Introduction
The Highlands of Scotland are widely recognized as supporting many important habitats and the Western Highlands, with their hyper-oceanic climate of high rainfall and low annual temperature fluctuation, are particularly important. The oceanic woodlands of the Western Highlands are internationally renowned for their important assemblages of bryophytes and lichens (Gilbert Reference Gilbert1984; Averis Reference Averis1991; Coppins & Coppins Reference Coppins and Coppins2012), and more recently the montane lichen biota has been shown to be equally important (Fryday Reference Fryday1997a, Reference Frydayb, Reference Fryday2001, Reference Fryday2002a). Previously, contributions by the authors (Fryday Reference Fryday2000, Reference Fryday2002b, Reference Fryday2005; Fryday & Coppins Reference Fryday and Coppins1996a, Reference Fryday and Coppinsb, Reference Fryday and Coppins2008) have added several new taxa to the lichen biota of the British Isles from this habitat, and here we add a further five new taxa and make other taxonomic innovations.
Materials and Methods
The study is based chiefly upon collections made by the first author during fieldwork leading to his PhD and now held in E or MSC. Type material and other collections were obtained on loan from GZU, STU and UPS.
Apothecial characteristics were examined by light microscopy on hand-cut sections mounted in water, 10% KOH (K), 15% HCl (H), 50% HNO3 (N) or 0·15% aqueous IKI. Thallus sections were investigated in water and 10% KOH (K). Ascus structure was studied in 0·15% aqueous IKI, both without prior treatment and after pretreatment with 10% KOH. Anatomical measurements were made in 10% KOH. For all fertile taxa at least 20 ascospores were measured. For C. subtenerum, where ascospore dimensions were critical for separation from C. tenerum, c. 50 ascospores were measured for each species.
Thin-layer chromatography followed the methods of Orange et al. (Reference Orange, James and White2001). Nomenclature for apothecial pigments follows Meyer & Printzen (Reference Meyer and Printzen2000).
Selected additional comparative material examined. Cliostomum tenerum (Nyl.) Coppins & S. Ekman. Great Britain:England: V. C. 3, South Devon: 1·5 km SW of Noss Mayo, Yealm Estate, coastline between Hilsea Point and Blackstone Point, 20/5.4, 1992, Coppins (15174) & A. M. O'Dare (E). Scotland: V.C. 103, Mid-Ebudes: Coll, A Chròic, 17/22(-3).62, 0–40 m, sheltered rock underhangs, 1983, Coppins 9678 (E). VC 91, Kincardine: Nr Nigg, Crombie (E–Lich. Brit. Exs. no. 68—isotype).
Fuscidea ‘alpina’ ad int.. Austria:Salzburg, Glockener-Gruppe: Madelz südlich über Rudolfshütte am Weißsee im Stubachtal, 2300–2500m, 8 ix 1973, J. Poelt (GZU).—Italy:Südtiroler Dolomiten: Passo di Rolle Quarzporphyr, c. 2000 m, 23 x 1976, J. Wetz (GZU). Trentino: Catena dei Lagora, Nordgrat des Berges Cavallazza S vom Pso di Rolle, an Porhyrblöcken, 2050–2250 m, 26 x 1984, J. Poelt (GZU).
Fuscidea ‘badensis’ ad int.. Germany:Baden, Nordschwarzwald: Lautenbach, Bad Sulzbach, Weg zum Pilatusfelsen, [41°31′N 8°9′E], 1989, V. Wirth 18561 (STU).
Fuscidea gothoburgensis (H. Magn.) V. Wirth & Vězda. Great Britain:Scotland: V. C. 92, South Aberdeen: Braemar, Glen Clunie, Creag nan Gabhar, 37/14.84, 450 m, rocks on scree slope, 1992, Fryday 3466 (E).
Fuscidea intercincta (Nyl.) Poelt. Great Britain:Wales: V. C. 49, Caernarvon: Bethesda, Carnedd Dafydd, Ysgolion Duon, below Grib Lem, 23/66.63, 525 m, side of giant boulder, 1994, Fryday 5332 (MSC). Scotland: V. C. 96, East Inverness: Cairngorm, Coire an Lochain, 28/98.04, 810 m, boulders, 1 x 1980, P. Topham s. n. (E). V. C. 97, West Inverness: Fort William, Glen Nevis, Meall Cumhann, 27/17.69, 500 m, vertical acid rock, 1990, Fryday (1239) & O. L. Gilbert (MSC). V. C. 106, East Ross: c. 17 km ESE of Ullapool, Seana Bhraigh, Loch Luchd Coire, 28/289.878, 1800 ft, N-facing vertical rocks, 1984, Coppins (10457) et al. (E).
Fuscidea kochiana (Hepp) V. Wirth & Vězda (sorediate). Great Britain:Scotland: V. C. 92, South Aberdeen: Braemar, between Meall Odhar and Glas Maol, top of siliceous rock, 37/15.77, 900 m, 1995, Fryday 6073 (E).
Fuscidea oculata Oberholl. & V. Wirth. Germany:Baden, Nordschwarzwald: Ottenhöfen, Melkereikopf, [48°33′N 8°2′W,] 880 m, Buntsandstein-Blockhalde am Nordwesthang, 1981, V. Wirth 9385, 9387 (STU–isotypes); ibid., 900 m, “Heidbeermauer”, Blockmeer, 1986, V. Wirth 15318, 9387 (STU).
Fuscidea lygaea (W. Mann) V. Wirth & Vězda (sorediate). Great Britain:Wales: V. C. 49, Caernarvon: Bethesda, Cwm Idwal, Clogwyn y Tarw, 23/64.59, 400 m, vertical face of rhyolite crag, 1994, Fryday 5316 (MSC).
Lecidea fuscoatra (L.) Ach. Great Britain:Scotland: V. C. 74, Wigtown: c. 0·6 km WSW of Glenluce, Torrs Warren, stable acid dune system, 25/13–4.55, on pebbles and stones, 1989, Coppins 13047 (E).
