Phylogeny, evolution and a re-classification of the Lichinomycetes

Abstract

The Lichinomycetes is an independent lichenized lineage within the Ascomycota comprising ca. 390 species and 50 genera. Very few studies have dealt with family and genus classification using molecular data and many groups are in need of thorough revision. Thus, we constructed a multilocus phylogeny (mtSSU, RPB2 and mcm7 gene regions) including 190 specimens of Lichinomycetes belonging to 126 species. Ancestral state reconstruction analyses were carried out to trace the evolution of selected characters. The current classification scheme of the Lichinomycetes based on morphological and anatomical characters is in great conflict with the phylogenetic relationships resulting from the present study. The results suggest substantial non-monophyly at the family and genus levels. A revised classification is proposed here and an overview of genera accepted in the Lichinomycetes is given. Ancestral Lichinomycetes are reconstructed as crustose with pycnoascocarps and octosporous asci. We used a combination of characters to delineate groups including the ascoma development and the type of asci. The revised classification includes 11 new genera, five resurrected genera, and 54 new combinations distributed in four families (three emended and one new). Three new species are also described. Taxonomic novelties: New family: Lichinellaceae M. Schultz & M. Prieto. New genera: Allopyrenis M. Schultz & M. Prieto, Gonotichia M. Schultz & M. Prieto, Lapismalleus M. Schultz & M. Prieto, Lingolemma M. Schultz & M. Prieto, Paludolemma M. Schultz & M. Prieto, Paracyphus M. Schultz & M. Prieto, Peltolemma M. Schultz & M. Prieto, Pseudocarpon M. Schultz & M. Prieto, Pseudotichia M. Schultz & M. Prieto, Pycnolemma M. Schultz & M. Prieto, Tichocyphus M. Schultz & M. Prieto. New species: Paracyphus gotlandicus M. Schultz & M. Prieto, Pseudocarpon persimile M. Schultz & M. Prieto, Tichocyphus gotlandicus M. Schultz & M. Prieto. New combinations: Allopyrenis grumulifera (Nyl.) M. Schultz & M. Prieto, Allopyrenis haemaleella (Nyl.) M. Schultz & M. Prieto, Allopyrenis impolita (Th. Fr.) M. Schultz & M. Prieto, Allopyrenis phaeococca (Tuck.) M. Schultz & M. Prieto, Allopyrenis reducta (Th. Fr.) M. Schultz & M. Prieto, Allopyrenis sanguinea (Anzi) M. Schultz & M. Prieto, Allopyrenis tenuis (Henssen) M. Schultz & M. Prieto, Cladopsis densisidiata (Aptroot et al.) M. Schultz & M. Prieto, Cladopsis foederata (Nyl.) M. Schultz & M. Prieto, Cladopsis guyanensis (M. Schultz et al.) M. Schultz & M. Prieto, Cladopsis palmana (J. Steiner) M. Schultz & M. Prieto, Cladopsis polycocca (Nyl.) M. Schultz & M. Prieto, Forssellia canariensis (Henssen) M. Schultz & M. Prieto, Forssellia concordatula (Nyl.) M. Schultz & M. Prieto, Gonotichia octosporella (Lettau) M. Schultz & M. Prieto, Lapismalleus lugubris (A. Massal.) M. Schultz & M. Prieto, Lemmopsis lutophila (Arnold) M. Schultz & M. Prieto, Lempholemma segregatum (Nyl.) M. Schultz & M. Prieto, Lichinella baicalensis (Makryi) M. Schultz, Lichinella etoshica (Brusse) M. Schultz, Lichinella lusitanica (Henssen) M. Schultz, Lichinella pulvinata (E. Dahl) M. Schultz, Lichinella schleicheri (Hepp) M. Schultz, Lichinella terrestris (Makryi) M. Schultz, Lingolemma lingulatum (Tuck.) M. Schultz & M. Prieto, Paludolemma syreniarum (C.J. Lewis & M. Schultz) M. Schultz & M. Prieto, Peltolemma socotranum (M. Schultz) M. Schultz & M. Prieto, Phylliscum aotearoa (Henssen & B. Bartlett) M. Schultz & M. Prieto, Phylliscum cylindrophorum (Vain.) M. Schultz, Phylliscum laatokkaense (Vain.) M. Schultz & M. Prieto, Phylliscum neglectum (Henssen) M. Schultz & M. Prieto, Phylliscum permiscens (Nyl.) M. Schultz & M. Prieto, Phylliscum rhodostictum (Taylor) M. Schultz & M. Prieto, Porocyphus antarcticus (Cromb.) M. Schultz & M. Prieto, Porocyphus macrosporus (Henssen et al.) M. Schultz & M. Prieto, Porocyphus minutissimus (Henssen) M. Schultz, Porocyphus rosulans (A. Henssen) M. Schultz, Porocyphus tasmanicus (A. Henssen) M. Schultz, Porocyphus willeyi (Tuck.) M. Schultz & M. Prieto, Pseudotichia vermiculata (Nyl.) Schultz & M. Prieto, Pycnolemma polycarpum (M. Schultz) M. Schultz & M. Prieto, Synalissina botryosa (A. Massal.) M. Schultz & M. Prieto, Synalissina cladodes (Tuck.) M. Schultz & M. Prieto, Synalissina condensata (Arnold) M. Schultz & M. Prieto, Synalissina degeliana (P.M. Jørg.) M. Schultz & M. Prieto, Synalissina dispansa (H. Magn.) M. Schultz & M. Prieto, Synalissina intricatissima (J. Steiner) M. Schultz & M. Prieto, Synalissina isidiodes (Nyl. ex Arnold) M. Schultz & M. Prieto, Synalissina vesiculifera (Henssen) M. Schultz & M. Prieto, Thelignya arnoldii (Frauenf.) M. Schultz & M. Prieto, Thelignya lacustris (P.M. Jørg. & R. Sant.) M. Schultz & M. Prieto, Thelignya neglecta (Erichsen) M. Schultz & M. Prieto, Thelignya obtenebrans (Nyl.) M. Schultz, Thyrea osorioi (Henssen) M. Schultz. New status and combination: Gonotichia depauperata (Servít) M. Schultz & M. Prieto. Emended description: Lempholemma Körb., Lichina C. Agardh, Thelignya A. Massal., Lichinaceae Nyl., Phylliscaceae Th. Fr., Porocyphaceae Körb. Resurrection: Cladopsis Nyl., Collemopsis Nyl. ex Crombie, Forssellia Zahlbr., Pleopyrenis Clem., Synalissina Nyl. Citation: Prieto M, Wedin M, Schultz M (2024). Phylogeny, evolution and a re-classification of the Lichinomycetes. Studies in Mycology 109: 595-655. doi: 10.3114/sim.2024.109.09.

