A review of the lichen family Parmeliaceae - history, phylogeny and current taxonomy

Arne Thell

Ana Crespo,1 Frank Kauff,2 Pradeep K. Divakar,1 Ruth del Prado,1 Sergio Pérez-Ortega,1 Guillermo Amo de Paz,1 Zuzana Ferencova,1 Oscar Blanco,3 Beatriz Roca-Valiente,1 Jano Núñez-Zapata,1 Paloma Cubas,1 Arturo Argüello,1 John A. Elix,4 Theodore L. ...

Phylogeny of the cetrarioid core (Parmeliaceae) based on five genetic markers Arne THELL, Filip HÖGNABBA, John A. ELIX, Tassilo FEUERER, Ingvar KÄRNEFELT, Leena MYLLYS, Tiina RANDLANE, Andres SAAG, Soili STENROOS, Teuvo AHTI and Mark RD SEAWARD ...

Recently, molecular phylogenetic studies have revolutionized the generic concepts in Parmeliaceae and in lichen forming fungi in general. In the present study, the generic delimitation in the Hypotrachyna clade is revised using a molecular phylogeny of nuclear ITS, LSU and mitochondrial SSU rDNA sequences of 88 hypotrachynoid taxa. Morphological and chemical features are also revised in each group. 118 sequences are newly generated for this study. Our phylogenetic analyses show the polyphyly of Hypotrachyna as currently circumscribed which falls into four well-supported and one unsupported clade. Cetrariastrum, Everniastrum and Parmelinopsis are nested within Hypotrachyna s. lat., Parmelinopsis being also polyphyletic and nested in one of the Hypotrachyna clades. Cetrariastrum is monophyletic but clustered within Everniastrum. Two alternative hypotheses tests significantly rejected the monophyly of these three genera. As a consequence, the genera Cetrariastrum, Everniastrum, and Parmelinopsis are reduced to synonymy with Hypotrachyna. Furthermore, we here propose an alternative classification to recognize the well-supported clades at subgeneric level and leave the remaining species unclassified within the genus. Five new subgenera are proposed: Hypotrachyna subgen. Cetrariastrum, Hypotrachyna subgen. Everniastrum, Hypotrachyna subgen. Longilobae, Hypotrachyna subgen. Parmelinopsis, and Hypotrachyna subgen. Sinuosae. Forty-nine new combinations are proposed.

Nordic Journal of Botany 30: 641–664, 2012 doi: 10.1111/j.1756-1051.2012.00008.x, © 2012 The Authors. Nordic Journal of Botany © 2012 Nordic Society Oikos Subject Editor: Torbjörn Tyler. Accepted 11 October 2012 A review of the lichen family Parmeliaceae – history, phylogeny and current taxonomy A. Thell (arne.thell@biol.lu.se) and I. Kärnefelt, Botanical Museum, Box 117, Lund Univ., SE-221 00 Lund, Sweden. – A. Crespo and P. K. Divakar, Depto de Biología Vegetal II, Facultad de Farmacia, Univ. Complutense de Madrid, ES-28040 Madrid, Spain. – S. D. Leavitt and H. T. Lumbsch, Dept of Botany, The Field Museum, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA. – M. R. D. Seaward, Dept of Archaeological, Geographical and Environmental Sciences, Univ. of Bradford, Bradford BD7 1DP, UK. The largest family of lichen-forming fungi, the Parmeliaceae, is reviewed. It includes 79 genera in current use and ca 2726 species, a large majority of which belong to one of five main clades: the parmelioid, cetrarioid, usneoid, alectorioid and hypogymnioid. However, 21 genera are positioned outside these clades, and four genera in current use still lack DNAdata. The family has been shown to be monophyletic and the generic classification is relatively well-settled compared with other lecanoralean families. Each clade and its genera are presented here with the latest results from phylogenetic analyses and current taxonomy. In addition, a historical outline of the family and its most prominent researchers is provided. Recently, the estimated number of lichen species worldwide has increased to ca 28 000, mainly due to increased research activity on their biodiversity in tropical regions (Lücking et al. 2009). The large majority of lichenized fungi are ascomycetes classified into ca 60 lichenized orders. The majority of these lichenized fungi belong to the almost entirely lichen-forming fungal order Lecanorales comprising 20 families. The Parmeliaceae is the largest family of lichenforming ascomycetes and includes well-known groups such as beard-lichens, Usnea, and shield-lichens, Parmelia et al. (Lumbsch and Huhndorf 2010). After two centuries of taxonomic splitting, which accelerated in the late 1960s, the systematics of Parmeliaceae is gradually stabilizing with help of DNA-based phylogenies. At present ca 2726 species are included in ca 79 genera (with three genera excluded here as discussed below) in current use (Table 1). The Parmeliaceae is primarily a Southern Hemisphere family with distributions centered in southern Africa, South America and Australia. However, the circumscriptions of several genera remain problematic and new species are still frequently discovered, mainly in the tropics, but also including several phenotypically cryptic and semi-cryptic lineages in groups which are generally considered wellinvestigated (Lumbsch and Leavitt 2011). Since the survey of Parmelia sensu lato by Elix (1993), no attempt has been made to circumscribe all genera within this family. The present review addresses the development from Acharius’s early systematics of Parmeliaceae based on morphology to today’s phylogenetic system based largely on DNA sequence data. The genera are briefly presented with comments on their features, relationships and distribution. Authors of genera of the Parmeliaceae are omitted in the text, as they are listed in Table 1. The delimitation of the family The family Parmeliaceae, as proposed by Eschweiler (1824), comprised six genera that are now understood to be only distantly related: Collema Wigg. (now in the Collemataceae), Cornicularia, Hagenia Eschw. (a synonym of Anaptychia Körb. in the Physciaceae), Lecanora Ach. (now in the Lecanoraceae), Parmelia and Sticta (Schreb.) Ach. (now in the Stictaceae). Of these, only Parmelia and Cornicularia belong to the Parmeliaceae in its modern sense (Lumbsch and Huhndorf 2010). When describing the family, Eschweiler (1824) mentioned the shield-like apothecia, surrounded by a thalline margin. The structure of the apothecia has been the most important character for delimiting the family, even in its modern sense (Henssen and Jahns 1973). The Usneaceae (Eschweiler 1824), later included in the Parmeliaceae, included the genera Cetraria, Evernia (with Alectoria and Ramalina Ach.) and Usnea. Hale (1983) excluded Cetraria but added Dactylina and Letharia among those genera accepted in the Parmeliaceae today. Additional families, now included in the Parmeliaceae, have been recognized over the years: Alectoriaceae (Hue) Tomas., Anziaceae M. Satô, Cetrariaceae Schaer., Corniculariaceae Schaer., Everniaceae (Hue) Tomas. and Hypogymniaceae Poelt ex 641 Special Invited Review Arne Thell, Ana Crespo, Pradeep K. Divakar, Ingvar Kärnefelt, Steven D. Leavitt, H. Thorsten Lumbsch and Mark R. D. Seaward Special Invited Review 642 Table 1. Genera of Parmeliaceae in current use, their authors, type species and number of species included. Genus Ahtiana Goward Alectoria Ach. Allantoparmelia (Vain.) Essl. Allocetraria Kurok. & M. J. Lai Anzia Stitzenb. Arctocetraria Kärnefelt & A. Thell Arctoparmelia Hale Asahinea W. L. Culb. & C. F. Culb. Austroparmelina A. Crespo et al. Brodoa Goward Bryocaulon Kärnefelt Bryoria Brodo & D. Hawksw. Bulborrhizina Kurok. Bulbothrix Hale Canoparmelia Elix & Hale Cetraria Ach. Cetrariastrum Sipman Cetrariella Kärnefelt & A. Thell Cetrelia W. L. Culb. & C. F. Culb. Cetreliopsis M. J. Lai Coelopogon Brusse & Kärnefelt Cornicularia (Schreb.) Hoffm. Dactylina Nyl. Davidgallowaya Aptroot Emodomelanelia Divakar & A. Crespo Esslingeriana Hale & M. J. Lai Evernia Ach. Everniastrum Hale ex Sipman Everniopsis Nyl. Flavocetraria Kärnefelt & A. Thell Flavoparmelia Hale Flavopunctelia (Krog) Hale Gowardia Halonen et al. Himantormia I. M. Lamb. Hypogymnia Nyl. Hypotrachyna (Vain.) Hale Imshaugia S. L. F. Mey Kaernefeltia A. Thell & Goward Letharia (Th. Fr.) Zahlbr. Lethariella (Motyka) Krog Masonhalea Kärnefelt Melanelia Essl. Melanelixia Blanco et al. Melanohalea Blanco et al. Menegazzia A. Massal. Reference Type No. species Bryologist 88 (1985, p. 370) In Luyken, Tent. Hist. Lich. (1809, p. 95) Mycotaxon 7 (1978, p. 46) Bull. Nat. Sci. Mus., Tokyo, ser. B 17 (1991, p. 60) Flora 44 (1861, p. 390) Bryologist 96 (1993, p. 402) Mycotaxon 25 (1986, p. 251) Brittonia 17 (1965, p. 183) Syst. Biodivers. 8 (2010, p. 216) Bryologist 89 (1986, p. 222) Opera Bot. 86 (1986, p. 17) Opera Bot. 42 (1977, p. 78) Acta Bot. Fenn. 150 (1994, p. 105) Phytologia 28 (1974, p. 479) Mycotaxon 27 (1986, p. 277) Meth. Lich. (1803, p. 292) Proc. Kon. Ned. Akad. Wetensch., ser. C, Biol. Med. Sci. 83 (1980, p. 335) Bryologist 96 (1993, pp. 402–403) Contr. U. S. Natl Herb. 34 (1968, p. 490) Quart. J. Taiwan Mus. 33 (1980, p. 218) Mycotaxon 42 (1991, p. 35) Descr. Adumb. Plant. Lich. 2 (1794, p. 36) Syn. Meth. Lich. 1 (1860, p. 286) Bibl. Lich. 95 (2007, p. 139) Taxon 59 (2010, pp. 1749–1750) Quart. J. Taiwan Mus. 33 (1980, p. 220) Lich. Univ. (1810, pp. 84, 441) Mycotaxon 26 (1986, p. 237) Syn. Meth. Lich. (1860, p. 374) Acta Bot. Fenn. 150 (1994, p. 81) Mycotaxon 25 (1986, p. 604) Mycotaxon 20 (1984, pp. 681–682) Bryologist 112 (2009, p. 141) Brit. Antarc. Surv. Sci. Rep. 38 (1964, pp. 17–18) Lich. Env. Paris (1896, pp. 39, 139) Phytologia 28 (1974, p. 340) Mycologia 77 (1985, p. 373) Bryologist 99 (1996, p. 125) Hedwigia 31 (1892, p. 34) Norw. J. Bot. 23 (1976, p. 88) Bot. Not. 130 (1977, pp. 101–102) Mycotaxon 7 (1978, p. 46) Mycol. Res. 108 (2004, p. 881) Mycol. Res. 108 (2004, p. 882) Neagen. Lich. (1854, p. 1) A. sphaerosporella (Müll. Arg.) Goward A. sarmentosa (Ach.) Ach. A. alpicola (Th. Fr.) Essl. A. stracheyi (Bab.) Kurok. & M. J. Lai A. colpodes (Ach.) Stitzenb. A. andrejevii (Oxner) Kärnefelt & A. Thell A. centrifuga (L.) Hale A. chrysantha (Tuck.) W. L. Culb. & C. F. Culb. A. pseudorelicina (Jatta) A. Crespo et al. B. oroarctica (Krog) Goward B. divergens (Ach.) Kärnefelt B. trichodes (Michx.) Brodo & D. Hawksw. B. africana Kurok. B. semilunata (Lynge) Hale C. texana (Tuck.) Elix & Hale C. islandica (L.) Ach. C. ecuadoriense (R. Sant.) Sipman C. delisei (Bory ex Schaer.) Kärnefelt & A. Thell C. cetrarioides (Delise) W. L. Culb. & C. F. Culb. C. rhytidocarpa (Mont. & v. d. Bosch) M. J. Lai C. abraxas Brusse C. normoerica (Gunnerus) DuRietz D. arctica (Richardson) Nyl. D. cornutispora Aptroot E. masonii (Essl. & Poelt) Divakar & A. Crespo E. idahoensis (Essl.) Hale & M. J. Lai E. prunastri (L.) Ach. E. cirrhatum (Fr.) Hale ex Sipman E. trulla (Ach.) Nyl. F. cucullata (Bell.) Kärnefelt & A. Thell F. caperata (L.) Hale F. flaventior (Stirton) Hale G. nigricans (Ach.) Halonen et al. H. lugubris (Hue) I. M. Lamb. H. physodes (L.) Nyl. H. brasiliana (Nyl.) Hale I. aleurites (Ach.) S. L. F. Mey. K. californica (Tuck.) A. Thell & Goward L. vulpina (L.) Hue L. intricata (Moris) Krog M. richardsonii (Hook.) Kärnefelt M. stygia (L.) Essl. M. glabra (Schaer.) O. Blanco et al. M. exasperata (De Not.) O. Blanco et al. M. terebrata (Hoffm.) A. Massal. 