Lecidea paupercula Th. Fr. Iceland:Austur-Barðastrandarsýsla: Kollabúðaheiði, an der Straße, c. 6 km nördlich Kollabúðir, 65°38′45″N, 22°02′W, 440 m, 22 vii 1979, H. Hertel (E–Lecid. Exs. 7).—Great Britain:Scotland: V. C. 90, Angus: Caenlochan Glen, Glasallt Burn, at base of W-facing cliffs, 37/18.7, c.750 m, on granitic rocks, 1989, Coppins (13348) & O. L. Gilbert (E). V. C. 106, East Ross: c. 17 km ESE of Ullapool, Seana Bhraigh, Loch Luchd Choire, 28/28.87, c. 550 m, on large boulder, 1984, Coppins 10451 (E).
Ropalospora lugubris f. lugubris (Sommerf.) Poelt. Great Britain:Scotland: V. C. 97, West Inverness: Sunart, Ben Resipole, Allt Mhic Chiarain, 56°43·5′N 5°40·0′W, 380 m, siliceous rock outcrop in acid moorland, 1992, Fryday 3196 (MSC). V. C. 98, Argyll Main: Glen Coe, Coire nam Beitheach, 27/142.545, 875 m, 1992, Fryday 3345, 3346 (MSC). V. C. 104, North Ebudes: Isle of Skye, Bla Bheinn, above Loch Fionna-choire, 18/53.21, c. 500 m, on NE–facing rocks, 1987, Coppins 12698 (E).
The Taxa
Cliostomum subtenerum Coppins & Fryday sp. nov.
MycoBank No: MB800464
Cliostomo tenero similis sed sorediis praesentibus, thallo zeorinum continentes, apotheciis et ascosporis maioribus.
Typus: Great Britain, Wales, V. C. 52, Anglesey, NE of Amlwch, cove E of Llam Carw, 23/460.936, on vertical siliceous (‘green’ schist) coastal rocks above HWM, 11 June 1995, S. P. Chambers s. n. (E—holotypus).
(Fig. 1)
Cliostomum subtenerum (S. Chambers s. n.—holotype). Scale=0·5 mm. In colour online.
Thallus effuse, thin and discontinuous, occasionally thicker (to 0·4 mm), white, non-corticate. Soralia pale green, covering most of the thallus, 0·1–0·3 mm diam., soon confluent and appearing effuse; soredia farinose c. 0·02 mm diam. Photobiont chlorococcoid, cells 8–12 µm diam.
Apothecia scattered, sessile, 0·4–0·8 mm diam., becoming tuberculate and then to 1·2 mm diam.; disc pinkish brown to pale brown with paler margin, slightly convex soon becoming tuberculate with excluded margin. Excipulum composed of conglutinate radiating hyphae 5 µm wide; internally colourless to yellow-brown with grey-brown granular intrusions not dissolving in K, outer cells with brown pigment, becoming colourless in K. Hymenium hyaline, I+ blue, 40–45 µm tall, epihymenium brown, granular, becoming colourless in K; paraphyses 1·5–2·0 µm wide, moderately branched and anastomosing, apices to 3 µm wide, pigmented cap absent. Hypothecium hyaline, composed of randomly orientated hyphae. Asci c. 30×12 µm, clavate, Bacidia-type. Ascospores hyaline, 0–1 septate, narrowly ellipsoid to fusiform, often slightly curved, (10–)12–15(–20)×3–4 µm.
Conidiomata pycnidia, rare; flesh-coloured to pale brown, immersed in thicker areas of the thallus. Conidia ellipsoid-bacilliform, 7–8×1·5–2·0 µm.
Chemistry. Thallus and soredia C−, K+ yellow, Pd+ orange, UV+ dull yellow; atranorin, stictic acid and zeorin by TLC.
Cliostomum subtenerum resembles C. tenerum (Nyl.) Coppins & S. Ekman in the apothecia having a thalline margin but the two species differ in a number of significant characters. The most obvious differences are that the thallus of the new species is sorediate and that the apothecia and ascospores are both larger [0·1–0·2(–0·5) mm diam. and 7–10(–15)×(1·5–)2·0–3·0 µm in C. tenerum]. Cliostomum subtenerum also has a wider ecological amplitude, occurring in underhangs in montane situations and in coastal habitats, as well as a different chemistry (zeorin absent in C. tenerum). An additional difference is that all collections of C. subtenerum are abundantly fertile and pycnidia are rare, whereas C. tenerum is rarely fertile and commonly pycnidiate. Reese Naesborg et al. (Reference Reese Naesborg, Ekman and Tibell2007) showed that C. tenerum was related to Lecania s. str., which suggests that the same probably also applies to C. subtenerum. However, as C. tenerum differed from that genus in a number of characters (e.g., anastomosing excipular hyphae, complex thalline chemistry), they did not make any taxonomic changes.
Cliostomum subtenerum is known only from Coire a' Bhathaich near the summit of Ben Lomond and Llam Carw on Anglesey; at both localities it is relatively frequent in shaded underhangs on schistose rock. Associated species are few but include Lecanora polytropa (Wales) and Acarospora smaragdula (Scotland).
Additional specimens examined. Great Britain:Scotland: V. C. 86, Stirlingshire: Ben Lomond, Coire a' Bhathaich, 27/368027, 925 m, in sheltered underhang on schistose rock, 1994, Fryday 5590 (E); ibid., 27/369027, 925 m, in sheltered underhang on schistose rock, 1994, Fryday 5663 (MSC).
Dactylospora suburceolata Coppins & Fryday sp. nov.
MycoBank No: MB800465
Dactylosporae urceolatae similis sed apotheciis maioribus, ascosporis 3-septatis (sine septatis longitudinalibus), pigmento interno brunneo (haud purpureo) differt.
Typus: Great Britain, Scotland, V. C. 88, Mid-Perthshire, Ben Lawers, An Stuc, north crags, 27/638.433, 1000 m, 18 July 1989, Fryday s. n. (E—holotypus).
