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Table 1 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Table 1. Strictly lichenicolous species of Arthrorhaphis with their host lichens and references.<table><tbody><tr><th>Taxon</th><th>Hosts</th><th>References</th></tr></tbody><tbody><tr><th>Arthrorhaphis aeruginosa</th><td>Cladonia spp. squamules, rarely podetia</td><td>Santesson & Tønsberg, 1994</td></tr><tr><th>Arthrorhaphis arctoparmeliae</th><td>Arctoparmelia incurva</td><td>Kocourková & Van den Boom, 2005</td></tr><tr><th>Arthrorhaphis grisea</th><td>Baeomyces rufus, B. placophyllus</td><td>Obermayer, 1994</td></tr><tr><th>Arthrorhaphis muddii</th><td>Dibaeis baeomyces</td><td>Obermayer, 1994</td></tr><tr><th>Arthrorhaphis olivaceae</th><td>Melanohalea olivacea</td><td>Santesson & Tønsberg, 1994</td></tr><tr><th>Arthrorhaphis phyllobaeis</th><td>Phyllobaeis imbricata</td><td>Etayo, 2017</td></tr></tbody></table>

Fig. 6. A in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 6. A, Arthrorhaphis aeruginosa (TØnsberg 19019, BG, holotype); B, A. arctoparmeliae (Kocourková & Kocourek JK5484 dpl., C); C, A. olivaceae (Santesson 11677, BG, isotype); D, A. muddii (Woods s.n., E, holotype); E, A. grisea (Th.M. Fries s.n., UPS, holotype); F, A. vulgaris (Norrlin s.n., H). — Scale: A–C, 0.5 mm; D–F, 1 mm. Photos: A. Frisch.

Fig. 5 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 5. Ascospore types distinguished in lichenised Arthrorhaphis (schematic redrawing based on illustrations in Obermayer, 1994, 1995; compare the cited literature for further details). A, Citrinella type; B, Alpina type; C, Vacillans type; D, Jungens type. — Ascospores of the citrinella type are arranged parallel, 1-seriate, while those of the alpina, vacillans, and jungens types are stacked in the asci. Ascospores of the variable jungens type are intermediary between the alpina and vacillans types, and either resembling the vacillans type but longer, or similar in size but 4–5(–7)-septate. — Scale: A–D, 10 μm.

Fig. 1. Bayesian 50 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 1. Bayesian 50% majority-rule consensus tree from analysis of MSA-1, showing the placement of Arthrorhaphis as sister to Ostropales, Ostropomycetidae. Branches supported by BPP ≥ 0.95 and ML BS ≥ 70% are indicated by bold black lines; branches supported only by BPP ≥ 0.95 are indicated by bold grey lines. Numbers in brackets represent clades discussed in the text.

Fig. 2. Bayesian 50 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 2. Bayesian 50% majority-rule consensus tree from analysis of MSA-2, showing the basal position of the exclusively parasitic species in Arthrorhaphis. The evolution of lichenised thalli containing pulvinic acid derivatives in the A. alpina- and the A. citrinella s.l. clades is indicated by an asterisk. Branches supported by BPP ≥ 0.95 and ML BS ≥ 70% are indicated by bold black lines; branches supported only by BPP ≥ 0.95 are indicated by bold grey lines. Numbers in brackets represent clades discussed in the text. Graphical representation of species delimitations in bGMYC, bPtP and bP&P: Colours represent delimited species for each species delimitation analysis independently, but have been selected to highlight delimitations congruent across analyses. White represents missing data. The colouring scheme applies only to the current figure.

Fig. 4 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 4. Partial representation of the Bayesian 50% majority-rule consensus tree from analysis of MSA-2, showing the Arthrorhaphis citrinella s.l. clade. Branches supported by BPP ≥ 0.95 and ML BS ≥ 70% are indicated by bold black lines; branches supported only by ML BS ≥ 70% are indicated by thin double lines. Numbers in brackets represent clades discussed in the text. Character states: 1 Life form: juvenile parasitism absent (light green), present (reddish brown), missing data (white). 2 Thallus areolae: present (yellow), absent (white); 3 Soredia: citrinella type (turquoise), farinosa type (light green), vulgaris type (dark green), absent (white); 4 Medulla: pale yellow (yellow), cavity (grey), absent (white); 5 Ca-oxalate crystals: absent (white); 6 Ascospores: citrinella type (blue), absent (white). Graphical representation of species delimitations in bGMYC, bPtP, and bP&P: Colours represent delimited species for each species delimitation analysis independently, but have been selected to highlight delimitations congruent across analyses. White represents missing data. The colouring scheme applies only to the current figure.