3 7 3 9 ca 34 2 5 2 13 3 4 ca 51 1 58 ca 40 15 5 4 18 7 2 1 2 1 1 1 10 ca 40 1 3 ca 32 5 2 2 ca 90 ca 188 7 2 6 11 2 ca 13 15 22 ca 70 Myelochroa (Asahina) Elix & Hale Nephromopsis Müll. Arg. Nesolechia A. Massal. Nipponoparmelia (Kurok.) K. H. Moon et al. in A. Crespo et al. Nodobryoria Common & Brodo Omphalodium Meyen & Flot. Omphalora T. H. Nash & Hafellner Mycotaxon 29 (1987, p. 240) Flora 74 (1891, p. 374) Misc. Lichenol. (1856, p. 43) Taxon 59 (2010, p. 1749) Oropogon Th. Fr. Pannoparmelia (Müll. Arg.) Darb. Parmelia Ach. Parmelina Hale Parmelinella Elix & Hale Parmelinopsis Elix & Hale Parmeliopsis (Nyl.) Nyl. Parmotrema A. Massal. Parmotremopsis Elix & Hale Platismatia W. L. Culb. & C. F. Culb. Pleurosticta Petr. Gen. Heterolich. Eur. recogn. (1861, p. 49) Wiss. Ergebn. Schwed. Südpol.-Exp. 1901–03 4 (1912, p. 11) Meth. Lich. (1803, p. 153) Phytologia 28 (1974, p. 481) Mycotaxon 29 (1987, p. 241) Mycotaxon 29 (1987, p. 242) Not. Sällsk. Fauna Flora Fenn. Förh. 8 (1866, p. 121) Atti Reale Veneto Sci. Lett. Arti, ser. 3, 5 (1860, p. 248) Mycotaxon 29 (1987, p. 243) Contr. U. S. Natl Herb. 34 (1968, pp. 524–525) Kryptog. Forsch. 2 (1931, p. 190) Protousnea (Motyka) Krog Pseudephebe M. Choisy Pseudevernia Zopf Pseudoparmelia Lynge Psiloparmelia Hale Punctelia Krog Relicina (Hale & Kurok.) Hale Relicinopsis Elix & Verdon Remototrachyna Divakar & A. Crespo Sulcaria Bystr. Tuckermanella Essl. Tuckermannopsis Gyeln. Usnea Adans. Usnocetraria M. J. Lai & J. C. Wei Vulpicida J.-E. Mattsson & M. J. Lai Xanthoparmelia (Vain.) Hale Norw. J. Bot. 23 (1976, p. 96) Icon. Lich. Univ-, ser. 2 (1930, sine p.) Beih. Bot. Centralbl. 14 (1903, p. 124) Ark. Bot. 13 (1914, p. 15) Mycotaxon 35 (1989, p. 42) Nord. J. Bot. 2 (1982, p. 290) Phytologia 28 (1974, p. 484) Mycotaxon 27 (1986, p. 281) Am. J. Bot. 97 (2010, p. 584) Ann. Univ. Mariae Curie Sklodowska, Sect. C, Biol. 26 (1971, p. 275) Mycotaxon 85 (2003, p. 135) Acta Fauna Flora Univ. 2, 1 (1933, p. 6) Fam. Pl. 2 (1763, p. 7) J. Nat. Taiwan Mus. 60 (2007, p. 45) Mycotaxon 49 (1993, p. 427) Phytologia 28 (1974, p. 485) Bryologist 98 (1995, p. 198) Nova Acta Acad. Caes. Leop. Carol. Nat. Cur. Dresden 19 (1843, p. 223) Lichenologist 22 (1990, p. 356) M. aurulenta (Tuck.) Elix & Hale N. stracheyi (C. Bab.) Müll. Arg. N. oxyspora (Tul.) A. Massal. N. laevior (Nyl.) K. H. Moon et al. ex A. Crespo et al. N. abbreviata (Müll. Arg.) Brodo & Common O. pisacomense Meyen & Flot. O. arizonica (Tuck. Ex Willey) T. H. Nash & Hafellner O. loxensis (Fée) Th. Fr. P. angustata (Pers.) Zahlbr. P. saxatilis (L.) Ach. P. tiliacea (Hoffm.) Hale P. wallichiana (Taylor) Elix & Hale P. horrescens (Taylor) Elix & Hale P. ambigua (Wulfen) Nyl. P. perforatum (Wulfen) A. Massal. P. antillensis (Nyl.) Elix & Hale P. glauca (L.) W. L. Culb. & C. F. Culb. P. lichenicola Petr. in Kryptog. Forsch. 2 (1931, p. 190) ( ! pycnidia of P. acetabulum (Neck.). Lumbsch & Elix in Lumbsch, Kothe & Elix in Mycotaxon 33 (1988, p. 453) P. magellanica (Motyka) Krog P. pubescens (L.) M. Choisy P. furfuracea (L.) Zopf P. cyphellata Lynge P. distincta (Nyl.) Hale P. borreri (Sm.) Krog R. eumorpha (Hepp) Hale R. intertexta (Mont. & Bosch) Elix & Verdon R. flexilis (Kurok.) Divakar & A. Crespo S. sulcata Bystrek T. weberi (Essl.) T. ciliaris (Ach.) Gyeln. U. florida (L.) F. H. Wigg. U. oakesiana (Tuck.) M. J. Lai & J. C. Wei V. juniperinus (L.) J.-E. Mattsson & M. J. Lai X. conspersa (Ehrh. ex Ach.) Hale ca 30 21 41 4 3 4 1 ca 40 5 ca 57 9 7 ca 25 3 ca 300 2 11 2 8 2 4 15 13 ca 45 ca 54 5 15 4 6 9 ca 350 1 6 ca 820 643 Special Invited Review Special Invited Review Elix. Among these family names, Alectoriaceae, Anziaceae, Hypogymniaceae and Usneaceae have been used in modern times (Poelt 1974, Elix 1979, Elix and James 1992, Galloway 1992, Kärnefelt et al. 1998). All these families were accepted by Hale (1983). Alectoriaceae, proposed by Tomaselli (1949), included three brown-spored, fruticose genera, Alectoria, Oropogon and Sulcaria (Kärnefelt and Thell 1992, Eriksson and Hawksworth 1998, Kärnefelt et al. 1998), until analyses of molecular data showed that these genera also belong in the Parmeliaceae (Mattsson and Wedin 1999). Parmeliaceae and its contributors Some genera today included in the Parmeliaceae are older than the family. Linnaeus [1707–1778] recognized 26 species in ‘Species Plantarum’, which today belong in the Parmeliaceae. Within the single genus Lichen (Linnaeus 1753), Linnaeus’ last pupil, Eric Acharius [1757–1819] (Fig. 1), known as the father of lichenology (Kärnefelt and Frödén 2007), attempted to arrange lichens into natural groups based on thallus morphology: Lichen crustacei, L. foliacei and L. caulescentes (Acharius 1794, 1795). Some years later he presented the lichens under new rules with species arranged under new generic names (Acharius 1803), the genera organized according to the form and position of apothecia. Six genera within the current circumscription of the Parmeliaceae predate the family, four of which were proposed by Acharius: Alectoria, Cetraria, Evernia and Parmelia (Acharius 1803, 1809, 1810). Two genera, Usnea (Adanson 1763) and Cornicularia (Hoﬀmann 1794), had been proposed earlier. William Nylander [1822–1899], who described several thousand species from a broad array of lichen groups (Kärnefelt 2009), contributed five genera to the Parmeliaceae (Nylander 1860, 1866, 1869, 1896), of which four are still in current use: Dactylina, Everniopsis, Hypogymnia and Parmeliopsis. The genus Chondropsis Nyl. ex Crombie, traditionally composed of two species, was synonymized with the younger Xanthoparmelia to avoid recombination of more than 800 species names (Hawksworth and Crespo 2002; see note under Xanthoparmelia below). Further genera that were proposed in the 19th century are Anzia, Letharia, Menegazzia, Nephromopsis, Nesolechia, Neuropogon Nees and Flot. (now included in Usnea), Omphalodium and Oropogon (Table 1). The Parmeliaceae as circumscribed by Zahlbruckner (1926) was an unnatural group, whereas Hillmann (1936) included four genera in his treatment of the family. Three of them, Cetraria, Parmelia and Parmeliopsis, belong to Parmeliaceae today. However, he also included Candelaria A. Massal., presently positioned in the Candelariaceae Hakul. Aino Henssen [1925–2011] (Fig. 2), a well-known professor in Marburg for many years (Kärnefelt 2009, Kärnefelt et al. 2012), circumscribed the higher taxa of lichenized ascomycetes based on ascocarp ontogeny. Members (in 18 genera) of the Parmeliaceae were characterized by an ascoma ontogeny forming a cupular exciple (Henssen and Jahns 1973), which is the single character that unites the whole family as circumscribed today (Crespo et al. 2007). 644 William L. Culberson [1929–2003] and Chicita F. Culberson [∗1931] (1965, 1968) first split the family into new and more natural genera, describing the new cetrarioid genera Asahinea, Cetrelia and Platismatia, a trend of splitting that continued for more than three decades. Mason E. Hale [1928–1990] (Fig. 3), a prominent systematist of the Parmeliaceae, travelled and collected extensively, visiting all continents (Kärnefelt 2009). He discovered and described hundreds of species and divided them into many new genera, based on external morphology, cortex anatomy and chemistry as diagnostic characters (summarized by Crespo et al. 2011). However, this subdivision of genera was controversial and not generally accepted (Poelt and Vezda 1981, Santesson 1984, Clauzade and Roux 1985, Purvis et al. 1992, Wirth 1995). One of William Culberson’s students, Theodore L. Esslinger [∗1944], in his monograph of the brown Parmeliae (Esslinger 1977), recognised three new segregates, Allantoparmelia, Melanelia and Neofuscelia (Esslinger 1978), then not generally accepted as separate from Parmelia. Today, these genera have been shown to belong to diﬀerent phylogenetic lineages within the family. Irwin Brodo [∗1935] and David Hawksworth [∗1946] showed the heterogenity of the alectorioid lichens, proposing the new genus Bryoria (Brodo and Hawksworth 1977) while Ingvar Kärnefelt [∗1944] arranged the cetrarioid group, segregating the genera Bryocaulon (Kärnefelt 1986) and Coelopogon (Brusse and Kärnefelt 1991). Hale frequently cooperated with an Australian professor of chemistry, John A. Elix [∗1941] (Fig. 4) (Kärnefelt 2009), with whom he described several new genera and many species in the Parmeliaceae in the 1980s. Elix had become interested in lichens when research on the secondary metabolic products was rising fast, mainly due to the work of the Culbersons. Elix background in chemistry was of profound importance in his analysis of lichen compounds through TLC, HPLC, mass spectrometry and NMR. The collaboration between Hale and Elix, included structural characters and new chemical data obtained from highly advanced methods, and resulted in several new genera (Elix 1993, Elix et al. 1986, Elix and Hale 1987). The frequent use of single characters to define parmelioid genera, frequently cortex structure or secondary compounds, and the lack of correlation between reproductive and vegetative characters, were the main arguments against the generic segregation presented by Hale and co-workers (Hawksworth et al. 1980, Nimis 1998). However, the genera created by Hale and Elix were later accepted, in part, in most floras and checklists (Santesson 1993, Feuerer 1998, Scholz 2000, Brodo et al. 2001, Coppins 2002, Santesson et al. 2004, Divakar and Upreti 2005a, Smith et al. 2009), and many of them have been supported by molecular data. Except