World survey of the genusLepraria(Stereocaulaceae, lichenized Ascomycota)

Lauri SAAG; Andres SAAG; Tiina RANDLANE

Abstract A comparative review of 57 Lepraria species and 2 varieties is provided together with species descriptions and a key. Lecanora leuckertiana is transferred to Lepraria. In addition some putative taxa by different authors are discussed.

The Lichenologist 41(1): 25–60 (2009) © 2009 British Lichen Society doi:10.1017/S0024282909007993 Printed in the United Kingdom World survey of the genus Lepraria (Stereocaulaceae, lichenized Ascomycota) Lauri SAAG, Andres SAAG and Tiina RANDLANE Abstract: A comparative review of 57 Lepraria species and 2 varieties is provided together with species descriptions and a key. Lecanora leuckertiana is transferred to Lepraria. In addition some putative taxa by different authors are discussed. Key words: taxonomy, new combination, sterile lichens, secondary substances, morphology Introduction The genus Lepraria Ach., with a worldwide distribution, comprises morphologically simple lichen-forming fungi that never develop fruiting bodies. Most species have a leprose thallus with the surface entirely composed of soredia (Laundon 1992; Tønsberg 1992), while a few taxa are squamulose or develop isidia-like structures (Tønsberg 2004; Wirth et al. 2004; Crespo et al. 2006). Their taxonomy is largely based on the chemistry of secondary metabolites as these lichens produce a wide variety of lichen substances, and other characters are often scarce. The genus Lepraria was described by Acharius (1803) for various sterile sorediate lichens. It is a nomen conservandum after the proposal by Laundon (1963). The genus was placed in Leprariaceae in Fungi Imperfecti (Reichenbach 1841). Later it was regarded as belonging to Ascomycota, incertae sedis by several authors (e.g. Henssen & Jahns 1974; Kirk et al. 2001). The modern treatment of this genus started when Laundon (1974, 1981) who transferred the bright-coloured species pro- L. Saag, A. Saag and T. Randlane: University of Tartu, Institute of Ecology and Earth Sciences, Department of Botany, Lai st. 38, 51005, Tartu, Estonia. Email: lauri.saag@ut.ee ducing anthraquinones to Caloplaca Th. Fr. and Leproplaca (Nyl.) Nyl. ex Hue (a current synonym of Caloplaca) and species containing pulvinic acid derivatives to Chrysothrix Mont. The number of species in Lepraria was further reduced when taxa producing dibenzofuranes were placed in Leproloma Nyl. ex Cromb. (Laundon 1989; Leuckert & Kümmerling 1991). The concept of the genus was developed further by Laundon (1992) who also moved some species from Crocynia (Ach.) A. Massal. into Lepraria, and by Tønsberg (1992), Lohtander (1995) and Leuckert et al. (1995). Lepraria lesdainii was transferred to a new monotypic genus Botryolepraria Canals, Hern.-Mar., GomezBolea & Llimona (Canals et al. 1997), while leprose, usnic acid-producing taxa were generally treated under Lecanora Ach. at that time. Later, Leproloma was synonymized with Lepraria by Kukwa (2002). Ekman & Tønsberg (2002) first showed the monophyly of Lepraria in a molecular study. In addition, the former Leproloma species nested in Lepraria in their analysis and the clade showed affinities with Stereocaulaceae. However, three species, L. flavescens, L. obtusatica and Botryolepraria lesdainii, were positioned outside the genus in the analysis by Ekman & Tønsberg. Later, L. flavescens was transferred to Lecanora as L. rouxii S. Ekman & Tønsberg (Grube et al. 2004) but the position of Lepraria obtusatica remains unclear. 26 THE LICHENOLOGIST Sipman (2003, 2004) also included usnic acid-containing species in Lepraria; an example that was followed by other authors (Kukwa 2006a; Knudsen & Elix 2008). Sterile crustose lichens have recently been rather intensely studied and the number of Lepraria species is constantly increasing. Since the year 2000, 35 taxa have been newly described (Orange 2001; Orange et al. 2001b; Aptroot 2002; Tønsberg 2002, 2004, 2007; Sipman 2003, 2004; Wirth et al. 2004; Bayerová et al. 2005; Orange & Wolseley 2005; Elix 2005, 2006a; Elix et al. 2005; Lendemer 2005; Crespo et al. 2006; Kantvilas & Kukwa 2006; SlavíkováBayerová & Orange 2006; Tønsberg & Zhurbenko 2006; Flakus & Kukwa 2007; Knudsen & Elix 2007, 2008; Knudsen et al. 2007; Lendemer & Harris 2007; Saag et al. 2007; Slavíková-Bayerová & Fehrer 2007; Lendemer et al. 2008) and 6 species have been transferred to Lepraria from the genera, Leproloma and Lecanora (Kukwa 2002, 2006a; Sipman 2004). In this paper 57 Lepraria species and 2 varieties are included. Today, the species concept is not uniform for different Lepraria species. Several chemotypes have been included in some species (e.g. L. caesioalba, L. nivalis), while the presence or absence of a compound is considered to justify the taxonomic rank of species in many other cases. The practical chemotaxonomy can be obscured by ‘mechanical hybrids’ i.e. specimens consisting of mixed soredia from more than one species, that are not rare in the genus. Additionally, chemical similarities may not necessarily indicate close phylogenetic relationships, as the production of the compounds can be phylogenetically homoplasious (Ekman & Tønsberg 2002; Nelsen & Gargas 2008). The purpose of this publication is to provide a comparative review of all 57 species and two varieties currently accepted in the genus in a standardized manner. We aim to summarize the present taxonomic situation, and hope the review will also be of practical help to lichenologists who are not specialists in Lepraria and the relatively rapid taxonomic changes in this genus. Vol. 41 Material and Methods The current review is mainly based on literature, but in many cases also on original observations. Material from B, C, H, LD, NMW, PH, TU, TUR, UPS, and the private herbarium of Š. Slavíková was examined to refine the species descriptions. Morphology and anatomy were studied using a stereomicroscope (Olympus SZ40) equipped with additional objectives and a camera mounting kit. Thin layer chromatography (TLC) was carried out according to Orange et al. (2001a). Some fatty acids that cannot be distinguished by TLC are reported as pairs, for instance ‘roccellic/ angardianic acid’. Stictic acid is usually accompanied by constictic and cryptostictic acids, while both pannaric acid 6-methylester and porphyrilic acid often occur together with related dibenzofurans (Elix & Tønsberg 2004). These complexes are referred to as ‘stictic acid complex’, ‘pannaric acid 6-methylester and accessories’, etc. The Species Lepraria Ach., nom. cons. Methodus Lichenum: 3 (1803); type species: Lepraria incana (L.) Ach. Thallus crustose to subfoliose or squamulose, with a powdery, granular, cottony, membranous or subsquamulose to subfoliose appearance; variously coloured, but not very bright, greyish, greenish and creamy hues prevalent; thin to thick, soft or hard; firmly or loosely attached to the substratum, sometimes parts of the thallus free from substratum, revealing lower surface; shape irregular or rosette-shaped (especially young thalli), individual thalli usually from few mm to 10 cm diam., thalli may fuse to form more or less continuous covers up to several metres diam.; margin diffuse or delimited, lobes absent or present, obscure to well-developed, mostly not wider than 2 mm, raised marginal rim may be present; cortex predominantly absent, but subcorticate areas may be present in a few species; medulla absent or present, thin to thick, sometimes soredia below the upper surface of sorediate thallus may be discoloured and/or poorly separated, forming a pseudomedulla; hypothallus absent or present, sometimes forming thick conspicous weft, white, grey or brown to black or orange in patches; prothallus rarely present; areoles sometimes present in thick specimens; 2009 World survey of Lepraria—Saag et al. marginal lobes can be squamulose; thallus surfaces without soredia sometimes present, medulla or hypothallus may be exposed or soredia may be poorly differentiated to form an almost smooth surface, or be distinguishable but not well separated from each other; soredia usually abundant, rarely absent, sometimes scarce on some parts of thallus or sparsely and evenly distributed throughout the thallus, often aggregated in consoredia; very fine to coarse, 10 µm to 1–3 mm diam., convex, ellipsoidal or irregular, loosely or densely packed; wall absent or present; hyphae projecting from soredia often present, very short (few µm) to very long (120 µm); isidia-like structures (large granules, warts or lobules) may be present, sometimes becoming sorediate. Photobiont trebouxioid green alga, most often Asterochloris (Hildreth & Ahmadjian 1981; Nelsen & Gargas 2006, 2008). Ascomata and conidiomata absent. Chemistry. Aliphatic acids, anthraquinones, benzyl esters, biphenyls, depsides, depsidones, dibenzofurans, terpenoids and usnic acids. Ecology and distribution. On various substrata, most often bark, mosses and rock; most species prefer places sheltered from rain and shaded from direct sunlight, often with high humidity, but several taxa grow on exposed and/or dry surfaces. Worldwide; according to current knowledge the highest number of species is found in temperate areas. 27 anthraquinones ± (2 of them major). K− or underside K+ purple in patches, C+ carmine red, KC+ red, PD− (Flakus & Kukwa 2007). Ecology and distribution. On humus, terricolous mosses, and rocks. Found in open areas of high Andean Puna vegetation and upper montane cloud forest. South America (Bolivia). Discussion. Lepraria neojackii sometimes has a very similar morphology but lacks lecanoric acid. Species that can produce lecanoric acid are discussed under L. cupressicola. Lepraria adhaerens K. Knudsen, Elix & Lendemer Opuscula Philolichenum 4: 5 (2007); type: USA (UCR— holotype; ASU, CANB, PH, SD, UGDA—isotypes). Thallus leprose, granular; margin diffuse, lobes absent; medulla absent, but older thalli forming a lower necral layer of gelatinized granules; soredia abundant, fine to medium, 40–100 µm diam., projecting hyphae absent, but frequently with thin colourless hyphae acting as anchors or rhizines; soredia clumped together. For further details see Knudsen et al. (2007). Chemistry. Pannarin and zeorin (major to trace, very rarely absent) and trace accessories including norpannarin, dechloropannarin, hypopannarin and atranorin. K−, C−, KC−, Pd+ orange (Knudsen et al. 2007). Lepraria achariana Flakus & Kukwa Lichenologist 39: 464 (2007); type: Bolivia (KRAM-L— holotype). Thallus leprose, powdery; margin diffuse, lobes absent; true medulla absent; hypothallus usually well-developed, lax, white or orange in places; soredia abundant, fine, up to 45 µm diam.; consoredia prevalent, up to 75(−150) µm diam. For more details see Flakus & Kukwa (2007). Chemistry. Lecanoric acid, roccellic/ angardianic acid ±, and 2–4 unidentified Ecology and distribution. On rocks (usually on mosses and lichens), rarely soil, in open habitats exposed to rain and sunlight, but in sheltered places in snowy areas. North America. Discussion. The occurrence of small attaching hyphae and the tendency of granules to adhere to one another and the substratum is characteristic (Knudsen et al. 2007). Unstratified, diffuse thalli are found in several other species, but L. santamonicae (argopsin and norargopsin) is the most similar. 28 THE LICHENOLOGIST Lepraria alpina (B. de Lesd.) Tretiach & Baruffo var. alpina in Baruffo et al., Nova Hedwigia 83: 395 (2006); type: USA (UPS—neotype; GZU, ASU—isoneotypes; distributed in Weber, Lichenes exsiccati, distr. by the Univ. of Colorado, Fasc. 16: no. 609). Crocynia alpina B. de Lesd., Bull. Soc. Bot. France 61: 85 (1914). Lepraria angardiana Øvstedal, Nova Hedwigia 37: 687 (1983). Lepraria caerulescens (Hue) Botnen & Øvstedal, Polar Research 6: 130 (1988). Leproloma cacuminum sensu J.R. Laundon, Lichenologist 24: 345 (1992).—Lepraria cacuminum sensu Loht. Ann. Bot. Fennici 32: 52 (1995)—non Diploicia cacuminum A. Massal., Symm. Lich. Nov.: 52 (1855). Thallus leprose, granular; margin usually delimited, lobes sometimes present, minute, often obscure, without raised rims; medulla usually present, not conspicuous, white; hypothallus sometimes present, poorly developed; soredia abundant, coarse or variably sized, typically 100–150 µm diam., projecting hyphae sometimes present, short; consoredia present from few to abundant, typically 200–300 µm. For further details see Lohtander (1995), Baruffo et al. (2006) and Saag et al. (2007). Chemistry. Atranorin, porphyrilic acid and a fatty acid (roccellic/angardianic or rarely rangiformic acid); very rarely fatty acids and/or atranorin can be absent (Leuckert et al. 1995; Saag et al. 2007), or another, unidentified fatty acid present (Flakus & Kukwa 2007). K− or + yellow, C−, KC− or + yellow, Pd− or + yellow. Ecology and distribution. On rock and saxicolous mosses, also on soil, lichens and rarely bark; mostly acidic substrata; usually exposed but also in shaded places, mostly cool climate. Europe, North and South America, Antarctica, Greenland. Discussion. According to molecular studies, L. alpina belongs to a monophyletic ‘L. neglecta group’ together with L. atlantica, L. borealis, L. caesioalba, L. elobata, L. granulata, L. humida and L. neglecta (Ekman & Tønsberg 2002; Slavílová-Bayerová & Vol. 41 Fehrer 2007). Morphologically similar species with a coarsely granular appearance include L. borealis, L. caesioalba, L. granulata and L. neglecta, but these taxa differ chemically. The chemotype with angardianic/ roccellic acid only was included in L. alpina (sub. nomine L. cacuminum) by Lohtander (1994, 1995), but in L. borealis by Kukwa (2006b); specimens with atranorin only or fatty acids only were placed in L. borealis by Saag et al. (2007). Thus specimens with a deficient chemistry can be identified as either L. alpina or L. borealis as there are no distinctive morphological differences between these two species (Prigodina-Lukošienë et al. 2003; Kukwa 2006b; see also Lohtander 1995). Lepraria atlantica has a similar chemistry but is usually powdery to cottony, and is only very rarely morphologically similar to L. alpina (Saag et al. 2007). Lepraria alpina var. zeorinica L. Saag Mycotaxon 102: 68 (2007); type: Greenland (C— holotype, TU—isotype). Thallus and chemistry as L. alpina var. alpina, but also produces zeorin. K− or + yellow, C−, KC− or + yellow, Pd− or + yellow. Ecoloy and distribution. On soil and mosses, and sometimes lichens. Greenland. Lepraria atlantica Orange Lichenologist 33: 462 (2001); type: Ireland (NMW— holotype, BG—isotype). Thallus leprose, powdery; margin diffuse or delimited, lobes absent; true medulla absent; hypothallus present in patches, of usually sparse, white to dull orange hyphae; soredia abundant, fine to rarely coarse, 40– 100 (160–220) µm diam., projecting hyphae rarely present, short; consoredia rare. For further details see Orange (2001), SlavíkováBayerová & Orange (2006) and Saag et al. (2007). Chemistry. The two common chemotypes contain (1) atranorin, porphyrilic acid and 2009 World survey of Lepraria—Saag et al. accessories, rangiformic acid; (2) atranorin, porphyrilic acid and accessories, roccellic/ angardianic acid. In addition, rare chemotypes have been reported with (3) porphyrilic acid and accessories only and (4) both fatty acids (rangiformic and roccellic/angardianic acids) besides atranorin and porphyrilic acid. The subthalline hyphae usually produce unidentified anthraquinones (Orange 2001; Slavíková-Bayerová & Orange 2006; Saag et al. 2007). K+ yellow, C−, Pd− or + yellow. Ecology and distribution. On siliceous rock (also on mosses), sometimes on soil and rarely bark; sheltered from rain. Europe, Australia, Greenland. Discussion. According to molecular studies, L. atlantica belongs to the monophyletic ‘L. neglecta group’ (Ekman & Tønsberg 2002; Slavíková-Bayerová & Fehrer 2007; see also the discussion under L. alpina). Fehrer et al. (2008) used a narrower definition of the ‘L. neglecta group’ and distinguished the ‘L. atlantica group’ including L. atlantica, L. humida and L. sp. H (discussed under L. humida). Other morphologically similar species are mentioned under L. jackii. Lepraria atlantica is chemically and morphologically rather similar to L. neojackii, which is described as having fine soredia only, and containing both roccellic/ angardianic and rangiformic acids together but no atranorin (Flakus & Kukwa 2007). See also the discussion under L. jackii. Lepraria atlantica is also chemically very similar to L. alpina (discussed under L. alpina var. alpina). Lepraria aurescens Orange & Wolseley Lichenologist 37: 247 (2005); type: Thailand (BM— holotype). Thallus leprose, powdery; margin usually diffuse, rim absent, lobes absent; medulla absent; hypothallus present in places, brown, well developed, lax; soredia abundant to sparse in few places, fine to medium, 40–100 µm diam., projecting hyphae usually absent, but present on marginal or solitary granules, 29 long, downward, mostly dark. For more details see Orange & Wolseley (2005). Chemistry. Thamnolic acid (Orange & Wolseley 2005). K+ bright yellow, C−, KC−, Pd+ orange-yellow. Ecology and distribution. On bark; in sheltered places in dry forest. Asia (Thailand). Discussion. Lepraria yunnaniana also has fine granules and a dark loose hypothallus, but produces divaricatic acid. Lepraria cupressicola develops a dark hypothallus but the thallus is blue-grey and has lobes, and it contains lecanoric acid, zeorin and atranorin. Species producing thamnolic acid include L. aurescens (usually thin pale grey to yellowish thallus with a brown well-developed hypothallus and a diffuse margin), L. nylanderiana (usually thick whitish thallus with a well-developed greyish hypothallus and a delimited margin, contains roccellic acid), L. pulchra (thin whitish to pale bluish grey thallus with a whitish medulla, delimited margin with a rim, and characteristic very short projecting hyphae on the soredia) and L. umbricola (usually thin green thallus with diffuse margin and no medulla or hypothallus). Lepraria bergensis Tønsberg Graphis Scripta 14: 47 (2002); type: Norway (BG— holotype; B, H, NMW—isotypes). Thallus leprose, membranous; margin usually delimited, sharply raised (in saxicolous specimens), lobes usually distinct, up to a few mm wide; medulla distinct in places, white; hypothallus absent or orange-brown to brown, inconspicuous, hyphae below thallus orange-brown to brown or colourless; lower surface present, white (at least along margin), pale yellowish or brown; soredia abundant, very fine, up to 25 µm diam., projecting hyphae present or absent, short; consoredia abundant, up to 200 (−340) µm. For more details see Tønsberg (2002). Chemistry. Atranorin, rangiformic acid, norrangiformic acid (trace), fragilin, 30 THE LICHENOLOGIST 7-chloroemodin, emodin, parietin and A01anthrone; the anthraquinones are located mostly in the subthalline hyphae. K+ faint yellow, subthalline hyphae K+ purple (in patches), C−, Pd− (Tønsberg 2002). Ecology and distribution. On siliceous rock (also on mosses); on shaded surfaces. Europe – Norway, Great Britain, Germany (Spribille & Tønsberg 2007). Discussion. The phylogenetic placement of this and other taxa with distinct lobes are mentioned under L. membranacea. Morphologically the most similar species include L. membranacea (pannaric acid), L. normandinoides (usually protocetraric acid and/or fumarprotocetraric acid in addition to roccellic acid and atranorin) and L. sipmaniana (pannaric acid 6-methylester and usually also anthraquinones). Anthraquinone producing species apart from L. bergensis and L. sipmaniana include L. goughensis (obscure minute lobes, lecanoric acid) and non-lobate L. atlantica (porphyrilic acid), L. incana chemotype 2 (divaricatic acid) and L. humida (atranorin and fatty acids). Lepraria borealis Loht. & Tønsberg in Lohtander, Ann. Bot. Fennici 31: 224 (1994); type: Norway (BG—holotype, H—isotype). Thallus leprose, granular; margin usually delimited, sometimes diffuse, obscure minute lobes sometimes present, without raised rim; medulla sometimes present, inconspicuous, white; hypothallus usually absent; soredia abundant, coarse or variably sized, typically 100–200 µm diam., projecting hyphae often present, usually short; consoredia sometimes present, typically 200– 300 µm, sometimes larger. For further details see the references below. Chemistry. The commonest chemotype (1) comprises atranorin, rangiformic and ± norrangiformic acid; rarely in (2) rangiformic acid is replaced by roccellic/angardianic acid or in (3) both fatty acids are present (Lohtander 1994; Tønsberg 2004; Kukwa 2006b and others). Saag et al. (2007) tenta- Vol. 41 tively also included specimens with fatty acids only (4) and atranorin only (5). K− or + faint yellow, C−, KC−, Pd−. Ecology and distribution. On acidic rock (also on mosses), rarely on soil and lichens; mostly exposed but also shaded places, mostly cool climate. Europe, North America, Antarctica, Greenland. Discussion. See the discussion under L. alpina. Lepraria caesiella R.C. Harris in Lendemer, Opuscula Philolichenum 2: 51 (2005); type: USA (NY— holotype; isotypes distributed as Lichens of Eastern North America Exsiccati, no. 172). Thallus leprose, powdery; margin diffuse to poorly delimited, lobes absent; medulla absent; hypothallus absent; soredia abundant, very fine, 20–30 µm diam., projecting hyphae present, short, 25–30 µm; consoredia present, up to 100 µm diam. For more details see Lendemer (2005). Chemistry. Atranorin and zeorin (Lendemer 2005). K− or + faint yellow, C−, KC−, Pd−. Saag et al. (2007) and Flakus & Kukwa (2007) included a few specimens with atranorin, zeorin and roccellic/angardianic acid or an unidentified fatty acid. Ecology and distribution. On bark, rarely on rock, lignum, soil and mosses. North America, South America (Chile), Greenland. Discussion. Lepraria caesiella was previously known as ‘‘Lepraria sp. 3’’ from North America (Harris 1977). Morphologically similar species include L. incana (divaricatic acid) and L. elobata (stictic acid complex). Lepraria pallida also contains atranorin and zeorin but always produces fatty acid(s) and possesses lobes; L. jackii s. lat. has been reported to contain zeorin as a rare accessory (Leuckert et al. 1995; Baruffo et al. 2006). Lendemer (2005) referred to an undescribed species that also produces atranorin and zeorin. 2009 World survey of Lepraria—Saag et al. Lepraria caesioalba (B. de Lesd.) J. R. Laundon var. caesioalba Lichenologist 24: 324 (1992).—Crocynia caesioalba B. de Lesd., Bull. Soc. Bot. France 61: 84 (1914); type: France (E—holotype; BM, GL, PC—topotypes). Thallus leprose, granular; margin usually delimited, sometimes diffuse, obscure minute lobes sometimes present, without raised rims; medulla present, inconspicuous, white; hypothallus usually absent; rarely small patches with exposed medulla present; soredia abundant, coarse or variably sized, typically 100– 150(−200) µm diam., projecting hyphae sometimes present, usually short; consoredia frequent, typically 200–300 µm. For further details see Laundon (1992), Lohtander (1994), Saag et al. (2007). Chemistry. This species comprises several chemotypes, e.g. Leuckert et al. (1995) distinguished three and Tønsberg (2004) five. In this paper, the classification of Leuckert et al. (1995), with additions, is followed, and specimens containing only atranorin and fatty acid(s) without depsidones are placed in L. borealis as described by Lohtander (1994) and Prigodina-Lukošienë et al. (2003). Tønsberg (2004) recognized such specimens as an additional chemotype of L. caesioalba. Chemotype (1): atranorin, fumarprotocetraric acid, protocetraric acid ± (in variable amounts), roccellic/angardianic or rangiformic acid; rare modifications of this chemotype include 1a) both fatty acids, 1b) no fatty acids, 1c) no fumarprotocetraric acid, protocetraric acid present and 1d) no atranorin. K− or + yellow, C−, KC−or + faint yellow, Pd+ orange. Chemotype (2): atranorin, stictic acid plus constictic and cryptostictic acids (in variable amounts, mostly minor), rangiformic or roccellic/angardianic acid; rare modifications include 2a) both fatty acids and 2b) no fatty acids. K− or + yellow, C−, KC− or + faint yellow, Pd+ orange. Chemotype (3): atranorin, psoromic acid, roccellic/angardianic or rangiformic acid. K− or + yellow, C+ red, KC−, Pd+ sulphur yellow. Chemotype (1) is the most frequent, (2) common and (3) is rare, but still widespread. 31 Ecology and distribution. On acidic rock (usually on mosses) and soil, rarely on bark, epiphytic mosses and lichens; in exposed places, mostly cool climate, montane-alpine in tropics. Europe, North and South America, Asia, Australasia, Antarctica, Greenland. Discussion. The phylogenetic placement of this and morphologically similar species is discussed under L. alpina. Lepraria caesioalba is not monophyletic (Ekman & Tønsberg 2002; Slavíková-Bayerová & Fehrer 2007), but no nomenclatural segregation has been proposed. Species producing stictic acid and atranorin include L. caesioalba (granular L. neglecta-type thallus), L. elobata (thin greenish thallus of relatively compact soredia, no medulla or lobes, contains zeorin), L. leprolomopsis (thick whitish thallus with medulla, no lobes, even lower surface, an unknown terpenoid), L. lobificans (greenish loosely packed soredia with long projecting hyphae, welldeveloped medulla, contains zeorin), L. multiacida (stictic acid complex mostly in traces, also salazinic acid and strepsilin derivates in minor to trace amounts and usually zeorin plus 1–2 unknown terpenoids), L. nivalis (some chemotypes; thick whitish thallus with medulla, no terpenoids), L. santosii (some chemotypes; membranous to granular thallus with well-developed lobes and marginal rim) and L. sp. 2 sensu Flakus & Kukwa (for this taxon, see Flakus & Kukwa 2007). The species that may contain protocetraric and/or fumarprotocetraric acids include L. caesioalba (granular L. neglecta-type thallus), L. eburnea (alectorialic acid, usually soft powdery to cottony thallus), L. friabilis (thin finely powdery thallus, often of only sparse soredia), L. isidiata and L. santosii (lobes with thick raised marginal rims and coarse, more or less isidia-like structures, discussed under L. isidiata), L. lanata (characteristic very large consoredia surrounded by woolly anastomosing hyphae), L. nivalis (thick whitish thallus with medulla, may be similar to L. eburnea but lacks alectorialic acid), L. normandinoides (dark thick hyphae on the lower side, lobes usually wide), L. squamatica 32 THE LICHENOLOGIST (contains squamatic acid, protocetraric acid trace accessory), L. toilenae (malonprotocetraric acid, thick light hypothallus). Lepraria caesioalba var. groenlandica L. Saag Saag et al. Mycotaxon 102: 73 (2007); type: Greenland (C—holotype; TU—isotype). Thallus identical to the main variety. For more details see Saag et al. (2007). Chemistry. Atranorin, stictic acid complex, zeorin (in variable amounts), roccellic/ angardianic acid; rarely roccellic/angardianic acid is replaced by rangiformic acid, or fatty acids are absent. Ecology and distribution. On soil and mosses, sometimes on lichens, rarely on rocks. Greenland. Discussion. Chemically similar to L. elobata, which differs in its thin powdery thallus composed of mostly fine soredia, rare consoredia and diffuse thallus margin. Vol. 41 Discussion. According to molecular studies, L. celata is a distinct monophyletic taxon (Fehrer et al. 2008). Morphologically similar species are mentioned under L. jackii. Chemically similar taxa include L. borealis (granular thallus), L. jackii s. lat. (discussed under L. jackii) and L. normandinoides (lobes, usually contains protocetraric or fumarprotocetraric acid). Lepraria coriensis (Hue) Sipman Herzogia 17: 28 (2004).—Crocynia coriensis Hue, Bull. Soc. Bot. France 71: 386 (1924); type: South Korea (KYO—isotype). Thallus leprose, powdery to membranous; margin delimited, lobes present, obscure or more often well-developed (0·5–2 mm wide) and with raised marginal rim; medulla usually present, thin to medium, white; hypothallus sometimes present, thin, brown to black; sometimes soredia sparse in places, exposing smooth ecorticate surface especially near margins; soredia fine to coarse, up to 300 µm diam., projecting hyphae usually absent. For more details see Laundon (2003) and Sipman (2004). Lepraria celata Š. Slavíková Chemistry. Atranorin ± (major to minor) and roccellic/angardianic acid (SlavíkováBayerová & Orange 2006). K− or + faint yellowish, C−, Pd−. Chemistry. Three chemotypes were distinguished by Elix (2006b): (1) usnic acid, zeorin, protodehydroconstipatic and constipatic acids (major to minor), isousnic acid ± (minor to trace), and atranorin ± (trace); (2) usnic acid, zeorin, protodehydroconstipatic and constipatic acids (minor to trace), argopsin (minor), norargopsin (minor to trace), isousnic acid ± (minor to trace) and atranorin ± (minor to trace); and (3) usnic acid, zeorin, protodehydroconstipatic and constipatic acids (minor to trace), caloploicin (minor), fulgidin (minor to trace), isousnic acid ± (minor to trace) and atranorin ± (trace). The third chemotype was considered rare. Laundon (2003) reported that rarely zeorin can be missing. K−, C−, KC−, Pd−. Ecology and distribution. Terricolous (soil and debris) and on mosses; open habitats, often in rock cervices. Europe (Bulgaria and Ukraine), montane; probably wider. Ecology and distribution. On rock (mostly siliceous), wood, bark, mosses and soil; shaded and sheltered places. Asia (India, South Korea), Australia; tropical to subtropical. in Slavíková & Orange Lichenologist 38: 504 (2006); type: Ukraine (PRA— holotype; BG, NMW—isotypes; GenBank accession no. DQ401100). Thallus leprose, powdery; margin diffuse or delimited, lobes absent; true medulla absent; hypothallus of sparse patches of white hyphae; prothallus absent; soredia abundant, fine, (20−)35–50(−60) µm diam., projecting hyphae rarely present, short. For further details see Slavíková-Bayerová & Orange (2006). 