Corresponding author: Juha Pykälä (
Academic editor: Cecile Gueidan
Species of
Pykälä J, Kantelinen A, Myllys L (2020) Taxonomy of
Species of
The taxonomy of this morphogroup is rather poorly known also in Fennoscandia and authors have treated the species somewhat differently (see, for example, Vainio 1921;
The phylogenetic position of large-spored (at least some spores exceeding 25 µm in length) species of
In this paper, we revise the Finnish species of
Perithecia and thalli were hand-sectioned with razor blades. The sections were examined and measured in tap water. Asci and ascospores were also studied in squash preparations of perithecia mounted in water. Sections and squash preparations of old herbarium specimens were studied using potassium hydroxide (KOH, 10% solution). Additionally, involucrellum characters and exciple colour and diameter were studied by cutting perithecia into two pieces and studying the pieces using a binocular microscope.
The range of ascospore size is indicated as arithmetic mean and standard deviation. Minimum and maximum values are given in parentheses. The size of the perithecia (in diameter) is given in surface view. The colour of the wall of the exciple is the colour of the base of the exciple.
Total genomic DNA was extracted from perithecia (1–3) of two- to six-year-old herbarium specimens. Most samples were placed in 96-well microplates and sent to the Canadian Centre for DNA Barcoding (
The DNA of 25 specimens (26865, 29589, 31528, 32606, 33120, 34601, 35326, 35361, 35857, 35920, 35922, 35930, 35933, 35965, 36222, 36244, 36245, 36254, 36294, 36304, 36308, 36335, 36371, 37331, 39475) was extracted using DNeasy Blood & Tissue kit by Qiagen following the protocol described in
The BLAST search facility in GenBank (
A total of 138 ITS sequences were aligned with MUSCLE v.3.8.31 (
Specimens used in the phylogenetic analyses. New sequences are in bold.
Species | Country | Voucher | GenBank accession numbers |
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Sweden | Tibell 23641 (UPS) |
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Sweden | Savić 3021 (UPS) |
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Sweden | Nordin 5466 (UPS) |
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Sweden | Palice 7666 (hb. Palice) |
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Sweden | Savić 3163 (UPS) |
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Sweden | Savić 3154 (UPS) |
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Sweden | Savić 3015 (UPS) |
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Sweden | Savić 3003 (UPS) |
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Sweden | Tibell 23959 (UPS) |
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UK | Orange 16318 (NMW) |
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Sweden | Tibell 23649 (UPS) |
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Sweden | Tibell 23525 (UPS) |
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UK | Orange 16278 (NMW) |
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UK | Orange 16309 (NMW) |
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UK | Orange 15898 (NMW) |
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UK | Orange 16103 (NMW) |
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Sweden | Orange 16377 (NMW) |
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Canada | McMullin (OAC) |
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Sweden | Savić 3063 (UPS) |
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Switzerland | Thues W1135 |
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Switzerland | Thues W1097 |
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We obtained 119 new nuITS sequences in this study (Table
Phylogenetic relationships of
Continued.
Habitus of the new
The monophyly of the ingroup was strongly supported, which suggests that all Finnish species in our study are members of the
All the studied species had relatively-low infraspecific genetic variation in their ITS sequences, but there seems to be species-specific variation (Table
Minimum infraspecific sequence similarity of the ITS region of the species. n = number of studied specimens.
n | Minimum sequence similarity | |
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4 | 99.6% |
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3 | 99.5% |
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16 | 98.9% |
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4 | 100% |
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12 | 98.5% |
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4 | 99.1% |
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18 | 99.8% |
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3 | 100% |
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18 | 98.7% |
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26 | 98.7% |
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10 | 98.6% |
Infraspecific morphological variation usually appeared to be rather high. For instance, in most study species, more than one major involucrellum type was detected and infraspecific variation of other perithecium characters was also considerable (Fig.
Schematic drawings of sections of perithecia of the study species
The main perithecium characters of the study species.
Species |
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0.13–0.26 | 0–60 | 0.18–0.27 | 21–26–30 × 9–11–13 | |
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0.15–0.28 | 30–60 | 0.16–0.32 | 23–28–34 × 10–12–14 | |
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0.13–0.40 | 0–80 | 0.20–0.35 | 20–27–35 × 10–13–16 | |
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0.18–0.36 | 40–70 | 0.16–0.28 | 23–27–34 × 10–11–13 | |
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0.11–0.42 | 0–60 | 0.19–0.42 | 24–30–37 × 10–13–17 | |
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0.11–0.26 |
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50–60 | 0.18–0.25 | 21–26–29 × 10–12–13 |
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0.07–0.37 | 50–70 | 0.21–0.28 | 23–28–31 × 10–12–14 | |
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0.17–0.45 | 30–80 | 0.19–0.29 | 21–28–34 × 9–12–14 | |
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0.21–0.42 | 30–80 | 0.21–0.34 | 23–28–35 × 11–13–15 | |
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0.16–0.45 | 40–100 | 0.20–0.36 | 23–30–40 × 12–14–17 | |
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0.15–0.44 | 30–80 | 0.16–0.33 | 20–25–31 × 8–10–13 | |
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0.15–0.47 | 30–90 | 0.15–0.26 | 18–25–32 × 8–12–15 |
Based on the Maximum Likelihood analysis, all the studied species in the morphogroup with large spores, perithecia leaving pits in the rock and a pale thin thallus belong to the
Molecular data show that the number of the Finnish species in the morphogroup is higher than previously expected. Similar results have often been obtained from other molecular studies in
Of the new species,
We included multiple specimens per species in our study to examine genetic and morphological infraspecific variation. Interestingly, in most of the species, we found one or a few specimens that differed morphologically from the other specimens and could not be reliably identified at species level. This suggests that a rather high number of specimens needs to be sequenced to cover the infraspecific morphological variation of the species. Even if the studied species are characterised by a high infraspecific morphological variation and even overlap in morphology, the infraspecific variation in the ITS sequence is rather low. This result is similar to the recently analysed
For example, the occurrence of dark lines between contiguous conspecific thalli varies between the species. Such lines are common in
The study group is characterised by a predominance of a dark exciple wall. In most species, pale exciple walls have not been seen. Pale exciples are rather common only in
The occurrence of a halonate perispore has been confirmed for all studied Finnish species, but
Most study species have a northern distribution in Finland. The result was unexpected, because most previously-described species in the morphogroup are from Central Europe. This result suggests that most Finnish species may be restricted to the boreal and arctic zones or be at least rare south of the boreal zone. Dolomite rocks in the Oulanka area in the biogeographical Province of Koillismaa have the highest species richness of large-spored
Species descriptions are based on the Finnish sequenced specimens.