Dactylospora suburceolata (Fryday s. n.—holotype). Scale=0·5 mm. In colur online.
Dactylospora suburceolata, ascospores in ascus (Fryday s. n.—holotype). Scale=10 µm. In colour online.
Thallus inapparent, lichenicolous on a whitish–yellowish muscicolous crustose lichen(s).
Apothecia 0·2–0·8 mm diam., black or with a dark brown margin; disc concave; margin prominent, persistent, smooth. Exciple 60–75 µm wide (lateral to hymenium); upper part, and especially outer part, dark brown, K−; lower part dilute brownish or ± hyaline, but often with a thin, dark brown outer edge; cells to 12 µm diam. Hymenium 70–95 µm tall, hyaline to dilute red-brown; epihymenium red-brown, K+ dulling [never purplish]; paraphyses mostly slender, 1·0–1·5 µm wide in mid-hymenium, apices only slightly widening to 3 µm at the apex, each with a distinct, dark brown apical cap. Occasional thicker filaments present, 2–3 µm wide, gradually widening to 4 µm towards apex. Hypothecium red-brown, K+ dulling or K+ dull olivaceous. Asci c. 70×12–14 µm, 8-spored. Ascospores (14–)17–21(–24)×(4·8–)6·0–7·0(–9·5) µm, narrowly ellipsoid, brown, smooth-walled but wall of old spores ± finely warted, (1–)3-septate.
Conidiomata not seen.
Dactylospora suburceolata occurs in similar habitats to D. urceolata (Th. Fr.) Arnold [incl. D. deminuta (Th. Fr.) Triebel], but has generally larger apothecia, a broader exciple, a taller hymenium, and persistently 3-septate ascospores (Fig. 3) that are slightly broader (Table 1). Furthermore, the purplish tinge (in water or K) of the internal brownish tissues of D. urceolata is not evident in D. suburceolata, and neither are the blue-violet, K+ green pigmented granules found in the exciple and hypothecium of that species. Dactylospora urceolata has 3–7-septate ascospores, with some becoming submuriform with 1–2 cells having a longitudinal or diagonal septum (Triebel Reference Triebel1989, fig. 32a as D. deminuta). However, it is variable with regard to ascospore septation, with some collections having a high proportion of 3-septate spores. Dactylospora frigida Hafellner also has 7-septate or submuriform ascospores, but differs from both D. urceolata and D. suburceolata in having a greenish brown epithecium, and is apparently confined to the thallus of Brigantiaea fuscolutea (Hafellner Reference Hafellner1985). To date, no Dactylospora species has been found on B. fuscolutea in the British Isles, and the listing of D. frigida in Hawksworth (Reference Hawksworth2003) refers to D. urceolata. Other species of Dactylospora with brown, 3-septate spores include D. borealis Holien & Ihlen, which occurs on Mycoblastus spp. and has a K+ purple epihymenium, and D. parasitica (Flörke) Zopf and D. attendenda (Nyl.) Arnold, both of which have shorter spores and occur on Ochrolechia and Pertusaria, and Porpidia and Amygdalaria spp. respectively. A key to the Scandinavian species of Dactylospora is provided by Ihlen et al. (Reference Ihlen, Holien and Tønsberg2004).
Dactylospora pseudourceolata Sarrión & Hafellner (2002), from SW Spain, is superficially similar to D. suburceolata, but has somewhat smaller apothecia and ascospores (Table 1) and is a saprobic species on the bark of Arbutus unedo L. Dactylospora rostrupii Alstrup (Alstrup et al. Reference Alstrup, Christensen, Hansen and Svane1994) is lichenicolous on the terricolous Pertusaria dactylina (Ach.) Nyl., but differs especially in its broader ascospores (Table 1).
In Scotland, D. suburceolata is so far known from only four collections from the Breadalbane mountains in Perthshire, with an altitude range of 750–1000 m. Here, it is lichenicolous on sterile, whitish to pale grey muscicolous crusts, which may belong to ‘Lecidea’ hypnorum Lib. One host thallus (from Coire Riadhailt) has pale yellow soralia and may be a species of Biatora. However, the single collection from Switzerland, at an altitude of 1518 m, is seemingly on the thallus of Mycobilimbia tetramera (De Not.) Vitik. et al., which was also parasitized by Zwackhiomyces cf. berengerianus (Arnold) Grube & Triebel. All British records of D. urceolata are also from the Breadalbane Mountains, with hosts being Megaspora verrucosa, Protopannaria pezizoides, Protothelenella sphinctrinoidella, and unidentified sterile crustose lichens.
Additional specimens examined. Great Britain:Scotland: V. C. 88, Mid-Perthshire: Meall na Samhna, 27/47.33, 750 m, damp N-facing, mica-schist crags, 1991, Fryday 2388 (E); Meall nan Tarmachan, crags on W side of Coire Riadhailt, 27/57.38, 800 m, 24 vii 1989, Fryday s. n. (E); Ben Ghlas, northern crags, 27/62.40, 1000 m, 4 viii 1989, Fryday s. n. (E).—Switzerland:Schwyz: Muotathal, Bödmerenwald forest reserve, Piceetum subalpinum (on limestone), with few, scattered beech and sycamore, c. 8°50′50″E, 46°58′58″N, 1518 m, over Mycobilimbia tetramera, with Zwackhiomyces aff. berengerianus on saxicolous mosses, 1992, U. Groner 1376 (hb. Groner).
Fuscidea oceanica Fryday & Coppins sp. nov.
MycoBank No: MB800466
Fuscidea gothoburgense similis sed soraliis minoribus, punctiformibus, 0·1–0·2 mm diam., sorediis caesiis, apotheciis frequentibus sessilibus differt.
Typus: Great Britain, Scotland, V. C. 88, Mid-Perthshire, Creag Mhòr, Coire-cheathaich, 27/40.34, 550 m, on quartzite intrusion on side of large schistose boulder, 1993, Fryday 2358 (E—holotypus).