2009 World survey of Lepraria—Saag et al. Discussion. Some authors have doubted the distinctiveness of L. usnica from L. coriensis (Orange & Wolseley 2005), however, chemical differences between these taxa were shown by Elix (2006b). In a molecular study Nelsen et al. (2008, in press) confirmed that L. coriensis and L. usnica are distinct species distant from one another and do not belong to Lepraria. Lepraria coriensis had an uncertain position outside Lecanorales in their analysis, while the placement of L. usnica in Pilocarpaceae (Lecanorales) was wellsupported. Other lobate species are discussed under L. membranacea and L. bergensis. Lepraria santamonicae also produces argopsin, but it has no medulla or lobes, does not contain usnic acid, and argopsin is produced in large amounts. Species producing usnic acid and usually zeorin include L. coriensis (usually welldeveloped lobes with a marginal rim, protodehydroconstipatic and constipatic acids), L. usnica (usually small lobes without a marginal rim and minor amounts of contortin), L. ecorticata (no lobes, predominantly no medulla, soredia regular and convex, wellseparated from each other), L. leuckertiana (obscurely and minutely lobed, cottony thallus with thick medulla, soredia not wellseparated from one another), L. straminea (granular, granules corticate, no lobes), L. texta (sometimes obscurely and minutely lobed, powdery thallus with hypothallus, some soredia not well-separated). Lepraria crassissima (Hue) Lettau Feddes Repert. 61: 125 (1958).—Crocynia crassissima Hue, Bull. Soc. Bot. France 71: 393 (1924); type: France (B—isotype). Thallus leprose, membranous to cottony; margin usually delimited, lobes sometimes present, poorly defined; medulla present, very thick, white; lower surface distinct, folded (if whole thallus folded), smooth to tomentose, white to brownish; often eroded patches with exposed medulla on older specimens; soredia abundant, mostly coarse, up to 300(−400) µm diam., projecting hyphae present, short to long; large granules or warts slightly re- 33 sembling isidia but lacking cortex may be present in well-developed thalli, may become sorediate. For more details see van den Boom et al. (1994). Chemistry. Divaricatic acid, nordivaricatic acid (major, rarely trace), zeorin ± (major to minor) (Laundon 1992; Tønsberg 1992; van den Boom et al. 1994). K−, C+ rose-red, KC−, Pd−. Ecology and distribution. On siliceous rock and epilithic mosses, also sometimes on bark or calcareous rock; shaded, humid. Europe, Australia. Discussion. Species producing divaricatic acid and zeorin include L. crassissima (thallus membranous to cottony, thick white medulla, nordivaricatic acid usually major), L. incana (thallus powdery, nordivaricatic acid ± trace) and L. yunnaniana (thallus cottony, dominant dark hypothallus, nordivaricatic acid ± trace). In addition, Baruffo et al. (2006) reported divaricatic acid as a very rare accessory in L. rigidula (lax cottony thallus, very long projecting soredial hyphae, atranorin and nephrosteranic acid). Lepraria multiacida, L. lobificans, and sometimes L. nivalis and L. vouauxii, may be morphologically similar, but differ in their chemistry and spot reactions (see descriptions of these species). Lepraria isidiata was historically described as a variety of L. crassissima, but today it is defined as differing in chemistry, morphology and ecology (see under L. isidiata). Lepraria crassissima was synonomized with L. incana by Kümmerling et al. (1991) and Leuckert et al. (1995), but was shown to be a distinct species by van den Boom et al. (1994) and Ekman & Tønsberg (2002). Other species producing isidia-like structures are mentioned under L. isidiata. Lepraria cupressicola (Hue) J. R. Laundon Lichenologist 40: 412 (2008).—Crocynia cupressicola Hue, Bull. Soc. Bot. France 71: 395 (1924); type: Japan (KYO—holotype). Lepraria atrotomentosa Orange & Wolseley, Biblioth. Lichenol. 78: 328 (2001); type: Sri Lanka (BM— holotype, PDA—isotype). 34 THE LICHENOLOGIST Thallus leprose, powdery to membranous; margin delimited, lobes absent or present, sometimes well-developed, 0·5-2 mm wide, with raised marginal rim, sometimes irregular and less than 0·5 mm wide, without a rim; medulla present, thin, white; hypothallus usually thick, sometimes thin, lax, dark brown, forms a tomentum under lobes; soredia abundant to sparse in places, fine to medium, sometimes coarse, 60–200 µm diam., projecting hyphae rarely present, short. For further details see Orange et al. (2001b) and Sipman (2004). Chemistry. Lecanoric acid, atranorin, zeorin and unidentified fatty acids (Orange et al. 2001b; Sipman 2004). K+ yellowish, C+ pink to red, KC+ more or less red, Pd+ yellow. Ecology and distribution. On siliceous rock or soil and bark; shaded, sheltered, damp. Japan, China (Hong Kong), Taiwan, Sri Lanka. Discussion. Other lobate species are discussed under L. membranacea and L. bergensis. Non-lobed specimens of L. cupressicola may resemble L. incana in appearance but L. cupressicola has a dark tomentum below the thallus and a different chemistry. A similar dark tomentum is also present in L. yunnaniana (divaricatic acid) and L. aurescens (thamnolic acid, tomentum less extensive). Species that can produce lecanoric acid include L. achariana (stratified thallus with fine soredia and a diffuse margin, roccellic acid ±), L. cupressicola (dark tomentose hypothallus, atranorin and zeorin), L. goughensis (very fine soredia, no hypothallus, gyrophoric acid and strepsilin in minor to trace amounts), L. impossibilis (pannaric acid 6-methylester) and L. lecanorica (usually thick thallus, no dark hypothallus). Lepraria diffusa (J. R. Laundon) Kukwa Ann. Bot. Fennici 39: 226 (2002).—Leproloma diffusum J. R. Laundon, Lichenologist 21: 16 (1989); type: Finland (BM—holotype). Lepraria diffusa var. chrysodetoides (J. R. Laundon) Kukwa, Ann. Bot. Fennici 39: 226 (2002).—Leproloma diffusum var. chrysodetoides J. R. Laundon, Lichenologist 21: 18 (1989). Vol. 41 Thallus leprose, powdery to cottony; margin diffuse, rarely delimited, lobes absent; medulla present, usually thick, sometimes thin, white; hypothallus sometimes present, weakly developed, whitish grey to brownish; medulla exposed but only in small patches or fissures, soredia abundant, coarse, up to 100 µm diam., projecting hyphae sometimes present, short. For more details see Laundon (1992) and Kukwa (2006b). Chemistry. 4-oxypannaric acid 2methylester; accessories are 4-oxypannaric acid (minor to trace), pannaric acid methylester, pannaric acid 2-methylester, pannaric acid and other dibenzofurans (traces) (Elix & Tønsberg 2004); rarely also atranorin and/or roccellic acid or very rarely rangiformic acid (Baruffo et al. 2006). K− or + yellow slowly becoming orange, C− or + yellow, KC− or + yellow, Pd+ reddish orange. Ecology and distribution. On calcareous rock (often on mosses), rarely on bark or soil; mostly in shaded and sheltered places. Europe, North America, Asia. Discussion. Lepraria diffusa var. chrysodetoides was subsumed within var. diffusa by Kukwa (2006b). Lepraria eburnea and L. lobificans and to a lesser extent also L. leprolomopsis, L. nivalis and L. nylanderiana may have a similar morphology but do not produce dibenzofurans. Lepraria vouauxii produces dibenzofurans and is sometimes also morphologically similar, but L. diffusa is more powdery, with less or no patches of exposed medulla and the diagnostic substance is different. Lepraria eburnea J. R. Laundon Lichenologist 24: 331 (1992); type: United Kingdom (BM—holotype). Lepraria frigida J. R. Laundon, Lichenologist 24: 332 (1992). Thallus leprose, powdery to cottony; margin diffuse, rarely delimited, lobes usually absent; medulla usually present, mostly thick, white; hypothallus usually not distinct; thallus surfaces without soredia often present, also 2009 World survey of Lepraria—Saag et al. soredia often embedded in a hyphal weft; soredia abundant, mostly fine, up to 60 µm diam., projecting hyphae usually present, short to long (to 100 µm); consoredia present, up to 200(−400) µm. For further details see Orange (1997). Chemistry. Orange (1997) distinguished 3 chemotypes: (1) alectorialic acid, protocetraric acid; (2) alectorialic acid, psoromic acid, 2’-O-demethylpsoromic acid; and (3) alectorialic acid only; accessories reported by other authors include barbatolic acid and other satellites of alectorialic acid (e.g. 5,7-dihydroxy-6-methylphthalide) and very rarely atranorin (all major to trace, Laundon 1992; Baruffo et al. 2006) or roccellic/ angardianic acid (Saag et al. 2007). K− or + yellow, C− or + reddish orange, KC+ pink or reddish orange, Pd+ lemon yellow or orange. Ecology and distribution. Substratum indifferent; on mosses, bark, wood, rock, soil and lichens. Europe, North America, Australasia, Greenland. Discussion. Lepraria frigida was synonymized with L. eburnea by Orange (1997) who showed chemical and morphological continuity between these taxa. Other morphologically similar species are discussed under L. leprolomopsis, and the species producing protocetraric and/or fumarprotocetraric acids under L. caesioalba (none of them produces alectorialic acid). Species producing alectorialic acid include L. eburnea (soft thallus, see chemistry above), L. gelida (usually soft thallus, porphyrilic acid) and L. neglecta (granular thallus, ± fatty acids and atranorin). Lepraria ecorticata (J. R. Laundon) Kukwa Mycotaxon 97: 64 (2006).—Lecanora ecorticata J. R. Laundon, Nova Hedwigia 76: 100 (2003); type: United Kingdom (BM—holotype). Thallus leprose, powdery; margin diffuse, lobes absent; medulla rarely present in places, weakly developed; soredia abundant, mostly fine to medium, 10–100 µm diam., well 35 separated from each other. For further details see Laundon (2003) and Kukwa (2006a). Chemistry. Usnic acid, zeorin ±, atranorin (in variable amounts) and unidentified terpenoids ±. Laundon (2003) reported that rarely zeorin can be missing and/or unidentified fatty acids present; Flakus & Kukwa (2007) detected porphyrilic acid in a few specimens. K± faint yellow, C−, KC−, Pd−. Ecology and distribution. On siliceous rock (also on mosses) and sometimes on lichens; in mostly shaded and sheltered places. Europe, North America, South America, Asia (China). Discussion. Taxa containing usnic acid and zeorin are compared under L. coriensis. Lepraria elobata Tønsberg Sommerfeltia 14: 197 (1992); type: Norway (BG— holotype). Thallus leprose, powdery; margin diffuse, lobes predominantly absent; medulla predominantly absent; hypothallus absent; soredia abundant, mostly fine, 20–45 µm diam., projecting hyphae absent, soredia well separated from one another; consoredia sometimes present, up to 100 µm. For more details see Tønsberg (1992). Chemistry. Atranorin, stictic acid, constictic acid, cryptostictic acid, zeorin and very rarely an unidentified fatty acid (major to trace) (Tønsberg 1992, 2004 and others). K− or + yellow, C−, KC−, Pd+ orange. Ecology and distribution. On bark, sometimes soil, wood, siliceous rock and mosses; in shady, humid places. Europe, North America, Greenland. Discussion. The phylogenetic placement is recorded under L. alpina. Lepraria lobificans (cottony appearance) and the rare L. caesioalba var. groenlandica (granular thallus) have similar chemistry to L. elobata. Morphologically, L. caesiella (atranorin and zeorin) and 36 THE LICHENOLOGIST L. incana (divaricatic acid and zeorin) may be similar to L. elobata; further morphologically similar species are mentioned under L. jackii. Species producing stictic acid complex are compared under L. caesioalba. Lepraria friabilis Lendemer, K. Knudsen & Elix Opuscula Philolichenum 5: 64 (2008); type: USA (NY— holotype; B, CANB, UCR, UDGA, hb. Lendemer— isotypes). Thallus leprose, powdery; margin diffuse, lobes absent; medulla absent; hypothallus present, thin, colourless, inconspicuous; soredia sparse to abundant, very fine, (10−)20–30 µm diam., projecting hyphae usually present, short; consoredia present, up to 60 µm. For more details see Lendemer et al. (2008). Chemistry. Two chemotypes: (1) fumarprotocetraric acid with minor amounts of protocetraric, succinprotocetraric and confumarprotocetraric acids; and (2) fumarprotocetraric acid only. Both chemotypes K−, C−, KC−, Pd+ orange or red (Lendemer et al. 2008). Ecology and distribution. On bark of conifers; in humid habitats. Southern North America. Discussion. Morphologically the most similar species is L. caesiella (thinner thallus, slightly smaller soredia, atranorin and zeorin). Species that may contain protocetraric and/or fumarprotocetraric acids are discussed under L. caesioalba. Lepraria gelida Tønsberg & Zhurb. Graphis Scripta, 18: 64 (2006); type: Svalbard (BG— holotype; NMW— isotype). Thallus leprose, powdery to cottony; margin diffuse to delimited, lobes usually absent; medulla present, thick, rarely thin, white; thallus surfaces without soredia present, can be relatively large, medulla exposed; soredia abundant to sparse in places, variable in size, Vol. 41 mostly 65–100(−200) µm diam., projecting hyphae present, short to medium. For more details see Tønsberg & Zhurbenko (2006) and Saag et al. (2007). Chemistry. Alectorialic and porphyrilic acids. K+ yellow or orange, C−, KC+ red, Pd+ yellow or orange (Tønsberg & Zhurbenko 2006; Saag et al. 2007). Ecology and distribution. On soil, mosses and sometimes lichens, rarely bark; in open habitats. Greenland, Svalbard and the Russian Arctic islands near the Taimyr peninsula (Tønsberg & Zhurbenko 2006; Saag et al. 2007). Discussion. Species producing alectorialic acid are compared under L. eburnea. Porphyrilic acid is also produced by L. alpina, L. atlantica and L. neojackii, which never contain alectorialic acid. Lepraria vouauxii is often morphologically quite similar but produces pannaric acid 6-methylester and atranorin. Lepraria goughensis Elix & Øvstedal Mycotaxon 93: 274 (2005); type: Gough Island (BG— holotype). Thallus leprose, powdery; margin delimited, usually poorly defined lobes present; medulla absent; hypothallus absent; soredia abundant, very fine, 20–26 µm diam., projecting hyphae present, numerous. For further details see Elix et al. (2005). Chemistry. Lecanoric acid, gyrophoric acid (minor to trace), strepsilin (minor to trace), fragilin (trace), 7-chloroemodin (trace), flavo-obscurin C (trace). K−, C+ red, KC+ red, Pd–(Elix et al. 2005). Ecology and distribution. On mosses and debris, peat; in shady, humid places. Gough Island (South Atlantic). Discussion. Species that can produce lecanoric acid are discussed under L. cupressicola. Lepraria multiacida and L. xerophila can produce strepsilin, but these taxa differ in both 2009 World survey of Lepraria—Saag et al. their morphology and chemistry. Anthraquinone producing species are mentioned under L. bergensis. Lepraria granulata Slav.