Species characterised by pale, usually endolithic thallus, perithecia leaving shallow to deep pits in the rock, very variable involucrellum appressed to the exciple and ascospores (21–)24–28(–30) × (9–)10–12(–13) mm, morphologically rather similar to the other Finnish species of the
Finland. Varsinais-Suomi, Länsi-Turunmaa (Parainen), Ersby, 150 m SE of Stormossen, abandoned lime quarry, quarry waste hill, S-slope, on pebbles, 27 m alt.,
Prothallus absent. Thallus white, grey or pale greyish-brown, endolithic or thinly epilithic, continuous or small patches surrounding perithecia, ca. 20–60 mm thick, algal cells (4–)5–8 mm. Perithecia 0.13–0.26 mm in diam., (1/2–)3/4(–1)–immersed, leaving shallow to deep pits in the rock, sometimes thinly thalline covered; 80–140 perithecia/cm2. Ostiole inconspicuous, tiny, pale or dark, plane or depressed, ca. 20–30 mm wide. Involucrellum absent, apical, to the exciple base level or enveloping the exciple, 20–60 mm thick, appressed to the exciple. Exciple 0.18–0.27 mm in diam., wall dark brown or black, ca. 20–30 mm thick. Periphysoids ca. 25–35 × 1.5–2.5 mm. Asci 60–104 × 22–33 mm, 8-spored. Ascospores 0-septate, (20.6–)23.8–25.8–27.8(–30.3) × (8.7–)10.0–11.0–12.0(–12.9) mm (n = 117), perispore 1–1.5 mm thick.
All finds are from lime quarries or road cuttings of calcareous rocks. The species seems to prefer pebbles and stones in lime quarries. It occurs both in sun-exposed and rather shady habitats. The specimens are from SW and SE Finland. This suggests that
The epithet refers to the dualistic nature of the involucrellum of the species: absent or short vs. long or enveloping the exciple.
Finland. Varsinais-Suomi, Särkisalo, Förby, E of Vähämaankaula, abandoned lime quarry, beneath NW-facing wall, on stone, 7 m alt.,
Species morphologically somewhat similar to
Finland. Koillismaa, Kuusamo, Oulanka National Park, Mataraniemi, shore of Oulankajoki river, treeless stony river shore, on dolomite stones, 145 m alt.,
Prothallus absent. Thallus grey to pale greyish-brown, endolithic or thinly epilithic, continuous, 20–80 mm thick, algal cells 5–8 mm. Perithecia 0.15–0.28 mm in diam., 1/2–1-immersed, leaving shallow to deep pits in the rock, often surrounded by thallus collar, few perithecia thinly thalline covered; 80–160 perithecia/ cm2. Ostiole inconspicuous, dark, plane or depressed. Involucrellum apical to covering half of the exciple, in one specimen also few longer involucrella almost reaching the exciple base level present, 30–60 mm thick, appressed to the exciple or slightly diverging from the exciple. Exciple 0.16–0.32 mm, wall dark brown or black, ca. 15–25 mm thick. Periphysoids (25–)30–40(–50) × 1.5–2.5 mm, branching. Asci 8-spored. Ascospores 0-septate, two 1-septate spores seen in one specimen, (23.1–)25.1–27.5–29.8(–34.1) × (9.8–)10.7–11.6–12.6(–13.7) mm (n = 111), perispore 1 mm thick.
Two specimens of the species are from SW Finland and one specimen from NE Finland. The three sequenced specimens are from different kinds of habitats: dolomite stone on river shore (apparently periodically submerged), calcareous rock on seashore (perhaps not submerged) and in a lime quarry on pebbles. The species may prefer more humid (but preferably sun-exposed?) habitats than the other species in the
The perithecia of the species leave shallow to deep pits in the rock when decayed.
Finland. Varsinais-Suomi, Raasepori (Karjaa), Knapsby, Mustio lime quarry, deciduous forest on lime quarry waste, on pebbles, 45 m alt.,
The species cannot be morphologically separated with certainty from the other species of the
[Russia,] Suojärvi, ad saxa calcarea Pöpönsaari, 1870, Norrlin (H!, H-NYL 3036a!, syntypes).
Prothallus absent. Thallus white, grey or pale brown, endolithic, rarely epilithic (two sequenced specimens), thin, continuous, algal cells 5–8 mm, occasionally (three sequenced specimens) contiguous conspecific thalli separated by dark brown lines, 0.13–0.22 mm wide. Perithecia 0.13–0.40 mm in diam., (1/4–)1/2–1-immersed, leaving shallow to deep pits in the rock, few perithecia occasionally not leaving pits, often surrounded by a thalline collar, sometimes thinly thalline covered; 50–140 perithecia/cm2. Ostiole usually inconspicuous, pale or dark, plane or depressed, ca. 20–50 mm wide. Involucrellum absent or apical, short, rarely covering half of the exciple (two sequenced specimens), (40–)50–80 mm thick, appressed to the exciple or diverging from the exciple. Exciple 0.20–0.35 mm in diam., wall dark brown or black, ca. 27–40 mm thick, apex thickened to ca. 50–100 mm thick if the involucrellum is absent. Periphysoids ca. 30–50(–60) × 1–2.5 mm, branching or branched-anastomosing. Ascospores 0-septate, (20.2–)24.6–27.4–30.2(–34.8) × (10.2–)11.7–12.6–13.5(–15.7) mm (n = 281), perispore 1 mm thick.
The species is a strict calcicole occurring on calcareous rocks. It may prefer fairly humid habitats.
Finland. Koillismaa, Kuusamo, Oulanka National Park, Pikkukönkäänkuru,
Based on the ITS phylogeny,
Specimens of
Species characterised by perithecia 1/4–3/4-immersed, leaving usually shallow pits, involucrellum covering half of the exciple or almost to the exciple base, ascospores (23–)25–29(–34) × (10–)11–12(–13) mm, morphologically rather similar to the other Finnish species of the
Finland, Varsinais-Suomi, Karkkila, Haavisto, 100 m S of E-part of Iitalammi, S-slope, clear cut herb-rich forest, on calcareous stone,
Prothallus absent. Thallus white or grey, inconspicuous, endolithic to thinly epilithic, continuous to irregularly rimose, ca. 20–80 mm thick, algal cells 5–7(–8) mm. Perithecia 0.18–0.36 mm in diam., 1/4–3/4(–1)–immersed, leaving shallow to more rarely deep pits in the rock, often thinly thalline covered except apex; 60–160 perithecia/cm2. Ostiole inconspicuous, tiny, pale to usually dark, plane or depressed, ca. 20–30 mm wide. Involucrellum covering half of the exciple or almost to the exciple base, 40–70 mm thick, appressed to the exciple or slightly or moderately diverging from it. Exciple 0.16–0.28 mm in diam., wall dark brown, ca. 20–25 mm thick. Periphysoids (20–)25–35(–40) × 1.5–2.5 mm, some branching. Asci 77–101 × 23–28 mm, 8-spored. Ascospores (22.7–)25.1–27.0–28.9(–33.6) × (9.6–)10.6–11.4–12.3(–13.3) (n = 78), perispore 1 mm thick.
Four sequenced specimens occur: two from SW Finland and two from NE Finland. The species grows on calcareous rocks and in lime quarries, on walls, boulders, stones and pebbles.