Keywords: Gloeoheppiaceae; Heppiaceae; Lichinaceae; Peltulaceae; ancestral reconstruction; ascoma ontogeny; new taxa.

Fig. 1

Main types of thallus growth forms in the Lichinomycetes. A. Pterygiopsis atra, crustose, margin slightly effigurate (Beeching 6938). B. Watsoniomyces obsoletus, endolithic with immersed, blackish apothecia (Powell). C. Peltula euploca, squamulose-peltate, concave with grey sorediate margin (Marques 1246). D. Digitothyrea polyglossa, foliose, divided into furcate, plicate lobes, surface isidiate (Schultz 16124). E. Ephebe lanata, filamentous, irregularly branched, shape determined by Stigonema cyanobiont (Schultz 16606a). F. Lichinella stipatula, fruticulose or dwarf-fruticose with short, erect branchlets (Schultz 16610b). A, D. Lichinaceae. B, E. Porocyphaceae. C. Phylliscaceae. F. Lichinellaceae. Scale bars = 1 mm.

Fig. 2

Ascoma ontogeny main pathways in the Lichinomycetes. A. Lichina confinis, spheroid tangle of generative hyphae in tip of thallus branch developing into an apothecium (Schultz 07195). B. Phloeopeccania pulvinulina, tangle of generative hyphae with densely coiled ascogones in LPCB (Brown 50910). C. Peccania fontqueriana, group of coiled ascogones developed directly from thallus hyphae in LPCB (Schultz 14102b). D. Psorotichia frustulosa, spheroid ascoma primordium formed by reticulate hyphae and first ascogenous hyphae at the base in LPCB (Palice 12703). E. Porocyphus antarcticus, early stage of pycnoascocarp with ascogones formed beneath a still fully functional pycnidium in LPCB (Eaton, holotype). F. Tichocyphus gotlandicus, somewhat later stage with trichogynes protruding into pycnidial cavity in LPCB (Schultz 05569, paratype). G. Collemopsis schaereri, medium stage of pycnoascocarp, juvenile asci and first paraphyses formed in pycnidial cavity and conidiophores still producing small conidia in LPCB (Schultz 03440b). H. Lichinella algerica aggr., earliest stage of thallinocarp development with coiled ascogones below thallus surface in LPCB (Groner 4354). I. Lichinella minnesotensis, young thallinocarp with sparse paraphyses and juvenile asci below almost continuous thallus cover (LaGreca 4245). J. Thallinocarp hymenium with strong blue staining in KOH/Lugol and sterile, yellowish brown thallus cover (LaGreca 4245). K. Lichinella myriospora, mature thallinocarp, irregular hymenium with sparse asci and paraphyses covered by almost continuous layer of sterile thallus plectenchyma containing coccoid cyanobionts (Palice 15858). L. Lichinella cribellifera, mature thallinocarp, hymenium divided into partial hymenia with punctiform discs and separated by intrusions of sterile thallus (Feuerer). Scale bars: A–H = 25 μm, I–L = 50 μm.

Fig. 3

Ascus types and ascospores in the Lichinomycetes. A. Peltula euploca, narrow clavate, polysporous, unitunicate-rostrate ascus in KOH/Lugol, tip thickened and distinctly amyloid (Marques 1247). B. Pyrenopsis furfurea/haematina, unitunicate-rostrate ascus, tip amyloid in KOH/Lugol with small ocular chamber. C. Emptied ascus with amyloid rostrum (Schultz 16930). D. Lichinella cribellifera, Lichinella type ascus, irregularly clavate, thin walled, no apical wall thickening, but outer wall coat distinctly staining in KOH/Lugol and tip with bluish outer gelatinous cap (Feuerer). E. Peccania coralloides, Peccania type ascus sensu Moreno & Egea (1991), broadly clavate, thin walled, with distinct amyloid outer cap, but no apical dome, KOH/Lugol (Schultz 08477). F. Anema tumidulum, Peccania type ascus sensu Moreno & Egea (1991) in KOH/Lugol (Henssen 17744b, isotype). G. Lichina confinis, Lichina subtype ascus, wall very thin, non amyloid in KOH (Schultz 05548). H. Psorotichia murorum, Psorotichia subtype ascus sensu Moreno & Egea (1991), 8-spored, wall thin with thin amyloid outer coat and only indistinct amyloid cap, KOH/Lugol (Schultz 19162). I. Collemopsis schaereri, Psorotichia subtype ascus sensu Moreno & Egea (1991), wall thin throughout, no distinct outer gelatinous cap in KOH (Schultz 05573). J. Wall non amyloid with indistinct, somewhat amyloid outer cap in KOH/Lugol (Schultz 05573). K. Synalissa ramulosa, Synalissa subtype ascus sensu Moreno & Egea (1991) in KOH, same as Psorotichia subtype but polysporous (Schultz 08946). L. Phylliscum demangeonii, Phylliscum type ascus, wall thin throughout, tip pointed, non amyloid, polysporous, in KOH (Schultz 16884). M. Phylliscum aotearoa, Phyllisciella type ascus, wall thick, with strongly amyloid outer coat and amyloid cap but no apical dome in KOH/Lugol (Kantvilas 113–16). Scale bars = 10 μm.

Fig. 4

Best tree from Maximum Likelihood (ML) with values from ML bootstrap followed by Posterior Probabilities from Bayesian analyses. New genera and species are depicted in bold. Previous family classifications are depicted in the tree with a lateral bar with increasing grey tones indicating Lichinaceae < Peltulaceae < Heppiaceae < Gloeoheppiaceae.

Fig. 4

Best tree from Maximum Likelihood (ML) with values from ML bootstrap followed by Posterior Probabilities from Bayesian analyses. New genera and species are depicted in bold. Previous family classifications are depicted in the tree with a lateral bar with increasing grey tones indicating Lichinaceae < Peltulaceae < Heppiaceae < Gloeoheppiaceae.

Fig. 5

Ancestral state reconstructions of selected morphological characters from stochastic mapping analyses. A. Growth form. 0: crustose, 1: endolithic, 2: squamulose, 3: foliose, 4: filamentose, 5: fruticulose. B. Ascoma development 0: typical apothecia, 1: pycnoascocarps, 2: thallinocarps. C. Type of asci. 0: unitunicate rostrate, 1: Lichinella, 2: Lichina, 3: Peccania, 4: Phylliscum, 5: Phyllisciella, 6: prototunicate. D. Spore number. 0: octosporous, 1: polysporous. For taxon names see Supplementary Fig. S2.