-Bay. In Slavíková-Bayerová & Fehrer, Lichenologist 39: 321 (2007); type: Bulgaria (PRA—holotype; BG—isotype; GenBank accession no. DQ914539). Thallus leprose, granular; margin delimited to diffuse, lobes sometimes present, obscure; medulla absent; hypothallus usually scarce, usually grey; soredia abundant, coarse, up to 0–2(–0·3) mm diam., projecting hyphae usually absent. For more details see Slavíková-Bayerová & Fehrer (2007). Chemistry. Atranorin ±, ‘granulata unknown 1’, ‘granulata unknown 2’ ±, unidentified anthraquinones ± (hyphae below thallus only). K+ yellowish, coloured hyphae below thallus K+ purple-red, other hyphae below thallus K−, C−, Pd− or + faint yellowish. The diagnostic feature of this recently described species is the production of two unknown fatty acids, ‘granulata unknown 1 & 2’ (Slavíková-Bayerová & Fehrer 2007). Ecology and distribution. On mosses on siliceous rock, sometimes also on soil. So far known from Central and Eastern Europe. Discussion. The phylogenetic placement is reported under L. alpina. In addition, Slavíková-Bayerová & Fehrer (2007) characterized L. sp. G that is chemically identical to L. granulata, and also is very close to that species in ITS sequences, but is morphologically more similar to L. humida. Morphologically similar species include L. alpina, L. caesioalba, L. neglecta and especially L. borealis, but L. granulata is chemically distinct by producing unknown fatty acids ‘granulata unknown 1 & 2’. Lepraria humida Slav.-Bay. & Orange Lichenologist 38: 505 (2006); type: United Kingdom (PRA—holotype; BG, NMW—isotypes; GenBank accession no. DQ401101). 37 Thallus leprose, powdery; margin diffuse or delimited, lobes absent; true medulla absent; hypothallus weakly developed, sparse, pale orange-brown; soredia abundant, mostly fine to medium sized, 40–100(−160) µm diam., projecting hyphae absent. For more details see Slavíková-Bayerová & Orange (2006). Chemistry. The usual chemotype contains (1) atranorin, jackinic/rangiformic acid, norjackinic/norrangiformic acid ± (minor), angardianic/roccellic acid ± (minor) and unknown anthraquinones ± (minor, in subthalline hyphae only); sometimes (2) stictic and constictic acids have also been found, but were interpreted as contamination by Slavíková-Bayerová & Orange (2006). K+ yellowish, hyphae below thallus K+ purplered, C−, Pd− or + yellow. Ecology and distribution. On siliceous rocks, often between mosses; on rain-sheltered damp surfaces. Europe (British Isles). Discussion. According to molecular studies, L. humida is a distinct taxon with considerable intraspecific variability. It is related to L. atlantica, thus, belonging to the extended L. neglecta group (Fehrer et al. 2008). Morphologically similar species are mentioned under L. jackii. Lepraria humida is distinguished by its chemistry, especially anthraquinones in subthalline hyphae, and ecology. Anthraquinone producing species are discussed under L. bergensis. Slavíková-Bayerová & Orange (2006) also reported L. sp. H which was said to be morphologically and chemically identical to L. humida but distinct in ITS sequences; according to Fehrer et al. (2008), it is monophyletic, closely related to L. humida and forms ‘L. atlantica group’ together with L. atlantica and L. humida. Lepraria sp. H has been reported growing on rain-sheltered siliceous rocks, moss and rarely bark in Western to Central Europe. Slavíková-Bayerová & Fehrer (2007) also added L. sp. BG with the same morphological and chemical features, growing on damp siliceous rock in Bulgaria. See also discussion under L. jackii. 38 THE LICHENOLOGIST Lepraria impossibilis Sipman Herzogia 17: 30 (2004); type: El Salvador (B—holotype; LAGU—isotype). Thallus leprose, powdery to membranous; margin delimited, lobes present, c. 1 mm wide and long, with raised marginal rim; medulla present, thin, white; hypothallus grey; soredia abundant, medium sized, c. 0·1 mm diam., projecting hyphae absent. For more details see Sipman (2004). Chemistry. Diagnostic substances are lecanoric acid and pannaric acid 6-methylester; accessories include 4-oxypannaric 6methylester, atranorin (trace), unidentified anthraquinones and ‘vouauxii unknown 1’. K− or + faint yellow, hyphae below thallus K+ purplish, C+ more or less red, KC+ more or less red, Pd− or + orange (Sipman 2004; Flakus & Kukwa 2007). Ecology and distribution. On bark, also on terricolous and saxicolous mosses, soil, siliceous rock and schist; in forest or open areas. South and Central America, Asia – Iran (M. Sohrabi, pers. comm.). Discussion. Lepraria cupressicola may be morphologically very similar but lacks pannaric acid 6-methylester and contains fatty acids. Species that can produce lecanoric acid are discussed under L. cupressicola. Species containing pannaric acid 6-methylester as major substance are compared under L. vouauxii. Distinctly lobate species are discussed under L. membranacea and L. bergensis. Lepraria incana (L.) Ach. Meth. Lich.: 4 (1803).—Byssus incana L., Sp. Pl. 2: 1169 (1753); type: United Kingdom, drawing in Dillenius, Hist. Musc. tab. I fig. 3 (1742) (holotype); typotype: hb Hist. Muse: tab. I no 3 (OXF). Thallus leprose, powdery; margin diffuse, lobes absent; medulla sometimes present, usually poorly developed; hypothallus scarce; soredia abundant, mostly fine, up to 50 µm diam., projecting hyphae sometimes present, short; consoredia sometimes present, up to Vol. 41 110 µm. For further details see Laundon (1992) and Tønsberg (1992). Chemistry. Common chemotype (1): divaricatic acid, nordivaricatic acid ± (trace), zeorin (rarely absent – Baruffo et al. 2006; Leuckert et al. 1995) and atranorin ± (major to trace); K− or + faint yellow, C−, KC−, Pd−. Rare chemotype (2): with anthraquinones in addition to the aforementioned substances: parietin, fallacinal, parietinic acid and citreorosein; K+ purple-red, C−, KC+ purple-red, Pd+ orange (Laundon 1992; Tønsberg 1992). Very rare accessories include gyrophoric acid, lecanoric acid, thamnolic acid and an unknown terpenoid (Laundon 1992; Leuckert et al. 1995; Baruffo et al. 2006; Makarova et al. 2006). Ecology and distribution. On bark, acidic rock, and sometimes on mosses, wood and soil; in more or less shaded places. Worldwide, except Arctic and Antarctic. Discussion. Lepraria incana can be morphologically quite variable, sometimes forming relatively loose cottony cushions and sometimes consisting of sparse or denser separate soredia only. Species producing divaricatic acid are discussed under L. crassissima. Lepraria caesiella and L. elobata are sometimes morphologically similar but both these species lack divaricatic acid. Lepraria isidiata (Llimona) Llimona & A. Crespo in Wirth et al., Guía de Campo de los Líquenes, Musgos y Hepáticas: 309 (2004).—L. crassissima var. isidiata Llimona, in Vězda, Lich. Sel. Exs. Fasc. 47: 7 (1973); type: Spain (BCC—holotype; isotypes distributed in Vězda, Lich. Sel. Exs. Fasc. 47: 7 (1973). Thallus crustose to subfoliose, membranous to granular; margin delimited, lobes present, usually well-developed, margin raised; hypothallus sometimes present, light brownish; subcorticate patches may be present; granules similar to true isidia abundant, sometimes not well separated at margin, very coarse, 0·32–0·54(−3) mm diam., 2009 World survey of Lepraria—Saag et al. compact outer layer present, projecting hyphae rare and short, granules often become sorediate. For more details see Wirth et al. (2004), Baruffo et al. (2006), Tretiach et al. (2008). Chemistry. Chemotype (1) with atranorin, fumarprotocetraric and protocetraric acids and roccellic acid ±; chemotype (2) with atranorin and fumarprotocetraric acid and roccellic acid ±; and chemotype (3) with atranorin, protocetraric acid and roccellic acid ± (Wirth et al. 2004; Crespo et al. 2006; Baruffo et al. 2006). Chemotype (3) is rare (Baruffo et al. 2006). K− or + yellow, C−, KC− or + yellow, Pd+ reddish orange, hypothallus Pd−. Ecology and distribution. On calcareous soil and rock (sometimes on mosses); in shaded but well-lit places. Europe (Mediterranean countries). Discussion. The phylogenetic placement of this taxon is mentioned under L. membranacea. Lepraria isidiata was historically described as L. crassissima var. isidiata Llimona (Vězda 1973). Specimens of L. isidiata have also been included in L. nivalis (Laundon 1992; Leuckert et al. 2004; Sipman & Raus 1999; Sipman 2004). Species that can produce structures more or less similar to isidia include L. crassissima (divaricatic and nordivaricatic acids), L. xerophila (pannaric acid 6-methylester or rarely norascomatic acid), L. isidiata and L. santosii. Lepraria isidiata and L. xerophila produce structures similar to true isidia having a compact outer layer. Some chemotypes of L. santosii are chemically distinguishable by the stictic acid complex, while others are chemically similar to L. isidiata. Lepraria santosii differs from L. isidiata morphologically (coarse soredia without compact outer layer) and ecologically (siliceous substrata). Species producing protocetraric and/or fumarprotocetraric acids are compared under L. caesioalba. Distinctly lobate species are discussed under L. membranacea and L. bergensis. 39 Lepraria jackii Tønsberg Sommerfeltia 14: 200 (1992); type: Norway (BG— holotype; BM—isotype). Lepraria toensbergiana Slav.-Bay & Kukwa, Bryologist 108: 132 (2005). Thallus leprose, powdery; margin diffuse or delimited, lobes absent; true medulla absent; hypothallus sparse to continuous and thick (epibryotic specimens), white; sometimes hypothallus exposed in places; soredia abundant, fine to coarse, up to 160 µm diam., projecting hyphae sometimes present, short; consoredia sometimes present, up to 160 µm diam. For further details see Tønsberg (1992), Bayerová et al. (2005) and Slavíková-Bayerová & Orange (2006). Chemistry. Lepraria jackii sensu SlavíkováBayerová & Orange (2006) comprises specimens with atranorin and jackinic/rangiformic acid as main compounds, and accessories roccellic acid, norjackinic/norrangiformic acid (minor) and toensbergianic acid (minor). Lepraria toensbergiana sensu Bayerová et al. (2005) contains atranorin and toensbergianic acid. In the earlier, broader sense, L. jackii included all chemotypes with atranorin and any of the fatty acid(s) mentioned above, plus rarely also zeorin and/or minor amounts of anthraquinones (Tønsberg 1992; Leuckert et al. 1995; Baruffo et al. 2006, and others). K− or + faint yellow, C−, KC−, Pd−. Ecology and distribution. On bark, sometimes mosses, wood, siliceous and neutral rock, rarely on soil, plant debris or lichens; mostly acidic substrata; usually in more or less sheltered, relatively humid places. Europe, North America, Asia, Australia (L. toensbergiana – Central Europe). Discussion. Recently L. toensbergiana was separated from the morphologically identical L. jackii on the basis of its chemistry (atranorin together with the newly described fatty acid toensbergianic acid), ITS sequences and montane distribution (Bayerová et al. 2005; Bayerová & Haas 2005). However, Baruffo et al. (2006) did not regard the differences 40 THE LICHENOLOGIST sufficient and L. toensbergiana was reduced to a synonym of L. jackii. Further evidence for synonymizing L. toensbergiana with L. jackii comes from the molecular phylogeny by Fehrer et al. (2008). The monophyly of L. toensbergiana and the small sequence difference between these taxa was confirmed (similar to the intraspecific variability of some other Lepraria species), however, the two taxa formed a single clade on a highly supported long branch of the phylogenetic tree. Three further species, L. celata, L. humida, and L. sylvicola were segregated from L. jackii by Slavíková-Bayerová & Orange (2006). According to molecular studies, these taxa are distinct and not closely related to L. jackii or to each other (Fehrer et al. 2008). They are all morpologically similar to L. jackii, the chemical separation being based on the contents of atranorin, different fatty acids and anthraquinones in the subthalline hyphae. Other morphologically similar species include L. atlantica, L. elobata and L. neojackii which are chemically easily separable (see the descriptions of these taxa). Lepraria jackii s. lat. can be chemically similar to several species: L. bergensis and L. lobata (delimited thallus margin with lobes, anthraquinones); L. borealis, L. caesioalba and L. granulata (granular L. neglecta type thallus); L. caesiella (zeorin, only rarely with fatty acids); L. nivalis (one chemotype, usually thicker, cottony and has a delimited thallus margin often with lobes); L. normandinoides (rare chemotype, marginal lobes, thick dark hyphae below thallus); L. pallida (lobes and well-developed dark hypothallus); L. rigidula (nephrosteranic acid, lax cottony thallus with long projecting hyphae from soredia). Vol. 41 but mostly not projecting hyphae, colourless or in external parts often brown; consoredia large, to 1 mm. For more details see Tønsberg (2007). Chemistry. Protocetraric acid, angardianic/ roccellic acid. Thallus K−, C−, KC−, PD+ orange (Tønsberg 2007). Ecology and distribution. Mostly on rock, and sometimes on soil or mosses; in dry to rather wet overhangs and rock walls. Eastern USA, montane. Discussion. Several species produce protocetraric and/or fumarprotocetraric acids (discussed under L. caesioalba). However, L. lanata can be distinguished by its characteristic consoredia. Lepraria lecanorica Tønsberg in Nash, T. H. III, Ryan, B. D., Diederich, P., Gries, C. & Bungartz, F. (eds), Lichen Flora of the Greater Sonoran Desert Region 2: 326 (2004); type: USA (BG—holotype; ASU—isotype). Thallus leprose, powdery to membranous; margin delimited, lobes sometimes present, obscure; medulla present, indistinct to distinct, white; lower surface sometimes present, white to pale brown, usually without tomentum; soredia abundant, coarse, up to 200 µm diam., projecting hyphae sometimes present, usually short. For more details see Tønsberg (2004). Chemistry. Lecanoric acid, atranorin (major to minor). K− or + yellowish, C+ red, KC+ red, Pd− (Tønsberg 2004). Ecology and distribution. On bark, rock, soil; in shaded places. North and South America. Lepraria lanata Tønsberg Opuscula Philolichenum 4: 51 (2007); type: USA (BG— holotype; ASU, DUKE, NMW, NY—isotypes). Thallus leprose, with characteristic woolly consoredia; margin diffuse, lobes absent; medulla absent; soredia abundant, all soredia aggregated into consoredia, surrounded by lax network of branching and anastomosing Discussion. Species that can produce lecanoric acid are discussed under L. cupressicola. Lepraria leprolomopsis Diederich & Sérus. in Aptroot, Diederich, Sérusiaux & Sipman, Bibl. Lichenol. 64: 76 (1997); type: Papua New Guinea (B— holotype; LG, hb. Diederich—isotypes). 