The species may be mixed up with several other species of
Finland, Koillismaa, Kuusamo, Oulanka National Park, Kiutaköngäs 400 m N,
Based on the ITS phylogeny,
A Genbank sequence
Not seen. Protologue: “auf Kalksteinen bei Rüdersdorf”.
Prothallus absent. Thallus white, grey or pale brown, endolithic, often inconspicuous, rarely thinly epilithic, algal cells 5–9 mm. Perithecia 0.11–0.42 mm, (1/2–)3/4–1-immersed in rock, leaving deep pits in the rock, commonly surrounded by a thallus collar, sometimes covered by a thin thalline layer except for the apex; (30–)60–120 perithecia/cm2. Ostiole usually inconspicuous, tiny, pale or dark, plane or depressed, ca. 20–40(–50) mm wide, wider ostiolar depression rarely present up to 80 mm wide. Involucrellum absent or apical, rarely covering half of the exciple, 40–60 mm thick. Exciple 0.19–0.42 mm in diam., usually round, but sometimes pear-shaped or at least longer than broad, medium brown (rarely), dark brown or black, ca. (20–)25–43(–60) mm thick, apex sometimes thickened to ca. 40–60 mm thick if the involucrellum is absent. Periphysoids ca. (30–)40–65 × 1–2(–3) mm, branching. Asci 78–102 × 27–39 mm, 8-spored. Ascospores 0-septate, rarely solitary spores 1-septate, (23.6–)27.4–30.5–33.7(–37.3) × (10.4–)12.1–13.4–14.6(–17.1) mm (n = 197), perispore 1–1.5 mm thick.
The species grows on calcareous rocks and in lime quarries, both on sun-exposed and shady rocks, both in southern and in northern Finland.
Finland. Varsinais-Suomi, Lohja, Torhola, 400 m E of Torhola cave, S-slope, calcareous rock outcrop, 40 m alt.,
Fennoscandian specimens of
Based on the ITS phylogeny,
Species characterised by dark lines between contiguous conspecific thalli, pale endolithic thallus, small perithecia leaving shallow to deep pits in the rock, involucrellum reaching the exciple base level and appressed to the exciple, ascospores measuring (21–)24–28(–29) × (10–)11–12(–13) mm.
Finland. Koillismaa, Kuusamo, Oulanka National Park, Pikkuköngäs, N shore of river Oulankajoki, dolomite rock outcrop, on SW-facing wall, 160 m alt.,
Prothallus not seen. Thallus pale grey, endolithic, dark lines between contiguous conspecific thalli, 0.21–0.35 mm wide. Perithecia 0.11–0.26 mm in diam., (1/2–)3/4–immersed, leaving shallow to deep pits in the rock, surrounded by a thallus collar; 120–140 perithecia/cm2. Ostiole inconspicuous, tiny, pale to dark, plane or depressed, ca. 30 mm wide. Involucrellum reaching the exciple base, 50–60 mm thick, appressed to the exciple. Exciple 0.18–0.25 mm in diam., wall dark brown to black. Periphysoids ca. 25–35 × 2–2.5 mm. Asci 8-spored. Ascospores 0-septate, (21.2–)24.5–26.5–28.4(–29.4) × (10.0–)10.9–11.7–12.5(–13.2) mm (n = 36), perispore 1 mm thick.
The only known specimen is from a dolomite rock on a river shore in north-eastern Finland, in Kuusamo.
The only specimen available is characterised by dark lines between contiguous conspecific thalli.
Russia, Karelia Onegensis, Mundjärvi, supra saxa
Prothallus absent. Thallus white, grey or pale greyish-brown, endolithic or thinly epilithic, farinose, algal cells 5–8 mm, contiguous conspecific thalli often separated by dark lines, 0.13–0.22 mm wide. Perithecia 0.07–0.37 mm, (1/2–)3/4–1-immersed, leaving shallow to usually deep pits in the rock, surrounded by a thalline collar; 40–80 perithecia/cm2. Ostiole pale or dark, plane or depressed ca. 20–40(–60) mm wide. Involucrellum apical or covering half of the exciple, 50–70 mm thick, appressed to the exciple or diverging from the exciple. Exciple 0.21–0.28 mm in diam., wall dark brown to black, ca. 20–31 mm thick. Periphysoids ca. 30–50 × 2–2.5(–3) mm. Asci ca. 66–84 × 26–33 mm, 8-spored. Ascospores 0-septate, (23.2–)26.2–27.9–29.5(–31.3) × (10.3–)11.7–12.3–13.0(–14.1) mm (n = 63), perispore not seen, but may have vanished during storage.
This species is known from Finland only from the Oulanka area in the biogeographical province of Koillismaa in NE Finland where it grows on dolomite rocks. It seems to occur in fairly shady habitats.
Finland. Koillismaa, Salla, Oulanka National Park, Savikoski 300 m W,
The type specimens of
Species characterised by pale, usually thinly epilithic thallus, rather large perithecia leaving shallow to deep pits in the rock, involucrellum usually covering more than half of the exciple, ascospores (21–)26–30(–34) × (9–)11–13(–14) mm, morphologically difficult to separate from
Finland. Koillismaa, Kuusamo, Juuma, Oulanka National Park, Hautaniitynvuoma, gorge, dolomite rock outcrop, on high NE-facing wall, 190 m alt.,
Prothallus absent. Thallus white, grey or more rarely pale brown, endolithic or usually thinly epilithic, continuous or rimose, often farinose, up to 0.2 mm thick, algal cells (4–)5–7 mm, contiguous conspecific thalli sometimes separated by a dark line, 0.12–0.35 mm wide, present in only few specimens. Perithecia 0.17–0.45 mm in diam., (1/4–)1/2–3/4(–1)-immersed, leaving shallow to deep pits in the rock, rarely few perithecia not leaving pits, often thinly thalline covered except apex; (30–)40–120 perithecia/cm2. Ostiole tiny, pale or dark, plane or depressed, ca. 20–40(–60) mm wide, occasionally wider ostiolar depression up to 110 mm wide. Involucrellum covering half of the exciple or to the exciple base level, rarely in few perithecia enveloping the exciple, (30–)40–70(–80) mm thick, appressed to the exciple or slightly or moderately diverging from it. Exciple 0.19–0.29 mm in diam., wall dark brown or black, rarely pale, ca. 20–42 mm thick. Periphysoids ca. (20–)25–40 × (1.5–)2–2.5(–3) mm. Asci 68–102 × 25–34 mm, 8-spored. Ascospores 0-septate, (21.4–)25.5–27.9–30.3(–34.5) × (9.3–)11.3–12.2–13.1(–14.2) mm (n = 312), perispore 1 mm thick.
Most specimens of the species originate from the Kuusamo area.