Fig. 6

Thallus growth forms in main clade 2 (Lichinaceae s. str.). A. Metamelanea umbonata, crustose, areoles angulate, thick with blackish, umbonate apothecia (Orange 15053). B. Forssellia affinis, crustose, margin effigurate, apothecia semi-immersed to sessile (Yoshi 100198). C. Gloeoheppia erosa, squamulose, margins coarsely sorediate, apothecia immersed to semi-immersed (Feuerer). D. Anema nummularium, squamulose, umbilicate-rosette shaped, pycnoascocarps at first punctiform and immersed, finally with widely opened, dark red discs (Marques 659). E. Lichina pygmaea, dwarf fruticose, branches palmate, flattened with globose pycnidia at tips (Schultz 17140). F. Zahlbrucknerella patagonica, filamentous, furcate, shape determined by Scytonema cyanobiont, apothecia lateral (Feuerer). Scale bars = 1 mm.

Fig. 7

Collemopsis schaereri. A. Thallus black, areoles with adnate apothecia (Schultz 08926b). B. Grey pruinose morphotype, apothecia with reddish brown, open discs (Teuber 1527). C. Apothecium with zeorine margin, subhymenium inversely cone shaped in LPCB (Massalongo Lich. Ital. 338, UPS lectotype). D. Zeorine apothecial margin (Schultz 03438). E. Paraplectenchymatous thallus anatomy in LPCB (Schultz 03440b). Scale bars: A, B = 1 mm; C = 25 μm; D, E = 10 μm.

Fig. 8

A. Forssellia affinis, areoles thick, slightly effigurate and pruinose, apothecia adnate (Palice 4034). B. Forssellia canariensis, areoles subsquamlose, epruinose, apothecia sessile with constricted base (Ertz 16309). C. Forssellia umbilicata, squamules imbricate with coarsely sorediate margin (Schultz 08907a). D. Forssellia concordatula, morphotype of “Pterygiopsis” coracodiza, areoles angulate, apothecia adnate (Schultz 08572a). E–G. Forssellia canariensis (Ertz 16309). E. Paraplectenchymatous thallus anatomy with anticlinal hyphae in LPCB. F. Paraphyses and polysporous asci in LPCB. G. Polysporous ascus and thick thalline margin. Scale bars: A–D = 1 mm; E–G = 10 μm.

Fig. 9

A. Lemmopsis arnoldiana, apothecia with distinct, brick coloured proper margin (Schultz 03446). B. Lemmopsis arnoldiana, apothecia with very thick proper, and receding thalline margin (Schultz 03441a). C. Lemmopsis pelodes, apothecia immersed in thallus, proper exciple distinct (Groner 4528, HBG). D. Lemmopsis lutophila, apothecia semi immersed with distinct proper and thin thalline margin (Vezda, GZU). E. Lemmopsis lutophila, zeorine apothecial margin in LPCB (Arnold Lich. Monac. 74a, M holotype). F. Lemmopsis lutophila, distinct proper exciple in KOH. G. Lemmopsis lutophila, asci and paraphyses in KOH (Zimmermann). Scale bars: A, C, D = 1 mm; E = 25 μm; F, G = 10 μm.

Fig. 10

Leprocollema americanum. A. apothecia with expanded, multiply umbonate discs, thalline margin receding (Vainio, TUR-VAIN11602, lectotype). B. Apothecial margin zeorine, becoming biatorine, hymenium divided by hyphal vertical bands into chambers in LPCB. C. Apothecia zeorine, discs multiply umbonate (van den Boom 36840). D. Zeorine apothecium, proper exciple distinct and apically reddish brown, hymenium divided into chambers (van den Boom 36843). Scale bars: A, C = 1 mm; B, D = 25 μm.

Fig. 11

Lingolemma lingulatum (Berger 25056a). A. Squamulose thallus with convex, lingulate lobules and numerous immersed apothecia. B. Anatomy with Nostoc cyanobionts, a thin central hyphal strand and two apothecia in LPCB. Scale bars: A = 1 mm; B = 25 μm.