Fig. 8. A in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Fig. 8. A, Arthrorhaphis alpina (Schaer., Lichenes Helvetici Exsiccati 532, G, lectotype); B, A. alpina var. jungens (Lichenotheca Graecensis 23, E, isotype); C, A. alpina (Ohmura 10119, TNS); D, A. alpina var. jungens (Kalb, Lichenes Neotropici 577, M); E, A. alpina (Brusse 4515, UPS); F, "A. septentrionalis" (Hansen 026, C). — Scale: A–F, 1 mm. Photos: A. Frisch.

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Table 2 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Table 2. Results from the bGMYC analyses for mrSSU, nrITS, and RPB1 gene loci.<table><tbody><tr><th></th><th>mrSSU</th><th>nrITS</th><th>RPB1</th></tr></tbody><tbody><tr><th>Likelihood null model</th><td>1231.294</td><td>1199.214</td><td>1160.623</td></tr><tr><th>Maximum likelihood GMYC model</th><td>1264.052</td><td>1230.614</td><td>1180.092</td></tr><tr><th>Likelihood ratio</th><td>65.51679</td><td>62.80141</td><td>38.93693</td></tr><tr><th>Result of likelihood ratio test</th><td>5.884182e-15***</td><td>2.309264e-14***</td><td>3.507144e-09***</td></tr><tr><th>Number of maximum likelihood clusters with confidence interval in brackets</th><td>10 (9–10)</td><td>15 (15–15)</td><td>12 (12–20)</td></tr><tr><th>Number of maximum likelihood entities with confidence interval in brackets</th><td>15 (14–15)</td><td>19 (19–20)</td><td>15 (15–30)</td></tr><tr><th>Threshold time</th><td>–0.002228722</td><td>–0.007533381</td><td>–0.002175746</td></tr></tbody></table>

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Table 3 in A phylogenetic survey of the ascomycete genus Arthrorhaphis (Arthrorhaphidaceae, Lecanoromycetes) including new species in Arthrorhaphis citrinella sensu lato

Table 3. Summary of species delimitations from the bGMYC, bPtP, and bP&P analyses.<table><tbody><tr><th></th><th>mrSSU</th><th>bGMYC nrITS</th><th>RPB1</th><th>bPtP</th><th>15</th><th>bP&P</th><th>22</th></tr></tbody><tbody><tr><th>A. aeruginosa</th><td>1</td><td>1</td><td>–</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. alpina s.l. (incl. var. jungens)</th><td>4</td><td>3</td><td>5</td><td>1</td><td>1</td><td></td><td>8</td></tr><tr><th>A. arctoparmeliae</th><td>1</td><td>1</td><td>–</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. bullata</th><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. catolechioides</th><td>–</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. citrinella</th><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. farinosa</th><td>1</td><td>3</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. grisea</th><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. muddii</th><td>1</td><td>1</td><td>1</td><td>2</td><td>1</td><td></td><td>1</td></tr><tr><th>A. olivaceae</th><td>–</td><td>1</td><td>–</td><td>2</td><td>1</td><td></td><td>1</td></tr><tr><th>“ A. septentrionalis ”</th><td>0</td><td>1</td><td>1</td><td>2</td><td>1</td><td></td><td>1</td></tr><tr><th>A. sp. 1</th><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. vulgaris</th><td>1</td><td>1</td><td>1</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>A. vacillans</th><td>1</td><td>1</td><td>1</td><td>0</td><td>1</td><td></td><td>1</td></tr><tr><th>Anzina carneonivea (outgroup)</th><td>1</td><td>1</td><td>–</td><td>1</td><td>1</td><td></td><td>1</td></tr><tr><th>Sum</th><td>15</td><td>19</td><td>15</td><td>17</td><td>15</td><td></td><td>22</td></tr></tbody></table>