Finland. Koillismaa, Kuusamo, Paljakka, E shore of Kuusinkijoki river, Kiukaankorva, dolomite rock outcrop, on overhanging NW-facing wall, scarce, 213 m alt.,
The exciple wall of
Differing from
Finland, Koillismaa, Kuusamo, Oulanka National Park, Taivalköngäs, shore of Oulankajoki river, dolomite rock outcrop, on gentle NE-slope, 165 m alt.,
Prothallus absent. Thallus white, grey, ochraceous or pale greyish-brown, endolithic to thinly epilithic, continuous to irregularly rimose, in one specimen contiguous conspecific thalli separated by a dark line. Perithecia 0.21–0.42 mm, 1/2–3/4-immersed, leaving shallow to deep pits in the rock, often surrounded by a thallus collar, in one specimen, thalline covered except apex, thalline cover 8–20 mm thick; 80–120 perithecia/cm2. Ostiole inconspicuous, tiny, pale to dark, plane or depressed, ca. 20–40 mm wide. Involucrellum covering half of the exciple or to the exciple base, in few perithecia may envelope the exciple, 30–80 mm thick, in one specimen, often apically thickened to 50–70 mm thick, appressed to the exciple. Exciple 0.21–0.34 mm in diam., wall blackish-brown, ca. 15–25 mm thick. Periphysoids ca. 25–50 × 1.5–2 mm. Asci 82–94 × 27–33 mm, 8-spored. Ascospores 0-septate, (23.0–)25.4–28.2–31.0(–34.9) × (11.2–)12.0–13.0–13.9(–15.2) mm (n = 83), perispore 1–1.5 mm thick.
All three finds are from the Oulanka area in NE Finland where the species grows on dolomite rock outcrops and on a dolomite boulder.
The species is close to
Finland. Koillismaa, Kuusamo, Oulanka National Park, Kiutaköngäs 400 m N,
This species is close to
[Russia,] Sibiria Septentrionalis: Si nus Konyam ad fretum Bering, 64°50' lat. bor., 173° long. occid. (Greenw.) 28–30.7.1879 E. Almquist (S-L46!, lectotype, designated here); Fretum Behring, Kongar Bay, E. Almquist (H-NYL 3512!, isolectotype).
Prothallus absent. Thallus white or grey, rarely pale ochraceous, endolithic or thinly epilithic, continuous or rimose, up to 0.1 mm thick, algal cells 5–8 mm. Perithecia (0.16–)0.23–0.45 mm in diam., (1/4–)1/2–3/4(–1)-immersed, not leaving pits or usually leaving shallow or deep pits in the rock, sometimes covered by a thin thalline layer except for the apex, often surrounded by a thalline collar; ca. (10–)30–100(–120) perithecia/cm2. Ostiole tiny, pale or dark, plane or depressed, ca. 20–40(–50) mm wide, ostiolar depression rarely wide, up to 130 mm wide. Involucrellum exceeding half of the exciple or reaching the exciple base level, rarely enveloping the exciple, (40–)50–100 mm thick, appressed to the exciple or slightly to moderately diverging from the exciple. Exciple 0.20–0.36 mm in diam., wall dark brown or black, ca. 22–45 mm thick. Periphysoids ca. 30–60 × (1–)1.5–2.5 mm, branching. Asci 84–109 × 32–40 mm, 8-spored. Ascospores 0-septate, rarely very few spores 1-septate, (23.4–)27.0–30.4–33.8(–40.1) × (11.7–)12.6–13.8–15.0(–17.4) mm (n = 242), perispore 1–2 mm thick.
The species occurs on calcareous rocks in both sun-exposed and shady sites. Most sequenced specimens are from the biogeographical province of Koillismaa. Three sequenced specimens (two localities) originate from eastern Finland (biogeographical Province of Pohjois-Karjala) and three (two localities) from southern Finland (biogeographical Province of Varsinais-Suomi). In southern Finland, the species seems to be very rare.
Finland. Varsinais-Suomi, Länsi-Turunmaa (Korppoo), Åfvensår, Kilamo, calcareous rock outcrop, on flat rock, scarce, 17 m. alt.,
This species has usually been treated as
[Switzerland] Bagnes-Thal, nördl. vom Hotel Monvoisin gegen den Plaine an Dolomitfelsen 16.9.1873 (G-00295028!, syntype); … Monvoisin & Bonat Mepa in Bagnes-Thal 1874 (G-00260361!, syntype?).
Prothallus absent. Thallus white, grey or pale brown, endolithic, or thinly epilithic, continuous to rimose, up to 0.1 mm thick. Perithecia 0.15–0.34(–0.44) mm in diam., (1/2–)3/4(–1)-immersed, leaving shallow to deep pits in the rock, few perithecia occasionally not leaving pits, sometimes covered by a thin thalline layer except for the ostiolar region; 40–160 perithecia/cm2. Ostiole inconspicuous, tiny, pale or dark, plane or depressed, in two specimens, several ostioles slightly projecting, ca. 20–40(–70) mm wide. Involucrellum apical or covering half of the exciple, rarely in few perithecia exceeding half of the exciple, 30–70(–80) mm thick, appressed to the exciple to clearly diverging from the exciple. Exciple 0.16–0.33 mm in diam., wall pale or pale brown (rather rare), usually dark brown or black, 18–30 mm thick. Periphysoids ca. 20–40(–50) × (1–)1.5–2.5(–3) mm, branching. Asci 58–84 × 22–28 mm, 8-spored. Ascospores 0-septate, (19.8–)22.9–25.2–27.4(–30.7) × (8.3–)9.6–10.5–11.4(–12.8) mm (n = 400), perispore 1 mm thick.
The species grows on various calcareous rocks and in lime quarries. It occurs both in sun-exposed and shady habitats. It is amongst the most common species of
Finland. Varsinais-Suomi, Lohja, Paavola, N of Rautaniemi, stony SE-slope, young
Salo (Kisko), Jyly, 200 m NE of Purslammi, calcareous rock outcrop, on NW-facing wall, 68 m alt.,
Vimpeli, Vimpeli, Ryytimaa, lime quarry, S-slope, on pebbles, 125 m alt.,
Species characterised by dark lines between contiguous conspecific thalli, pale usually endolithic thallus, perithecia leaving shallow to deep pits in the rock, very variable involucrellum, ascospores (18–)23–28(–32) × (8–)11–13(–15) mm, morphologically rather similar to the Finnish species of the
Finland. Enontekiön Lappi, Enontekiö, Porojärvet, Toskalharji, Toskalpahta, fell, SW-slope, scree, on dolomite boulder, 795 m alt.,
Prothallus absent. Thallus white, whitish grey or pale brownish, mainly endolithic to thinly epilithic, 20–170 mm thick, algal cells 5–10 mm, contiguous conspecific thalli separated by dark lines, 0.21–0.41 mm wide. Perithecia 0.15–0.47 mm in diam., 1/4–3/4-immersed, usually leaving shallow to fairly deep pits in the rock, rarely few perithecia not leaving pits, often surrounded by a thalline collar, 60–160(–200) perithecia/cm2. Ostiole tiny or conspicuous, pale to dark, plane or depressed, ca. 20–40(–60) mm wide, wider ostiolar depression occasionally present, up to 160 mm wide. Involucrellum apical, covering half of the exciple, exceeding half of the exciple or rarely to the exciple base, 30–70(–90) mm thick, appressed to the exciple, moderately diverging from the exciple, strongly diverging from the exciple or even spreading outwards away from the exciple. Exciple 0.15–0.26 mm, wall dark brown or black, 17–35 mm thick. Periphysoids ca. 25–40(–50) × 1.5–2.5 mm, branching. Asci 67–84 × 27–28 mm, 8-spored. Ascospores 0-septate, (18.1–)22.7–25.3–28.0(–31.7) × (8.3–)10.8–11.9–13.1(–15.2) mm (n = 228), perispore 1–1.5 mm thick.