2009 World survey of Lepraria—Saag et al. Thallus leprose, powdery to cottony; margin usually delimited, lobes absent; medulla present, distinct, white; hypothallus usually present, poorly developed, white, lower surface present, even, white, not or poorly tomentose; soredia abundant, mostly medium sized, 75–125 µm diam., projecting hyphae sometimes present; consoredia often present, 150–300 µm. For more details see Aptroot et al. (1997). Chemistry. Atranorin, stictic, constictic, cryptostictic, norstictic and connorstictic acids and an unknown terpenoid. K+ yellow, C−, KC−, Pd+ orange (Aptroot et al. 1997). Ecology and distribution. On bark (also on mosses). Australasia (Papua New Guinea). Discussion. Species producing the stictic acid complex and atranorin are discussed under L. caesioalba. Lepraria leprolomopsis may be morphologically similar to L. diffusa (4-oxypannaric acid 2-methylester), L. eburnea (alectorialic acid), L. lobificans (stictic acid complex plus zeorin), L. nylanderiana (thamnolic acid), L. vouauxii (pannaric acid 6-methylester) and L. nivalis (discussed under L. nivalis). Lepraria leuckertiana (Zedda) L. Saag comb. nov. Lecanora leuckertiana Zedda, Nova Hedwigia 71: 108 (2000); type: Italy (B—holotype). Thallus leprose, cottony and powdery to granular; margin diffuse or delimited, lobes usually minute, often obscure; medulla present, well-developed, white; patches with exposed medulla present, also medullary hyphae intermixed with soredia; soredia abundant, fine to coarse, up to 500 µm diam., not well-separated from each other. For more details see Zedda (2000a). Chemistry. Usnic acid, zeorin and isousnic acid (trace). K−, C−, KC−, Pd− (Zedda 2000a). Ecology and distribution. On bark, sometimes on soil; in humid, open, well-lit places. 41 Europe, North Africa, South America (Peru). Discussion. Zedda (2000a) described Lecanora leuckertiana as very similar to Lepraria vouauxii (pannaric acid 6-methylester, no usnic acid), but provisionally included the new species in Lecanora because of its chemistry. At that time, following the concept of Laundon (1992, 2003), the genus Lepraria did not contain any usnic acid-producing species. However, Sipman (2003) placed the newly described L. usnica with usnic acid in Lepraria arguing that this character alone is not sufficient for discrimination at the generic level. Subsequently L. coriensis and L. ecorticata were transferred from Lecanora (Sipman 2004; Kukwa 2006a), and Lepraria texta described as new (Knudsen & Elix 2008). Taking into account these developments, the authors considered the transfer of L. leuckertiana appropriate. Even so, the phylogenetic relationships of the aforementioned species should be clarified using molecular methods in the future. Species producing usnic acid and zeorin are compared under L. coriensis. Lepraria lobata Elix & Kalb in Elix, Mycotaxon 94: 220 (2005); type: Australia (PERTH—holotype). Thallus leprose, granular, partly membranous; margin usually delimited, lobes often present and well-defined, 1–2 mm wide, slightly to distinctly raised at margins; medulla present, distinct, white; hypothallus absent; sometimes only sparse soredia on exposed medulla; soredia sparse to abundant, fine, 20–75 µm diam., projecting hyphae usually present, usually short; consoredia common, up to 350 µm diam. For more details see Elix (2005). Chemistry. Atranorin, zeorin and a fatty acid (rangiformic/jackinic or roccellic/ angardianic acid); rarely zeorin is present in only a minor amount or very rarely it is absent. Accessories include norrangiformic, pallidic, conpallidic and ursolic acids and an unknown dibenzofuran (minors), and 42 THE LICHENOLOGIST 3,7-di-O-methylstrepsilin, fragilin, 7chloroemodin and 3’-demethylatranorin (traces). K+ yellow, C−, Pd+ pale yellow (Elix 2005). Ecology and distribution. On bark, mosses on rock and on soil; in sheltered places. Australia. Discussion. Lepraria lobata and L. pallida both produce atranorin, zeorin and fatty acids, and have lobes, but L. pallida has a more membranous and rough appearance, a well-developed ±dark hypothallus and thin medulla, and contains unidentified fatty acids; whereas L. lobata has a slightly thicker thallus with a thicker medulla, is more green in colour and has finer individual soredia and larger consoredia, produces roccellic or rangiformic acid and sometimes small amounts of anthraquinones and unknown dibenzofurans. However, the characters of these species seem relatively similar. Other species containing atranorin and fatty acids are mentioned under L. jackii and taxa with well-developed lobes under L. membranacea. Lepraria lobificans Nyl. Flora 56: 196 (1873); type: France (H—lectotype; BM—topotype). Thallus leprose, cottony to rarely powdery; margin usually diffuse, lobes rarely present, weakly developed; medulla present, usually thick, white; hypothallus rarely present, scarce, pale brown; thallus surfaces without soredia sometimes present, then medulla exposed; soredia abundant, fine, up to 60 µm diam., projecting hyphae present, long; consoredia abundant, up to 100(−200) µm. For further details see Laundon (1992). Chemistry. Atranorin, stictic acid, constictic acid, cryptostictic acid ± (variable amounts, often major), norstictic acid ± (trace) and zeorin; rare accessories include roccellic/angardianic acid, an unidentified fatty acid and an unidentified terpenoid (Laundon 1992; Baruffo et al. 2006 and others). Laundon (1992) reported another, Vol. 41 rare chemotype that lacks zeorin. K− or + yellow, C−, KC−, Pd+ orange. Ecology and distribution. On bark or rock (usually on mosses) and various other substrata; in shaded, sheltered places. Worldwide. Discussion. Species producing the stictic acid complex and atranorin are discussed under L. caesioalba. For morphologically similar species, see the discussion under L. leprolomopsis. Lepraria incana (divaricatic acid) is sometimes also morphologically similar as both species can be quite variable. Lepraria membranacea (Dicks.) Vain. Acta Soc. Fauna Flora Fennica 49(2): 265 (1921).— Lichen membranaceus Dicks., Fasc. Pl. Crypt. Brit. 2: 21 (1790); type: United Kingdom (BM ex K ex D Turner—holotype). Thallus crustose to subfoliose, leprose, membranous; margin delimited, lobes present, well-developed, wide, margin raised or flat; medulla present, distinct, white; hypothallus present, usually well-developed, dark, sometimes white along margin; thallus surfaces without soredia sometimes present; soredia abundant, sometimes not wellseparated at margin, fine to medium, up to c. 100 µm diam., projecting hyphae sometimes present, short; consoredia often present, up to c. 200 µm. For more details see Laundon (1989). Chemistry. Pannaric acid (major) with satellite dibenzofurans (minors to traces), roccellic/angardianic acid (rarely absent), atranorin ± (major to trace) and very rarely norstictic acid or zeorin (Laundon 1989; Baruffo et al. 2006 and others). K− or + yellow, C−, Pd+ reddish orange. Ecology and distribution. On rock (also on mosses), rarely on bark or soil; shaded to sun-exposed, sheltered from rain. Worldwide but scattered. Discussion. According to molecular studies L. membranacea, L. vouauxii, L. xerophila, L. 2009 World survey of Lepraria—Saag et al. bergensis, L. isidiata, L. santosii form a monophyletic group in Lepraria based on ITS (Crespo et al. 2006; Tretiach et al. 2009), most of these species having membranous and lobed thalli (L. vouauxii only rarely develops small lobes). Lepraria membranacea is the oldest name and the most widely distributed of the strongly lobate Lepraria species. Species that can form prominent lobes include L. bergensis (anthraquinones, relatively small thallus), L. coriensis (usnic acid and zeorin), L. cupressicola (lecanoric acid, atranorin and zeorin, well-developed dark hypothallus), L. impossibilis (lecanoric and pannaric acids), L. isidiata (protocetraric and/or fumarprotocetraric acids, coarse granules similar to true isidia), L. lobata and L. pallida (atranorin, zeorin and fatty acids, see the discussion under L. lobata), L. membranacea (pannaric acid), L. normandinoides (usually protocetraric and/or fumarprotocetraric acids, soredia fine), L. sipmaniana (pannaric acid 6-methylester, thallus usually rather bright yellowish), L. santosii (stictic or protocetraric and/or fumarprotocetraric acids, coarse soredia slightly similar to isidia), L. squamatica (squamatic acid, only rarely lobate) and L. xerophila (usually pannaric acid 6-methylester, very coarse structures similar to true isidia). Lepraria multiacida Aptroot Fungal Divers. 9: 20 (2002); type: Brazil (SP—holotype, ABL—isotype). Thallus leprose, granular; margin delimited, minute lobes often present, up to 0·5 mm wide, thinner than the rest of the thallus, without raised rims; medulla present, distinct, white, sometimes blackened; sometimes medullary hyphae also form hypothallus, white to black; soredia abundant, coarse, c. 100–200 µm diam., projecting hyphae present, long. For more details see Aptroot (2002). Chemistry. Atranorin; usually zeorin together with 1–2 unknown terpenoids; constictic acid (major to minor); stictic, cryptostictic and norstictic acids (all minor to trace); salazinic acid (minor to trace); 43 3,7-O-methylstrepsilin (minor to trace); 7-O-methylstrepsilin, strepsilin and an unidentified dibenzofuran (all trace). K+ yellow to orange, C−, KC− (Aptroot 2002; Elix & Tønsberg 2004; Elix 2006a). Ecology and distribution. On sandstone and soil. South America (Brazil). Discussion. The chemically and morphologically similar L. salazinica contains large quantities of salazinic acid and lacks other substances present in L. multiacida. Lepraria crassissima and sometimes L. nivalis and L. squamatica may be morphologically similar, but differ in their chemistry. Species producing the stictic acid complex are discussed under L. caesioalba (none of them contains salazinic acid). Lepraria neglecta (Nyl.) Erichsen in Lettau, Feddes Repert. 61: 127 (1958).—Lecidea neglecta Nyl., Not. Skällsk. Fauna Fl. Fenn. Förh. 4: 233 (1859); type: Finland (H—lectotype). Thallus leprose, granular; margin usually delimited, obscure minute lobes sometimes present; medulla sometimes present, inconspicuous, white; hypothallus sometimes present, poorly-developed, grey to brown; rarely small patches with exposed medulla present; soredia abundant, mostly coarse, typically 100–150(−200) µm diam., projecting hyphae usually absent; consoredia frequent, typically 200–300 µm. For more details see Laundon (1992). Chemistry. Alectorialic acid and roccellic/ angardianic acid; rarely rangiformic acid replaces roccellic/angardianic acid or very rarely both fatty acids are present; rarely atranorin (in variable amounts) occurs; other accessories include norrangiformic acid (minor to trace), 5,7-dihydroxy6-methylphthalide (minor to trace) and barbatolic acid (trace). K− or + yellow, C− or + reddish orange, KC+ pink or reddish orange, Pd+ lemon yellow or orange (Laundon 1992; Leuckert et al. 1995 and others). Kukwa (2006b) reported a very rare chemotype without alectorialic acid, 44 THE LICHENOLOGIST containing an unknown substance ‘neglecta unknown’. Ecology and distribution. On acidic rock (usually on mosses), soil and rarely on bark, epiphytic mosses and lichens; in exposed places, mostly cool climate, montane-alpine in tropics. Europe, North and South America, Asia, Australasia, Antarctica, Greenland. Discussion. According to molecular studies, L. neglecta belongs to the monophyletic ‘L. neglecta group’, see the discussion under L. alpina. Morphologically the most similar species include L. borealis, L. caesioalba and L. granulata, but these all differ chemically. Alectorialic acid-producing species are discussed under L. eburnea. Lepraria neojackii Flakus & Kukwa Lichenologist 39: 468 (2007); type: Bolivia (KRAM-L— holotype; LPB, UGDA—isotypes). Thallus leprose, powdery; margin diffuse, lobes absent; medulla absent but sometimes with pseudomedulla of bleached soredia mixed with hypothallus hyphae; hypothallus usually present, whitish or orange in places; soredia abundant, very fine, up to 20 µm diam., projecting hyphae usually absent; consoredia present, not prevalent, up to 40 µm diam. For further details see Flakus & Kukwa (2007). Chemistry. Porphyrilic acid, roccellic/ angardianic acid, rangiformic/jackinic acid and an unidentified anthraquinone (Flakus & Kukwa 2007). Ecology and distribution. On mosses, humus and rocks; in mountain cloud forest. South America (Bolivia). Discussion. Lepraria atlantica is chemically and morphologically the most similar species, but it contains atranorin together with porphyrilic acid. Lepraria achariana is also morphologically close but contains lecanoric acid. Other similar species are mentioned under L. jackii. Vol. 41 Lepraria nivalis J. R. Laundon Lichenologist 24: 327 (1992); type: France (MARSSJ— holotype). Thallus leprose, cottony to powdery or membranous; margin delimited or diffuse, lobes absent or present; sometimes welldeveloped and with raised marginal rim; medulla usually present, often exposed in places, usually thick, white; lower surface sometimes with distinct brown tomentum; soredia abundant, variably sized, up to 300 µm diam., projecting hyphae usually present, short to long. For more details see the references below. Chemistry. Leuckert et al. (2004) distinguished 6 chemotypes in L. nivalis s. lat; a classification was complemented by Baruffo et al. (2006). Chemotype (1) atranorin, protocetraric acid; (2) atranorin, protocetraric acid, roccellic acid; (3) atranorin, fumarprotocetraric acid (major to trace), protocetraric acid (major to trace); (4) atranorin, stictic and constictic acids, cryptostictic acid ± (trace), norstictic acid ± (trace); (5) atranorin, stictic, constictic and roccellic acids, cryptostictic acid ± (trace), norstictic acid ± (trace); (6) atranorin, psoromic acid, 2’-Odemethylpsoromic acid (trace). In addition, Zedda (2000b) reported a chemotype (7) with atranorin and roccellic acid only. Laundon (1992) reported gyrophoric acid as a very rare accessory; Baruffo et al. (2006) detected atranorin, protocetraric acid, virensic acid and strepsilin dimethyl ester in a single specimen; Sipman (2004) reported a specimen with protocetraric and roccellic acid without atranorin. Chemotypes 1–5: K− or + yellow, C−, KC− or + yellow, Pd+ reddish orange; chemotype 6: K− or + yellow, C−, KC− or + yellow, Pd+ yellow; chemotype 7: K− or + yellow, C−, KC−, Pd−. Ecology and distribution. On limestone (also on mosses), sometimes on soil, rarely on bark or siliceous rock; in mostly shaded places. Europe, North America, Asia, Australasia (Papua New Guinea), Greenland (tentatively reported by Saag et al. 2007). Most 2009 World survey of Lepraria—Saag et al. 45 common in Mediterranean areas and southern Europe. 