for some new cetrarioid genera erected largely during the first half of the 1990s, only a few new genera were segregated in the 1990s, of which Omphalora (Nash et al. 1990) and Nodobryoria (Common and Brodo 1995) are still in current use. The monotypic Bulbothricella (Marcano et al. 1996) was almost immediately reduced to synonymy with Bulbothrix (Lumbsch 1997). Davidgallowaya and Usnocetraria are the genera most recently described Special Invited Review Figure 1–4. Parmeliaceae researchers. (1) Eric Acharius [1757–1819], the father of lichenology and creator of the well-known genera of the Parmeliaceae Alectoria, Cetraria, Evernia and Parmelia, (2) Aino Henssen [1925–2011], discoverer of the cupular exciple, the single non-molecular character uniting Parmeliaceae as circumscribed today, (3) Mason Hale [1928–1990] described hundreds of species in Parmeliaceae, (4) Jack Elix [∗1941] cooperated intensely with Mason Hale with whom he discovered and described a large number of species and genera. without DNA-data (Aptroot 2007, Lai et al. 2007). Many of the most recently described genera appear as monophyletic clades and generally have correlating morphological and or geographical features (Crespo et al. 2010a, 2010b, Divakar et al. 2010a). Nevertheless, several genera known to be polyphyletic still remain and additional molecular data will doubtlessly lead to further taxonomic changes in the family (Divakar et al. 2006, Thell et al. 2009, Crespo et al. 2010b). Morphology, chemistry and distribution The three main growth forms in lichens, crustose, foliose and fruticose, proposed by Acharius were soon abandoned as systematic entities. However, until recently these morphotypes were continuously used for practical reasons and have gradually also been further subdivided. In the Parmeliaceae several major groups have been circumscribed according to the appearance of the thallus, including: alectorioid, parmelioid, cetrarioid and usneoid lichens, named after four genera of the family. Alectorioid lichens are fruticose, often beard-like, pendent or caespitose. Cetrarioid lichens are erect foliose with marginal apothecia and pycnidia. Parmelioid lichens are mainly foliose, often adnate to the substratum and possess laminal apothecia and pycnidia. Usneoid lichens comprise a heterogeneous group of fruticose and strapshaped lichens (Kärnefelt et al. 1998, Crespo et al. 2007). 645 Special Invited Review Allantoparmelia (3) Anzia (ca 34) Asahinea (2) Bulborrhizina* (1) Bryocaulon (4) Coelopogon (2) Cornicularia (1) Davidgallowaya* (1) Evernia (10) Himantormia (2) Imshaugia (7) Menegazzia (ca 70) Nodobryoria (3) Omphalodium (4) Omphalora (1) Oropogon (ca 40) Parmotremopsis* (2) Platismatia (11) Pseudoparmelia* (15) Pannoparmelia (5) Protousnea (8) Usnea (ca ( 350)) Arctoparmelia (5) Brodoa (3) Hypogymnia (ca 90) Pseudevernia (4) Bryoria (ca 51) Alectoria (7) Gowardia (2) Pseudephebe (2) Sulcaria (4) Letharia (6) Lethariella (11) ( ) Everniopsis (1) Psiloparmelia (13) Melanelia† (ca 13) Dactylina (2) Esslingeriana (1) Masonhalea (2) Cetrariella (4) Allocetraria (9) Vulpicida (6) Cetraria (15) Usnocetraria (1) Arctocetraria (2) Kaernefeltia (2) Tuckermanella (6) Tuckermannopsis (9) Cetreliopsis (7) Nephromopsis (21) Ahtiana (3) Flavocetraria (3) Nipponoparmelia (4) Parmelia (ca 57) Parmeliopsis (3) Relicina (ca 54) Relicinopsis (5) Pleurosticta (2) Emodomelanelia (1) Melanelixia (15) Melanohalea (22) Myelochroa (ca 30) Parmelina (9) Bulbothrix (58) Parmelinella (7) Remotrachyna (15) Cetrariastrum (5) Everniastrum (ca 40) Hypotrachyna (ca 188) Parmelinopsis (ca 25) Cetrelia (18) Xanthoparmelia (ca 820) Canoparmelia (ca 40) Flavopunctelia (5) Nesolechia (41) Punctelia (ca 45) Austroparmelina (13) Parmotrema (ca 300) Flavoparmelia (ca 32) Genera with uncertain affinities (A) –21 genera –ca 226 species (B) (C) Usneoid clade Hypogymnioid clade –4 genera –ca 102 species Alectorioid clade (D) –5 genera –ca 66 species Letharioid clade Psiloparmelioid clade Cetrarioid clade –17 genera –ca 97 species (E) (F) Parmelioid clade –27 genera –ca 1854 species (G) (H) (I) Figure 5. Constructed phylogeny of Parmeliaceae based on studies by Crespo et al. (2010b) and Thell et al. (2009). Selected species representing clades and genera in Parmeliaceae: (A) Asahinea chrysantha (uncertain aﬃnity), (B) Usnea hirta (usneoid clade), (C) Hypogymnia imshaugii (hypogymnioid clade), (D) Alectoria sarmentosa (alectorioid clade), (E) Letharia lupina ined. (letharioid clade), (F) Cetraria islandica (cetrarioid clade), (G) Vulpicida tilesii (cetrarioid clade), (H) Parmelia saxatilis (parmelioid clade), (I) Xanthoparmelia chlorochroa (parmelioid clade). Photos kindly provided by Curtis Björk, Jason Hollinger, and the ‘Ways of enlichenment’ (" www. waysofenlichenment.net/ #). ∗ ! genera still not investigated for DNA, † ! Melanelia s.s. belongs to the cetrarioid clade (4 species) and ca 9 Melanelia s.l. species belong to the parmelioid clade, see text. 646 A new classification based on DNA sequence data Several phylogenetic analyses have supported the circumscription of Parmeliaceae in its broad sense (Crespo and Cubero 1998, Mattsson and Wedin 1998, 1999, Wedin et al. 1999, Crespo et al. 2001, 2007, 2010b, Blanco et al. 2004b, Thell et al. 2004). The closest relatives of the family Parmeliaceae are presumably found in the Gypsoplacaceae Timdal and in Protoparmelia M. Choisy and allied genera. The crustose genus Protoparmelia, placed either in the Lecanoraceae Körb. or the Parmeliaceae, has occasionally been used as an outgroup of the Parmeliaceae in phylogenetic studies. However, according to a study by Arup et al. (2007), its taxonomic position cannot be supported without also including the family Gypsoplacaceae. Sequences from several loci have been used in studies of Parmeliaceae phylogeny: nucITS, nucSSU, nucLSU, mitSSU rDNA, beta tubulin, glyceraldehyde-3-phosphate dehydrogenase, RPB1, and MCM7 (Crespo et al. 2001, 2010b, Myllys et al. 2002, Leavitt et al. 2011, 2012). Almost all genera in current use in the Parmeliaceae have been studied using molecular data and the phylogeny of the family is well-settled. Some genera have isolated positions in the family but most genera belong to either of the more or less well supported clades discussed below (Fig. 5). Since the DNA-based phylogenetic analyses started in the late 1990s, the number of accepted genera has decreased from ca 90 to ca 79. However, nine genera have been described during the same period. Seven of them (Austroparmelina, Emodomelanelia, Gowardia, Melanelixia, Melanohalea, Nipponoparmelia and Remototrachyna) are based mainly on DNA-evidence corroborated by phenotypic features and/or biogeography (Table 1). As a result of wide collaboration among researchers, workshops and joint publications, there is a large degree of concordance in Parmeliaceae taxonomy. The new genera described and listed in Table 1 constitute monophyletic clades. However, some polyphyletic genera remain in the family, often because of a hesitation to draw immediate taxonomic conclusions from phylogenetic analyses in addition to a resistance against erecting genera without any known morphological or chemical diagnostic characters. The number of species described annually within the Parmeliaceae has increased since the turn of the millennium, partly as a result of more extensive research activity in tropical and subtropical areas, partly through phylogenetic analyses (Pooprang et al. 1999, Elix et al. 2000, Kurokawa and Moon 2000, Louwhoﬀ and Elix 2000, Elix and Schumm 2001, Divakar et al. 2001, 2005, Marcelli and Ribeiro 2002, Divakar and Upreti 2003, 2005b, Elix 2003a, Sinha and Elix 2003). A new category of species with subtle diagnostic characters, and even in some cases species lacking diagnostic morphological or chemical characters have been discovered and described as the result of molecular analyses. Most of these phenotypically cryptic or semicryptic species cannot be distinguished without DNAdata. Previously unknown morphological characters have occasionally been discovered when the phylogenetic results are known (Feuerer and Thell 2002, Molina et al. 2004, Divakar et al. 2005, 2010b, Crespo and Perez-Ortega 2009). Recent examples of cryptic or semicryptic species in Parmeliaceae include Parmelia encryptata A. Crespo et al. (morphologically indistinguishable from P. sulcata Taylor), Parmelia mayi Divakar et al. (visibly identical with P. saxatilis, but with a diﬀerent ecology, distribution and secondary chemistry) (Molina et al. 2011a, 2011b) and Parmelina cryptotiliacea (similar to P. tiliacea but with a restricted geographical distribution and showing subtle morphological diﬀerences in excipular tissues and spores) (Núñez-Zapata et al. 2011). Parmeliaceae phylogeny and its genera in current use The Parmeliaceae includes approximately one tenth of all lichen species. Most species and genera belong to either of 647 Special Invited Review No exact limits exist between these groups and they do not represent evolutionary lines, although some correlation can be recognized with the alectorioid, cetrarioid, parmelioid and usneoid clades (Fig. 5). In addition to these main forms, subcrustose and peltate lichens occur in the genus Xanthoparmelia (Amo de Paz et al. 2010a, 2010b). The parasitic genus Nesolechia, with vegetative hyphae that are almost completely hidden in the host thallus, was shown to belong in the Parmeliaceae using DNA-analyses (Peroh and Rambold 2002), a conclusion later supported by Crespo et al. (2007, 2010b). Elix (1993) summarized the characters most frequently used in Parmeliaceae systematics, and Crespo et al. (2011) reviewed the taxonomic significance of main characters in parmelioid lichens based on molecular phylogeny. In addition to the growth form, considerable focus has been paid to the nature of the cortex and pseudocyphellae, position and structure of the apothecia, shape and size of the ascospores, pycnidial and conidial characters, chemical compounds, biogeography and ecology. The appearance of the asci has been considered of particular importance in lichen taxonomy (Bellemère and Letrouit-Galinou 1981, Hafellner 1984, Kärnefelt et al. 1992, Thell et al. 1995a). However, when it became possible to perform phylogenetic analyses based on DNA-sequences