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(Published Version): Phylog Dieses Dokument ist eine Zweitveröffentlichung (Verlagsversion) / This is a self-archiving document (published version): Cécile Gueidan, Truong Van Do, Ngan Thi Lu Phylogeny and taxonomy of Staurothele (Verrucariaceae, lichenized ascomycetes) from the karst of northern Vietnam Erstveröffentlichung in / First published in: The Lichenologist. 2014, 46(4), S. 515 – 533 [Zugriff am: 11.03.2020]. Cambridge University Press. ISSN 1096-1135. DOI: https://doi.org/10.1017/S0024282914000048 Diese Version ist verfügbar / This version is available on: https://nbn-resolving.org/urn:nbn:de:bsz:14-qucosa2-390096 „Dieser Beitrag ist mit Zustimmung des Rechteinhabers aufgrund einer (DFGgeförderten) Allianz- bzw. Nationallizenz frei zugänglich.“ This publication is openly accessible with the permission of the copyright owner. The permission is granted within a nationwide license, supported by the German Research Foundation (abbr. in German DFG). www.nationallizenzen.de/ The Lichenologist 46(4): 515–533 (2014) 6 British Lichen Society, 2014 doi:10.1017/S0024282914000048 Phylogeny and taxonomy of Staurothele (Verrucariaceae, lichenized ascomycetes) from the karst of northern Vietnam Ce´cile GUEIDAN, Truong VAN DO and Ngan Thi LU Abstract: The crustose genus Staurothele (Verrucariaceae, Ascomycota) is a common component of the lichen flora from subneutral to alkaline silicate rocks in temperate to cold-temperate climates. Our field study in the karst system of northern Vietnam showed that it is also common on dry to humid limestone in the wet tropics. Molecular data revealed that species of Staurothele from Vietnam belong to an unnamed clade sister to the genus Endocarpon, together with the tropical Australian species Staurothele pallidopora and Staurothele diffractella, a North American species recently transferred to Endocarpon based on molecular data. The genus Willeya is here resurrected for this clade of crustose epilithic Staurothele with pale ascospores. Eight new combinations are proposed and three new species of Willeya are described from Vietnam. Sampling tropical members of a lichen family previously mostly known from temperate areas contributed significantly to improving its generic classification. Key words: classification, Endocarpon, generic delimitation, lichens, Verrucariales, Willeya Accepted for publication 9 January 2014 Introduction Alps). It has occasionally been reported from wetter and warmer areas of the world, Staurothele Norman is a genus of lichenized for example, Indonesia, (Groenhart 1954), fungi forming crustose thalli on subneutral southern China (Harada & Wang 1996, to alkaline silicate rocks. The species are 2006), but its diversity in subtropical and found both in dry and amphibious condi- tropical regions remains understudied. tions and are common in habitats including Staurothele currently comprises 72 species dry limestone outcrops or alkaline rocks (Kirk et al. 2008), all characterized by a crus- along rivers and streams. This genus belongs tose thallus and muriform ascospores. They to the Verrucariaceae Zenker, a fungal family also all possess a peculiar feature: algal cells, whose members are predominantly lichen- usually present only in the thallus in other forming. Although members of Verrucaria- lichens, are also found in the perithecia, ceae occur on various substrata, rock surfaces between the asci. These stichococcoid algae, constitute the most common habitat. As with recently shown with molecular data to be- many genera within Verrucariaceae, Stauro- long to the green-algal genus Diplosphaera thele is most diverse in temperate climates, Bial. (Thu¨s et al. 2011), are co-dispersed including the Mediterranean region, cold- with large muriform fungal ascospores. This temperate to subarctic regions (e.g., Alaska, character is not unique to this genus, but is Scandinavia) and high elevations (e.g., the also found in Endocarpon Hedw., another genus of Verrucariaceae. Staurothele is mor- C. Gueidan: Natural History Museum, Department of phologically similar to Endocarpon, but dif- Life Sciences, Cromwell Road, London, SW7 5BD, fers in the habit (squamulose in Endocarpon UK. Email: [email protected] and crustose in Staurothele) and the structure T. Van Do and N. T. Lu: Vietnam National Museum of Nature, Vietnam Academy of Science and Technology, of the upper cortex (eucortex in Endocar- 18 Hoang Quoc Viet, Hanoi, Vietnam. pon and pseudocortex in Staurothele). Within T. Van Do: Institute of Botany, Technical University of Staurothele, species-diagnostic features in- Dresden, Zellescher Weg 20b, 01062 Dresden, Germany. Downloaded from https://www.cambridge.org/core. SLUB Dresden, on 11 Mar 2020 at 