60 µm diam., projecting hyphae absent. For more details see Lendemer & Harris (2007). Discussion. Lepraria nivalis is very heterogeneous both chemically and morphologically, and possibly includes different taxa (Baruffo et al. 2006; Crespo et al. 2006). Chemotype (4) represents ‘isidioid’ morphology and agrees with the description of some chemotypes of L. santosii. Chemotype (5) includes both ‘isidioid’ and ‘non-isidioid’ morphs and the ‘isidioid’ morphotype also has an affinity with L. santosii. The ‘isidioid’ representatives of the chemotype (3) belong to L. isidiata according to the present taxonomy (Baruffo et al. 2006) while the less warted and softer specimens are left in L. nivalis. Lepraria crassissima (divaricatic and nordivaricatic acids), L. lobificans (stictic acid complex and zeorin) and especially L. nylanderiana (thamnolic acid) are morphologically similar. Chemotype 7 included by Zedda (2000b) is chemically identical to L. borealis, which has a granular thallus. Morphological characters for separating non-lobate forms of L. nivalis and L. leprolomopsis seem to be limited. Lepraria leprolomopsis has been found on mossy bark, has a white lower surface and produces a terpenoid, whereas L. nivalis is associated with calcareous rock and soil, sometimes has a brown tomentum under the thallus and terpenoids are absent. Other similar species are mentioned under L. leprolomopsis. Species producing the stictic acid complex and atranorin, or protocetraric and/or fumarprotocetraric acids are discussed under L. caesioalba. Chemistry. Common chemotype (1): atranorin, protocetraric acid, virensic acid (trace), roccellic acid; rare chemotype (2): atranorin, fumarprotocetraric acid, protocetraric acid (minor to trace), roccellic acid; very rare chemotype (3): atranorin, roccellic acid. Chemotypes (1) & (2): K+ yellow, C−, KC+ yellowish, Pd+ orange; chemotype (3): K+ yellow, C−, KC+ yellowish, Pd− (Lendemer & Harris 2007). Lepraria normandinoides Lendemer & R. C. Harris Opuscula Philolichenum 4: 45 (2007); type: USA (NY— holotype; isotypes distributed in Lichens of Eastern North America V: 221). Thallus leprose, membranous; margin delimited, lobes usually well developed, with marginal rim; medulla present, whitish; lower surface pale, with thick brown tomentum of thick hyphae; soredia abundant, fine, 30– Ecology and distribution. On acidic rock and bark; in semi-shaded more or less humid places. Eastern North America. Discussion. Morphologically similar species are discussed under L. bergensis and L. membranacea. Species producing protocetraric and/or fumarprotocetraric acids are compared under L. caesioalba. Lepraria nylanderiana Kümmerl. & Leuckert Biblioth. Lichenol. 58: 250 (1995); type: Italy (B— holotype). Thallus leprose, cottony to powdery or membranous; margin delimited, minute lobes present in well-developed specimens, without raised rims; medulla usually present, whitish; conspicuous greyish hypothallus present in well-developed specimens; thallus surfaces without soredia often present, then medulla exposed; soredia abundant, medium to coarse, up to 300 µm diam., projecting hyphae usually present, short. For more details see Leuckert et al. (1995). Chemistry. Thamnolic acid, decarboxythamnolic acid (trace), roccellic acid and rarely atranorin (trace). K+ lemon yellow, C−, KC−, Pd+ orange-yellow (Leuckert et al. 1995; Baruffo et al. 2006 and others). Ecology and distribution. On soil, mosses, siliceous rocks and sometimes bark. Europe (Mediterranean area, Central Europe, Great Britain). 46 THE LICHENOLOGIST Discussion. Morphologically the most similar species is L. nivalis but it contains no thamnolic acid. Further morphologically similar species are discussed under L. leprolomopsis. For species producing thamnolic acid, see under L. aurescens. Lepraria obtusatica Tønsberg Sommerfeltia 14: 204 (1992); type: Norway (BG— holotype; DUKE, E—isotypes). Thallus leprose, powdery; margin diffuse, lobes absent; medulla usually absent; rarely soredia mixed with some medullary hyphae; soredia abundant, very fine, up to 35 µm diam., very fragile; consoredia sometimes present, up to 50 µm diam. For more details see Tønsberg (1992). Vol. 41 unidentified substances ± (traces) (Sipman 2004; Flakus & Kukwa 2007). K+ pale yellow, C−, KC−, Pd+ yellow to orange. Ecology and distribution. On bark, sandstone and soil. South America, Africa (Seychelles, Madagascar). Discussion. Lepraria cupressicola (lecanoric acid) and L. impossibilis (dibenzofurans) also have a developed hypothallus and lobes but L. pallida is more granular and harder. Lepraria lobata has a similar chemistry and lobes, see under L. lobata. Other species with distinct lobes are discussed under L. membranacea and taxa with atranorin, zeorin and fatty acids under L. jackii. Lepraria pulchra Orange & Wolseley Chemistry and distribution. Obtusatic acid, an unidentified pigment, barbatic acid ± (trace). K−, C−, KC−, Pd+ yellow (Tønsberg 1992; Makarova et al. 2006). Ecology and distribution. On bark; in shaded places. Europe, Australia; scattered. Discussion. According to molecular studies, L. obtusatica does not belong to the genus Lepraria (Ekman & Tønsberg 2002). It is chemically unique by producing obtusatic acid. Lepraria pallida Sipman Herzogia 17: 33 (2004); type: Brazil (B—holotype, SP— isotype). Thallus leprose, granular to partly membranous; margin usually delimited, lobes present in places, often well-developed, 0·52 mm wide and long, with raised marginal rim; medulla present, thin, white; hypothallus sometimes well-developed, tomentose, grey to black; soredia sometimes not well-separated from each other, forming a smoother surface towards margin, abundant, medium, c. 0·1 mm diam. For more details see Sipman (2004). Chemistry. Atranorin, zeorin, unidentified fatty acids, unidentified terpenoids ± and Lichenologist 37: 249 (2005); type: Thailand (BM— holotype). Thallus leprose, powdery to membranous; margin delimited, with raised irregularly indented rim, sometimes small and indistinct lobes present, without raised rims; medulla present, thin to medium, whitish; lower surface smooth, white; soredia abundant to sparse in places, medium to coarse, 80– 140 µm diam., projecting hyphae usually present, abundant, very short (10–20 µm, rarely to 60 µm), often soredia not separated from each other, especially in thallus centre. For further details see Orange & Wolseley (2005). Chemistry. Thamnolic acid. K+ bright yellow, C−, KC−, Pd+ orange-yellow (Orange & Wolseley 2005). Ecology and distribution. On bark; in sheltered places, dry forest. Asia (Thailand). Discussion. Species producing thamnolic acid are compared under L. aurescens. Lepraria rigidula (B. de Lesd.) Tønsberg Sommerfeltia 14: 205 (1992).—Crocynia rigidula B. de Lesd., in Hue, Bull. Soc. Bot. France 71: 331 (1924); type: United Kingdom (E—holotype). 2009 World survey of Lepraria—Saag et al. Thallus leprose, cottony to powdery; margin diffuse, lobes absent; medulla sometimes present, poorly to well developed, lax, white; soredia abundant, mostly fine, sometimes coarse, up to 60(−100) µm diam., projecting hyphae present, very long or long at least on some soredia (up to 120 µm); consoredia sometimes present, up to 300 µm diam. For more details see Tønsberg (1992) and Kukwa (2006b). Chemistry. Atranorin and nephrosteranic acid (Tønsberg 1992 and others). Very rarely unidentified anthraquinones (Flakus & Kukwa 2007). K− or + faint yellow, C−, KC−, Pd−. Ecology and distribution. On bark, also on mosses, rarely on rock, soil, wood and lichens; in both shaded and open places. Europe, North America, Asia, northern Africa. Discussion. Taxa with atranorin, zeorin and fatty acids are discussed under L. jackii. Lepraria rigidula is unique in producing a rare fatty acid, nephrosteranic acid, and characteristic soredia. It was regarded as a synonym of L. alpina [sub nomine Leproloma cacuminum by Laundon (1992)], but was resurrected as a separate species by Tønsberg (1992). Lepraria salazinica Tønsberg Opuscula Philolichenum 4: 52 (2007); type: USA (BG— holotype; DUKE—isotype). Thallus leprose, powdery to granular; margin diffuse, lobes absent; medulla absent; soredia abundant to scattered, very fine, to 30 µm diam., projecting hyphae absent; consoredia present, up to 50 µm, a few aggregations larger. For more details see Tønsberg (2007). Chemistry. Atranorin, salazinic acid, roccellic/angardianic acid. K+ yellow turning red, C−, KC−, Pd+ orange (Tønsberg 2007). Ecology and distribution. On rock; under rock overhangs. Eastern USA, montane. 47 Discussion. Lepraria salazinica is best characterized by the production of salazinic acid. Lepraria multiacida also produces salazinic acid (only minor amounts in addition to other substances), but has a thick thallus with a dark hypothallus. Lepraria santamonicae K. Knudsen & Elix Bryologist 110: 115 (2007); type: USA (holotype—UCR; isotypes—ASU, BM, CANB, H, NY, hb. Lendemer). Thallus leprose, powdery (minutely granular); margin diffuse, lobes absent; medulla absent, but sometimes a white pseudomedullary layer of gelatinized hyphae present; soredia abundant, fine, 30–50 µm diam., projecting hyphae absent, but occasionally with thin colourless hyphae, some soredia not well-separated from each other. For more details see Knudsen & Elix (2007). Chemistry. Argopsin, norargopsin (major to minor). K−, C−, KC−, Pd− or Pd+ orange to orange-red (Knudsen & Elix 2007). Ecology and distribution. On rock or soil, mostly siliceous substrata; in open, sometimes exposed places; a pioneer species. North America (California, USA). Discussion. Lepraria santamonicae is unique in containing argopsin as the major secondary compound. Leparaia coriensis may also contain argopsin, but only in minor amounts in addition to usnic acid. Lepraria santosii Argüello & A. Crespo in Crespo et al., Lichenologist 38: 218 (2006); type: Canary Islands (MAF—holotype; BG, GZU— isotypes). Thallus leprose to subfoliose, membranous to granular; margin delimited, lobes present, distinct, with raised marginal rim; hypothallus sometimes present, usually poorly developed, light brownish; soredia abundant, fine to very coarse, (20–)220–340(−650) µm diam., projecting hyphae present, can be long, very coarse soredia slightly resemble 48 THE LICHENOLOGIST isidia but lack compact outer layer and have projecting hyphae instead. For more details see Crespo et al. (2006) and Tretiach et al. (2008). Chemistry. Crespo et al. (2006) reported chemotype (1) atranorin, stictic acid, constictic and norstictic acids (traces), zeorin and roccellic acid. K+ yellow to brownish, C−, KC−, Pd+ orange. Tretiach et al. (2009) added more chemotypes: (2) atranorin, stictic and constictic acids and an unknown substance (UV 366 pink); (3) atranorin, stictic and constictic acids and roccellic acid; (4) atranorin, stictic and constictic acids, protocetraric or fumarprotocetraric acid (not both) and roccellic acid; (5) atranorin, protocetraric acid and roccellic acid; (6) atranorin, protocetraric and fumarprotocetraric acids and roccellic acid; (7) atranorin, fumarprotocetraric acid and roccellic acid. In addition, they found thamnolic and/or gyrophoric acids as rare accessories in chemotype 3. Ecology and distribution. On siliceous soil and rock and also basalt; in shaded places. Europe (Mediterranean countries) and Canary Islands. Discussion. According to molecular studies, L. santosii is close to L. isidiata and L. bergensis, and has considerable intraspecific variability (Crespo et al. 2006; Tretiach et al. 2009), see also the discussion under L. membranacea. Previously, the specimens corresponding to the description of L. santosii were included in L. nivalis (discussed under L. nivalis). Other species producing more or less isidia-like structures are compared under L. isidiata. Several species produce the stictic acid complex and atranorin, or protocetraric and/or fumarprotocetraric acids, see under L. caesioalba. Lepraria sipmaniana (Kümmerl. & Leuckert) Kukwa Ann. Bot. Fenn. 39: 226 (2002).—Leproloma sipmanianum Kümmerl. & Leuckert, in Leuckert & Kümmerling, Nova Hedwigia 52: 27 (1991); type: South Africa (B—holotype). Vol. 41 Thallus leprose to subfoliose, membranous; margin delimited to diffuse in places, lobes present, well developed, often over 2 mm wide, with raised marginal rim; medulla present, thin, white; hypothallus sometimes present, inconspicuous, brownish; soredia sparse in places, exposing smooth ecorticate surface, especially near margins, soredia fine to coarse, 40–200 µm diam., projecting hyphae rarely present, short. For more details see Leuckert & Kümmerling (1991). Chemistry. The diagnostic substance is pannaric acid 6-methylester; accessories include oxypannaric acid 6-methylester, 4-oxypannaric acid 6-methylester, pannaric acid, 7-chloroemodin, fragilin, A01anthrone, parietin, emodin and ‘vouauxii unknown 1’ sensu Tønsberg (1992) (Leuckert & Kümmerling 1991; Flakus & Kukwa 2007 and others). Thallus K+ reddish brown or yellowish, hyphae below thallus K+ purple to brownish red, C± yellowish, KC+ reddish brown or yellowish, Pd+ pink. Ecology and distribution. On soil, rock, bark and mosses; in mostly exposed, well-lit places. South America, Central America, Asia (Sri Lanka, Taiwan), Africa. Discussion. Morphologically similar species are discussed under L. bergensis and L. membranacea. Species with pannaric acid 6-methylester as a major substance are compared under L. vouauxii. Lepraria squamatica Elix Australasian Lichenology 58: 20 (2006); type: Australia (CANB—holotype). Thallus leprose, powdery; margin delimited to diffuse, lobes absent or present, sometimes well-defined, up to 1 mm wide and with raised margin; true medulla absent; hypothallus sometimes present, thin, lax, white, exposed in places; soredia abundant to sparse in places, very fine, 15–40 µm diam., projecting hyphae present, variable, some very long, soredia well separated from each other; consoredia common to scarce, 150– 200 µm. For more details see Elix (2006a). 2009 World survey of Lepraria—Saag et al. Chemistry. Elix (2006a) described a chemotype (1) with squamatic and baeomycesic acids (major to minor), an unknown fatty acid (major to minor) and trace accessories barbatic, protocetraric, subsquamatic, subbaeomycesic and hypothamnolic acids. Flakus & Kukwa (2007) reported additional accessory traces of unidentified substances that were interpreted as contamination. K+ yellow, C−, Pd+ yellow. Ecology and distribution. On bark, wood and rock; in more or less shaded, mostly humid places. South America (Bolivia – Flakus & Kukwa 2007), Australia. Discussion. Lepraria squamatica does not have very characteristic morphology but is unique in producing squamatic and baeomycesic acids. Species producing protocetraric and/or fumarprotocetraric acids are discussed under L. caesioalba. Species with well-defined lobes are compared under L. membranacea. Lepraria straminea Vain. in Résult. Voyage S. Y. Belgica, Botan.: 40 (1903); type: Antarctica (TUR—syntypes). Thallus crustose-leprose, granular; margin mostly delimited; lobes absent; cortex present on soredia; medulla absent; soredia abundant, coarse, 130–160 µm diam., projecting hyphae absent. For more details see Øvstedal & Lewis-Smith (2001). 