Fig. 12

Pseudotichia vermiculata. A. Thallus areoles angulate, becoming eroded, apothecia immersed (H-NYL42517, lectotype). B. Older hymenium divided by sterile hyphae in LPCB (H-NYL42517, lectotype). C. Section of thick areole, top paraplectenchymatous, vertical hyphae forming thick medulla below in LPCB (H-NYL42517, lectotype). D. Thick, blackish thallus areoles becoming superficially eroded (Lojka 4, B, isolectotype). E. Thick, angulate areoles with two large apothecia, discs multiply umbonate (Palice 14422). F. Thick thallus areoles with slightly elevated, large apothecium (Prieto SMP52B). Scale bars: A, D–F = 1 mm; B, C = 25 μm.

Fig. 13

A. Psorotichia murorum, granulose crustose thallus with indistinct areoles and grouped apothecia (Massalongo Lich. Ital. 300, M, isolectotype). B. Psorotichia columnaris, apothecia distinctly sessile, thalline margin bulging (Feuerer). C. Psorotichia diffracta, thallus pruinose, apothecia adnate with finally expanded discs (Groner 4157). D. Apothecium with thick, paraplectenchymatous thallus margin, hymenium with straight, septate paraphyses and juvenile asci in LPCB (Claudel & Harmand Lich. Gall. 485, B). E. Psorotichia frustulosa, thallus areolate, somewhat pruinose, apothecia with concave discs and thick thallus margin (Palice 12703). F. Apothecium with thick thalline margin and inversely cone shaped subhymenium (Prieto SL96). Scale bars: A–C, E = 1 mm; D, F = 25 μm.

Fig. 14

Pyrenocarpon thelostoma (Hyerczyk 2400). A. Hemispherical apothecium with narrow discs resembling fish eyes. B. Zeorine apothecial margin with distinct thalline and proper margin in KOH. Scale bars: A = 0.5 mm; B = 10 μm.

Fig. 15

A. Thelignya lacustris, thallus rimose areolate with immersed, black apothecia (Gilbert). B. Thelignya lignyota, thallus squamule with numerous immersed, black apothecia (Schultz 16867). C. Apothecium with distinct green colouration of exciple and central umbo (Prieto Pirineos 51). D. Thelignya neglecta, (semi-)immersed apothecia in granulose areoles with brownish, umbonate discs (Schultz 07143b, epitype). E. Zeorine apothecial margin, proper exciple pale, paraphyses lax (Schultz 07226, epitype). F. Ascus with simple ascospores in small hymenial chamber surrounded by robust excipular hyphae, with unknown lichenicolous fungus (Malíček & Kulíková 7316). G. Thelignya arnoldii, thallus crustose with sessile apothecia, discs black (Arnold, W1913-5405, holotype). H. Flat thallus areoles with semi immersed to sessile apothecia and black discs (Schultz 08933). I. Hymenium with polysporous asci, upper part of proper exciple and central umbo distinctly greenish (Schultz 08933). Scale bars: A = 1 mm; B, D, G, H = 0.5 mm; C = 25 μm; E, F, I = 10 μm.

Fig. 16

Thallus growth forms in main clade 5 (Lichinellaceae). A. Lichinella myriospora, crustose, areoles fully occupied by mature thallinocarps (Palice 15858). B. Lichinella algerica, squamules becoming convex and ascending, thallinocarps inconspicuous submarginal swellings (Feuerer). C. Lichinella cribellifera, foliose, lobes rounded, margin down rolled, surface folded (Ertz 16312). D. Synalissina condensata, squamules bluish grey pruinose, stout, erect and densely aggregated, tips with small apothecia and concave discs (Groner 4014). E. Synalissina vesiculifera, lower surface of fruticulose cushion, branches cylindrical, furcate (Rosentreter 18798). F. Gonotichia octosporella, crustose, areoles irregular, some with thallinocarps resembling gall like swelling (Schultz 05576b). Scale bars = 1 mm.

Fig. 17

Gonotichia octosporella. A. Areoles with small, gall like thallinocarps (Prieto SL28). B. Areoles with slightly lifted margins and some thallinocarps (Prieto SMP137). C. Mature thallinocarp with scattered patches of sterile thallus on the hymenium and 8-spored ascus (Lettau, B, holotype). D. Thallinocarp with patches of sterile thallus on hymenium in LPCB (Prieto SL28). Gonotichia depauperata. E. Irregularly areolate thallus with convex, roundish thallinocarps (Nadvorník, PRM-634147, syntype). F. Thallus areoles of variable size, large areoles filled by thallinocarps (Schultz 08299). Scale bars: A, B, E, F = 0.5 mm; C, D = 10 μm.