The species is restricted in Finland to the calcareous mountains (Scandes) in NW Finland above the tree level. It always grows on dolomite. It grows on rock outcrops, boulders, stones and pebbles.
The specific epithet refers to the high morphological variation in the involucrellum from apical to (rarely) reaching the exciple base level, from being appressed to the exciple to spreading outwards away from the exciple and from fairly thin to thick.
Finland. Enontekiön Lappi, Enontekiö, Porojärvet, Toskalharji, Toskalpahta, fell, SW-slope, scree, on dolomite pebbles, 785 m alt.,
Based on ITS sequences,
France, Cantal: Auf hartem Kalk bei St. Santin, 1886, F. Adelminien (B600191351!, syntype).
The specimen in B is tiny with ca. 10 perithecia, of which all but two are covered by glue. The specimen is not identifiable and the species is better to be treated as a species with unresolved status (
[Slovenia] Carniola, Mojakrana, Aljazev dom, 1100 m, 1931, Servít (PRM-858477!, holotype?).
Prothallus not seen. Thallus white, endolithic. Perithecia 0.11–0.36 mm, immersed, leaving deep pits in the rock. Involucrellum absent. Exciple ca. 0.25 mm in diam., wall dark. Periphysoids ca. 25–35 × 2–3 mm, sparsely branching,
According to the protologue (
Austria, Niederösterreich, Voralpen, Bez. Lilienfeld, Gem. Kleinzell, SE von Salzerbad, Weg von Reintal zum Kruckensattel, 550–650 m alt., 29.3.2002, O. Breuss (8060) 19.990 (LI-01763881!, holotype).
Prothallus rather weakly developed, medium brown, weakly fimbriate. Thallus pale greyish-brown with frequent medium brown flecks, rimose, ca. 0.05–0.15 mm thick. Perithecia 0.22–0.38 mm, 1/2–3/4-immersed, not leaving pits to leaving shallow pits in the rock, thinly thalline covered except apex; ca. 80–100 perithecia cm2. Ostiole pale brown, plane, ca. 20–60 mm wide. Involucrellum to the exciple base level, occasionally enveloping the exciple, ca. 40–60 mm thick, appressed to the exciple. Exciple 0.21–0.24 mm in diam., wall pale to dark brown. Ascospores 0-septate, (22.7–)26.1–28.1– 30.9(–33.6) × (12.1–)12.4–13.5–14.5(–15.8) mm (n = 20).
This species was erroneously reported from Finland by
[Germany,] Alg. Alpen, Britzelmeyer (PRM-858488!, holotype?).
Prothallus not seen. Thallus whitish grey, endolithic. Perithecia 0.22–0.36 mm, 3/4–1-immersed, leaving deep pits in the rock. Involucrellum apical, ca. 60 mm thick, appressed to the exciple. Exciple ca. 0.24 mm in diam., wall dark. Ascospores 0-septate, 23–31 × 11–13 mm.
The specimen is small and only one perithecium was dissected. Our spore measurements match well with the original description (26–32 × 10–12(–14) mm, according to
[Switzerland] ad saxa
Prothallus not seen. Thallus inconspicuous, endolithic. Perithecia 0.15–0.25 mm, 3/4–1-immersed, leaving deep pits in the rock. Involucrellum absent. Exciple ca. 0.25–0.3 mm in diam., wall black. Periphysoids ca. 30–40 × 2 mm. Ascospores 0-septate, 17–23 × 11–12(–14) mm.
The species differs from
[Germany] an einer Sandsteinmauer in den Weinbergen bei Neuenheim, Febr. 1863, W. von Zwackh (M-0023494!, syntype?).
Prothallus not seen. Thallus pale grey, rimose to areolate, areoles 0.3–0.7 mm. Perithecia 0.22–0.26 mm, immersed in thallus. Involucrellum absent. Exciple wall pale. Periphysoids ca. 50–80 × 2.5–3 mm, branching. Ascospores 0-septate (only few seen), 20–28 × 13–14 mm.
[France,] Saléve (H-NYL3038!, UPS!, probably syntypes).
Prothallus not seen. Thallus pale greyish-brown, thinly epilithic, continuous. Perithecia 0.38–0.61 mm, 1/2–3/4-immersed, leaving fairly deep to deep pits in the rock; ca. 30–60 perithecia/cm2. Ostiole plane to depressed, ca. 20–60 mm wide. Involucrellum covering half of the exciple, ca. 70–180 mm thick. Exciple 0.23–0.38 mm in diam., wall black. Periphysoids long, ca. 1.5–3 mm thick. Ascospores 0-septate, 30–38 × 12–15 mm.
The species may be related to
[France] Calcaire argileux enposé au N, á 100 m au NE du pas du Bourreau Allaunch, 7.7.1951, Clauzade (PRM-858628!, syntype?).
Prothallus not seen (but, according to the protologue, “linea nigra marginatus”). Thallus grey with tiny brown flecks, thinly epilithic, continuous. Perithecia 0.25–0.45 mm, 3/4–1-immersed, leaving deep pits in the rock; ca. 70–80 perithecia/cm2. Involucrellum covering half of the exciple, ca. 60–80 mm thick. Exciple ca. 0.25 mm in diam., wall black. Periphysoids ca. 35–50 × 2–2.5 mm. Ascospores 0-septate, 28–34(–38) × 12–13 mm.
The studied specimen is tiny and better material is needed to solve the identity of the species. The specimen matches in most respects with
[Italy] An Kalksteinen einer Schutthalde unterhalb der Kalkwände an der Südseite des Latemar –Gebirges oberhalb Predazzo, Südtirol, 21. Aug. 1883, Arnold (H-NYL 7009!, H!, UPS-L-169663!, isotypes).
Prothallus absent. Thallus endolithic, grey. Perithecia 0.15–0.39 mm, (3/4–)1-immersed in rock, leaving deep pits in the rock. Involucrellum absent or possibly in some perithecia, small apical involucrellum. Exciple ca. 0.25–0.40 mm in diam., apex thickened, wall black, ca. 30 mm thick. Periphysoids ca. 50–70 × 2 mm, branched-anastomosing. Ascospores 0-septate, 25–30(–32) × (12–)13–16(–17) mm, perispore 1(–1.5) mm thick, persistent.
