Biatora
Crust lichens containing a trebouxioid alga; thallus various including smooth, areolate, granular, verrucose small-squamulose and sorediate; apothecia of various colours, generally pale or medium tones of tans, brownish pink or orangish brown, black or white in some species; apothecial sections lacking POL+ crystals in most species; rim a non-algal exciple formed of cylindric hyphae embedded in gel, arranged fan-like as seen in cross section; hymenium of straight, mostly unbranched paraphyses embedded in gel, often distinctly pigmented in upper portions, or streaky-pigmented throughout; hypothecium colourless or, in a few species, pigmented; asci clavate, with a thick IKI+ tholus with an axial mass that is, in most species, blunt at the tip and lined with darker blue in iodine stain; spores 8 per ascus in most species, simple or tansverse-septate, elliptic, oblong or fusiform, colourless.
References: Printzen 1995; Printzen & Tønsberg 1999, 2003; Spribille et al. 2009.
| 1a. | Thallus sorediate or formed of isidioid granules …2 |
| 2a. | Thallus of weakly corticate granules about 150-300 µ wide, mustard yellow to yellow-green, all spot tests negative; apothecia unknown …Biatora sp. nov. “gold nuggets” |
| 2b. | Thallus sorediate or, if formed of corticate granules, then these more green than yellow and PD+ orange-red …3 |
| 3a. | Thallus olive-green, grass-green or dark green, formed of tiny, easily detached squamules that often have isidioid branchlets or, rarely, sorediate patches, C- or C+ reddish, K-, P+ orange-red, UV+ dull whitish; spores 8-15 × 3-5 µ; rare …Biatora fallax |
| 3b. | Thallus of whitish or dingy colours, never grass-green nor dark green, lacking squamules, soredia common, sometimes leprose, spot test reactions various; spores various; common …4 |
| 4a. | Thallus on rough bark or on moss; soralia sometimes quickly confluent and the thallus becoming leprose; regionally common species found in both cold and warm forest types …5 |
| 5a. | Spores 12-15.5 × 3.5-6 µ; apothecia in section C-; soredia generally more abundant than corticate portions of the thallus …Biatora chrysantha |
| 5b. | Spores 9-13 × 3-3.5 µ; apothecia in section C+ reddish; soredia generally less abundant than corticate portions of the thallus …Biatora chrysanthoides |
| 4b. | Thallus on smooth bark of acid-bark shrubs, especially of Ericaceae …6 |
| 6a. | Soredia C+ pink or red, K-, P-; mostly on ericaceous shrubs at upper forested elevations; spores 7.5-14 × 2.5-4 µ …Biatora vacciniicola |
| 6b. | Soredia C-, K- or + yellowish, P- or + orange-red; substrate and elevations various; spores 7.5-22 × 2.5-5 µ …7 |
| 7a. | Soredia P+ orange-red, K-; spores, if present, 9-22 × 3-5 µ; upper hymenium lacking granules; typically growing at lower elevations …Biatora efflorescens |
| 7b. | Soredia P-, K+ dingy yellow-orange; spores, if present, 7.5-13.5 × 2.5-3.5 µ; typically growing at higher elevations …Biatora flavopunctata |
| 1b. | Thallus lacking soredia and isidioid granules …8 |
| 8a. | Hymenium blue-greenish or purple; apothecia grey to black, sometimes with a greenish or bluish tone as seen under the dissecting scope …9 |
| 9a. | Pigments of the hymenium limited to the upper 5-10 µ, unpigmented below that; spores only 2-2.5 µ wide …Biatora globulosa |
| 9b. | Upper and main portions of the hymenium pigmented with dark blue-green pigment streaks; spores 3-4 µ wide …Biatora hypophaea |
| 8b. | Hymenium colourless or at any rate lacking blue-green pigments …10 |
| 10a. | Hypothecium obviously darker than the hymenium; spores mostly simple, 8-17 × 3-6 µ …Biatora subduplex |
| 10b. | Hypothecium unpigmented or only very weakly so; spores various …11 |
| 11a. | Upper hymenium with abundant POL+ granules; spores 9-18 × 2-3 µ, 3-septate …Biatora pallens |
| 11b. | Upper hymenium POL-, lacking any distinct thick layer of granules; spores various …12 |
| 12a. | Upper 5-10 µ or so of the hymenium with a layer of obviously darker pigments, K+ greenish; apothecia blackish; spores 2-2.5 µ wide; asci Bacidia-type …Biatora globulosa |
| 12b. | Upper hhymenium without any obviously darker pigmented layer, any pigments not confined to a distinct layer; apothecia mostly pale colours: pinks, tans, orangish, red-browns; spores more than 2.5 µ wide; asci various …13 |
| 13a. | Spores mostly 3-septate …14 |
| 14a. | Apothecia up to 0.7 mm wide; exciple IKI+ purplish; spores 11-18 × 3.5-6.5 µ (mostly 4-5.5 µ) wide; common in wet forests …Biatora rufidula |
| 14b. | Apothecia up to 1.5 mm wide; exciple IKI-; spores 12-19 × (5‑) 5.5-8.5 µ; occasional in wet forests in inland BC, to be sought …[Biatora nobilis Printzen & Tønsberg] |
| 13b. | Spores mostly 0- to 1-septate …15 |
| 15a. | Thallus mostly granular, growing within the winter snowpack over mosses or spreading onto adjacent bark or wood …16 |
| 16a. | Thallus greenish grey; subhymenium and exciple not especially pigmented; spores (10‑) 13-20 (‑25.5) × (4‑) 4.5-6 (‑7); rare …Biatora vernalis |
| 16b. | Thallus whitish; subhymenium and exciple often of an obviously darker brown (sometimes also the hypothecium); spores (8‑) 9.5-15 (‑21) × (3‑) 3.5-5 (‑6) µ.; common …Biatora subduplex |