49 Lepraria sylvicola Orange in Slavíková-Bayerová & Orange, Lichenologist 38: 507 (2006); type: United Kingdom (NMW—holotype; BG, PRA—isotypes; GenBank accession no. DQ401102). Thallus leprose, powdery; margin diffuse or delimited, lobes absent or obscure and poorly developed; true medulla absent; hypothallus sparse, pale orange-brown; soredia abundant, fine to coarse, 40–160 µm diam., projecting hyphae rarely present, short. For further details see Slavíková-Bayerová & Orange (2006). Chemistry. Atranorin, roccellic/ angardianic acid, toensbergianic acid, an unknown anthraquinone ± (minor, in subthalline hyphae only). K+ yellowish (subthalline hyphae K+ purple-red), C−, Pd− or + yellow (Slavíková-Bayerová & Orange 2006). Ecology and distribution. On neutral to slightly acidic bark, especially oak, sometimes on rock; on more or less sheltered surfaces, especially in mature well-lit woodland. Europe (British Isles, France) (Kukwa & Diederich 2007). Discussion. According to molecular studies, L. sylvicola is a distinct monophyletic taxon (Fehrer et al. 2008). Apart from L. sylvicola, only L. jackii also produces toensbergianic acid. For other similar species, see the discussion under L. jackii. Chemistry. Usnic acid and zeorin. K−, C−, KC−, Pd− (Øvstedal & Lewis Smith 2001). Lepraria texta K. Knudsen, Elix & Lendemer Ecology and distribution. On mosses and peaty soil; in open habitats. Antarctica, endemic. in Nash III, T. H., Gries, C. & Bungartz, F. (eds), Lichen Flora of the Greater Sonoran Desert Region 3: 387 (2008); type: USA (UCR—holotype; ASU, B, CANB, H, PH, UGDA—isotype). Discussion. Lepraria straminea is characterized by a granular, L. neglecta-like thallus without lobes and medulla, and also by the corticate soredia. According to Øvstedal & Lewis-Smith (2001), it only dubiously belongs to Lepraria. Species with usnic acid and zeorin are discussed under L. coriensis. Thallus leprose, powdery; margin diffuse, lobes absent; medulla absent; hypothallus sometimes present, distinct, whitish; soredia abundant, fine, up to 50 µm diam., projecting hyphae present, short, some soredia not well separated from each other. For more details see Knudsen & Elix (2008). 50 THE LICHENOLOGIST Chemistry. Usnic acid, zeorin (minor), atranorin (major to minor) and roccellic/ angardianic acid (minor). K± yellow, C−, KC± yellow, Pd− (Knudsen & Elix 2008). Ecology and distribution. On rock. North America (California, USA). Discussion. Taxa that produce usnic acid and zeorin are discussed under L. coriensis. Lepraria toilenae Kantvilas & Kukwa Muelleria 23: 3 (2006); type: Tasmania (HO—holotype; BG, BM, UGDA—isotypes). Thallus leprose, powdery to cottony; margin diffuse (except young colonies), lobes absent; true medulla absent; hypothallus very well-developed, thick, white or rarely pale greyish; sometimes only sparse soredia on exposed hypothallus; soredia sparse to abundant, fine, 16–40 µm diam., projecting hyphae usually absent; consoredia sometimes present, up to c. 80 µm diam. More details in Kantvilas & Kukwa (2006). Chemistry. Malonprotocetraric, fumarprotocetraric and roccellic acids, protocetraric acid (minor) and confumarprotocetraric acid (trace). K−, C−, KC−, Pd+ red (Kantvilas & Kukwa 2006). Ecology and distribution. On bark, often slightly burnt; in high humidity, old forest, dry bark. Australia (Tasmania). Discussion. The production of the rare substance malonprotocetraric acid in large quantities makes L. toilenae unique. Characteristic morphological features are the fine soredia on thick whitish hypothallus and the lack of lobes. Species producing protocetraric and/or fumarprotocetraric acids are compared under L. caesioalba. Lepraria umbricola Tønsberg Sommerfeltia 14: 206 (1992); type: Norway (BG— holotype). Vol. 41 Thallus leprose, powdery; margin diffuse, lobes absent; medulla very rarely present, thin, white; hypothallus absent; soredia abundant to scattered, fine, up to 60 µm diam., projecting hyphae rarely present, short. For more details see Tønsberg (1992). Chemistry. Thamnolic acid, atranorin ±, roccellic/angardianic acid ± (major to trace) and decarboxythamnolic acid ± (trace). K+ lemon yellow, C−, KC−, Pd+ orange-yellow (Tønsberg 1992; Leuckert et al. 1995; Baruffo et al. 2006 and others). Ecology and distribution. On bark, rock, mosses and soil, mostly acidic substrata; shaded, sheltered, humid, sometimes extremely shaded. Europe, Africa (Macaronesia). Discussion. Lepraria umbricola may be similar to several powdery, unstratified and green-coloured species, for example, L. ecorticata, but is distinguished by producing thamnolic acid. Species producing thamnolic acid are discussed under L. aurescens. Lepraria usnica Sipman Biblioth. Lichenol. 86: 179 (2003); type: Singapore (B— holotype, SINU—isotype). Thallus leprose, powdery; margin diffuse or delimited, lobes sometimes present, mostly less than 0·5 mm wide, usually without a rim; medulla usually present, thin to medium, white to pale yellowish; soredia abundant, mostly medium sized, 70–100 µm diam., projecting hyphae usually absent. For more details see Sipman (2003, 2004). Chemistry. Three chemotypes were distinguished by Elix (2006b): (1) usnic acid, zeorin, contortin (minor), placodiolic acid (trace), hopane-16,22-diol (major to minor) and isousnic acid (minor to trace); (2) usnic acid, zeorin, contortin (minor), placodiolic acid (trace), isousnic acid ± (trace) and roccellic acid ± (trace); (3) usnic acid, zeorin, contortin (minor), placodiolic acid (trace), isousnic acid (minor to trace), atranorin 2009 World survey of Lepraria—Saag et al. 51 (minor) and chloratranorin (minor). In addition, Sipman (2003) reported 5-chloro3–0-methylnorlichexanthone and 5,7dichloro-3–0-methylnorlichexanthone as trace accessories. K−, C± yellow to orange, KC± yellow to orange, Pd−. (e) very rarely unidentified anthraquinones; (f) ‘vouauxii unknown 1’ and very rarely ‘vouauxii unknown 2’ (Tønsberg 1992; Elix & Tønsberg 2004 and others). K− or + faint yellow, C−, KC−, Pd+ reddish orange or Pd− (soredia). Ecology and distribution. On rock, bark and soil; in shaded places. Central and South America, Australia, southern and south-eastern Asia (Singapore, Sri Lanka), southern Africa, tropical. Ecology and distribution. Substratum indifferent, on mosses, bark, rock, soil and lichens; in mostly shaded and sheltered, but also open and exposed places. Worldwide. Discussion. Taxa that produce usnic acid and zeorin are discussed under L. coriensis. In a molecular study Nelsen et al. (2008, in press) showed that L. usnica belongs to the Pilocarpaceae (Lecanorales), thus being distant from Lepraria. Lepraria vouauxii (Hue) R. C. Harris In Egan, Bryologist 90: 163 (1987).—Crocynia vouauxii Hue, Bull. Soc. Bot. France 71: 392 (1924); type: France (BM—lectotype). Thallus leprose, cottony to powdery; margin diffuse to delimited, sometimes obscure lobes present, without raised rims; medulla usually present, usually thick, white; hypothallus sometimes present, brownish, often poorly developed; often medulla exposed between soredia; soredia abundant, mostly coarse, up to 100 µm diam., projecting hyphae often present, short; consoredia often present, up to 300 µm; thallus characters rather variable. More details in Laundon (1989) and Tønsberg (2004). Chemistry. The diagnostic substance is pannaric acid 6-methylester. Accessories include (a) dibenzofurans (mostly minor to trace): oxypannaric acid 6-methylester, 4-oxypannaric acid 6-methylester, 4oxypannaric acid 2-methylester, pannaric acid, pannaric acid 2-methylester, methyl porphyrilate and porphyrilic acid; (b) fatty acids (major to trace): roccellic/angardianic or rarely rangiformic acid; (c) p-depsides: rarely atranorin (major to trace), very rarely gyrophoric or lecanoric acids (mostly minor to trace); (d) terpenoids: very rarely zeorin; Discussion. Phylogenetic placement is mentioned under L. membranacea. Several species may have similar thalli, for example, L. crassissima, L. diffusa, L. eburnea, L. gelida, L. leuckertiana, L. leprolomopsis, L. lobificans, L. nivalis and L. nylanderiana, for differences see under these taxa, especially L. leprolomopsis. Species containing pannaric acid 6-methylester as major substance include L. impossibilis (lecanoric acid, distinct lobes, relatively soft thallus), L. sipmaniana (distinct lobes, relatively soft thallus), L. vouauxii (sometimes obscure lobes, relatively soft thallus) and L. xerophila (distinct lobes and isidia-like structures, soredia absent or few, thallus hard). Lepraria xerophila Tønsberg In Nash, T. H. III, Ryan, B. D., Diederich, P., Gries, C. & Bungartz, F. (eds.), Lichen Flora of the Greater Sonoran Desert Region 2: 326 (2004); type: Mexico (ASU— holotype; BG, DUKE—isotypes). Thallus crustose to subfoliose or squamulose, membranous, slightly granular; margin delimited, obscurely to distinctly lobed, with marginal rim up to 0·5 mm thick; subcorticate patches may be present; medulla usually present, distinct in thick specimens, white; hypothallus absent, lower surface whitish grey; soredia few or absent; numerous large granules or lobules similar to isidia present, compact outer layer present. For more details see Tønsberg (2004). Chemistry. Common chemotype (1): pannaric acid 6-methylester, rangiformic and/or roccellic acid; atranorin (minor to trace), 52 THE LICHENOLOGIST methyl porphyrilate ± (minor to trace), porphyrilic acid ± (trace), pannaric acid ± (trace) and an unknown dibenzofuran (trace); K−, C−, KC−, Pd− or + orange; rare chemotype (2): norascomatic acid plus minor to trace amounts of strepsilin, isostrepsilic acid, atranorin and chloratranorin; K− or + pale yellow, C− or + pale yellow, KC− or + green becoming brown, Pd− (Tønsberg 2004, Elix & Tønsberg 2004). Ecology and distribution. On soil, rarely rock or wood; in dry places. Europe, North America, arid areas. Discussion. Phylogenetic placement and other lobate species are mentioned under L. membranacea. Species containing pannaric acid 6-methylester as major substances are discussed under L. vouauxii. Species producing isidia-like structures are compared under L. isidiata. Lepraria yunnaniana (Hue) Zahlbr. in Handel-Mazzetti, Symbolae Sinicae 3: 244 (1930).— Crocynia yunnaniana Hue, Bull. Soc. Bot. France 71: 396 (1924); type: China (PC−holotype; BM—isotype). Lepraria nigrocincta Diederich, Sérus & Aptroot, in Aptroot, Diederich, Sérusiaux & Sipman, Bibl. Lichenol. 64: 78 (1997). Vol. 41 Thallus leprose, of cottony hypothallus and powdery soredia; margin usually delimited, lobes usually absent, rarely indistinct; medulla sometimes present, white; hypothallus thick, very lax, blackish brown or whitish, of thick hyphae; hypothallus exposed between soredia; soredia scattered, fine, 30–70 µm diam., projecting hyphae sometimes present; consoredia often present, 75–125 µm. For further details see Aptroot et al. (1997) and Laundon (2008). Chemistry. Divaricatic acid and nordivaricatic acid (trace). K−, C− or + pinkish, KC+ pink, Pd− (Aptroot et al. 1997). Ecology and distribution. On bark (also on mosses). South America (Ecuador, Nöske et al. 2007), Asia (China), Africa (Burundi), Australasia (New Guinea), montane. Discussion. Lepraria yunnaniana usually looks different from other Lepraria species because of the very thick and loose, dark hypothallus, which is the dominant part of the thallus, with only a loose cover of scattered soredia. Species producing divaricatic acid are discussed under L. crassissima. Species with a similar dark hypothallus are mentioned under L. cupressicola. Key to the species of Lepraria* 1 Protocetraric acid and/or fumarprotocetraric acid present . . . . . . . . . . . . . 2 Protocetraric acid and/or fumarprotocetraric acid absent . . . . . . . . . . . . . 9 2(1) All soredia aggregated into coarse consoredia (up to 1 mm diam.), surrounded by a conspicuous lax network of branching and anastomosing but mostly not projecting hyphae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lanata Soredia different . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3(2) Thick weft of thick brown hyphae on the lower surface of thallus; soredia fine (30– 60 µm diam.), lobes distinct . . . . . . . . . L. normandinoides chem. 1 & 2 Thick weft of thick brown hyphae absent (however, hypothallus or medulla may be well developed); soredia fine or coarse, lobes absent or present . . . . . . . . 4 4(3) Medulla or hypothallus well developed, thallus soft . . . . . . . . . . . . . . . . 5 Medulla and hypothallus poorly developed or absent, thallus soft or hard . . . . 6 *chem. – chemotype; couplets 68–70 adapted from Slavíková-Bayerová & Orange (2006). 2009 World survey of Lepraria—Saag et al. 53 5(4) Malonprotocetraric acid present, soredia and consoredia fine (up to c. 80 µm diam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. toilenae Malonprotocetraric acid absent, soredia variably sized (up to c. 300 µm diam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. nivalis chem. 1–3 6(4) Thallus powdery, soredia very fine (up to 30 µm, consoredia up to 60 µm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. friabilis Thallus granular or membranous, soredia or thallus granules (similar to isidia) mostly coarse (predominantly >100 µm diam.) . . . . . . . . . . . . . . . . . 7 7(6) Thallus distinctly lobate, with thick raised marginal rim, lobes >0·5 mm wide . . 8 Thallus without lobes or obscure minute lobes present without thick raised rim . . . . . . . . . . . . . . . . . . . . . . . . L. caesioalba var. caesioalba chem. 1 8(7) Structures similar to true isidia with compact outer layer present, projecting hyphae absent; granules mostly 320–540 µm diam. . . . . . . . . . . . . . .L. isidiata Structures slightly similar to isidia may be present but compact outer layer absent, numerous projecting hyphae present, soredia mostly 220–340 µm diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. santosii chem. 4–7 9(1) Stictic acid complex present . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Stictic acid complex absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 10(9) Terpenoids present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Terpenoids absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 11(10) Thallus distinctly lobate, with raised marginal rim, lobes over 0·5 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. santosii chem. 1 Thallus without lobes or lobes poorly developed, without marginal rim . . . . 12 12(11) More than one terpenoid and minor amounts of salazinic acid present; soredia coarse (up to 200 µm diam.), with long projecting hyphae . . . . . . . L. multiacida Mostly only one terpenoid present, salazinic acid absent; soredia not coarse or if coarse then projecting hyphae short or absent . . . . . . . . . . . . . . . . 