Fig. 18

A. Synalissina botryosa, thallus button like with some emptied hormocystangia (Schultz 05517a). B. Synalissina isidiodes, thallus with minute, isidioid outgrowths (Schultz 05517b). C. Synalissina condensata, densely aggregated, erect squamules, basally pruinose, small apothecia at tips (Schultz 08216a). D. Synalissina intricata, much elongated, intricately furcate branches (Schultz 08148c). E. Synalissina cladodes, branches shortly furcate, somewhat bent, surface striate (Schultz 08779). F. Synalissina degeliana, tufted, caespitosely branched lobes with coarsely granulose tips (Schultz 05544a). G. Synalissina botryosa, apothecium in lobe tip with thick thalline margin and rudimentary proper exciple (Schultz 08406a). H. Synalissina vesiculifera, pycnidium hidden in lobe tip with small, simple pycnospores (Rosentreter 18798). For Synalissina condensata and S. vesiculifera see also Fig. 16 D, E. Scale bars: A–F = 1 mm; G = 25 μm; H = 10 μm.

Fig. 19

Thallus growth forms in main clade 4 (Phylliscaceae). A. Allopyrenis sanguinea, crustose, areoles irregular, granulose, pycnoascocarps small with thick thalline margin narrow, concave disc (Schultz 16954a). B. Phylliscum aotearoa, crustose-squamulose, margin slightly effigurate, pycnoascocarps numerous, semi-immersed to adnate, discs narrow (Kantvilas 113-16). C. Peccania cernohorskyi, squamulose, margin becoming lobulate, center isidiate (Candan 22). D. Phylliscum demangeonii, squamulose, umbilicate-rosette shaped, pycnoascocarps remaining immersed and with pin hole discs (Schultz 16868). E. Peccania coralloides, foliose, lobes conspicuously grey pruinose and the marginal apothecia (TARI-3677). F. Peltula tortuosa, subfruticose with tufted, contorted lobes (Büdel 24058). Scale bars = 1 mm.

Fig. 20

A. Allopyrenis sanguinea, areoles thick, slightly granulose, apothecia semi immersed to sessile and almost globose, discs narrow, later somewhat opened, thalline margin thick (Anzi Lich. Rar. Langob. 474, W2009-00156, type). B. Allopyrenis haemaleella, thallus areoles thin, smooth, apothecia sessile with open discs and surrounded by thin, elevated thalline margin (Schultz 05594). C. Allopyrenis reducta, thallus areoles small granulose, very thin, apothecia very small (Schultz 16950). D. Allopyrenis grumulifera, thallus very thin, granulose, apothecia very small with discs remaining punctiform (Prieto SL39). E. Allopyrenis haemaleella, lecanorine apothecium, thick-walled asci (Schultz 05595). F. Allopyrenis grumulifera, lecanorine apothecium lacking proper exciple, clavate asci with distinct gelatinous cap and apical thickening in KOH/Lugol (Malmgren, H-NYL42947, holotype). Scale bars: A–D = 0.5 mm; E, F = 10 μm.

Fig. 21

A. Phylliscum neglectum, thallus very thin with hemispherical apothecia (Blomberg, B600132196). B. Phylliscum rhodostictum, thallus areolate, perithecioid apothecia semi-immersed (Øvstedal, BG L-34197). C. Phylliscum permiscens, thallus with minute, inconspicuous apothecia (Schultz 16877). D. Phylliscum granuliforme, thallus effigurate, apothecia semi-immersed (Lewis 1303). E. Phylliscum demangeonii, thallus rosulate, apothecia semi-immersed (Schultz 16386). F. Phylliscum japonicum, thallus lobate, apothecia (semi-)immersed (Ogata 519, W1927-373, holotype). Scale bars = 1 mm.