in the late 1990s, it was evident that the conidial shape was the morphological character that correlated best with molecular phylogenetic reconstructions (Thell et al. 2004). In cetrarioid lichens the two main clades correlate almost perfectly with the two main types of conidia found in Parmeliaceae, namely conidia with a single swelling in the Cetraria clade and conidia with double swelling in the Nephromopsis clade (Thell et al. 2002). Bottle- or lemon-shaped conidia have a single swelling that is terminally or centrally positioned. Conidia with double-swelling are thickened more or less terminally, being dumbbell-shaped. Biogeography also shows considerable correlation with DNA data. Members of the Parmeliaceae are primarily found in the Southern Hemisphere where the species-rich parmelioid and Usnea clades are represented by large genera, such as Hypotrachyna, Menegazzia, Parmotrema, Usnea and Xanthoparmelia, having their main distributions in subtropical to temperate areas. However, in contrast, species belonging to cetrarioid and hypogymnioid clades are mainly distributed in the Northern Hemisphere. Special Invited Review the seven main clades in the family. The parmelioid clade is the largest with ca 1854 species (two thirds of the species in the family) within 27 of the ca 79 currently recognized genera (Table 1). The second largest is the Usnea-clade with ca 350 species, all classified within the genus Usnea, whereas the cetrarioid clade includes ca 97 species in 17 genera (Fig. 5). The alectorioid and hypogymnioid clades include five and four genera respectively. However, 21 of the genera are still not aﬃliated to any well supported clade (Fig. 5). Seventy-nine accepted genera in current use are briefly discussed below in terms of their current phylogenetic position, aﬃnities, size and distribution. 1. The parmelioid clade The parmelioid clade, composed of 27 genera and ca 1854 species, is subdivided into nine subclades (Fig. 5), and largely overlaps the morphological group of parmelioid lichens (Crespo et al. 2010b). Several of the most speciose genera included here, such as Hypotrachyna, Parmotrema and Xanthoparmelia, have distribution centres in the Southern Hemisphere. The phylogeny of the parmelioid lichens has been frequently investigated during the past 15 years (Crespo and Cubero 1998, Crespo et al. 2001, 2007, 2010b, Blanco et al. 2004b, Divakar et al. 2006, 2010a). The taxonomy has gradually settled with the proliferation of molecular phylogenetic studies, but many genera are still in need of attention (Crespo et al. 2010b, 2011). 1.1. The Cetrelia clade Cetrelia This genus, earlier included in Cetraria because of the submarginal apothecia, is related to the parmelioid lichens according to molecular data (Crespo et al. 2007). This is consistent with the latter’s broadly lobed thalli and the presence of isolichenan in the cell walls which occurs in some groups of parmelioid genera but is absent among cetrarioid lichens (Elix 1993). Some species of Cetrelia are spread over eastern and Pacific North America and Eurasia, whereas others have a more restricted distribution in eastern Asia. Five distinct morphologies have been recognized within this usually broadly foliose genus. However, based on diﬀerences in secondary compounds, in combination with morphology, 18 species are generally recognized (Randlane and Saag 1991, Czeczuga et al. 2000, Obermayer and Mayrhofer 2007, Otnyukova et al. 2009). Molecular analyses based on DNA-sequences have not yet been carried out in order to evaluate the species delimitations in the genus. 1.2. The Hypotrachyna clade The phylogeny of the Hypotrachyna clade is currently being studied in order to determine the taxonomic position of Cetrariastrum, Everniastrum and Parmelinopsis (Divakar et al. 2010a). Cetrariastrum This pantropical genus of five strap-shaped species was segregated from Everniastrum, which diﬀers in having more regularly branched lobes, a hollow stipe, larger asci and a 648 thinner hypothecium (Sipman 1980, 1986, Wei and Jiang 1982, Kurokawa 1999a). However, the distinction between the two genera is still uncertain (Crespo et al. 2010b), because two of the species (C. andense Kärnefelt ex Sipman and C. dubitans Sipman) form a clade together with Everniastrum lipidiferum (Hale and M. Wirth) Hale ex Sipman. However, the type species of Cetrariastrum has not yet been investigated, and hence, taxonomic conclusions have not been drawn. Everniastrum The distinction of this strap-shaped, pantropical genus from Cetrariastrum and parts of the polyphyletic Hypotrachyna requires further studies, as discussed above. The genus, characterized by linearly elongate, dichotomously divided and marginally ciliate lobes, includes ca 40 species distributed in South America and Asia (Chen et al. 1989, Elix 1993, Pooprang et al. 1999, Boonpragob 2002). Hypotrachyna This is one of the most speciose genera in the family, with ca 188 foliose species worldwide, of which only 18 are present in Europe. It is mainly distributed in montane tropical and subtropical regions. Molecular data indicate that the genus is polyphyletic (Divakar et al. 2006, 2010a). The species have characteristically narrow, dichotomously branched lobes (Fig. 6). A group of 15 species with broad lobes, distributed in Asia, were shown to be unrelated and consequently segregated as a new genus, Remototrachyna (Divakar et al. 2010a). Hypotrachyna is a genus where new species are being continuously discovered (Krog 2000, Kurokawa and Moon 2000, Louwhoﬀ and Elix 2000, 2002, Moon et al. 2000, Wang et al. 2000, Marcelli and Ribeiro 2002, Nash et al. 2002, Divakar et al. 2006, Yánez-Ayabaca and Eliasaro 2009, Lumbsch et al. 2011). Parmelinopsis This genus of ca 25 pantropical to temperate foliose species has traditionally been distinguished from Hypotrachyna based on the presence of cilia and sparsely branched rhizines (Elix and Hale 1987), characters that do not correlate with phylogenies based on DNA sequences. It remains to be seen whether this paraphyletic genus can be kept separate from Hypotrachyna. To date, the taxonomy of the Hypotrachyna clade has not been fully revised using molecular data (Divakar et al. 2006, 2010a). 1.3. The Melanohalea clade Emodomelanelia This newly described genus includes a single foliose species, E. masonii (Essl. & Poelt) Divakar & A. Crespo, with some characters typical of Parmelia s.s. such as eﬃgurate pseudocyphellae and large ascospores, frequently with an olive-brown to brown, not grey, upper surface due to absence of atranorin. It is an Asian endemic, known from rocks at high altitudes of China, India, Nepal and Taiwan (Esslinger and Poelt 1991, Crespo et al. 2010b). According to Crespo et al. (2010b), the species is a sister clade of Melanelixia and Melanohalea. Special Invited Review Figure 6–21. Morphology in Parmeliaceae. (6)–(9) the parmelioid clade: (6) Hypotrachyna sinuosa, Tønsberg 5028 (BG), (7) Pleurosticta acetabulum, Thell 11003 (LD1449012), (8) Punctelia jeckeri, Lund, Sweden, photographed in situ, (9) Xanthoparmelia protomatrae, Timdal 9920 (O). (10)–(13) the cetrarioid clade: (10) Dactylina arctica, Hansen, Lichenes groenlandici exsiccati 820 (LD1076368), (11) Melanelia stygia, Westberg 2927 (LD1189807), (12) Cetraria islandica, Kärnefelt 2339 (LD1006514), (13) Masonhalea inermis, Kärnefelt 2367 (LD1035952). (14) The usneoid clade: Usnea florida, Thell 9606 (LD1054321). (15)–(16) The hypogymnioid clade: (15) Arctoparmelia centrifuga, Falk (LD1006017), (16) Brodoa oroarctica, Hansen, Lichenes groenlandici exsiccati 953 (LD1094200). (17)–(18) the alectorioid clade: (17) Bryoria fuscescens, Velmala 10 (H), (18) Gowardia nigricans, Kärkkäinen 45 (H). (19) The letharioid clade: Letharia vulpina, Hasselrot 1936 (LD1044386). (20)–(21) species with uncertain aﬃnities: (20) Bryocaulon divergens, Westberg 2809b (LD1189669), (21) Imshaugia aleurites, Almborn (LD1036226). Photos: Patrik Frödén. 649 Special Invited Review Figure 6–21. (continued). 650 Melanelixia Melanohalea Similar to Melanelia and the closely related Melanelixia, the species included in Melanohalea are primarily distributed in the Northern Hemisphere where they occur on bark and wood. The 22 foliose species included in the genus are characterized by their pseudocyphellae on warts or isidial tips, a non-pored epicortex, and a medulla containing depsidones or lacking secondary compounds. Both Melanelixia and Melanohalea were segregated from the distantly related Melanelia based on DNA-evidence and morphological data (Blanco et al. 2004a). Pleurosticta This genus of only two species is characterized by a broadly foliose thallus with a dark olive-green upper surface (Fig. 7). It is distributed in temperate regions of Eurasia and North Africa (Lumbsch et al. 1988). Its relationships with other groups of brown parmelioid lichens requires further study, but the ‘Melanelia’ disjuncta group is probably the closest relative (Crespo et al. 2010b). 1.4. The Nipponoparmelia clade Nipponoparmelia This genus of four foliose species constitute a group of east Asian species previously placed in Parmelia s.s. from which it diﬀers by its marginal punctate pseudocyphellae. Nipponoparmelia was proposed as a subgenus by Kurokawa (1994b), later raised to the generic rank (Crespo et al. 2010b) based on molecular evidence. 1.5. The Parmelia clade Parmelia In the traditional circumscription, this foliose genus included almost the whole of the parmelioid clade. Presently it contains ca 57 species, but is still paraphyletic (Crespo et al. 2010b). The core of the genus includes a dozen temperate species in the Northern Hemisphere. Parmelia saxatilis (L.) Ach. and P. sulcata Taylor are the most widespread species present in cold-temperate areas of both hemispheres. Some of the other species have restricted or poorly known distributions. Molecular data have repeatedly demonstrated the presence of several new, cryptic or semi-cryptic species within this genus (Feuerer and Thell 2002, Molina et al. 2004, 2011a, 2011b, Divakar et al. 2005). Relicina This tropical genus of ca 54 species has its centres of diversity in eastern Asia and Australasia (Crespo et al. 2007). It is similar to Bulbothrix in having a foliose thallus and bulbate cilia, but diﬀers in the yellow–green, not grey, upper surface and conidia with a double swelling (Elix 1993). Another similar genus is Relicinopsis, which diﬀers in having an eciliate margin and conidia with a single swelling (Elix et al. 1986). DNA-studies support that the morphologically very similar genera Relicina and Relicinopsis may be synonymous, but additional studies are needed to clarify if they are better not kept separate (Crespo et al. 2010b). Relicinopsis The five species of this foliose genus have a southern Asian to Australasian distribution. In spite of a diﬀerent conidial shape, having a single instead of double swelling, and the absence of cilia, the genus may be congeneric with Relicina, see above (Elix et al. 1986, Crespo et al. 2010b). 