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Systemic infection of Bryoria (Lecanoromycetes, Ascomycota) by Athelia (Agaricomycetes, Basidiomycota) in western North America

Bryoria (Parmeliaceae, Ascomycota) is one of the dominant genera of hair lichens in western North America and is characteristic of high-elevation conifer forest ecosystems. In areas where Bryoria is abundant, it is common to find thalli in which the thalline filaments become conglutinated, forming brittle dead zones. After sampling Bryoria thalli across western Canada and the northwestern United States at different times of the year, we found this dieback phenomenon is associated with the winter growth of a mold-forming basidiomycete. We report that this fungus belongs to Athelia (Atheliaceae, Basidiomycota) a genus known to contain lichen pathogens, most notably A. arachnoidea. By sequencing a combination of genetic markers – nuc rDNA ITS1-5.8S-ITS2 (ITS), partial nuc 28S rDNA (28S), and partial translation elongation factor 1-α (TEF1) – paired with morphometric analyses, we reveal the involvement of at least three additional lineages of lichen-associated Athelia and describe one as a new species, Athelia abscondita. A. abscondita is morphologically distinguished from other Athelia species by its basidia and basidiospores, was found to frequently infect members of Bryoria sect. Implexae, and was occasionally on other foliose and fruticose species within Parmeliaceae.

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Systemic infection of Bryoria (Lecanoromycetes, Ascomycota) by Athelia (Agaricomycetes, Basidiomycota) in western North America

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FIGURE 2. Ochrolechia incarnata. A in Ochrolechia incarnata comb. nov. (Lecanoromycetes, Ascomycota), a distinct species of the O. parella group from Europe and Macaronesia

FIGURE 2. Ochrolechia incarnata. A, holotype of Pertusaria incarnata; B, close up of the thallus of the holotype; C–D, morphology of apothecia and thallus (Ertz 10572). Scale bars: C = 1 mm, D = 500 μm.

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FIGURE. 1. Phylogenetic relationships among 25 in Ochrolechia incarnata comb. nov. (Lecanoromycetes, Ascomycota), a distinct species of the O. parella group from Europe and Macaronesia

FIGURE. 1. Phylogenetic relationships among 25 samples of Ochrolechia inferred from Bayesian analysis of nuITS sequences. Posterior probabilities are shown above internal branches, and Maximum Likelihood bootstrap values obtained from a RAxML analysis are shown below internal branches. Internal branches strongly supported in both analyses, are represented by thicker lines. The clade of O. incarnata, a species resurrected here, is highlighted. Corticolous samples of O. parella are marked with an asterisk (*).

FIGURE 20 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 20. Species of Vainionora. A–B. Thallus variation of Vainionora aemulans. A. Granular-leproid thallus growing on smooth bark of Zanthoxylon (Aptroot, A. 65060, CDS 31642). B. Granular-leproid thallus growing on coarse basalt lava (Aptroot, A. 65170, CDS 31754). C–D. Vainionora nugrae (Nugra, F. 279, CDS 33195–holotype). C. Thallus coarsely granular, pale yellowish green, with crenate lecanorine apothecia with blackened disc covered in a fine yellowish green pruina. D. Apothecial section (in water, DIC): exciple with large crystals only; epihymenium yellowish brown, lacking crystals; subhymenium well developed, distinctly yellowish brown. Scales: A, B & C = 5 mm; D =100 µm.

FIGURE 18 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 18. Species in the L. varia-group. A–E. Lecanora strobilina. A–B. Thallus variation. A. thick rimose-areolate, pale yellow thallus with adnate apothecia, farinose soredia developing irregularly from the thallus surface and along the apothecial margin, discs epruinose, golden yellow (Nugra, F. 124, CDS 32778). B. scant rimose thallus on a black hypothallus, farinose soredia forming irregular from the surface and apothecial margin, discs epruinose, golden yellow (Bungartz, F. 7859, CDS 38368). C–E. Apothecial section (Nugra, F. 124, CDS 32778). C. Exciple abundantly filled with minute crystals and yellowish brown pigment, pigment and crystals extending across epihymenium, hypothecium hyaline (in water, DIC). D. After treatment with 10% KOH, crystals and pigments dissolved, trebouxioid photobionts more easily distinguished (in KOH, DIC). E. Characteristically small, narrowly oblong ascospores (in KOH, DIC). F. Leproid thallus of Lecanora terpenoidea, sorediate granules embedded among cottony thallus hyphae, with prolonged herbarium storage forming abundant terpenoid crystals that give the thallus a conspicuous 'fluffy' appearance (Aptroot, A. 65410, CDS 31996–holotype). Scales: A, B & F= 5 mm; C & D =100 µm; E = 10µm.