Fig. 22

Thallus growth forms in main clade 3 (Porocyphaceae). A. Lempholemma chalazanum, crustose, film-like thin to somewhat lobate at margin and numerous laminal apothecia (Feuerer). B. Pyrenopsis conferta, crustose, granules becoming coralloid with terminal, globose apothecia (van den Boom 26043). C. Thyrea plectopsora, squamulose, margin incised forming lobules (Schultz 18162a). D. Thyrea girardii, foliose, divided into few broadly rounded, conspicuously bluish grey pruinose lobes (Urbanavichus 0905071a). E. Thermutis velutina, filamentous, branches furcate, shape determined by Scytonema cyanobiont, biatorine apothecia lateral (Lich. Fenn. 51b, W1903-638). F. Porocyphus willeyi, dwarf-fruticose, branches furcate with terminal apothecia (Beeching). For endolithic growth see Watsoniomyces obsoletus in Fig. 1B. Scale bars = 1 mm.

Fig. 23

A. Cladopsis triptococca, thallus areoles granulose, apothecia with thick thalline margin and concave discs (Marques 928). B. Cladopsis polycocca, areoles abundantly fertile, becoming subquamulose (Beeching 3152). C. Cladopsis palmana, thallus areoles angulate, plane to slightly convex, apothecia small, discs punctiform to sunken (Feuerer). D. Cladopsis guyanensis, crustose, areoles angulate and effigurate at margin, isidiate towards the center (Schultz 20004). E. Cladopsis portoricensis, pycnidium with tangle of generative hyphae, ascogones and trichogynes beneath in LPCB (Fink 690, MICH, isosyntype). F. Cladopsis polycocca, juvenile and mature ascus lacking apical thickening but covered by distinct gelatinous cap in KOH/Lugol (Feuerer). G. Cladopsis triptococca, two thin-walled asci with distinct outer gelatinous cap in KOH/Lugol (Schultz 16587a). Scale bars: A–D = 1 mm; E–G = 10 μm.

Fig. 24

Lapismalleus lugubris. A. Areoles becoming eroded, apothecia almost lecideine (Hepp Lich. Eur. 728, M0140086, topotype). B. Apothecium with thick, brownish proper exciple and simple ascospores in LPCB (Arnold 6, M0140087, syntype). C. Thallus black, areoles thin with whitish base, apothecia almost lecideine (Psorotichia lugubris f. atrata Arnold 40, M0140084, type). D. Apothecium with thick, brownish proper exciple, indistinct thalline margin and simple ascospores in LPCB (Psorotichia lugubris f. atrata Arnold 40, M0140084, type). E. Areoles thick, irregularly polygonal, apothecia (semi)immersed with widely opened discs surrounded by thin, blackish proper exciple, thalline margin indistinct (Schultz 05566). F. Apothecium with thick, brownish proper exciple, indistinct thalline margin (Schultz 05559). Scale bars: A, C, E = 1 mm; B, D, F = 10 μm.

Fig. 25

Paracyphus gotlandicus. A. Irregularly shaped, thick thallus areoles with numerous apothecia (Prieto SL19, holotype). B. Thick thallus areoles with coarsely warty surface, mature apothecia adnate with pale brownish excipulum proprium and reddish brown, finally umbonate discs (Schultz 05554, paratype). C. Juvenile apothecia emerging in thallus warts in the center, mature apothecia with distinct proper and receding thalline margin at the top and bottom right (Schultz 05553, paratype). D. Apothecium with zeorine margin, asci and paraphyses (Prieto SL134, paratype). E. Margin of old apothecium, proper exciple pale, only at the top reddish brown, thalline margin receding (Prieto SL19, holotype). F. Margin of younger apothecium with less distinct proper exciple and thalline margin still well developed. G. Mature apothecium with zeorine margin, asci and paraphyses in LPCB (Schultz 05554, paratype). H. Tip of ascus and released ascospores in KOH (Prieto SL134, paratype). I. Ascus with uniformly thickened walls and septate paraphyses in KOH. J. Ascus wall uniformly thickened and thick outer coast staining deeply blue in KOH/Lugol (Prieto SL19, holotype). Scale bars: A–C = 1 mm; D = 25 μm; E–J = 10 μm.

Fig. 26

Pleopyrenis picina. A. Thallus thin, areolate, apothecia (semi-)immersed, discs remaining very narrow (Schultz 08573). B. Thallus thin, continuous to rimose (Wirth 1832). C. Polysporous asci with distinct apical wall thickening in KOH (Schultz 08609). D. Ascus with amyloid tip in KOH/Lugol (Schultz 08573). E. Two immature asci with amyloid tips and an emptied ascus between them with stretched rostrum in KOH/Lugol (Schultz 08609). Scale bars: A, B = 1 mm; C–E = 10 μm.