1.6. The Parmelina clade Bulbothrix Elix (1993) listed ca 32 foliose species in this genus, however, Bulbothrix belongs to a tropical–subtropical group of genera in which new species are frequently discovered (Louwhoﬀ and Elix 2000, Wang et al. 2000, Marcelli and Ribeiro 2002, Spielmann and Marcelli 2008, Jungbluth et al. 2008, Benatti 2011). By 2011, the number of species had reached 36 within Brazil alone and 58 worldwide (Benatti pers. comm.). In current phylogenies Bulbothrix appears as paraphyletic in the Parmelina clade, including the genus Parmelinella (Divakar et al. 2006, 2010a, Crespo et al. 2010b, Benatti 2012). However, the type species of the genus has not yet been sequenced and the taxon sampling needs to be widened to address the monophyly of the genus. A polysporous member, Bulbothrix amazonensis V. Marcano et al., was segregated as a monospecific genus, Bulbothricella (Marcano et al. 1996), which was subsequently reduced to synonymy (Lumbsch 1997). Myelochroa This genus of ca 30 foliose species is characterized by the presence of yellow–orange pigments in the medulla and simple to squarrosely branched rhizines. It is morphologically similar to the closely related Parmelina, but the latter diﬀers in having a white medulla and lacking hopane triterpenes. Interestingly, these two genera have diﬀerent distributions with little overlap: Myelochroa has a centre of distribution in eastern Asia where two species were recently described (Otnyukova et al. 2009), whereas Parmelina is particularly common in western Europe and North America (Divakar et al. 2006). Parmelina This foliose genus, characterized by broad lobes with a smooth upper surface, diﬀers from Myelochroa by its white medulla. The present circumscription of Parmelina diﬀers considerably from how the genus originally was described by Hale (1974c). In its restricted sense it contains nine species, mainly distributed in North America and Europe. The sister group Myelochroa is mainly distinguished by chemical characters (Crespo et al. 2010b). 651 Special Invited Review The 15 foliose species of this genus are segregated from Melanelia, being mainly distributed in temperate regions, predominantly in the Northern Hemisphere, where they grow on bark and wood. The genus lacks pseudocyphellae and lecanoric or gyrophoric acids are present in the medulla (Blanco et al. 2004a). Melanelixia is superficially similar to Pleurosticta, which diﬀers in having broader lobes, often with a bluish-green, not entirely brown, upper surface. However, Melanelixia appears to be more closely related to Melanohalea (Crespo et al. 2010b). Special Invited Review Parmelinella Flavopunctelia Under the current generic circumscription, Parmelinella is composed of seven foliose species (Benatti 2012). The genus was originally distinguished from Bulbothrix by the absence of bulbate cilia and presence of secalonic acid derivatives (Elix and Hale 1987), characters that are not supported as taxonomically important in molecular phylogenies (Divakar et al. 2006). Furthermore, in the past 20 years, a broader range of variation has been documented within Parmelinella, including characters that originally separated it from other genera. Parmelinella appears to be nested within Bulbothrix and its relationship with the latter is currently being studied. This genus of only five foliose species occurs in temperate and tropical regions on all continents except Australia. Originally described as a subgenus of Punctelia (Krog 1982), it was later recognized as a separate genus based on a diﬀerent shape of the conidia, appearing dumbbell- or thread-shaped, rather than comma-shaped as in Punctelia (Krog 1982, Hale 1984, Kurokawa 1999b). Molecular studies confirmed the distinction of these two genera which both have roundish pseudocyphellae (Thell et al. 2005a, Crespo et al. 2010b). Remototrachyna This recent segregate from Hypotrachyna includes 15 foliose species with a centre of distribution in southeast Asia (Divakar et al. 2010a). It appears most closely related to Bulbothrix but has broader lobes and shorter rhizines compared to Bulbothrix and Hypotrachyna. 1.7. The Parmeliopsis clade Parmeliopsis This genus of three small, adnate foliose species is distributed in temperate boreal areas (Ahti et al. 2011). Previously it included Imshaugia, which is superficially similar, but diﬀers in the shape and size of spores and conidia and the presence of thamnolic acid. DNA data confirmed that the two genera are only distantly related (Crespo et al. 2010b). 1.8. The Parmotrema clade Austroparmelina The 13 foliose species of Austroparmelina were previously placed in the genera Canoparmelia and Parmelina. Characteristic features include the adnate, grey thalli with narrow lobes and they have Australasian distributions. The genus is sister group to Flavoparmelia and Parmotrema (Crespo et al. 2010a, 2010b). Canoparmelia After the segregation of the genus Austroparmelina and the inclusion of the Canoparmelia crozalsiana group in Parmotrema as the subgenus Crespoa D. Hawksw, Canoparmelia became a monophyletic sister group to the remaining genera in the Parmotrema clade (Crespo et al. 2010b). The genus includes ca 40 foliose species, generally distributed in South America and Africa, with broad and large ascospores, lacking depsidones and having a maculate upper surface (Crespo et al. 2010a, 2010b, Hawksworth 2011). Flavoparmelia This cosmopolitan, mainly pantropical genus of ca 32 species was segregated from Pseudoparmelia for species with broad lobes, containing usnic acid, isolichenan in cell walls, large ascospores and eciliate lobes (Hale 1986b, Elix 1993, Elix et al. 2010). The phylogeny of Flavoparmelia is currently under investigation (Blanco pers. comm.). It is positioned as a monophyletic sister group of Parmotrema from which it diﬀers in having dumbbell-shaped conidia and by always containing usnic acid (Moberg et al. 2011). 652 Nesolechia This widely distributed genus of 41 species of obligately lichenicolous fungi was treated as a synonym of Phacopsis Tul. by Triebel and Rambold (1988). This was not followed by Alstrup and Hawksworth (1990) and Doré et al. (2006), but supported by Diederich (2003). The molecular study of Phacopsis by Peršoh and Rambold (2002) revealed that Nesolechia was polyphyletic and positioned in Parmeliaceae, results later confirmed by Crespo et al. (2007). Thus, we prefer to use the name Nesolechia for this sister clade of Punctelia, until further investigations are undertaken. Parmotrema This large genus of ca 300 species that are often broadly foliose, is mainly distributed in tropical and subtropical areas, being particularly common in the Pacific Islands and South America (Hale 1965, 1974b, Benatti et al. 2008). Based on molecular evidence, the genera Canomaculina, Concamerella, Parmelaria and parts of Canoparmelia have been included in Parmotrema (Blanco et al. 2005, Crespo et al. 2010b, Hawksworth 2011). Punctelia This monophyletic genus includes ca 45 foliose species, commonly with punctiform pseodocyphellae (Fig. 8), with its highest diversity in South America and Africa. Flavopunctelia is morphologically similar, but diﬀers by its usually dumbbell-shaped, not comma-shaped, conidia and the presence of usnic acid (Krog 1982, Hale 1984). However, Punctelia is a well-supported sister group to Nesolechia (Crespo et al. 2010b). 1.9. The Xanthoparmelia clade Xanthoparmelia This large genus comprising ca 820 species is characterized by the presence of Xanthoparmelia-type of lichenan in the cell-walls and an arachnoid vacuolar body in the ascospores. The species within this clade are more frequently found with spores compared to other parmelioid lichens (Fig. 9), which may explain its evolutionary success (Del Prado et al. 2007). The huge morphological variation within the genus has been over-emphasized in previous classifications and in the present sense it includes nine genera that were formerly recognized as separate: Almbornia Essl., Chondropsis Nyl., Namakwa Hale, Neofuscelia Essl. Paraparmelia Elix & J. Johnston and Xanthomaculina Hale (Elix 2003b, Blanco et al. 2004b, Thell et al. 2006), Karoowia Hale, Omphalodiella Henssen and Placoparmelia Henssen (Amo de Paz et al. 2010a, 2010b). The diﬀerent morphologies represented within the monophyletic Xanthoparmelia clade do not correlate with monophyletic groups within the clade, preventing the recognition of smaller genera. Chondropsis, composed of two erect, foliose and vagrant species, was the oldest genus name, but the name Xanthoparmelia has been conserved to avoid recombination of all the other species (Hawksworth and Crespo 2002). The cetrarioid clade contains 17 genera and 97 species, including four species in Melanelia s.s. belonging to this clade (Fig. 5). About 148 species, currently spread over 25 genera, belong to the morphological group of cetrarioid lichens, which is a polyphyletic group characterized by the submarginal position of the apothecia and pycnidia (Randlane et al. 2010). Similar to the parmelioid group, the cetrarioid group had a phase of frequent taxonomic splitting, starting with Asahinea, Cetrelia and Platismatia, segregated from Cetraria in the 1960s (Culberson and Culberson 1965, 1968), three genera now known to be unrelated to the cetrarioid clade (Fig. 5). Only three genera, Cetraria, Cornicularia and Dactylina, had been in common use until Lai (1980) reinstated the genus names Nephromopsis and Tuckermannopsis within which he proposed further combinations. Several genera and species with a cetrarioid morphology are positioned outside the cetrarioid clade. The genus Cetrelia and two Parmotrema-species, P. subthomsonii (D. D. Awasthi) A. Crespo et al. and P. thomsonii (Stirt.) A. Crespo et al., formerly forming the genus Parmelaria D. D. Awasthi (Crespo et al. 2010b), belong to the parmelioid clade (Fig. 5), whereas the genera Asahinea, Bryocaulon, Coelopogon, Cornicularia, Platismatia and the species Himantormia deusta (Hook. F.) A. Thell & Søchting, formerly constituting the monotypic genus Nimisia Kärnefelt & A. Thell, have uncertain aﬃnities. In contrast, the cetrarioid clade includes taxa that were considered to be parmelioid based on morphology, including Ahtiana sphaeropsorella and Melanelia stygia (Randlane et al. 2010). 