The species is rather similar to
Prothallus not seen. Thallus grey, pale brown, medium brown or rarely dark brown, with a violet tinge, continuous, rimose or areolate, thallus colour may be variable within specimen, 0.05–0.2(–0.3) mm thick, contiguous conspecific thalli separated by dark lines. Perithecia 0.26–0.52 mm, (1/2–)3/4-immersed, leaving shallow to deep pits in the rock; ca. 60–120 perithecia/cm2. Involucrellum apical, strongly diverging from the exciple (mainly spreading outwards away from the exciple), (40–) 50–90 μm thick. Exciple 0.25–0.4 mm in diameter, pale or dark. Periphysoids ca. 40–60 × (1–)1.5–2 μm. Ascospores 0-septate, (24–)27–35(–45) × 10–18(–20) μm, few spores 1-septate.
Based on morphology,
[Italy] ad saxa calcarea prope Veronam Mass., Anzi, Lich. Exs. minus rari Italiae superioris 377 (UPS!, syntype).
Prothallus not seen. Thallus grey, rimose to areolate, 0.2–0.4 mm thick. Perithecia 0.18–0.23 mm, immersed in thallus. Involucrellum apical, ca. 30–40 mm thick. Exciple ca. 0.4–0.45 mm in diam., pear-shaped, wall black. Ascospores 0-septate, 27–32 × 13–15 mm.
The studied specimen is conspecific with
[Italy,] in op. Giazza ad saxa
Prothallus not seen. Thallus pale greyish-cream, endolithic to thinly epilithic surrounding perithecia, slightly rimose, thalli bordered by a blackish-brown line. Perithecia 0.26–0.53 mm, (1/2–)3/4-immersed, leaving deep pits in the rock; 70–100 perithecia/cm2. Ostiole, pale, plane or depressed, ca. 40–150 mm wide. Involucrellum apical, 50–80 mm thick. Exciple 0.24–0.42 mm in diam., wall medium brown to blackish-brown, pale in one studied perithecium. Periphysoids ca. 40–50 × 2 mm. Ascospores 0-septate, 26–37 × 11–18 mm.
[Switzerland] Helvetia (H-ACH 686!, holotype?, piece on the upper left).
Prothallus not seen. Thallus pale grey, thin, continuous, up to 0.1 mm thick. Perithecia 0.26–0.41 mm, 1/4–1/2-immersed in thallus, sometimes with thin irregular thalline cover; ca. 70–100 perithecia/cm2. Ostiole inconspicuous, dark, plane or depressed, ca. 20–70 mm wide. Involucrellum slightly exceeding half of the exciple or reaching the exciple base level, 50–70 mm thick, appressed to the exciple or slightly diverging from it. Exciple 0.25–0.32 mm in diam., wall pale. Periphysoids ca. 30–40 × 1.5–2 mm, branching. Asci 75–117 × 25–37 mm, 8-spored. Ascospores 0-septate, (25.3–)26.8–29.6–32.3(–34.4) × (10.4–)11.8–12.5–13.2(–13.4) mm (n = 37).
[Greece,] Euboea: Berg Xerowuni, ca. 1100 m alt., 1931, Rechinger (PRM-858655!, isotype).
Prothallus not seen. Thallus white to whitish-grey, endolithic. Perithecia 0.2–0.35 mm, 3/4–1-immersed, leaving deep pits in the rock. Involucrellum covering half of the exciple, ca. 100 mm thick, appressed to the exciple. Exciple ca. 0.35–0.45 mm in diam., wall black. Periphysoids ca. 40–50 × 2–2.5 mm. Ascospores 0-septate, 25–30 × 12–16 mm.
The species may be conspecific with
[Russia,] Lapponia Rossica, 1843, F. Nylander (H!, holotype or syntype).
Prothallus not seen. Thallus grey, rimose. Perithecia ca. 0.4–0.6 mm, 1/2-immersed, leaving deep pits in the rock. Involucrellum enveloping the exciple, ca. 50–100 mm thick, thicker at apex. Exciple ca. 0.3–0.4 mm in diam., wall black. Ascospores in very poor condition, 0-septate, ca. 22–25 × 10 mm.
The specimen in H is small and in a very poor condition. Vainio (1921) reported the spore size of 15–24 × 10–18 mm.
[Montenegro] Dalmatia mer., Herceg Novi, 80 m, 1929, M. Servít (PRM-760604!, holotype).
Prothallus not seen. Thallus white to grey, endolithic. Perithecia 0.12–0.26 mm, immersed, leaving deep pits in the rock; ca. 30–40 perithecia/cm2. Involucrellum absent. Exciple ca. 0.4 mm in diam., wall black. Periphysoids ca. 35–50 × 1.5–2 mm, branched-anastomosing. Ascospores 0-septate (only few seen), 20–23 × 10–11 mm.
The spore size given in the protologue (
Germania: Regni Würtemberg, Blabyrae, ad rupes calcareas, Hochstetter (UPS-L-708716!, holotype).
Prothallus not seen. Thallus light grey, endolithic, dark lines between contiguous conspecific thalli present. Perithecia immersed, leaving deep pits in the rock. Involucrellum absent. Exciple 0.32–0.4 mm in diam., longer than wide, wall black, rather thin. Asci ca. 110–125 × 30–38 mm. Ascospores 0-septate, (25–)26–30(–35) × 16–20 mm.
The specimen is small and the description above is based on only one perithecium dissected.
Not in H-NYL, protologue: “in Gallia passim (Ejus statum ochraceo-tinctum, E Cebennis inferioribus in hb. Mougeot vidi)”.
The type material has not been located (possibly in Paris). Nylander had a very wide circumscription for
[Switzerland] ad dolomit supra Poschiavo, Anzi (H-NYL 3384!, syntype).
Prothallus absent. Thallus inconspicuous, endolithic. Perithecia 0.18–0.23 mm, 3/4–1-immersed, leaving deep pits in the rock; ca. 100–110 perithecia/cm2. Ostiole depressed, ca. 20–50 mm wide. Involucrellum absent (?). Exciple ca. 0.2 mm in diam., wall dark. Ascospores 0-septate, ca. 17–21 × 11–12 mm.
The studied specimen may be a tiny syntype. Nylander has annotated to the specimen: spores 18–24 × 11–14 mm.
[Germany] an Dolomitfelsen in Laubwäldern bei Eichstätt (Baiern), F. Arnold, Hepp, Flechten Eur. 692 (UPS-L-069713!, syntype).
Prothallus not seen. Thallus white, endolithic to thinly epilithic. Perithecia 0.2–0.3 mm, 3/4–1-immersed, leaving deep pits in the rock, surrounded by a thalline collar, ca. 50–120 perithecia/cm2. Ostiole inconspicuous, dark, depressed, ostiolar depression up to 100 mm wide. Involucrellum apical, 40–70 mm thick, appressed to the exciple. Exciple 0.17–0.25 mm in diam., wall dark. Periphysoids ca. 30 × 1.5 mm. Ascospores 0-septate, (17–)18–21 × (7–)8 mm.