FIGURE 17 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 17. Species in the L. varia-group. A–E. Lecanora pyrrhosporoides. A–B. Thallus variation on different substrates. A. Thick, granular leproid thallus with ±immersed to sessile apothecia, disc brown to darkened, coarsely pruinose, lecanorine margin either dissolved into blastidiate granules or excluded below; on decorticated branches of Prosopis (Aptroot, A. 64140, CDS 30703). B. Thallus of scattered clusters of coarse coralline blastidiate granules, with scarce apothecia; on smooth bark of Bursera (Bungartz, F. 5359, CDS 28443). C–D. Apothecial sections (in water, DIC). C. Exciple appearing biatorine, almost completely reduced to a layer of pale brown cells extending from the epihymenium to the outer exciple; lecanorine part not developed (Aptroot, A. 64140, CDS 30703). D. Exciple ±lecanorine, with a thin, inner, proper exciple, surrounded by a thick, disintegrating, outer, thalline part (Aptroot, A. 64140, CDS 30703). E. Filiform conidia (Aptroot, A. 64140, CDS 30703). F–G. Lecanora subaureoides (Aptroot, A. 65158, CDS 31741–holotype). F. Dispersed thallus squamules with marginal soredia extruding granular soredia, apothecia malformed, not clearly differentiated from thallus squamules. G. Section of poorly developed lecanorine apothecium lacking asci or ascospores. Scales: A, B & F = 5 mm; C, D & G = 100 µm; E = 20 µm.

FIGURE 14 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 14. Sorediate species in the L. subfusca-group; epihymenium at least in part aeruginose. A–C. Lecanora floridula A–B. Morphological variation. A. Thallus pale greenish gray, rimose-areolate, with pustulate yellow green soralia and broadened apothecia with deep brown disc (Bungartz, A. 8616, CDS 41262). B. Thallus rimose, yellow green, with pustulate yellow green soralia, apothecia indistinct, disc barely expanded, dark brown (Bungartz, F. 6474, CDS 34691). C. Apothecial section (in water, DIC): exciple with large crystals (pulicaris-type), epihymenium mostly brown, with few crystals (chlarotera-type), but in parts also aeruginose (Aptroot, A. 64915, CDS 31494). D–G. Lecanora malagae (Bungartz, F. 10352, CDS 52326–holotype). D. General thallus aspect, composed of inflated, closely aggregated, pale gray areoles with pustulate pale yellowish green soralia, apothecia with a thick margin, disc bluish blackened, faintly pruinose. E. Close-up of the thallus. F. Lecanora-type ascus, tholus with wide, apically open, unstained central cone and deeply stained ascus flanks (in Lugol's, DIC). G. Apothecial section (in water, DIC): exciple with minute crystals only (allophana-type), epihymenium aeruginose, lacking crystals (gangaleoides-type). Scales: A, D, B & E = 5 mm; F = 10 µm; C & G = 100 µm.