2.1. Esslingeriana, Dactylina and Melanelia The three genera, Dactylina, Esslingeriana and Melanelia are strongly supported as early-diverging members of the cetrarioid clade (Crespo et al. 2010b, Nelsen et al. 2011), outside the two major subclades, named after the largest and oldest genera of each clade, Cetraria (Acharius 1803) and Nephromopsis (Müller 1891). Esslingeriana This monotypic genus with a set of unique characters reported from North America (Lai 1980) is described in detail by Esslinger (1971). It is characterized by a broadly foliose thallus with a grey upper cortex due to the presence of atranorin. The apothecia develop submarginally, but become laminal, whereas the pycnidia remain marginal. Dactylina This genus forms a small clade of two bulbose, arctic–alpine species, morphologically characterized by an unbranched or Melanelia This is a polyphyletic genus of ca 13 adnate foliose species. However, Melanelia in a strict sense is probably a genus of four species distributed in the Northern Hemisphere, namely the DNA-analysed M. hepatizon (Ach.) A. Thell and the type species M. stygia (L.) Essl. (Fig. 11), and, in addition, M. agnata (Nyl.) A. Thell and M. culbersonii (Hale) A. Thell, formerly combined in Cetraria, which appear to be closely related to M. hepatizon according to their morphology (Thell 1995, Westberg and Thell 2011). The M. disjuncta group, including M. disjuncta (Erichsen) Essl., M. predisjuncta (Essl.) Essl., M. panniformis (Nyl.) Essl., M. sorediata (Ach.) Goward & Ahti and M. tominii (Oxner) Essl., represents a diﬀerent evolutionary line that belongs to the parmelioid clade (Blanco et al. 2004a, Crespo et al. 2010b). 2.2. The Cetraria and Nephromopsis clades – the cetrarioid core The term cetrarioid core was originally used for the Cetraria and Nephromopsis clades (Thell et al. 2009). The generic concept in this clade is narrow and is not always supported by molecular data. An alternative would be to allow a broader generic concept allowing a large morphological variation within genera similar to the case of Xanthoparmelia. 2.2.1. The Cetraria clade Allocetraria This monophyletic genus containing nine species mainly occurs at high altitudes, and seven of the species are endemic to the Himalayas (Kurokawa and Lai 1991, Randlane and Saag 2003). The morphology ranges from foliose to fruticose, and two species with a finger-like morphology, earlier accommodated in Dactylina, are placed here (Kärnefelt and Thell 1996). The genus is characterized by its unusually long and narrow conidia, slightly thickened at one end (Thell et al. 1995b). The phylogenetic position of the genus, close to Vulpicida and Cetraria, is well supported (Thell et al. 2009). Cetraria The 15 species of Cetraria are common elements in arctic, alpine and boreal vegetation in the Northern Hemisphere, but a few species also occur in cold areas of the Southern Hemisphere (Kärnefelt 1979, 1986). However, Cetraria is paraphyletic as currently circumscribed. Cetraria s.s. only includes the erect foliose C. islandica (Fig. 12) and the fruticose C. aculeata groups, both characterized by the presence of fatty acids and conidia with a single swelling (Thell et al. 2009, Nelsen et al. 2011). Cetrariella This Cetraria segregate was proposed for two erect foliose, closely related species, C. delisei (Schaer.) Kärnefelt & A. Thell, the type of the genus, and C. fastigiata (Nyl.) Kärnefelt & A. Thell, being characterized by bottle-shaped 653 Special Invited Review 2. The cetrarioid clade weakly dichotomously branched thallus with a hollow medulla (Kärnefelt and Thell 1996). Apothecia are rare and positioned on lateral branches (Fig. 10). Special Invited Review conidia and the presence of gyrophoric and hiascic acids. Two closely related species with an adnate foliose morphology, C. commixta (Nyl.) A. Thell & Kärnefelt and C. sorediella (Lettau) V. J. Rico & A. Thell, were transferred to Cetrariella from Cetraria and Melanelia respectively (Thell et al. 2004, Nelsen et al. 2011). Cetrariella occurs in the Northern Hemisphere: C. delisei and C. fastigiata are circumpolar, C. commixta has a wide arctic–alpine distribution, and C. sorediella is restricted to the Iberian Peninsula, Schwarzwald and the Alps (Rico et al. 2005). Usnocetraria Thell et al. (2009) regarded Usnocetraria as monospecific after a revision of this recent segregate of originally 11 species (Lai et al. 2007). The single species, U. oakesiana (Tuck.) M. J. Lai & J. C. Wei, has an adnate foliose habit and a disjunct alpine distribution in the Northern Hemisphere (Randlane and Saag 2004, Klepsland and Timdal 2010). Vulpicida The six foliose or subfruticose species in this genus have intensely yellow to greenish–yellow thalli, due primarily to the presence of pinastric and vulpinic acids, and conidia with a single swelling either centrally or terminally positioned (Mattsson 1993, Mattsson and Lai 1993). Vulpicida appears paraphyletic in current phylogenetic analyses (Thell et al. 2009). The Eurasian species form one clade, whereas the species endemic to diﬀerent parts of North America form a separate clade. However, the delimitations of species in this genus are in need of revision (Mark et al. 2012). 2.2.2. The Nephromopsis clade Ahtiana This genus was proposed as monospecific for Parmelia sphaerosporella Müll. Arg. (Goward 1985), but two additional species were transferred from Cetraria by Thell et al. (1995c): A. aurescens (Tuck.) Randlane & A. Thell and A. pallidula (Tuck. ex Riddle) Goward & A. Thell. The three species have diﬀerent North American distributions. According to recent phylogenetic analyses (Thell et al. 2009, Nelsen et al. 2011, unpublished), the genus is polyphyletic and in need of taxonomic revision. Lai and Elix 2002, Randlane and Saag 2003). This originally monospecific genus (Lai 1980) forms a monophyletic clade nested within Nephromopsis as presently circumscribed. However, only three of the species have so far been included in molecular analyses. Flavocetraria This small, but polyphyletic genus occurs mainly in the Northern Hemisphere and diﬀers from Cetraria by its yellow colour, thinner cortex and dumbbell-shaped conidia (Kärnefelt et al. 1994). Flavocetraria seems to be most closely related to Tuckermannopsis and Arctocetraria (Thell et al. 2002). However, Flavocetraria minuscula (Elenkin & Savicz) Ahti et al. is not closely related to the two other species of the genus (Thell et al. 2009). Kaernefeltia Two species form the genus, namely K. merrilli (Du Rietz) A. Thell & Goward, a morphologically variable but often narrowly erect foliose species, and K. californica (Tuck.) A. Thell & Goward, the fruticose type species. The former has a remarkable disjunct distribution, occurring in western North America and central Spain, whereas K. californica occurs exclusively along the Pacific coast in North America (Thell and Goward 1996). Kaernefeltia is paraphyletic, but forms a monophyletic clade together with the genus Tuckermanella (Nelsen et al. 2011). Masonhalea Originally monotypical, including M. richardsonii, it is characterized by a terete foliose, cartilaginous thallus with large pseudocyphellae in the form of decorticate patches on the lower side and small conidia without swellings (Kärnefelt 1977). The new combination M. inermis (Nyl.) Lumbsch et al. recently provided for Tuckermannopsis inermis (Nyl.) Kärnefelt, is based on DNA-evidence (Nelsen et al. unpublished); this species has pseudocyphellae in form of a continuous line close to the margin of the lower side (Fig. 13). Both species have a northern Beringian distribution, but Masonhalea inermis also occurs in Svalbard. Nephromopsis This arctic genus of two erect foliose species was segregated from Cetraria on the basis of deviating characters of the asci and the presence of rangiformic and norrangiformic acids (Kärnefelt et al. 1993). The most frequent shape of conidia is with a central swelling, as in Cetraria s.s., but more or less dumbbell-shaped conidia have been observed very occasionally. DNA-studies confirmed a close relationship between the two species and a position in the Nephromopsis clade, which are thus only distantly related to Cetraria s.s. (Nelsen et al. 2011). This originally monotypic genus (Müller 1891) was rarely accepted until it was re-established by Lai (1980). The genus was re-delimited several times by Randlane et al. (1995) and Randlane and Saag (1998) based on morphology, and by Thell et al. (2005b), based on DNA evidence; it currently includes 21 species, distributed mainly in eastern Asia. The most closely related genus is Cetreliopsis, which is distinguished by the presence of pseudocyphellae on both upper and lower surfaces, and fumarprotocetraric acid in the medulla. As currently delimited, the two genera constitute a single clade with Cetreliopsis nested within Nephromopsis (Thell et al. 2009, Nelsen et al. 2011). Cetreliopsis Tuckermanella This genus includes seven foliose species with pseudocyphellae on both surfaces, dumbbell-shaped conidia, and containing fumarprotocetaric acid. The species occur in eastern and southeastern Asia (Randlane et al. 1995, 2001, This genus was described to accommodate six small, brown, adnate foliose species, endemic to southern and western inland areas of North America (Esslinger 2003). The presence of mostly ellipsoid ascospores, submarginal and Arctocetraria 654 laminal pycnidia and continuous marginal pseudocyphellae distinguish it from Tuckermannopsis in which most of the species had previously been accommodated. Three of the six species have been included in phylogenetic analyses and form a well supported clade nested within Kaernefeltia in the Nephromopsis clade (Nelsen et al. 2011). Tuckermannopsis 3. The usneoid clade In spite of its remarkable size, viz. 350 species, the usneoid clade contains only the genus Usnea, after the segregates Dolichousnea (Y. Ohmura) Articus, Eumitria Stirt. and Neuropogon Nees & Flotow were revealed as nested within Usnea and consequently regarded as synonyms (Ohmura and Kanda 2004, Wirtz et al. 2006, 2008). The morphological group of usneoid lichens, and the now synonymized family Usneaceae, used to be a very diverse group, including genera now positioned in several clades, such as Evernia or Letharia (Hale 1983, Kärnefelt et al. 1998). Usnea Although this cosmopolitan genus is a typical element of northern boreal forests, its highest species diversity is in tropical and subtropical regions. The genus is recognized by a fruticose thallus with an elastic central cord in the medulla (Fig. 14) (Randlane et al. 2009, Clerc 2011a, 2011b, Saag et al. 2011). Much of the species taxonomy is poorly known due to the enormous morphological and chemical infraspecific variation. Several revisions of Usnea in South America are ongoing or have been finished recently (Rodriguez et al. 2011, Truong et al. 2011, Truong and Clerc 2012). The morphologically similar Protousnea is only distantly related in the phylogeny (Crespo et al. 2010b). 