This species differs from
[Italy,] in op. Tregnago – Viacara (VER!, syntype).
Prothallus not seen. Thallus pale brownish-grey, continuous to rimose. Perithecia 0.12–0.21 mm, 3/4–immersed in thallus. Involucrellum apical, ca. 40–50 mm thick, appressed to the exciple. Exciple 0.27–0.33 mm in diam., wall black. Periphysoids ca. 40–45 × 2 mm. Ascospores 0-septate, 28–31 × 12–15 mm.
The syntype specimen studied is probably conspecific with
?, protologue: “Jugoslavia, Pulac pr. Rijeka (Fiume), 250 m, dolom., Schuler (P)”.
The type material was not located.
[Montenegro,] Lovcen, Veterni mlin, 1400 m, 1929, M. Servít (PRM-859152!, holotype).
Prothallus not seen. Thallus grey with frequent tiny brown flecks, endolithic. Perithecia 0.18–0.26 mm, 3/4(–1)-immersed, leaving deep pits in the rock; ca. 60–80 perithecia/cm2. Involucrellum reaching the exciple base or enveloping the exciple, in the latter case diffusely pigmented under the exciple, ca. 70–110 mm thick, appressed to the exciple. Exciple ca. 0.20–0.22 mm in diam., wall dark. Periphysoids ca. 20–25 × 2.5–3 mm. Ascospores 0-septate (only few seen), 20–25 × 11–14 mm.
The species differs from the species of the
[Czech Republic,] Moravia, Kopřivnice, Piskovnice, 490 m alt., 1922, Suza (PRM-760594!, syntype).
Prothallus not seen. Thallus whitish-grey with abundant medium greenish-brown punctae, endolithic, a dark line between contiguous conspecific thalli. Perithecia 0.23–0.35 mm, 3/4–1-immersed, leaving deep pits in the rock, surrounded by a thalline collar; ca. 40–60 perithecia/cm2. Involucrellum apical. Exciple ca. 0.26 mm in diam., wall brown. Ascospores 0-septate, 20–25 × 9–12 mm.
Perithecia are mostly over-mature. One perithecium was sectioned.
[France] Salève, J. Müller (M-0193432!, holotype).
Prothallus not seen. Thallus pale brown with a violet tinge, continuous, hemi-endolithic, contiguous conspecific thalli separated by dark lines. Perithecia 0.38–0.46 mm, 3/4-immersed, leaving deep pits in the rock; ca. 30–50 perithecia/cm2. Involucrellum apical, ca. 50–60 mm thick, appressed to the exciple. Exciple ca. 0.34–0.35 mm in diam., wall black, ca. 25 mm thick. Periphysoids ca. 50–80 × 1–1.5 mm. Ascospores 0-septate, 32–41 × 12–15 mm.
The species is morphologically close to
[France] Gallia, Sevres (M-0193237!, holotype).
Prothallus not seen. Thallus greenish-grey with green flecks, continuous, ca. 0.1–0.3 mm thick. Perithecia 0.12–0.32 mm, immersed, leaving deep pits in the rock; ca. 80–100 perithecia/cm2. Involucrellum absent. Exciple ca. 0.27–0.41 mm in diam., higher than broad, often pear-shaped, wall dark brown. Periphysoids ca. 40–60 × 2–2.5 mm, branched-anastomosing. Asci 85–106 × 25–29 mm, 8-spored. Ascospores 0-septate, 18–23 × 12–14 mm.
The species differs from
[Germany] Eichstätt, ober dem Tiefenthale, 2. 1887, Boll (M-0204594!, holotype).
Prothallus not seen. Thallus grey, endolithic. Perithecia 0.08–0.21 mm, immersed, leaving deep pits in the rock; ca. 60–100 perithecia/cm2. Involucrellum absent. Exciple ca. 0.19–0.24 mm in diam., wall medium brown to dark brown, apex often thickened. Periphysoids ca. 15–30 × 2–2.5 mm. Asci ca. 61–69 × 20–21 mm, 8-spored. Ascospores 0-septate, 18–23 × 8–11 mm.
The species may differ from
Jugoslavia, Lovčen, Sanatorium, 1240 m, 1929, M. Servít (PRM-858454!, holotype?).
Prothallus not seen. Thallus grey with brown punctae, endolithic, contiguous conspecific thalli separated by dark lines. Perithecia 0.15–0.2 mm, 3/4(–1)-immersed, leaving deep pits in the rock; ca. 80–240 perithecia/cm2. Involucrellum absent. Exciple ca. 0.21–0.24 mm in diam., wall pale brown to medium brown, apex thickened to ca. 40–50 mm thick. Ascospores 0-septate 16–24 × 10–13(–14) mm.
The species is rather similar to
[Austria,] Dolomit … Grosser Rettenstein bei Kizbühel im Tirol, 1869, Arnold (PRM-858456!, isotype).
Prothallus not seen. Thallus pale brown, epilithic, thin, continuous. Perithecia 0.15–0.23 mm, (3/4–)1-immersed, leaving deep pits in the rock. Involucrellum absent or apical, ca. 70–90 mm thick. Exciple ca. 0.22–0.25 mm in diam., wall blackish-brown, the apex is strongly thickened when the involucrellum is absent. Ascospores 0-septate, 27–37 × 12–15 mm.
The species may fall within the variation of
[Croatia] Dalmatien: Schlossruine Vrlika a.d. … Granuga, an Kalk… c. 550 m, 5.7.1911, J. Baumgartner 4250 (W-4250!).
Prothallus not seen. Thallus endolithic, grey. Perithecia 0.15–0.34 mm, immersed, leaving deep pits in the rock; ca. 100–130 perithecia/cm2. Involucrellum absent. Exciple ca. 0.35–0.5 mm in diam., longer than wide, often pear-shaped, apex thickened, wall black. Periphysoids ca. 50–80 × 2 mm. Ascospores 0-septate, 26–35 × 12–14 mm.
Material similar to
[Italy] 87. Dolomitfelsen in der Schlernklamm ober … in Süd Tirol, 7.1867, Arnold (H-NYL 3208!, H-NYL 3209!, syntypes).
Prothallus absent. Thallus endolithic, grey with a violet tinge, a dark line between contiguous conspecific thalli present, 0.15–0.22 mm wide. Perithecia 0.21–0.44 mm, immersed in rock, leaving deep pits in the rock. Ostiolar depression large. Involucrellum absent or possibly in some perithecia, small apical involucrellum. Exciple ca. 0.4 mm in diam., apex thickened to ca. 60–80 mm, wall black. Periphysoids ca. 50–60 × 2 mm. Ascospores 0-septate, ca. 26–34 × 16–20 mm, perispore ca. 1–1.5 mm thick.
The perithecia of the syntypes in H-NYL are mainly over-mature. The spore size annotated by Nylander to the specimen is larger (40–48 × 23–26 mm) than the few spores measured by us.
Slovakia, in valle fl. Hnilec, pr. R. Ztratená, 800 m alt., calc., 1933, Suza (PRM 858074!, syntype).