(Fig. 4)
Fuscidea oceanica (Fryday 2358—holotype). Scales: A=0·5 mm; B=0·2 mm. In colour online.
Thallus usually wide-spreading, cracked-areolate, pale grey to (rarely) brown, areoles flat, 0·15–0·40 mm diam. Soralia mostly punctiform arising from the centre of each areole, 0·1–0·2 mm diam., less often following cracks in the thallus and becoming stellate; soredia blue-grey (K−, N−), 20–25 µm diam. Photobiont chlorococcoid, cells thick-walled, 7–16(–19)×8–12(–13) µm, often dividing to give 2–4 daughter cells (Chlorella-type).
Apothecia black, lecideine, initially innate with a white ring around the inner edge of the exciple but soon becoming sessile, 0·5–1·0 mm diam., proper exciple persistent, becoming flexuose; disc occasionally umbonate. Excipulum of branched, anastomosing hyphae, colourless with dark brown outer cells, reflexed below apothecium. Hymenium 70–90 µm tall, upper third with brown pigment; paraphyses simple, 2·5 µm wide, swelling to 5 µm at apex, with dark brown hood. Hypothecium hyaline. Asci c. 45×20 µm, Teloschistes-type; ascospores hyaline, simple, broadly ellipsoid, 7–9×5–6 µm.
Conidiomata not seen.
Chemistry. Thallus C−, K−, Pd−, UV−; soredia and medulla C−, K−, Pd−, UV+ white; divaricatic acid detected by TLC.
Fuscidea oceanica is unusual amongst lichenized fungi in regularly producing both sexual and asexual propagules. It is the only sorediate member of the genus occurring in the British Isles that is also usually fertile. However, as the soredia of F. oceanica are very small and inconspicuous they are easily overlooked in the field and this species may then be mistaken for other, non-sorediate, members of the genus (e.g., F. lygaea). Other sorediate saxicolous species of Fuscidea containing divaricatic acid are F. gothoburgensis (H. Magn.) V. Wirth & Vězda and F. recensa (Stirt.) Hertel et al., both of which are very rarely fertile. Fuscidea recensa is a very different species with a thicker thallus, more wide-spreading, often convex soredia, and curved ascospores, whereas F. gothoburgensis differs mainly in thallus morphology and ecology, having a much more dispersed thallus and usually occurring in shaded underhangs. The sorediate counterpart of F. intercincta, which has been recognized as F. oculata Oberholl. & V. Wirth, differs in having ± innate apothecia with a white margin and erumpent, cream-coloured soralia.
Two further sorediate species were recognized by Clauzade & Roux (Reference Clauzade and Roux1985) but have not been formally described. Fuscidea ‘alpina’, known from four collections from the Austrian and Italian Alps, is similar to F. oceanica but has brownish soralia. In three of the collections these tend to be scattered over the thallus as individual soredia, often appearing as minute coralloid isidia, whereas the other collection, which is from the same locality and date as one of the previous collections, has more discrete soralia and is very similar in appearance to F. oceanica. Unfortunately, all four collections lack apothecia and, given the geographic and environmental separation of the two populations, we have decided not to include these collections in our new species. Fuscidea ‘badensis’, the other species mentioned by Clauzade & Roux (Reference Clauzade and Roux1985), is referable to F. recensa (Stirt.) Hertel et al.
Fuscidea oceanica is locally common on exposed, hard acidic rocks in the NW Scottish Highlands and also occurs at scattered localities elsewhere in the Highlands. It is particularly common on Cambrian quartzite, and on the Beinn Eighe NNR in West Ross it covers large areas and is the most frequent species. Associated species are few but include Allantoparmelia alpicola, F. gothoburgensis, F. intercincta, Ionaspis odora and Rhizocarpon geographicum aggr.
Specimens examined. Great Britain:Scotland: V. C. 92, South Aberdeen: Braemar. Creag Choinnich, west side, on top of ‘tor-like’ outcrop by path, 37/1596.9173, 500 m, on granite, 2004, Coppins 21467 (E). V. C. 97, West Inverness: c. 7 km NE of Strontian, NE of Ariundle Wood, disused mine, 17/86.66, 240–260 m, top of large boulder, 1992, Coppins (15346) et al. (E). V. C. 98, Argyll Main: Ben Starav, 27/13.42, 850 m, shaded side of granite boulder, 1991, Fryday (2341) & P. Goddard (MSC); Glen Coe, Coire Gabhail (below ‘Lost Valley’) 27/16.55, 450 m, vertical acid rocks in damp Betula wood, 1992, Fryday 3281 (E); Aonach Eagach, S side of E ridge of Sgurr nam Fiannaidh, 27/14.58, 900 m, top of exposed granite rock, 1992, Fryday 3327 (MSC); Creag nan Gabhar, 27/13.57, 600 m, top of acid boulder, 1992, Fryday 3329 (MSC). V.C. 105, West Ross: Beinn Eighe NNR, ridge between Spidean Coire nan Clach and Coinneach Mhòr, 18/96.59, c. 950 m, Cambrian quartzite crags, 1991, Fryday 2734 (E); ibid., Fryday 2725 (MSC).
Lecania granulata Coppins & Fryday sp. nov.
MycoBank No: MB800467
Lecaniae subfusculae similis sed thallo blastidiato granulato, crassiore, apotheciis initio cum excipulo thallino crenulato vel granulato, disco semper pallido.
Typus: Great Britain, Scotland, V. C. 111, Orkney, Hoy, coast by Wastlee Moor, Kilns of Hawick, ND/251.886, on coastal turf ledges, 21 August 2008, S. G. Price [Coppins 22942] (E—holotypus).
Lecania granulata [S. G. Price (Coppins 22942—holotype)]. A, mature apothecia (with Lecania subfuscula bottom left); B, immature apothecia. Scales=1·0 mm. In colour online.
Lecania granulata, ascospores [S. G. Price (Coppins 22942—holotype)]. Scale=10 µm.