4. The hypogymnioid clade This clade constitute a comparatively less investigated part of the Parmeliaceae. In the most recent phylogeny by Crespo et al. (2010b), it is a weakly supported clade that includes four genera (Fig. 5), three of which are entirely confined to the Northern Hemisphere. The clade corresponds reasonably well to the morphological group of hypogymnioid lichens or the synonymized family Hypogymniaceae. Poelt (1974) proposed to include four genera in the Hypogymniaceae, namely Cavernularia Degel., Hypogymnia, Menegazzia and Pseudevernia. Brodoa and Arctoparmelia have been segregated from Hypogymnia and Xanthoparmelia Arctoparmelia This arctic–boreal, foliose genus of five species is characterized by a sparse development of rhizines (Hale 1986a). Although the yellow–green thalli are reminiscent of some Xanthoparmelia species (Fig. 15), molecular analyses support a position in the hypogymnioid clade (Thell et al. 2004, Crespo et al. 2007, 2010b). Brodoa This genus of three foliose arctic–alpine species has swollen, often convex lobes which are characterized by a greyish or brownish colour due to atranorin in the cortex (Fig. 16). The genus was segregated from Hypogymnia based on the compact medulla, larger spores and diﬀerent cortical structure (Goward 1986). A molecular study revealed that Brodoa is a sister group to the three other genera in the hypogymnioid clade (Crespo et al. 2010b). Hypogymnia This foliose genus, characterized by swollen, hollow lobes, includes ca 90, predominantly corticolous, species. It is one of the few large Parmeliaceae genera with its main distribution in the Northern Hemisphere (Hansen and McCune 2010, Elvebakk 2011, Westberg et al. 2011). Hypogymnia is probably monophyletic. New species supported by morphology and DNA evidence are continuously being described in the genus (McCune 2008, 2009, 2011, McCune and Schoch 2009). Pseudevernia This genus includes four foliose species with narrow lobes occurring in temperate to tropical North America and boreal to subtropical western Eurasia and adjacent Africa. Although the bottle-shaped conidia indicate a relationship with e.g. Letharia, Platismatia or Usnea, phylogenetic analyses place Pseudevernia in the hypogymnioid clade (Crespo et al. 2010b). 5. The alectorioid clade The alectorioid group as treated by Brodo and Hawksworth (1977) included five fruticose genera: Alectoria, Bryoria, Oropogon, Sulcaria and Pseuephebe. Except for Oropogon, the alectorioid clade based on molecular studies contains the same group (Fig. 5), 66 species in all. Gowardia is a recently proposed segregate and sister group of Alectoria (Halonen et al. 2009). The relationship between Bryoria and the other genera in the clade is weakly supported (Crespo et al. 2010b). 655 Special Invited Review This genus, originally monospecific (Gyelnik 1933), has contained many cetrarioid species with a dubious generic position (Randlane and Saag 2003). Kärnefelt and Thell (2001) attempted to delimit the genus based on morphology and DNA sequences, and listed seven species in Tuckermannopsis s.s. with distribution centres in western North America and Japan. However, the genus as currently circumscribed is still polyphyletic. The North American species T. platyphylla does not belong to this clade (Thell et al. 2002). respectively (Goward 1986, Hale 1986a), whereas Cavernularia is currently regarded as a synonym of Hypogymnia (Miadlikowska et al. 2011, Westberg et al. 2011). The position of the mainly Southern Hemisphere genus Menegazzia among hypogymnioid lichens was rejected by Kärnefelt and Thell (1992) based on the large-sized ascospores and frequently branched paraphyses. Molecular phylogenetic studies have not resolved the postion of this genus (Crespo et al. 2010b). Alectoria Special Invited Review Currently this genus of seven species occurs in the Northern Hemisphere or has a bipolar distribution, its most characteristic feature being the large brown ascospores (Kärnefelt and Thell 1992, Thell et al. 1995a). Its closest relative Gowardia was accepted by Velmala and Myllys (2011a, 2011b), but proposed as a synonym of Alectoria by Lumbsch and Huhndorf (2010). clade with Everniopsis and Psiloparmelia, and the letharioid clade with Letharia and Lethariella (Fig. 5). In other cases the groupings have weak support and may be the result of too few taxa included in the analyses. Additional work is needed to elucidate the phylogenetic relationships of these genera. 6.1. The psiloparmelioid clade According to Crespo et al. (2007) this clade includes two genera, Everniopsis and Psiloparmelia. Bryoria This genus of fruticose, often pendent species includes ca 51 species, mainly confined to boreal regions, diﬀering from Alectoria by having small hyaline spores compared to the large brown spores of Alectoria, and from Usnea in lacking a medullary cord (Fig. 17). Together with Usnea, it must be regarded as the taxonomically most diﬃcult genus of the family. In the absence of apothecia in most species, secondary chemistry has been important in the delimitation of the species. Molecular studies indicate that the current species concept needs revision (Velmala et al. 2009, Myllys et al. 2011a, 2011b). The phylogenetic position of Bryoria has diﬀered between diﬀerent phylogenetic analyses, but according to Crespo et al. (2010b) it tentatively belongs in the alectorioid clade. Gowardia According to Halonen et al. (2009), Alectoria was not monophyletic, but consisted of two separate genera, Alectoria and Gowardia, the latter with two circumpolar species: G. arctica Halonen et al., growing in arctic regions of Canada and Russia, and the more widely distributed G. nigricans (Ach.) Halonen et al., which also occurs in the Nordic countries. However, the position of Pseudevernia between Alectoria and Gowardia seemed doubtful according to Lumbsch and Huhndorf (2010) who did not recognize the genus, but it was accepted by Velmala and Myllys (2011b). Species in Gowardia diﬀer from Alectoria s.s. by the greyish thallus (Fig. 18). Everniopsis This is presumably a monospecific genus from Africa and South America. Everniopsis trulla (Ach.) Nyl. has a shrubby foliose thallus, attached by a basal holdfast (Elix 1993) and is superficially similar to Evernia and Everniastrum, but is much more closely related to the genus Psiloparmelia with which it forms a separate clade with uncertain aﬃnities (Crespo et al. 2010b). Psiloparmelia This genus of 13 species is distributed in the Southern Hemisphere, being particularly common in the high Andes in South America. However, one species, P. arhizinosa, is endemic at a high altitude in Lesotho in southern Africa (Hale 1989). The genus is characterized by its velvety lower surface without rhizines and the presence of isolichenan in the cell walls (Elix and Nash 1992, Lumbsch et al. 1992). Psiloparmelia is morphologically similar to Arctoparmelia, but closely related to Everniopsis (Crespo et al. 2010b). 6.2. The letharioid clade The genera Letharia and Lethariella form a strongly supported clade according to several molecular studies (Thell et al. 2004, Crespo et al. 2010b). Letharia This genus includes only two adnate, fruticose and saxicolous species with a bipolar (boreal)–arctic–alpine distribution. It belongs to the alectorioid clade (Crespo et al. 2010b), but diﬀers from other alectorioid genera by the absence of lichen products and the lack of isidia, soralia and pseudocyphellae (Brodo and Hawksworth 1977). This fruticose genus of six phylogenetic, semi-cryptic species has been identified by phylogenetic analyses based on DNA sequences. Only L. vulpina (Fig. 19) is widely distributed in the Northern Hemisphere (Kroken and Taylor 2000). One of the semi-cryptic species, L. gracilis Kroken ex McCune & Altermann, has been formally described (McCune and Altermann 2009). Letharia has apical or subapical apothecia, not lateral as in the related Lethariella, and has a distribution centre in western North America (Krog 1976). Sulcaria Lethariella Pseudephebe This fruticose genus of four species distributed in east Asia and western North America is characterized by the large, ellipsoid septate ascospores, becoming yellowish or brown when mature (Brodo and Hawksworth 1977, Brodo 1986). According to Crespo et al. (2010b), Sulcaria and Pseudephebe are sister groups. Krog (1976), who segregated Lethariella from Usnea, considered it to be most closely related to Letharia, which is strongly supported by molecular data (Crespo et al. 2010b). Lethariella diﬀers from Letharia by the position of the apothecia and a main distribution in central Asia (above). Eleven species are currently included in the genus. 