13:20:01, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282914000048 516 THE LICHENOLOGIST Vol. 46 clude the structure of the thallus, which can (Gueidan et al. 2007, 2009; Savic´ et al. either be epilithic (growing on the rock sur- 2008). One well-supported monophyletic face) or endolithic (growing within the super- lineage composed exclusively of crustose epi- ficial layer of the rock), and the ascospore lithic species of Staurothele is closely related colour, which varies from pale to darkly pig- to the genera Catapyrenium and Placidiopsis. mented. Other differences include the size This lineage includes Staurothele clopima and structure of the perithecia and the size (Wahlenb.) Th. Fr., the type of the genus, of the ascospores. The shape of the algal cells and is now considered as Staurothele s. str. present in the hymenium has also been used Crustose endolithic species of Staurothele for species delimitation in the past (Malme for which molecular data were obtained [S. 1919; Vainio 1921). However, culture ex- immersa (A. Massal.) Dalla Torre & Sarnth. periments showed that their shape may vary and S. rupifraga (A. Massal.) Arnold] did depending on the culture conditions and de- not cluster in this lineage, but in another velopmental stages (Ahmadjian & Heikkila¨ group including species of Thelidium A. 1970). This character was therefore suggested Massal. and Polyblastia A. Massal. Finally, as being inappropriate for species delimita- one crustose epilithic species, Staurothele dif- tion in Staurothele and Endocarpon (Thomson fractella, was sister to the genus Endocarpon. 1991). This species, found in North America, was In the past, a number of genera have been transferred to the genus Endocarpon.Asa segregated from Staurothele.Mu¨ller Argo- result, the definition of this genus was viensis (1883) placed the species Staurothele emended to include both squamulose and diffractella (Nyl.) Tuck. in the new genus crustose species (Gueidan et al. 2007). Willeya Mu¨ll. Arg. as its ascospores remain The genus Staurothele has been studied in pale at maturity whereas they become dark various parts of the world. In North America, brown in many other species of Staurothele. Thomson provided the most complete re- Later, the same author described a second vision of the temperate and arctic species species of Willeya, W. rimosa Mu¨ll. Arg., (Thomson & Murray 1988; Thomson 1991, based on a specimen with pale ascospores 2002). It has also been well studied in Japan collected by the French botanist P. Hariot in (Bouly de Lesdain 1921; Harada & Iwatsuki Tonkin, northern Vietnam (Mu¨ller Argo- 1989; Harada 1992), Australia (McCarthy viensis 1889). Clements (1909) described 1995, 2001) and Europe (Malme 1919; Ser- the genus Phalostauris Clem. for Staurothele vı´t 1955; Swinscow 1963; Clauzade & Roux with pale ascospores. These two genera were, 1985; Thu¨s & Schultz 2009). Few species of however, not accepted by most authors and Staurothele have been described from South- were considered as synonyms of Staurothele East Asia: S. australis Groenh. from Java by Thomson (1991) and Brodo et al.(2001). (Groenhart 1954), S. malayensis Zahlbr. from Other synonyms of Staurothele listed in Spe- Java and Sumatra (Zahlbruckner 1934), and cies Fungorum (http://www.speciesfungorum. S. rimosa (Mu¨ll. Arg.) Zahlbr. from Vietnam org, 27/05/2011) are Goidanichia Tomas. & (Mu¨ller Argoviensis 1889). Several species Cif., Goidanichiomyces Cif. & Tomas., Para- have been described from karst in southern physorma A. Massal., Polyblastiomyces E. A. China: S. chlorospora Zahlbr., S. honghensis Thomas, Sphaeromphale A. Massal. and Stig- H. Harada & Li S. Wang, S. kwapiensis matomma Mu¨ll. Arg. Zahlbr., S. microlepis Zahlbr., S. muliensis More recently, molecular phylogenetic Zahlbr., S. ochroplaca Zahlbr., S. sinensis studies have allowed the traditional generic Zahlbr., and S. yunnana H. Harada & Li S. delimitation in Verrucariaceae to be tested Wang (Zahlbruckner 1930; Harada & Wang (Gueidan et al. 2007, 2009; Savic´ & Tibell 1996, 2006). However, the species diversity 2008; Savic´ et al. 2008; Muggia et al. 2010; of the genus in this part of the world remains Prieto et al. 2010, 2012). In studies sup- understudied, as only a few localities have ported by a good sampling of Staurothele spe- been explored. In the large karstic system of cies, the genus was shown to be polyphyletic northern Vietnam, exposed calcareous out- Downloaded from https://www.cambridge.org/core. SLUB Dresden, on 11 Mar 2020 at 13:20:01, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282914000048 2014 Staurothele in Vietnam—Gueidan et al. 517 crops and cliffs are very abundant and con- our material did not match any previously described
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