Thallus whitish to pale buff, densely granular-blastidiate, to 1·5 mm thick; granules 30–80(–100) µm diam. Photobiont chlorococcoid, cells 5–12 µm diam. or ellipsoid to 14×12 µm.
Apothecia (0·2–)0·3–0·7 mm diam., plane or becoming convex and up to 0·9 mm, each developing within a thalline wart then becoming urceolate with a crenulate thalline margin and concave disc (Fig. 5), later expanding further with the thalline margin becoming granulate or receding to reveal a proper margin, and with the disc becoming plane to convex. Disc pale, pinkish or pallid, often with patchy brown pigment towards the margin in older apothecia, with paler proper margin. Thalline exciple c. 40–60 µm wide. Proper exciple lateral to hymenium 35–60 µm wide, but not clearly delimited from hypothecium below, hyaline or dilute reddish brown, in outer half composed of ± radiating hyphae with lumina to 5·0×2·5 µm, cortical cells c. 5–6 µm diam. Hymenium 50–65 µm tall, hyaline or tinged dilute reddish brown in part in older apothecia; epihymenium hyaline or with patchy dilute reddish brown pigment, especially towards the margin; subhymenium 20–35 µm tall, K+ dilute yellow, with cells of ascogenous hyphae to 5 µm wide; paraphyses simple, c. 1·5 µm wide in mid-hymenium, but upper 2–4 cells gradually widening to 4(–6) µm wide, ± lax in K. Hypothecium massive, of irregularly orientated, slender hyphae c. 1·0–1·7 µm wide. Asci narrowly clavate, 53–60×9–11 µm, Bacidia-type; ascospores narrowly clavate-fusiform to shortly acicular, 20–33×(2·0–)2·5–2·7(–3·0) µm, 3–5(–7)-septate.
Conidiomata not seen.
Chemistry. Thallus C−, KC−, K−, Pd−, UV−; no substances detected by TLC.
Because of its shortly acicular ascospores and Bacidia-type asci, this species was originally filed under Bacidia. However, the presence of a thalline margin indicates a more appropriate placement in the genus Lecania, as circumscribed by Ekman (Reference Ekman1996: 41). Reese Naesborg et al. (Reference Reese Naesborg, Ekman and Tibell2007) introduced a revised circumscription for Lecania s. str. that included a crustose thallus, with or without blastidia, that usually lacks lichen substances, and lecanorine to biatorine apothecia; branched, but not anastomosing excipluar hyphae; an unevenly distributed epihymenial pigment that is usually yellowish or brown; gradually widening paraphyses that are lax in K; and 0–3(–5)-septate, and often curved, ellipsoid, fusiform, or bacilliform ascospores. Lecania granulata is consistent with all these characters and so clearly belongs in Lecania s. str..
Lecania granulata has somewhat similar ascospores to L. subfuscula (Nyl.) S. Ekman, which can also inhabit nutrient-enriched, coastal turf. Lecania granulata differs in having a thickly granular (blastidiate) thallus and a crenate to granular thalline margin. Lecania subfuscula (Fig. 5a) has a generally thinner and warty granular thallus, with coherent ‘granules’ 40–200 µm diam., and usually darker pink to grey-brown or even blackish apothecia, with little hint of a thalline margin (Fletcher et al. Reference Fletcher, James, Purvis, Smith, Aptroot, Coppins, Fletcher, Gilbert, James and Wolseley2009). Lecania cuprea van den Boom & Coppins is another species with narrow, multiseptate ascospores, but it has a thinner thallus, no thalline margin, smaller paraphysis-apices, and inhabits shaded limestone rocks.
This distinctive species was first discovered by Oliver Gilbert during expeditions to the remote Hebridean islands of North Rona and the Flannan Isles and reported as “Bacidia sp.” (Gilbert et al. Reference Gilbert, Holligan and Holligan1973; Gilbert Reference Gilbert1976; Gilbert & Wathern Reference Gilbert and Wathern1976). It grows on coastal, peaty turf near cliff tops or on cliff ledges. At the Orkney locality it was associated with Anaptychia runcinata, Lecania subfuscula and Lecanora zosterae. In the cliff-top maritime grassland on Foula, its associated species included Anisomeridium polypori, Catapyrenium cinereum, Lecania subfuscula, Lecidea berengeriana, Leptogium imbricatum and Psoroma hypnorum.
Additional specimens examined. Great Britain:Scotland: V. C. 110, Outer Hebrides: North Rona, on peat near the top of the north slope, HW/83, 1972, O. L. Gilbert (E); ibid., above Leac na Sgrob, HW/83, bare peat, rare, vi 1976, O. L. Gilbert (E); Flannan Isles, Eilean Mòr, NA/7246, collected once on peat near cliff top, vi 1975, O. L. Gilbert (E). V. C. 112, Shetland: Foula, Stremness, HT/9693 4116, in cliff-top maritime grassland, 27 vii 2011, A. Acton (hb. Acton).
Lecanora viridiatra (Stenh.) Nyl.
MycoBank No: MB389573
Flora 55: 251 (1872).
Biatora viridiatra Stenh. in E. Fries, Lichenogr. Europ. Reform.: 277 (1831).—Lecidea viridiatra (Stenh.) Schaer. non (Wulfen) Ach. (1803), in Enumer. Critic. Lich. Europ.: 108 (1850).
Lecidea luteoatra Nyl., Flora 61: 299 (1873).
This species is locally frequent on the Cambrian quartzite of the NW Highlands and in the British Isles is also known from several localities in the Cairngorm Mountains, as well as Ben Lawers in the Central Highlands. The Lecanora-type asci and filiform conidia strongly suggest that this species belongs in Lecanora, and the lecideine apothecia, blue epihymenium, and thalline chemistry (usnic acid) refer it to the L. marginata group.
Stenhammar's name is the earliest available for this species but was not available in Lecidea because of Lecidea viridiatra (Wulfen) Ach. (1803), which is Rhizocarpon viridiatrum (Wulfen) Körb. However, it becomes available in Lecanora and so is taken up here.
