Blackwell Publishing LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066© 2007 The Linnean Society of London? 2007 91? 455467 Original Article QUERCINA WORLDWIDE VARIABILITY A. ARGÜELLO ET AL

Biological Journal of the Linnean Society, 2007, 91, 455–467. With 1 figure

Parmelina quercina (, ) includes four phylogenetically supported morphospecies

ARTURO ARGÜELLO, RUTH DEL PRADO, PALOMA CUBAS and ANA CRESPO*

Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, 28040 Madrid, Spain

Received 23 November 2005; accepted for publication 10 August 2006

Morphological and phylogenetic relationships of the worldwide Mediterranean forming , Parmelina quercina, have been studied. Specimens from western Europe, western North America and southern Australia were analysed using molecular data (nuITS rDNA, nuLSU rDNA and mtSSU rDNA) and selected morphological features (upper cortex maculae, scanning electron microscopy examination of the epicortex, ascospores and conidia shape and size, and amphithecial retrorse rhizines). The results conclusively reveal that: (1) there is not one single species but four separate species in the Mediterranean or sub Mediterranean areas of the world. Parmelina quercina and Parmelina carporrhizans (Euroasiatic species), Parmelina coleae sp. nov. (North America) and Parmelina elixia sp. nov. (Australia); (2) largely debated P. carporrhizans is not a synonym of P. quercina but supported as a valid spe- cies circumscribed to Macaronesic relict sites; (3) the geographical isolation of the Australian population is correlated with a large genetic distance; (4) morphological characters (ascospores and conidial variability and thallus epicortex) correlate with the phylogenetic hypothesis; (5) the new or revalidated species within Parmelina quercina are not cryptic species but morphologically recognizable taxa. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 455–467.

ADDITIONAL KEYWORDS: cryptic species – Macaronesic species – Mediterranean species – Parmelina carporrhizans – Parmelina coleae – Parmelina elixia.

INTRODUCTION northern Africa and western India is particularly common in the Mediterranean region of the Iberian The genus Parmelina Hale (Parmeliaceae) currently Peninsula and Morocco. Parmelina pastillifera, includes 13 species distributed in the temperate another isidiate taxon, is an oceanic species scattered regions of the northern and southern Hemisphere throughout Europe and Britain (Schauer, 1965), (Elix, 1993). For the most part, three species are rec- locally abundant in humid oceanic areas of the west- ognized in Europe: Parmelina quercina (Willd.) Hale, ern Mediterranean region. (Hoffm.) Hale, and Parmelina pas- Parmelina quercina is a sexual, fertile species that tillifera (Harm.) Hale, although some authors consider grows on evergreen, sclerophyllous and marcescent that Parmelina carporrhizans (Taylor) Poelt & Vezda vegetation associated with mediterranean climates represents a fourth (Poelt & Vezda, 1977; Nimis, and has been cited from four disjunctive areas: (1) 1993). However, the latter is frequently considered a around the Mediterranean basin extending to the synonym of P. quercina (Culberson, 1961; Hale, 1976; Canary Islands, but also occurring in mild microcli- Clauzade & Roux, 1985; Purvis et al., 1992). A further matic sites in oceanic and interior regions of northern related species (Parmelina gyrophorica Elix, Sheng L. and central Europe; currently reported as extinct from Wang & J. B. Chen) has been described from China Danish Islands (Sochting & Alstrup, 2002; Thell, (Wang, Chen & Elix, 2001; Chen, Wang & Elix, 2003). Westberg & Kärnefelt, 2004), and with scattered The isidiate species P. tiliacea known from Europe, reports as far east as Japan (Hale, 1976; Kurokawa,

. 2003), in China (Hale, 1976; Wei, 1991) and Thailand (Wolseley, Aguirre-Hudson & McCarthy, 2002); *Corresponding author. E-mail: [email protected] (2) North America (southern California and Baja

© 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 455–467 455