FIGURE 13 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 13. Species in the L. subfusca-group; superficially similar species with pale to dark brown apothecial disc and melacarpella- or allophana-type exciple (coarse and fine or fine crystals only). A–B. Lecanora darwiniana (Bungartz, F. 4859, CDS 29055–holotype). A. Thallus rimose, with abundant, coarsely granular soredia all across its surface and along the apothecial margin, apothecial disc deep brown, epruinose. B. Apothecial section: exciple with both coarse and fine crystals (melacarpella-type), epihymenium lacking crystals, brownish pigment insoluble in K (glabrata-type), exciple breaking apart into granular soredia (in water, DIC). C–D. Lecanora cactacea (Bungartz, F. 8178, CDS 40824–holotype). C. Thallus areolate verrucose, growing on the flaky surface of cactus pads, apothecia large, distinctly sessile, margin soon conspicuously fissured, disc pale to deep brown, with age occasionally blackened. D. Apothecial section (in water, DIC): exciple evenly yellow brown throughout, filled with abundant, minute crystals that completely dissolve in KOH (allophana- type), epihymenium yellowish brown, not dissolving in KOH, lacking crystals (glabrata-type). E–F. Lecanora subcrenulata (Bungartz, F. 7617, CDS 38113). E. Thallus saxicolous, areolate, with large, distinctly sessile apothecia, margin moderately to strongly undulate with age, disc pale orange-brown, epruinose. G. Apothecial section (in water, DIC): exciple with both coarse and fine crystals (melacarpella- type), epihymenium with minute crystals and brown pigment (chlarotera-type). Scales: A, C, & E = 5 mm; B, D & F = 100 µm.

FIGURE 12 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 12. Species in the L. subfusca-group; superficially similar species with deep brown apothecial disc and pulicaris-type exciple (large crystals only). A–B. Lecanora schindleri (Weber, W. A. s.n., L-40337, COLO 188861). A. Thallus continuous to barely rimose with fimbriate whitish prothallus, apothecia adnate to sessile, margin entire, disc deep reddish brown to almost blackish brown, epruinose or with very faint, whitish pruina. B. Apothecial section (in water, DIC), exciple with abundant minute, soluble crystals obscuring few large, insoluble ones (melacarpella-type), epihymenium with reddish brown, insoluble pigment and minute soluble crystals (pulicaris-type). C– D. Lecanora tropica. C. Thallus rimose-areolate, slightly verrucose, delimited by a darkened prothallus line, apothecia adnate to sessile, margin entire to ±crenulate, disc deep reddish brown, epruinose (Bungartz, F. 8425, CDS 41071). D. Apothecial section (in water, DIC): exciple with conspicuous, large, insoluble crystals and few minute, soluble ones (pulicaris-type), epihymenium with insoluble brown pigment, lacking crystals (glabrata-type) (Dawson, E.Y. s.n., L-40089, COLO 188062). E–F. L. subimmergens (Bungartz, F. 7600, CDS 38096). E. Thallus areolate, apothecia adnate to sessile, margin entire, slightly undulate, disc dark reddish to blackish brown, epruinose. F. Apothecial section (in water, DIC): exciple with conspicuous, large, insoluble crystals and few minute, soluble ones (pulicaris-type), epihymenium with insoluble brown pigment, lacking crystals (glabrata-type). Scales: A, C & E = 5 mm; B, D & F = 100 µm.

FIGURE 11 in Lecanoroid lichens in the Galapagos Islands: the genera Lecanora, Protoparmeliopsis, and Vainionora (Lecanoraceae, Lecanoromycetes)

FIGURE 11. Species in the L. subfusca-group; superficially similar species with brown apothecial disc and pulicaris-type exciple (large crystals only). A–C. Ecological thallus variation of Lecanora sulfurescens. A. Scantly developed, thin thallus with net-like areoles, apothecia with pale greenish yellow disc; developed below shaded overhang on coarse lava (Bungartz, F. 4386, CDS 28471). B. Thallus of minute, scattered, isolated, granular areoles, apothecial disc chestnut brown; growing on strongly weathered rock with ±disintegrating mineral surface (Bungartz, F. 6416, CDS 34631). C. Smooth, contiguous areolate thallus delimited by blackened prothallus line, apothecial disc chestnut brown; growing on smooth basalt surface (Bungartz, F. 4822, CDS 28988). D.–F. Lecanora galactiniza. D. Exuberant, thick areolate thallus with marginal areoles becoming subsquamulose, apothecial disc chestnut brown; growing at a nutrient-rich site (Bungartz, F. 4706, CDS 28804). E. Thallus of discrete, dispersed squamules, apothecial disc dark brown; growing on coarse, irregular lava (Bungartz, F. 5207, CDS 29420). F. Section of apothecium with pulicaris-type exciple (with large crystals only) and pulicaris-type epihymenium (deep brown pigment not dissolving, but minute crystals dissolving in KOH; in KOH, DIC). Scales: A–E = 5 mm; F = 100 µm.