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Taxonomic Revision of West-Alpine Cushion Plant Species Belonging to Androsace Subsect

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Taxonomic Revision of West-Alpine Cushion Plant Species Belonging to Androsace Subsect Botany Letters ISSN: 2381-8107 (Print) 2381-8115 (Online) Journal homepage: http://www.tandfonline.com/loi/tabg21 Taxonomic revision of West-Alpine cushion plant species belonging to Androsace subsect. Aretia Cédric Dentant, Sébastien Lavergne & Valéry Malécot To cite this article: Cédric Dentant, Sébastien Lavergne & Valéry Malécot (2018) Taxonomic revision of West-Alpine cushion plant species belonging to Androsace subsect. Aretia , Botany Letters, 165:3-4, 337-351, DOI: 10.1080/23818107.2018.1450784 To link to this article: https://doi.org/10.1080/23818107.2018.1450784 Published online: 27 Apr 2018. Submit your article to this journal Article views: 29 View Crossmark data Citing articles: 1 View citing articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tabg21 BOTANY LETTERS 2018, VOL. 165, NOS. 3–4 , 337–351 https://doi.org/10.1080/23818107.2018.1450784 ARTICLE Taxonomic revision of West-Alpine cushion plant species belonging to Androsace subsect. Aretia Cédric Dentanta , Sébastien Lavergneb and Valéry Malécotc aParc national des Ecrins, Domaine de Charance, Gap, France; bLaboratoire d’Écologie Alpine (LECA), CNRS – University Grenoble Alpes – University Savoie Mont-Blanc, Grenoble, France; cIRHS, Agrocampus Ouest, INRA, Université d’Angers, Beaucouzé, France ABSTRACT ARTICLE HISTORY High-altitude rockjasmines (genus Androsace) are a paramount example of evolutionary Received 29 January 2018 radiations in temperate mountains of the Northern Hemisphere. Yet, we show here that their Accepted 1 March 2018 taxonomy is incomplete and has been subject to many historical mistakes, probably due to the KEYWORDS lack of exploration of mountains by the classical botanists who described these species. Here Androsace; trichomes; we wish to clarify the application of names with regard to four defined morphotypes, typical lectotype; epitype; of high-elevation zones of the Western Alps, in order to set a definitive basis for morphological phylogeny; high mountain; delimitation of a likely new species discovered in the Mont Blanc range (species not described Alps in this work). To do so, we review the historical taxonomic treatments and positions, carefully reconsider types and species names for these morphotypes, and designate lectotypes and epitypes for each of them. In particular, we confirm the validity of names commonly used to refer to the taxa Androsace alpina, Androsace helvetica, and Androsace pubescens. We show that Androsace vandellii is an invalid name and that Androsace argentea should be used instead. Our work illustrates the utility of historical herbaria to clarify the taxonomy of complex groups of plants growing in inaccessible environments such as high-altitude regions. Introduction pubescens, A. vitaliana, A. obtusifolia, and A. lactea. Here we focus on four cushion species of this Western group Androsace sect. Aretia subsect. Aretia, [Clade]/Eu- which present taxonomic and nomenclatural issues for Aretia is an infrageneric group of alpine species com- taxonomists and field botanists, namely A. helvetica, A. posed of plants occurring in southwestern European alpina, A. vandellii, and A. pubescens. Clarifying taxon- mountains (Alps and Pyrenees): Androsace alpina (L.) omy is a prior requirement for on-going research on Lam., Androsace brevis (Hegetschw.) Ces., Androsace the phylogeny of this subsection (Boucher et al. 2012), cylindrica DC., Androsace komovensis Schönswetter & and more generally for understanding the origin and Schneeweiss, Androsace helvetica (L.) All., Androsace structure of high-alpine plant communities (Marx et al. hausmannii Leyb., Androsace lactea L., Androsace mathil- 2017). Recently, a field assessment of endangered plants dae Levier, Androsace obtusifolia All., Androsace pubes- of the Northern French Alps listed a morphologically cens DC., Androsace vandellii (Turra) Chiov., Androsace distinct Androsace taxon, which could be described as a vitaliana (L.) Lapeyr., and Androsace wulfeniana (Sieber new species (Jordan 2015). As this new Androsace taxon ex Koch) Rchb. (Schneeweiss et al. 2004; Schönswetter shares features with both A. pubescens and A. alpina, and Schneeweiss 2009). it is of utmost importance to set a clear and definitive Three of these species are strictly endemic to moun- basis for the morphological delimitation of the four tain ranges outside the Alps: A. cylindrica (Pyrenees), A. aforementioned species. Here we propose to reconsider komovensis (Balkans), and A. mathildae (Apennines). the nomenclature and taxonomy of these species and to Three of the remaining 10 species have a distribution select proper