Selected specimens examined. Austria:Salzburg: Radstädter Tauern, Hänge östlich oberhalb der Hundsfeldalpe, NE oberhalb Obertauern, am Radstädter Tauern paß, 47°15′N, 13°35E, 1900–2000 m, S-W exponierte, steile, mitunter kurzzeitig wasserüberrieselte Gneiswand, 6 ix 1981, J. Poelt (E; Hertel: Lecid. Exsic. nr. 72).—Great Britain:Scotland: V. C. 89, East Perthshire: The Cairnwell, 37/1.7, c. 900 m, on mountain top on quartzite boulders, 23 viii 1990, C. J. B. Hitch (E). V. C. 94, Banff: Ben Avon, Lochan nan Gabhar, 38/14.03, 2500 ft, rock above corrie, 1975, Coppins (4618) & P. Harrold (E).
Lecidea herteliana Fryday & Coppins sp. nov.
MycoBank No: MB800468
Lecideae pauperculae similis sed thallo minore pallidore et substantiis lichenicis absentibus.
Typus: Great Britain, Scotland, V. C. 97, West Inverness, c. 7 km NE of Strontian, NE of Ariundle Wood, disused mine, 17/86.66, 240–260 m, on stones, 1992, Coppins (15340) et al. (E—holotypus).
(Fig. 7a)
Lecidea herteliana (Coppins 15340 et al.—holotypus). B, Lecidea paupercula (Coppins 10451). Scales: A & B=0·5 mm. In colour online.
Thallus effuse, rarely more than 1–2 mm across; areolate, areoles flat to slightly convex, irregular, 0·2–0·4 mm across, grey to pale brown often with a paler margin, contiguous or somewhat dispersed on a black hypothallus; cortex 10–20 µm thick with pale brown (rarely red-brown or ± hyaline) pigmented upper layer 7–10 µm thick; epinecral layer thick, 45–95 µm, 20–25 µm at edge of areoles; medulla and upper cortex I+ blue. Photobiont chlorococcoid 9–15 µm.
Apothecia black, lecideine, 0·4–0·6 mm diam., sessile, slightly convex with a thin (0·05 mm), barely raised margin. Excipulum dark blue-black with swollen (to 5 µm) cortical cells. Hymenium 90–105 µm tall; epihymenium c. 10 µm tall, sharply delimited, blue-black (H+ blue, N+ red; cinereorufa-green); paraphyses sparingly branched and anastomosing, c. 1·5–2·0 µm thick swelling at apex to 5 µm with a dark blue-black cap. Hypothecium dark brown. Asci cylindrical to slightly clavate, 45–50×12–15 µm, Lecidea–type; ascospores simple, hyaline, 12–14×5–6 µm.
Conidiomata not observed.
Chemistry. All spot tests negative; no substances by TLC.
Etymology. The specific epithet honours our friend and colleague Professor Dr Hannes Hertel (München) for his outstanding contributions to the understanding of the genus Lecidea over the past 45 years.
Lecidea herteliana is known from several sites in the Scottish Highlands (Isle of Mull, Rannoch, Strontian, East Perthshire, Angus) and is also present in north-eastern North America (Maine). It has a small (usually <2 cm diam.), pale grey-brown, atrobrunnea-type thallus with an I+ blue medulla but all other spot tests are negative. It most closely resembles L. paupercula Th. Fr. (Fig. 7b) but that species has a more wide-spreading, red-brown to dark grey-brown thallus that contains stictic acid (K+ yellow, Pd+ orange). Lecidea paupercula also differs in having flat to slightly concave areoles with a dark margin, red-brown pigmented cortical cells and a much thinner epinecral layer (20–35 µm), as well as flat ± innate apothecia. Associated species include Porpidia flavocruenta and Pertusaria corallina.
The two collections from NE North America agree well with British material, but have a somewhat thinner epinecral layer (25–50 µm).
Additional specimens examined. Great Britain:Wales: V. C 49, Caernarvon: Carnedd Llewellyn, Clogwynyreryr, 23/71.66, c. 450 m, boulder by track, ix 1997, Fryday s. n. (MSC). Scotland: V. C. 90, Angus: Caenlochan Glen, Glasallt Burn, 37/183.767(–772), 600–700 m, 1989, Coppins (13334) O. L. Gilbert & R. K. Brinklow (E). V. C. 98, Argyll Main: Bridge of Orchy, Forest Lodge, Coire Toaig, 27/44.25, 400 m, top of schistose boulder, 1992, Fryday 3040 (M). V. C. 103, Mid Ebudes: Isle of Mull, Ardmeanach, Bearraich, 17/41.27, 250 m, basalt outcrops and boulders, 1993, Fryday 4440 (E).—USA:Maine: Piscataquis Co., Baxter State Park, Katahdin, North Basin, between ponds and ‘calcareous seep’, 45°55·75′N, 68°55·10′W, 1020 m, on upper surface of granitic boulder, 2003, Fryday 8580 (MSC); ibid., South Basin, bottom of Chimney Gully, 45°54·4′N, 68°54·7′W, 1120 m, on granitic pebbles, 2004, Fryday 8830 (MSC).
Roccellographa sorediata (Sparrius, P. James & M. A. Allen) Coppins & Fryday comb. nov.
MycoBank No: MB800470
Sclerophytomyces circumscriptus var. sorediatus Sparrius et al., Lichenologist 37: 285 (2005) [Basionym].—Peterjamesia sorediata (Sparrius et al.) D. Hawksw., Lichenologist 38: 189 (2006).
In their landmark paper on the phylogeny of the Arthoniales, Ertz & Tehler (Reference Ertz and Tehler2011) found that the genus Peterjamesia D. Hawksw. should be included under the earlier generic name Roccellographa J. Steiner. They transferred the type species of the former genus, P. circumscripta [syn. Sclerophyton circumscriptum (Leight.) Zahlbr.] into Roccellographa but did not do so for the related, sterile taxon, P. sorediata. This is rectified here.
