0009">2002</xref>; GBIF <xref ref-type="bibr" rid="cit0010">2016</xref>; SEV 225757&#x0021; from the southern Algeria is a misidentification of <italic>H. lippii</italic> (L.) Dum.-Cours.). Taxonomically, <italic>H. pomeridianum</italic> was ascribed by Dunal to <italic>H</italic>. sect. <italic>Halimium</italic> Dunal, but Pomel (<xref ref-type="bibr" rid="cit0014">1860</xref>: 11) formulated the species sub <italic>Rhodax pomeridianus</italic> Pomel. Willkomm (<xref ref-type="bibr" rid="cit0018">1856</xref>) and Grosser (<xref ref-type="bibr" rid="cit0012">1903</xref>) re-ascribed the species to <italic>H</italic>. sect. <italic>Eriocarpum</italic> Dunal <italic>(H</italic>. subg. <italic>Helianthemum),</italic> but Qu&#x00E9;zel &#x0026; Santa (<xref ref-type="bibr" rid="cit0016">1962</xref>) included it in <italic>H</italic>. sect. <italic>Chamaecistus</italic> Willk. of <italic>H</italic>. subg. <italic>Plectolobum</italic>. Phylogenetic data (Aparicio &#x0026; al. <xref ref-type="bibr" rid="cit0001">2017</xref>) has revealed an unexpected close relationship between <italic>H. pomeridianum</italic> and <italic>H. lunulatum</italic> DC., a dwarf shrub found only in the Maritime and Ligurian Alps (Barbero <xref ref-type="bibr" rid="cit0003">1966</xref>) that is the only member of <italic>H</italic>. sect. <italic>Macularia</italic> Dunal in <italic>H</italic>. subg. <italic>Plectolobum</italic>. This finding triggers further biogeographical interest since it may reveal a major intercontinental disjunction during the early diversification of <italic>H</italic>. subg. <italic>Plectolobum</italic> in the Upper Pliocene and that geographical vicariance might represent an important speciation force in this lineage; indeed, the first chromosome count for <italic>H. pomeridianum</italic> obtained from plants from this new Moroccan locality (2<italic>n</italic> = 22; <xref ref-type="fig" rid="f0003">fig. 3</xref>) provides additional support and emphasizes the relationship of this species to <italic>H</italic>. subg. <italic>Plectolobum</italic> (cf. Rice &#x0026; al. <xref ref-type="bibr" rid="cit0017">2015</xref>).</p>
<fig id="f0003">
<label>Fig. 3</label>
<caption>
<p>Mitotic metaphase of <italic>Helianthemum pomeridianum</italic> Dunal (SEV 286762).</p>
</caption>
<graphic xlink:href="AJBM201713-2472-g003.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<p>Finally, we were unable to extract DNA from a leaf sample of the type specimen of <italic>H. mathezii</italic> provided by herbarium G despite we followed two standard &#x2014;and slightly modified&#x2014; DNA extraction protocols: Invisorb Spin Plant Mini Kit &#x2014;Stratec&#x2014; and Isolate II Plant DNA-kit &#x2014;Bioline&#x2014;. Consequently, this accession could not be included in the phylogenetic analysis of <italic>Helianthemum,</italic> but in the phylogenetic analyses our samples collected in type locality &#x2014;SEV 286762&#x2014; clustered with other Moroccan accessions of <italic>H. pomeridianum</italic> Dunal and maintained the supported sister-taxa relationship between <italic>H. pomeridianum</italic> and <italic>H. lunulatum</italic> (Aparicio &#x0026; al. <xref ref-type="bibr" rid="cit0001">2017</xref>).</p>
</sec>
</body>
<back>
<ack>
<title>ACKNOWLEDGEMENTS</title>
<p>We acknowledge Alan Dobignard and the Herbarium Genavense for cooperation in this study. The project has been funded by the Spanish Ministerio de Econom&#x00ED;a y Competitividad (CGL2014&#x2013;52459-P). We also thank Mike Lockwood for reviewing the English style and an anonymous reviewer for helpful commentaries.</p>
</ack>
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<app-group>
<app id="app01">
<title>APPENDIX 1. Studied material of <italic>H. pomeridianum</italic>.</title>
<p>ALGERIA. Oran: <italic>Balansa</italic> (P 06686325 photo&#x0021;); ibid., 1842, <italic>Cosson</italic> (P 06686319 photo&#x0021;); ibid., <italic>Durando</italic> (P 06686313 photo&#x0021;, P 06686330 photo&#x0021;); ibid., 1842, <italic>Durieu</italic> (P 06686306 photo&#x0021;, P 06686307 photo&#x0021;); ibid., 1849, <italic>Reuter</italic> (P 06686308 photo&#x0021;); ibid., 1850, <italic>Gouget</italic> (P 06686309 photo&#x0021;); ibid., 1842, <italic>Pomel</italic> (P 06686326 photo&#x0021;); ibid., 1861, <italic>Pomel</italic> (P 06686310 photo&#x0021;, P06686303 photo&#x0021;, P 04747844 photo&#x0021;, P 06686327 photo&#x0021;); ibid., 1842, <italic>Spach</italic> (P 06686311 photo&#x0021;); ibid., Santa Cruz, 1905, s. coll. (P 04750429 photo&#x0021;); ibid., 1947, <italic>Chevassut</italic> (P 04637966 photo&#x0021;); ibid., 1918, <italic>D&#x0027;Alleizette</italic> (P 04637967 photo&#x0021;); ibid., 1912, <italic>D&#x0027;Alleizette</italic> (P 06686322 photo&#x0021;); ibid., 1887, <italic>Luizer</italic> (P 06686316 photo&#x0021;); ibid., 1930, <italic>Faure</italic> (P 06686328 photo&#x0021;); ibid., 1930, <italic>Senay</italic> (P 04729081 photo&#x0021;); ibid., Djebel Murdjadjo, 1894, <italic>Doumergue</italic> (P 04637877 photo&#x0021;); ibid., 1905, <italic>Faure</italic> (P 06686304 photo&#x0021;, P 06686331 photo&#x0021;, P 06686332 photo&#x0021;, P 06686336 photo&#x0021;); ibid., 1906, <italic>Faure</italic> (P 06686337 photo&#x0021;); ibid., 1952, <italic>Santa, Daumas</italic> &#x0026; <italic>Retz</italic> (P 04637876 photo&#x0021;, P 04729083 photo&#x0021;, P 04637968 photo&#x0021;); ibid., Djebel Santo, 1852, <italic>Balansa</italic> (P 06686317 photo&#x0021;, P 06686318 photo&#x0021;, P 06686334 photo&#x0021;, P 06686324 photo&#x0021;); ibid., 1882, <italic>Debeaux</italic> (P 06686296 photo&#x0021;, P 06686333 photo&#x0021;); ibid., 1887, <italic>Garrigues</italic> (P 06686320 photo&#x0021;, P 06686321 photo&#x0021;); ibid., 1850, <italic>Munby</italic> (P 06686312 photo&#x0021;, P 06686315 photo&#x0021;); ibid., <italic>Munby</italic> (P 06686323 photo&#x0021;); ibid., 1849, <italic>Reuter</italic> (P 06686329 photo&#x0021;); ibid., 1914, <italic>Rotereau</italic> (P 06686314 photo&#x0021;).</p>
</app>
</app-group>
</back>
</article>