Fig. 27

A. Porocyphus coccodes, irregular, thick areoles with small, almost globose apothecia (Schultz 08895a). B. Zeorine apothecial margin with equally thick thalline and proper exciple, thin walled ascus with ascospores (Schultz 16918c). C. Porocyphus rehmicus, zeorine apothecial margin, slender paraphyses and ascospores (Schultz 03471a). D. Areoles with rough surface and adnate apothecia (Schultz 03456). E. Porocyphus antarcticus, fragment of areole composed of erect branchlets with terminal pycnoascocarps (Eaton, BM000022209, holotype). F. Hyphae reticulate and fountain like surrounding Rivulariaceae cyanobiont in LPCB (for pycnoascocarp see Fig. 2E). G. Porocyphus macrosporus, thallus fruticulose with terminal pycnoascocarps (Ertz 12986). H. Mature apothecium (pycnoascocarp) with zeorine margin, thin walled, Lichina type asci and ascospores (Büdel & Wessels 14312a). Scale bars: A, D, E, G = 1 mm; B, C, F, H = 10 μm.

Fig. 28

Pseudocarpon persimile. A, B. Thallus small areolate, thin, apothecia at first hemispherical, later sessile, discs narrow, plane to slightly concave and resembling fish eyes (Schultz 05557, holotype). C. Section of mature apothecium showing zeorine margin, asci and paraphyses, hypothecium roundish lacking a stipe in LPCB (Prieto SL76, paratype). D. Ascogones in tangle of generative hyphae beneath a pycnidium in LPCB (Groner 4588). E. Mature ascus with ellipsoid spores in LPCB (Prieto SL76, paratype). F. Thin-walled ascus with spores in KOH (Schultz 05557, holotype). G, H. Ascospore walls staining blue in KOH/Lugol, ascus wall not amyloid. Scale bars: A, B = 0.5 mm; C–H = 10 μm.

Fig. 29

A. Pyrenopsis furfurea, juvenile to mature apothecia, hymenia partly fallen off (Schultz 16925). B. Ascus with ascospores appearing 2-celled due to plasmatic bridges, thalline margin thick, proper margin thin in LPCB (Jones, H-NYL42916, lectotype). C. Pyrenopsis subareolata, thallus areoles angulate, plane to slightly convex, apothecia remaining immersed and discs punctiform (Westberg, S). D. Immersed apothecium, thallus anatomy compact paraplectenchymatous (Halda 17352). E. Ascus with distinct amyloid tip and outer gelatinous cap in KOH/Lugol (Halda 17352). F. Pyrenopsis fuscatula, ascus with thickened apex, simple ascospores in KOH (Holmes, BM00073226). G. Pyrenopsis conferta, asci with amyloid tip in KOH/Lugol (Prieto SL42). H. Pyrenopsis haematina, sterile thallus with coralloid-granulose areoles partly overgrowing other crustose lichens (Prieto SMP249). Scale bars: A, C, H = 1 mm; B, E–G = 10 μm; D = 25 μm.

Fig. 30

Tichocyphus gotlandicus. A. Thallus areoles angulate with finely granulose surface texture, mature apothecia with pale brownish discs (Schultz 05538b, holotype). B. Juvenile, semi-immersed apothecia with narrow, concave discs (Schultz 05534, paratype). C. Mature apothecium top left with thalline margin and juvenile apothecium center right with still indistinct thalline margin and narrow, concave disc (Schultz 055569, paratype). D. Thallus compact paraplectenchymatous in upper part and loose hyphal reticulum towards the base in LPCB (Schultz 05538b, holotype). E. Marginal section of pycnidium with small conidia released and tangle of generative hyphae formed at the base of pycnidial cavity as earliest stage of ascoma development in LPCB. F. Juvenile, thin-walled asci in KOH. G. Section of mature apothecium with low thalline margin and rather distinct, pale excipulum proprium, subhymenium and hypothecium extending downwards as a stipe in LPCB (Schultz 055569, paratype). H. Section of hymenium, clavate asci with ascospores and septate, sparsely branched paraphyses in LPCB (Schultz 05538b, holotype). I. Simple, ellipsoid ascospores in KOH. Scale bars: A–C = 0.5 mm; D–F, H, I = 10 μm; G = 25 μm.