Ropalospora lugubris f. sorediata Fryday & Coppins forma nov.
MycoBank No: MB800469
Ropalosporae lugubri f. lugubri similis sed sorediis praesentibus.
Typus: Great Britain, Scotland, V. C. 98, Argyll Main, Glen Coe, Coire nam Beitheach 27/142.545, 875 m alt., on east-facing rhyolite crag in dry underhang, 15 July 1992, A. M. Fryday 3347 (E—holotypus).
Lecidea atroumbrina H. Magn, Bot. Notiser 1930: 468 (1930).—Ropalospora atroumbrina (H. Magn.) S. Ekman, Bryologist 96: 590 (Reference Ekman1993); type: Sweden, Bohuslän, Forshälla, Stora Hasselön, on a gently sloping rock facing the north, 26 July 1930, A. H. Magnusson (UPS!—lectotype; Magnusson, Lich. sel. Scand. exs. Exsic.-Nr 88).
(Fig. 8)
Ropalospora lugubris f. sorediata. A, farinose soralia (with apothecia; Fryday 3347—holotype); B, granular soralia (Fryday 2237). Scales: A=0·5 mm; B=0·1 mm. In colour online.
Thallus crustose, dark grey (often purple tinged) or pale to dark brown, cracked areolate, areoles plane to verrucose. Soralia c. 0·1–0·5 mm diam., initially discrete but becoming confluent, soredia initially concolorous with, or darker than the thallus, coralloid-granular, becoming pale yellow-brown, farinose. Photobiont chlorococcoid, cells thick-walled, 7–16(–19)×8–12(–13) µm, often dividing to give 2–4 daughter cells (Chlorella-type).
Apothecia very rare, mature apothecia known only from the type collection. Identical to R. lugubris f. lugubris.
Conidiomata pycnidia, frequent, dark brown to black, 0·2–0·3 mm diam., immersed in the thallus; conidia bacilliform, 6–8×0·8 µm.
Chemistry. C−, K−, KC−, Pd−; TLC revealed an unidentified substance (?terpenoid) at Rf 4 in A, 5 in C, and 4 in G that is colourless, UV+ white, brownish pink, UV+ cream-pink after developing, accompanied by two accessories; one with the same characteristics at about Rf 3 in A, 3–4 in C and G, and the other UV+ blue-white after developing at Rf 4–5 in C. Traces of atranorin or another unidentified substance at Rf 3–4 in solvent C that is yellow, UV+ dark after developing were also rarely present.
Species that usually reproduce by apothecia also occasionally produce soredia, but these are usually rare, occur only within normal fertile populations, and the sorediate specimens are always fertile. Some of these are recognized taxonomically [e.g., Fuscidea cyathoides var. sorediata (H. Magn.) Poelt,] whereas others are not [e.g., the sorediate form of Ochrolechia parella (L.) A. Massal.]. Ropalospora lugubris f. sorediata, however, is very rarely fertile and is also more frequent and widespread than the non-sorediate form. Because of this we consider it appropriate to afford it some taxonomic recognition, and that of forma seems most suitable.
Ekman (Reference Ekman1993) separated R. atroumbrina, which is not known fertile, from sorediate R. lugubris by the different ontogeny of the soralia and also by the presence of atranorin in R. atroumbrina. However, the much larger number of collections of R. lugubris f. sorediata available to us indicates that, although there appear to be two different types of soredia, those of ‘atroumbrina’ usually being composed of dark, granular isidia whereas those of R. lugubris f. sorediata are often cream-coloured and farinose, the two forms intergrade. Initially the soredia of both ‘taxa’ are granular-coralloid and concolorous with, or darker than, the thallus and often coalesce to cover areas up to 5 mm in diameter. However, these later ‘burst' (or abrade) giving rise to smaller cream-coloured farinose soredia. In older soralia there is sometimes very little evidence of the original form, but careful examination usually reveals some darker, granular soredia around the edges of at least some soralia. This ‘bursting’ of the granular soredia is most evident in R. lugubris f. sorediata but can also be observed in some collections of ‘atroumbrina’, including the type specimen. The chemical difference reported by Ekman (Reference Ekman1993) is also not consistent. We performed TLC on four collections each of R. atroumbrina, R. lugubris f. lugubris and R. lugubris f. sorediata and found no significant differences. One collection of R. lugubris f. sorediata contained atranorin, whereas all the other collections of the three taxa lacked this substance. Atranorin has also been reported from R. lugubris f. lugubris (A. Orange, pers. comm.), indicating that this substance is occasionally present in all three ‘taxa’. The main chemical constituent (see above) was present in all samples analyzed and the only other difference was that the accessory at Rf 4·5 was sometimes feint or apparently absent in some collections of all three taxa, and the accessory at Rf 3·5 was yellow-orange in R. atroumbrina and mauve in both forms of R. lugubris, although one collection of f. sorediata had both yellow and mauve spots.
Consequently, as we can find no significant morphological or chemical differences between the two taxa, R. atroumbrina is included here in the synonym of R. lugubris f. sorediata. We have decided to introduce the new epithet ‘sorediata’ rather than take up ‘atroumbrina’ because that epithet describes the colour of both forms.
This taxon was first reported from Scotland (as Fuscidea sp. A) by Gilbert et al. (Reference Gilbert, Coppins and Fox1988: 238) and has subsequently been found to be widespread on acidic rocks throughout the western Highlands. Similarities of thallus morphology and chemistry, and ecology, suggested that it may be the sorediate counterpart of R. lugubris, and the discovery of a population with mature apothecia confirmed this opinion. It is generally more frequent than R. lugubris f. lugubris and has also been reported from North Wales, Scandinavia [as Ropalospora atroumbrina (H. Magn.) S. Ekman], and NE North America. A report of R. lugubris from the English Lake District supported neither apothecia nor soralia (A. Orange, pers. comm.) and so it is not possible to ascertain to which forma it should be referred.
Species associated with Scottish collections include Coccotrema citrinescens, Fuscidea gothoburgensis, F. intercincta, Lepraria lobificans, Pertusaria corallina, Porpidia spp., and Rhizocarpon geographicum aggr.