types for each of them. restricted to the Central Alps: A. hausmannii, A. brevis, and A. wulfeniana. The species A. pubescens was long thought to occur in the Pyrenees, but it was recently Features historically associated to species argued that it has been mixed up with the morphologi- names in the genus Androsace cally similar, but genetically differentiated, A. cylindrica Morphological features (Schneeweiss et al. 2017) – thus A. pubescens seems to be endemic to the Western Alps. Considering historical species’ features cited in the pro- Overall, only seven species of this clade occur in tologues, four morphological characters are relevant to Western Alps: A. helvetica, A. alpina, A. vandellii, A. be exposed: CONTACT Cédric Dentant [email protected] © 2018 Société botanique de France Published online 27 Apr 2018 .C. DENTANT ET AL 338 (i) leaf shape: short or long, apex pointed or round called “Aretia villosa scapis unifloris”. Later, Linnaeus (ii) leaf organization: imbricated or loose (1753) provided these two morphotypes with bino- (iii) stalk size: visible stalk or very short stalk (flowers mials, namely Diapensia helvetica L. for the Helvetica more or less sessile) morphotype and Aretia alpina L. for the Alpina mor- (iv) corolla colour: white or reddish photype. However, herbarium samples used in Linnaeus’ works show that under the name Diapensia helvetica These features are generally not sufficient to distin- Linnaeus merged individuals belonging to the Helvetica guish species. However, they are good complements to morphotype and individuals with radiate hairs assigned the last and ultimate feature: the shape of leaf trichomes to the Vandellii-Argentea morphotype. Similarly, most (simple, furcate, branched, or radiate). Villars (1787) was available specimens collected by Allioni (herbarium the first author to consider the shape of trichomes as a of Torino – TO) under the name Androsace helvetica relevant character to distinguish species in the group. are in fact morphologically similar to what we call here While merging Aretia in Androsace, he wrote:“pour dis- the Vandellii-Argentea morphotype, i.e. bearing leaves tinguer les Aretia des Androsace, il falloit ou avoir recours with dense radiate trichomes. Furthermore, in Allioni’s aux tiges uniflores, ou à la forme des poils. […] Dans le herbarium, a sheet with a short note written by Allioni second [cas], nous n’aurions eu que l’Androsace villosa L. and reproduced in his Flora pedemontana (1785) holds […] car elle a des poils simples, articulés ou séparés par under the same name specimens of both morphotypes: des diaphragmes, caractère qui nous a paru lui être par- Helvetica (simple trichomes, plant growing exclusively ticulier” [To distinguish Aretia from Androsace, it would on limestone) and Vandellii-Argentea (radiate trichomes, have been necessary to refer either to single-flowered plant growing exclusively on siliceous substratum). stems or to hair shape, […]. In the second [case] we These observations clearly highlight the fact that two would have had only Androsace villosa L. […] because it different morphotypes were actually merged under the has simple hairs, articulated or separated by diaphragms, name Androsace helvetica ever since its first use. a character that seemed restricted to this species]. This Consequently, it is not surprising that successive key feature was usually absent from early protologues authors gave species descriptions mixing both morpho- and was not necessarily cited in later ones. types or concerning just one or the other. The first case of such mixed description is the one of Villars (1787), Locus classicus where he gave the following description under the name When the locus classicus of a described species is availa- Diapensia helvetica: “poils ramifiés en Y & en goupil- ble, it may be decisive to scrutinize the plants occurring lon” [ramified Y-shaped and bottlebrush-like hairs], in this location. where “ramified hairs” corresponds to the Vandellii- Argentea morphotype. Similarly, in the second edition of the Flore Française (1805), Lamarck and de Candolle Features of the four cushions of Androsace described the Vandellii-Argentea morphotype using The four aforementioned Androsace taxa (currently a name – Androsace imbricata Lam. – directly linked named Androsace helvetica, A. alpina, A. vandellii, and to Diapensia helvetica L. They even provided a precise A. pubescens) correspond to four morphotypes that can description: “feuilles […] tapissées d’un duvet court, be summarized as in Table 1. blanc et serré, composé de poils rameux et rayonnants” [leaves […] lined with a short, tied and white fluff, composed of ramified and radiant hairs], which are History of taxa description definitely the specific features of the Vandellii-Argentea Since Haller (1742), at least two west-alpine cushion
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