13 13(12) Medulla distinct, conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Medulla absent or inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . 15 14(13) Soredia with long projecting hyphae, loosely packed and soft, mostly fine to medium sized (30–100 µm diam.); thallus greenish; zeorin present . . . . L. lobificans Projecting hyphae short or absent, soredia harder, medium sized to coarse (75–300 µm diam.); thallus yellowish; an unknown terpenoid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. leprolomopsis 15(13) Soredia are mostly fine (20–45 µm diam.), consoredia are rare, thallus margin diffuse; thallus greenish . . . . . . . . . . . . . . . . . . . . . . . .L. elobata Soredia are coarse (100–200 µm diam.), consoredia abundant (up to 300 µm diam.), thallus margin usually delimited; thallus grey or bluish grey, not greenish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. caesioalba var. groenlandica 54 THE LICHENOLOGIST Vol. 41 16(10) Well-developed medulla present, thallus with membranous appearance, lobes usually distinct or rarely absent . . . . . . . . . . . . . . . . L. nivalis chem. 4–5 Well-developed medulla absent, thallus with granular or membranous appearance, lobes present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 17(16) Thallus distinctly lobate, with thick raised marginal rim, lobes over 0·5 mm wide; projecting hyphae from soredia present and mostly long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. santosii chem. 2 & 3 Thallus without lobes or obscure minute lobes present without thick raised rim; projecting hyphae from soredia mostly short or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. caesioalba var. caesioalba chem. 2 18(9) Alectorialic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Alectorialic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 19(18) Porphyrilic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . L. gelida Porphyrilic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 20(19) Thallus soft, soredia loosely packed, medulla distinct and usually thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. eburnea Thallus hard, granular, soredia densely packed, medulla inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. neglecta 21(18) Divaricatic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Divaricatic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 22(21) Nordivaricatic acid present as major substance, medulla well-developed, lower surface distinct . . . . . . . . . . . . . . . . . . . . . . . . . . L. crassissima Nordivaricatic acid absent or in trace amounts, medulla absent or less developed, lower surface not distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23(22) Thick lax hypothallus, usually brown, soredia sparsely distributed on hypothallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. yunnaniana Hypothallus absent, soredia abundant . . . . . . . . . . . . . . . . . . L. incana 24(21) Pannaric acid or one of pannaric acid derivatives present as main substance . . 25 Pannaric acid and its derivatives absent or present in trace to minor amounts besides other major compound(s) . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 25(24) 4-oxypannaric acid 2-methylester present as only major dibenzofuran . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. diffusa 4-oxypannaric acid 2-methylester absent or present in trace to minor amounts besides other major dibenzofurans . . . . . . . . . . . . . . . . . . . . . . . 26 26(25) Pannaric acid present as only major dibenzofuran . . . . . . . L. membranacea Pannaric acid absent or present in trace to minor amounts besides other major dibenzofurans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 27(26) Lecanoric acid present . . . . . . . . . . . . . . . . . . . . . . . L. impossibilis Lecanoric acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28(27) Soredia absent or few, isidia-like structures present, lobes well-developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. xerophila chem. 1 Soredia numerous, isidia-like structures absent, lobes well or poorly developed or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 2009 World survey of Lepraria—Saag et al. 55 29(28) Lobes well-developed, with raised marginal rim . . . . . . . . . . L. sipmaniana Lobes absent or poorly-developed, without marginal rim . . . . . . . L. vouauxii 30(24) Porphyrilic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 Porphyrilic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 31(30) Thallus granular, hard, soredia coarse (100–300 µm diam.) . . . . . . . . . . . 32 Thallus relatively soft, powdery (or rarely cottony), soredia medium to very fine (20–100, rarely some up to 200 µm diam.) . . . . . . . . . . . . . . . . . . 33 32(31) Zeorin present . . . . . . . . . . . . . . . . . . . . . . .L. alpina var. zeorinica Zeorin absent . . . . . . . . . . . . . . . . . . . . . . . . L. alpina var. alpina 33(31) Atranorin present, one or very rarely two fatty acids present . . . . . L. atlantica Atranorin absent, two fatty acids present . . . . . . . . . . . . . . . .L. neojackii 34(30) Lecanoric acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Lecanoric acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 35(34) Zeorin present, dark brown hypothallus present (usually thick, rarely thin) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. cupressicola Zeorin absent, hypothallus absent or present and white or orange in patches . 36 36(35) Atranorin present, soredia coarse (up to 200 µm diam.) . . . . . . . L. lecanorica Atranorin absent, soredia fine (up to c. 50 µm diam.) . . . . . . . . . . . . . . 37 37(36) Thallus margin delimited, hypothallus absent, soredia very fine (up to c. 30 µm diam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. goughensis Thallus margin diffuse, hypothallus usually present, soredia fine (up to c. 50 µm diam.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. achariana 38(34) Thamnolic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Thamnolic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 39(38) Thallus unstratified – medulla and hypothallus absent, colour green to rarely whitish green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. umbricola Thallus stratified – medulla or hypothallus present, colour not green . . . . . . 40 40(39) Roccellic acid present; thallus thick, usually with well-developed greyish hypothallus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. nylanderiana Roccellic acid absent; thallus thin, hypothallus absent or present and brown . . 41 41(40) Thallus margin diffuse without raised rim; brown hypothallus present, usually welldeveloped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. aurescens Thallus margin delimited, with raised rim; hypothallus absent or poorly-developed, white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. pulchra 42(38) Usnic acid present together with zeorin . . . . . . . . . . . . . . . . . . . . . 43 Usnic acid absent, zeorin absent or present . . . . . . . . . . . . . . . . . . . 48 43(42) Thallus margin distinctly lobed, lobes with raised rim, predominantly over 0·5 mm wide; protodehydroconstipatic and constipatic acids present (in major to trace amounts) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. coriensis Thallus margin diffuse or lobed; if lobes present, without the rim, less than 0·5 mm wide; protodehydroconstipatic and constipatic acids absent . . . . . . . . . 44 56 THE LICHENOLOGIST Vol. 41 44(43) Medulla thick and cottony . . . . . . . . . . . . . . . . . . . . .L. leuckertiana Medulla absent or thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 45(44) Lobes present, obscure to distinct, small, less than 0·5 mm wide . . . L. usnica Lobes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 46(45) Thallus granules coarse (up to 160 µm diam.), corticate . . . . . . L. straminea Cortical granules absent, soredia smaller (up to 100 µm diam.) . . . . . . . . . 47 47(46) Soredia regular, spherical, well separated from each other; hypothallus absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. ecorticata Soredia irregularly shaped, not well separated from each other in places; whitish hypothallus may be present . . . . . . . . . . . . . . . . . . . . . . . L. texta 48(42) Squamatic acid present . . . . . . . . . . . . . . . . . . . . . . . L. squamatica Squamatic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 49(48) Argopsin present . . . . . . . . . . . . . . . . . . . . . . . . . L. santamonicae Argopsin absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50 50(49) Pannarin present . . . . . . . . . . . . . . . . . . . . . . . . . . . L. adhaerens Pannarin absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 51(50) Psoromic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52 Psoromic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53 52(51) Thallus granular, hard, medulla inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. caesioalba var. caesioalba chem. 3 Thallus softer, with well-developed medulla . . . . . . . . . . L. nivalis chem. 6 53(51) Salazinic acid present . . . . . . . . . . . . . . . . . . . . . . . . . L. salazinica Salazinic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 54(53) Obtusatic acid present . . . . . . . . . . . . . . . . . . . . . . . . L. obtusatica Obtusatic acid absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 55(54) Norascomatic acid present; soredia absent or few, isidia-like structures (lobules) present, lobes well-developed . . . . . . . . . . . . . . .L. xerophila chem. 2 Norascomatic acid absent; thallus different . . . . . . . . . . . . . . . . . . . 56 56(55) Distinct well-developed marginal lobes present, over 0·5 mm wide and mostly with marginal rim . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 Marginal lobes absent or obscure . . . . . . . . . . . . . . . . . . . . . . . . . 60 57(56) Zeorin present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58 Zeorin absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 58(57) Hypothallus present, grey to black; unidentified fatty acids present . . . L. pallida Hypothallus absent (medulla present); rangiformic/jackinic or roccellic/angardianic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lobata 59(57) Anthraquinones present (mostly in subthalline hyphae) . . . . . . . L. bergensis Anthraquinones absent . . . . . . . . . . . . . . . L. normandinoides chem. 3 60(56) Zeorin present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 Zeorin absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 2009 World survey of Lepraria—Saag et al. 57 61(60) Fatty acids present; medulla or hypothallus present . . . . . . . . . . . . . . . 62 Fatty acids absent; medulla and hypothallus absent . . . . . . . . . . L. caesiella 62(61) Thallus relatively hard, soredia densely packed, consoredia up to 350 µm diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. lobata Thallus soft, soredia loosely packed, consoredia up to 160 µm diam. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. jackii s. lat 63(60) Nephrosteranic acid present; soredia with very long projecting hyphae (60–120 µm), very loosely packed . . . . . . . . . . . . . . . . . . . . . . . . . . L. rigidula Nephrosteranic acid absent; projecting hyphae absent or shorter, soredia densely or loosely packed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 64(63) Soredia up to 300 µm diam., thallus granular (L. neglecta type) . . . . . . . . . 65 Soredia up to 160 µm diam., thallus powdery or slightly cottony . . . . . . . . 66 65(64) Fatty acid ‘granulata unknown 1’ present, rangiformic/jackinic and roccellic/ angardianic acids absent . . . . . . . . . . . . . . . . . . . . . . L. granulata ‘Granulata unknown 1’ absent, rangiformic/jackinic or roccellic/angardianic acid present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. borealis 66(64) Jackinic/rangiformic acid present as the only major fatty acid . . . . . . . . . . 67 Other fatty acids present as major compounds (jackinic/rangiformic acid also present in one species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 67(66) Anthraquinones present on subthalline hyphae (local and often inconspicuous) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. humida Anthraquinones absent . . . . . . . . . . . . . . . . . . . . . . . L. jackii s. str. 68(66) Roccellic/angardianic acid present as the only major fatty acid . . . . . L. celata Toensbergianic or jackinic/rangiformic acid present as major fatty acids (roccellic acid may be present in addition) . . . . . . . . . . . . . . . . . . . . . . . . 69 69(68) Toensbergianic acid present as the only major fatty acid . . . .L. toensbergiana Other fatty acids present as major substances (toensbergianic acid present in combination with roccellic/angardianic acid) . . . . . . . . . . . . . . . . . . . 70 70(69) Jackinic/rangiformic acid present as major fatty acid, present) lacking anthraquinones . . . . . . . . . . . Toensbergianic and roccellic/angardianic acids present thalline hyphae (when present) with anthraquinones The authors are grateful to the curators of the herbaria who made their specimens available for us; to H. Sipman, L. Zedda, A. Orange and S. Ekman for valuable comments on the manuscript, to M. Tretiach, K. Knudsen and M. Nelsen for insightful discussions on some species, and to the reviewers of the manuscript. Financial support was received from the Estonian Science Foundation (grant no. 5823 to Andres Saag, and nos 5505 and 7470 to Tiina Randlane); from the Estonian Ministry of Education and Research (targeted financing no. 0153 to Urmas Kõljalg; grant to Andres Saag within national research and development programme ‘‘Collections of Humanities and Natural Sciences’’), and from the European Union through the subthalline hyphae (when . . . . . . L. jackii s. str. as major compounds, sub. . . . . . . . L. sylvicola European Regional Development Fund (Centre of Excellence FIBIR). REFERENCES Acharius, E. (1803) Methodus qua omnes detectos lichenes secundum organa carpomorpha ad genera, species et varietates redigere atque observationibus illustrare tentavit Erik Acharius (Methodus Lichenum). Cum tab. aen. Stockholm. Aptroot, A. (2002) New and interesting lichens and lichenicolous fungi in Brazil. 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M. & Werner, O. (2004) Guía de Campo de los Líquenes, Musgos y Hepdticas. Barcelona: Ediciones Omega. 60 THE LICHENOLOGIST Zedda, L. (2000a) Lecanora leuckertiana sp. nov. (lichenized Ascomycetes, Lecanorales) from Italy, Greece, Morocco and Spain. Nova Hedwigia 71: 107–112. Vol. 41 Zedda, L. (2000b) The lichen genera Lepraria and Leproloma in Sardinia (Italy). Cryptogamie, Mycologie 21: 249–267. Accepted for publication 15 October 2008

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