Additional specimens examined. Sweden:Halland: Onsala par., Gottskär, on oxidated rock, 1921, A. H. Magnusson 5271 (UPS). Bohuslän: Långelanda par., Gömme, on steep rock, N side, 1926, A. H. Magnusson 10046 (UPS). Gotland: Hörsne, NÖ hörnet vid gränsen till Vallstena, exponerat urbergsblock nära landsvägen, riklig, 13 vi 1943, G. Degelius (UPS). Södermanland: Nämdö, Nämdö Böke, öppen leptitklippa på nordsidan, 1 ix 1943, G. Degelius (UPS). Uppland: Danmark, Karlsro, Nordsiden av klippvägen, samhällsbildande, 14 x 1945, G. Degelius (UPS). Gästrikland: Hille, 350 m SSE of the parish church, 10 m 60°43′52·1″N 17°11′10·2″E (RT90), on N side of roadside boulder, 2007, A. Nordin 6456 (UPS).—Great Britain:Wales: V. C. 49, Caernarvon: Carnedd Llewellyn, Foel Grach, 23/68.65, c. 950 m, shaded vertical side of boulder, ix 1997, Fryday s. n. (MSC); Glyder Fawr, near Llyn y Cwn, 23/639.586, 700 m, on shaded rock, 1998, A. Orange 12068 (NMW). Scotland: V. C. 88, Mid-Perthshire: Glen Lyon, Cairn Gorm, 27/63.48, c. 2700 ft., ±underhang, dry, mica-schist, 1977, Coppins 3055 (E); Blair Atholl, Tulach Hill, 27/8.6, N-facing side of old wall, 1988, Coppins (12905) & O. W. Purvis (E); Ben Ghlas, on coll, 27,620.408, 20 vii 1986, B.W. Fox, s. n. (E); ibid., crags at head of Coire Odhair, 27/62.40, 900 m, on west-facing mica-schist crags, 3 vii 1989, Fryday s. n. (E); Beinn Heasgarnich, Creag na h-Acharich, 27/42.38, 750 m, exposed acidic crag, 1991, Fryday 2237 (E), 2238–9 (MSC) [with immature apothecia]; ibid., north ridge, 27/42.38 950 m, mica-schist boulder, 1991, Fryday 2834 (MSC); ibid., 27/4204.3884, 950 m, on vertical siliceous rocks, 2006, A. Orange 16514 (NMW); ibid., S of Meall a' Chall, 27/4357.4004, 710 m, 2006, A. Orange s. n. (NMW); Creag Mhòr, N-E Crags, 27/39.36, 800 m, mica-schist underhang, 1991, Fryday 2332 (MSC); Ben Vorlich, 27/62.19, c. 600 m, on underside of schistose boulder, 1992, Fryday 3052 (MSC). V. C. 90, Angus: Caenlochan Glen, N-facing cliffs below Little Glas Maol, 37/17.76, c. 800 m, sheltered, vertical, siliceous cliff face, in gulley, 1989, Coppins (13405 ) & O. L. Gilbert (E); Coire na Berrach, 37/442.772, slightly underhanging rock-face, 2000, R. C. Munro s. n. (E). V.C. 97, West Inverness: Fort William, Glen Nevis, Meall Cumhann, 27/17.69, 500 m, exposed vertical acid rock, 1990, Fryday 1240 (E); Sunart, Strontian River, 17/86.66, 250 m, acid rock in ravine below disused metal mine, 1992, Fryday 3174 (E); ibid., Beinn Resipol, 17/75.65, 450 m, vertical, shaded acidic rock, 1992, Fryday 3193 (E). V. C. 98, Argyll Main: Glen Coe, Achnambeithach, 27/13.56, 200 m, underside of ?rhyolite boulder, 1992, Fryday 3348 (E). V. C. 104, North Ebudes: Isle of Skye, Bla Bheinn, An Carnach, 18/55.20, 200 m, east-facing basalt crag, 1991, Fryday 2390 (MSC), 2391 (E); ibid., Quiraing (Cuithearaing), 18/45.69, 250 m, exposed basalt crags, 1991, Fryday 2452 (E). V. C. 105, West Ross: Kinlochewe, Beinn Eighe NNR, Coille na Glas-leitir, valley of Allt Doire Daraich, 28/00.62, c. 250 m, upper end of rock outcrop by stream, 2000, Coppins (19766) & A. M. Coppins (E); Lochcarron, c. 1 km W of village, NE-facing crags of hill “212”, 18/885.399, c. 200 m, on vertical rocks, 2001, Coppins (20997) & A. M. Coppins (E). V.C. 108, West Sutherland: Creagan Meall Hourn, NE-facing crags above An Dubh Loch, 29/34.45, c. 475 m, shaded underhang, 1991, Fryday 2952 (E). V. C. 110, Outer Hebrides: West Lewis, Glen Raonasgill, Teinnasval, 19/03.26, shaded acidic (?granite) boulder, 1991, Fryday 2507 (E); North Harris, Glen Ulladale, Creagan Leathan, 19/07.12, exposed east-facing crag, 1991, Fryday 2612 (E).—USA:Maine: Piscataquis Co., Baxter State Park, Katahdin, north basin, east of the ‘calcareous seep’, 45°55·8−N, 68°55·5−W, 1120 m, on exposed vertical, granitic rock-face, 2004, Fryday 8866, 8868 (MSC).
The first author acknowledges the receipt of a BLS Overseas Members' Travel Fund award, which allowed him to spend two weeks in Edinburgh working with the second author. We thank Alan Orange (Cardiff) for sharing the results of his investigations into the chemistry of Ropalospora lugubris with us, and the curators of GZU, STU and UPS for the loan of comparative material of Fuscidea ‘alpina’, F. ‘badensis’, F. oculata, and Ropalospora atroumbrina. Andy Acton and Urs Groner are also thanked for the loan of material from their private herbaria.