6. Small clades 6.3. Genera with uncertain affinities Allantoparmelia Some genera form small clades of two or three genera in the phylogeny presented by Crespo et al. (2010b). In two cases they are strongly supported, namely the psiloparmelioid The three foliose species from boreal and arctic–alpine regions of the Northern Hemisphere included in the genus are characterized by swollen lobes with a brown upper 656 with Pseudevernia and Ramalina, but the former has a black lower surface and the latter has a shiny cortex (Moberg and Thell 2011). Evernia is polyphyletic according to recent phylogenetic studies (Thell et al. 2004, Crespo et al. 2010b). Anzia Himantormia This genus of ca 34 foliose species is characterized by its multispored asci, small, curved ascospores, and a grey upper cortex (Jaylal et al. 2012). The species occur in mountainous areas in both hemispheres, most frequently in eastern Asia, Australasia, and Central and South America. Molecular studies on Anzia have largely focused on species delimitation (Fujiwara et al. 2004, Elix 2007). The genus is probably monophyletic but isolated in the family (Crespo et al. 2010b). Morphologically and anatomically, Pannoparmelia appears closely related, although this genus diﬀers in having a yellow–green upper cortex and 8-spored asci with spherical ascospores. This Patagonian–Antarctic genus comprises two blackish species, the fruticose H. lugubris (Hue) I. M. Lamb, described in detail by Hue (1915) and Lamb (1964), and the strap-shaped H. deusta (Hook. f.) A. Thell and Søchting, described by Kärnefelt and Thell (1993) as Nimisia fuegiae Kärnefelt & A. Thell. Fryday (2005) corrected the name to N. deusta (Hook. f.) Friday, whereafter Thell et al. (2007) showed using DNA-sequence data that its closest relative is Himantormia lugubris. Asahinea This arctic–alpine genus of two species is restricted to the Northern Hemisphere. It is distinguished by a foliose thallus with marginal apothecia and pycnidia, and presence of isolichenan in the cell walls (Culberson and Culberson 1965, Gao 1991). Molecular studies have not revealed any close relatives to Asahinea (Thell et al. 2002, Crespo et al. 2010b). Bryocaulon This fruticose genus of four species with an arctic–alpine distribution is common in coniferous forests of eastern Asia and northwestern North America. The type species of the genus, B. divergens (Fig. 20), is widespread in the Northern Hemisphere, but the other species have much more restricted distributions (Kärnefelt 1986, Øvstedal et al. 2009, Thell and Kärnefelt 2011). Previously, these species were combined in Alectoria and Cornicularia, but they are unrelated to these genera (Thell et al. 2007). Coelopogon This genus of two species from southern Africa and southern South America has an erect fruticose habit, and medullary bundles of periclinal hyphae, and lacks pseudocyphellae (Kärnefelt 1986, Brusse and Kärnefelt 1991). Although it was originally a segregate of Cetraria, DNA-analyses show there is no close relationship between the two genera (Thell et al. 2004). Cornicularia This is a monotypic, fruticose genus. Cornicularia normoerica grows on rocks in coastal mountain areas in the Northern Hemisphere (Kärnefelt 1986). It was assumed to be related to the superficially and ecologically similar Southern Hemisphere genus Himantormia (Henssen and Jahns 1973), but DNA-data do not support this (Thell et al. 2007). Evernia This is primarily a Northern Hemisphere genus with 10 strapshaped to fruticose species. They have some resemblance Imshaugia This genus (Fig. 21) of seven foliose species from boreal and temperate regions of the Northern Hemisphere and from cool-montane regions in the Southern Hemisphere (Meyer 1985) is similar to Parmeliopsis, but the conidia are ellipsoid to dumbbell-shaped in Imshaugia and curved in Parmeliopsis; furthermore, phylogenetic analyses based on molecular data do not confirm a close relationship between these two genera (Thell et al. 2004, Crespo et al. 2010b). Menegazzia This genus of ca 70 foliose species worldwide, but mainly in the temperate and montane areas of the Southern Hemisphere (Bjerke 2001, 2004, 2005, Bjerke and Elvebakk 2001), is easily distinguished by the scattered perforations on the upper surface. According to DNA-based phylogenetic studies, Menegazzia may have aﬃnities with the hypogymnioid clade (Crespo et al. 2010b). Nodobryoria This North American genus of three fruticose species with a characteristic cortex anatomy was segregated from Bryoria by Common and Brodo (1995). According to phylogenetic analyses, its phylogenetic position in the family is uncertain (Thell et al. 2002). Omphalodium This genus of four species distributed in North and South America is distinguished by an umbilicate thallus with a brown upper surface. The similarly umbilicate Omphalora has a yellow–green upper surface and is distributed in southwestern North America (Nash et al. 1990). Henssen (1992) pointed out important diﬀerences in the ontogeny of the apothecia between these two umbilicate genera and supported the idea of keeping them separate; this view was recently confirmed by a molecular study (Crespo et al. 2010b). Omphalora This monotypic genus from southwestern North America is one of the morphologically most spectacular genera in Parmeliaceae with its large umbilicate thallus with a bright yellow–green upper surface and tuberculate pseudocyphellae 657 Special Invited Review surface, lack of rhizines, and direct attachment to substrata via the lower cortex or by loboid holdfasts (Esslinger 1977). Allantoparmelia may be related to Brodoa, a morphologically similar genus which diﬀers in having a grey upper cortex (Thell et al. 2004). on the lower surface (Nash et al. 1990, Henssen 1992). It diﬀers from the morphologically similar Omphalodium which has a densely pseudocyphellate, brown upper surface and larger ascospores. Special Invited Review Oropogon This fruticose genus of ca 40 species has characteristic large, brown, multi-celled, muriform ascospores. Its distribution largely overlaps with the genus Anzia, being most frequent in eastern and southern Asia, Central and South America and the West Indies (Esslinger 1989). The appearance of the hymenium is reminiscent of Alectoria and Sulcaria and the three genera were believed to be closely related, belonging to a separate family Alectoriaceae (Kärnefelt and Thell 1992). However, molecular phylogenetic studies neither support a separate family nor a close relationship between Oropogon and the two other genera (Crespo et al. 2007, 2010b). Pannoparmelia This foliose genus of five species occurs in Australasia and southern South America. It is morphologically most similar to Anzia, a genus that diﬀers by its multi-spored asci (Beltman 1978, Calvelo and Adler 1992, Elix 1993). A close relationship between these two genera has not been confirmed by molecular phylogenetic studies (Crespo et al. 2010b). Platismatia This well-defined genus of 11 foliose species was segregated from Cetraria by Culberson and Culberson (1968). The distribution of the genus is centred around the Pacific Ocean, with some endemic species restricted to western North America and northeastern Asia. A new species, P. wheeleri Goward et al., has recently been decribed (Lumbsch et al. 2011). DNA-data indicate a weakly supported relationship of Platismatia with Omphalodium and Imshaugia (Crespo et al. 2010b). Protousnea This genus of eight fruticose, often pendulous species, including P. fibrillatae Calvelo et al., is distributed in southern South America (Krog 1976, Calvelo et al. 2003). Protousnea was originally described as a subgenus of Usnea, but the two genera do not seem to be closely related according to phylogenetic analyses (Thell et al. 2004, Crespo et al. 2007). The type species, P. magellanica (Mont.) Krog, forms a weakly supported branch together with Pannoparmelia angustata with the genera Letharia and Lethariella as a sister group (Crespo et al. 2010b). 7. Genera without DNA-data Four genera in current use have not yet been studied using molecular data, and thus, their phylogenetic relationships are uncertain. Bulborrhizina This monospecific genus, consisting of B. africana, occurs only in Mozambique (Kurokawa 1994a). 658 Davidgallowaya This monospecific genus, consisting of D. cornutispora Aptroot, is restricted to Papua New Guinea where it was found on a single tree with 173 lichen species (Aptroot 2007). The foliose thallus with hollow lobes suggests its position in the hypogymnioid group. However, the bicornute and curved ascospores are similar to those in Bulbothrix. It has not been possible to extract DNA from the single existing sample. Parmotremopsis This genus of two Central American species was segregated from Parmotrema mainly on the basis of its smaller ascospores and shorter conidia (Elix and Hale 1987). Pseudoparmelia This genus, proposed by Lynge (1914), was based on characteristic pseudocyphellae that Santesson (1942) considered to be an artefact. However, Hale (1974a) redefined the genus, to include 76 species, which was clearly a polyphyletic group; this was recircumscribed later by Hale (1986b), who restricted it to four species characterized on the basis of a few morphological characters, of which the simple rhizines and the high concentration of secalonic acid in the medulla were considered the most important. Elix and Nash (1997) described seven additional species in the genus and proposed four new combinations. Recently, P. subambigua Hale was transferred to Flavoparmelia by Crespo et al. (2010b). Thus, 15 species with a distribution centre in tropical South America, currently belong to Pseudoparmelia. 8. Uncertain genera Although Coronoplectrum Brusse, Neopsoromopsis Gyeln. and Psoromella Gyeln. have been listed under Parmeliaceae (Köfaragó-Gyelnik 1940, Brusse 1987, Lumbsch and Huhndorf 2010), it now appears unlikely that these genera belong to the family, given the increased understanding of the circumscription of Parmeliaceae. While it is evident that these genera do not belong to Parmeliaceae, their aﬃnity to other families remains uncertain and additional studies are required to ascertain their relationship within the Lecanorales. Conclusions The phylogeny of the Parmeliaceae is well-known compared to other families of lichen-forming ascomycetes. The family is monophyletic and the delimitation of genera has made enormous progress as a result of DNA-based phylogenies. 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