Prothallus not seen. Thallus grey, endolithic to semi-endolithic. Perithecia 0.25–0.33 mm, 3/4-immersed, leaving deep pits in the rock, usually surrounded by a thalline collar or is covered by a thin thalline layer except for the apex. Ostiole pale, plane, ca. 20–50 mm wide. Involucrellum covering half of the exciple, ca. 50–70 mm thick, diverging from the exciple. Exciple ca. 0.3–0.33 mm in diam., wall pale brown. Periphysoids ca. 25–30 × 2–2.5 mm. Ascospores 0-septate, 27–38 × 12–15 mm.
[France] Reculet, Jan. 1855, J. Müller (M-0220250!, holotype).
Prothallus not seen. Thallus pale brown, epilithic, thin, continuous. Perithecia 0.38–0.55 mm, 1/2–3/4-immersed, leaving deep pits in the rock; ca. 30–40 perithecia/cm2. Ostiole tiny, inconspicuous, dark, plane, often surrounded by a projecting neck up to ca. 150 mm wide. Involucrellum absent. Exciple ca. 0.38–0.45 mm in diam., wall dark, ca. 30–40 mm thick, apex thickened to 70–100 mm thick. Periphysoids ca. 50–60 × 1.5–2 mm, branched-anastomosing. Ascospores 0-septate, 25–30 × 13–16 mm.
The species is rather similar to
[Greece,] Samos, Vathy, Rechinger (PRM-858434!, holotype).
Prothallus not seen. Thallus whitish-grey, endolithic to thinly epilithic, occasionally irregularly rimose around perithecia. Perithecia 0.22–0.28 mm, (1/2–)3/4-immersed, leaving shallow to deep pits in the rock; ca. 70–80 perithecia/cm2. Involucrellum enveloping the exciple, 40–50 mm thick. Exciple 0.19–0.28 mm in diam., wall black. Periphysoids ca. 50–60 × 2–2.5 mm. Ascospores 0-septate, ca. 21–25 × 11–13 mm.
According to the protologue, the spores may be larger: 20–29 × 9–15 mm (
[Italy,] avi in op. Avesa ([Monte] Ongarine) ad saxa putrida eocenica (VER!, syntype).
Prothallus not seen. Thallus whitish-grey, endolithic to semi-endolithic, a black line between thalli present. Perithecia 0.16–0.23 mm, immersed, leaving deep pits in the rock. Involucrellum absent. Exciple ca. 0.26 mm in diam., wall brown. Ascospores 0-septate, 24–33 × 12–18 mm.
Not seen. Protologue: “Italia, in Valle Bisagno prope Genuam, loco Prato, supra rupem calcaream colore fuscorufo tinctam. leg. Sbarbaro, 1946”.
The type material of
[Austria], Kalksteine des Serlosgipfels 8200’ Matrei-Tirol, 7. 1869, Arnold (M-0193173!, holotype).
Prothallus not seen. Thallus grey to pale brown, endolithic, somewhat inconspicuous. Perithecia 0.12–0.22 mm, immersed, leaving deep pits in the rock; ca. 60–100 perithecia/cm2. Involucrellum absent. Exciple ca. 0.2 mm in diam., wall pale to pale brown, apex dark, thickened. Ascospores 0-septate, (23.2–)23.9–24.7–25.4(–25.5) × (12.7–)12.8–13.7–14.6(–15.1) mm (n = 15).
The specimen is rather poor. The species differs from
Slovakia, Liptovskě hole, Zuberec, Osobita, 1650–1680 m, 1935, Suza (PRM-765231!, syntype).
Prothallus not seen. Thallus white or grey, endolithic. Perithecia 0.25–0.35 mm, 1/2–3/4(–1)-immersed, leaving shallow to deep pits in the rock. Involucrellum reaching the exciple base, ca. 70–90 mm thick, appressed to the exciple or slightly diverging from the exciple. Exciple ca. 0.15–0.25 mm in diam., wall pale. Periphysoids ca. 25 × 2–2.5 mm. Ascospores 0-septate, few spores 1-septate, ca. 20–27 × 9–10 mm, not well developed.
[Italy], Auf Kalkconglommerat in Villa Lagarina bei Roveredo in Südtirol, 1.5.1883, P. Strasser (M-02039301!, holotype).
Prothallus not seen. Thallus whitish-grey, endolithic. Perithecia 0.15–0.38 mm, (3/4)–1-immersed, leaving deep pits in the rock. Involucrellum apical, ca. 50–90 mm thick, appressed to the exciple. Exciple ca. 0.22–0.26 mm in diam., wall pale brown to dark brown. Periphysoids ca. 30–50 × 1.5–2.5 mm, branching. Asci ca. 88–95 × 26–28 mm, 8-spored. Ascospores 0-septate, (23.6–)24.5–26.6–28.8(–30.2) × (10.1–)10.4–11.4–12.4(–13.7) mm (n = 17).
The species may differ from the
Deutschland. Thüringen, Jonastal bei Arnstadt, alt. 350–400 m, co-ord.
For the description of the species, see
[Italy,] S. Leonardo, L. Tonini (VER!, syntype); ad saxa eocenica circa urbem Veronam (S. Leonardo), leg. Tonini, Massalongo Lichenes Ital. Exsiccatae 8 (VER!, syntype); Massalongo, Lich. Ital. exs. 8 (UPS!, syntype).
Prothallus absent. Thallus greenish-grey or grey with some brown pigmentation, epilithic, rimose, ca. 0.2–0.3(–0.4) mm thick. Perithecia 0.12–0.32 mm, 3/4–1-immersed in thallus. Involucrellum apical, ca. 60–70 mm thick. Exciple ca. (0.2–)0.3–0.5 mm in diam., often longer than broad, wall dark. Ascospores 0-septate, 27–35 × 11–15 mm.
The type material of the species is morphologically similar to
The fieldwork was mainly done during the research project “Threatened lichens of calcareous rocks”, which belonged to the research programme of deficiently known and threatened forest species (PUTTE) financed by the Ministry of the Environment. The Kone Foundation and Finnish Cultural Foundation are thanked for their financial support through the FinBOL project to the Finnish Museum of Natural History. We are grateful to Diana Weckman and Laura Häkkinen for their laboratory work. We wish to express our appreciation to Seppo Huhtinen for arranging the possibility to use the photography equipment and focus stacking programme, Combine ZP, in TUR and to Nelly Llerena Martinez and Timo Kosonen in advising and helping with the photo processing. Herbarium visits were made possible by a grant from Societas pro Fauna et Flora Fennica. Othmar Breuss, Claude Roux and an anonymous referee are thanked for their useful comments on the manuscript. Curators of the herbaria B, G, M, PRM, S, UPS and W are thanked for granting loans of specimens. We also would like to thank Andreas Beck, František Bouda, Francesco Di Carlo and Martin Westberg for their hospitality during visits by the first author to M, PRM, VER and UPS, respectively. Sonja Virta corrected the English.