| 15b. | Thallus mostly not granular, not on mosses, some species growing above the winter snowpack …17 |
| 17a. | Thallus and apothecia in section C+ red; thallus mostly vivid dark green (at least when fresh) formed of piled-up granules, often forming distinct squamules or soredia; rare …Biatora fallax |
| 17b. | Thallus and apothecia C-; thallus various, but not vivid green, never forming distinct squamules or soredia; common …18 |
| 18a. | Thallus PD+ orange-red; apothecia mostly orange to red-brown; spores 11-17 × 3.5-5 µ; to be sought on ericaceous shrubs, especially in subalpine habitats …[Biatora toensbergii Holien & Printzen in Printzen] |
| 18b. | Thallus PD-; apothecia mostly pinkish or tan; spore size various …19 |
| 19a. | Cell lumina of excipular hyphae cylindrical, up to 2.5 µ wide; paraphyses at most slightly thickened; spores 9-17 × 3.5-5 µ …Biatora subduplex |
| 19b. | Cell lumina of excipular hyphae appearing elliptical, up to 6 µ wide; many paraphyses swollen upwards to as much as 4 µ; spores 9-15 × 3-4 µ …20 |
| 20a. | Thallus containing xanthones …Biatora meiocarpa var. tacomensis |
| 20b. | Thallus containing no detectable lichen substances; to be sought …[Biatora meiocarpa (Nyl.) Arnold var. meiocarpa] |
Biatora chrysantha (Zahlbr.) Printzen in V. Wirth
Syn.: Biatora gyrophorica, Lecidea gyrophorica
Thallus expansively spreading, often as wide as 1 dm, pale greenish yellow, producing soralia that eventually cover the entire thallus forming a leprose crust; soredia grouped into tight clusters up to 40 µ across; apothecia rare; disc pale to medium reddish brown, strongly convex when mature, margined by a concolourous proper rim that becomes obscure with maturity; hypothecium colourless; asci 8-spored; spores colourless, non-septate or 1-septate, 10-16.5 × 3.5-6 µ.
Reactions: Thallus C+ pink-red (quickly fading), UV+ dull yellow.
Contents: Gyrophoric acid plus minor amounts of lecanoric acid.
Habitat: On moss and bark usually on tree bases and fallen logs, always within the winter snowpack. All forested elevations.
Similar Species: Biatora chrysanthoides is very similar, but tends to have a thinner thallus and soredia usually limited to round soralia. Fertile material is required for definitive identification, but apothecia in both species are rare. Biatora vacciniicola has a relatively bluish-green colour and grows on smooth bark of shrubs.
Specimens: Björk 8551, 12212.
Local Status: Common.
References: Printzen & Tønsberg (2003).
Biatora chrysanthoides Printzen & Tønsberg
Thallus expansively spreading, often as wide as 1 dm, with a thin, pale greyish-green or pale yellowish green thallus and scattered soralia; apothecia rare, up to 0.5 mm across; disc flesh-coloured to reddish brown, strongly convex, margined by a concolourous proper rim that becomes obscure with maturity; hypothecium colourless; asci 8-spored; spores colourless, ellipsoid, non-septate or (less often) 1-septate, 10-15 µm × 3 µm.
Reactions: Thallus C+ pink-red (quickly fading), UV+ dull yellow.
Contents: Gyrophoric acid and minor amounts of lecanoric acid.
Habitat: On bark and moss on tree bases and fallen logs. All forested elevations.
Similar Species: See notes under Biatora chrysantha.
Specimens: Björk 12829.
Local Status: Occasional.
References: Printzen & Tønsberg (2003).
Biatora efflorescens (Hedl.) Räsänen
Thallus greyish-green, areoles about 0.2 mm wide, producing light yellowish green or pure green soredia in soralia 0.2-0.7 mm wide; soredia 15-30 µ wide, with a compact, subcorticate wall and indistinct medulla, the thallus often becoming almost entirely sorediate; apothecia uncommon, 0.4-0.9 mm wide, pale pinkish brown, glossy, the rim slightly paler and receding with age, the disc flat to strongly convex; hymenium 40-70 µ high, it, like the upper hymenium and hypothecium, lacks pigmentation; spores lacking or sometimes with a single crosswall, 12-19 × 3-5 µ.
Reactions: Thallus PD+ orange-red.
Contents: Argopsin, and sometimes norargopsin.
Habitat: Bark of Shepherdia in dripzone of Populus tremuloides. Lower to middle elevations.
Similar Species: In B. vacciniicola the thallus gives a C+ pink-red and PD- reaction. It grows on shrubs with acid-bark at upper forested elevations.
Specimens: Björk 15181.
Local Status: Occasional.
Biatora fallax Hepp
Thallus small to medium-sized, dull greyish green, consisting of coarse, discrete granules mostly 100-200 µm across, these often with scattered patches of powdery, bluish-tinged soredia; apothecia common, 0.4-0.6 (‑1.2) mm wide; disc convex, red-brown, compound or lumpy at maturity (often appearing paired), margin obscure when dry, but apparent when moist, darker than the disc; hypothecium colourless; asci 8-spored; spores colourless, thin-walled, ellipsoid, non-septate or (rarely) 1-septate, 8-15 × 3-5 µm.
Reactions: C- or + pink-red, PD+ orange-red.
Contents: Argopsin, small amount of gyrophoric acid.
Habitat: On rough bark of conifers within the winter snowpack. Middle to upper forested elevations.
Similar Species: Biatora vernalis has smaller spores, gives a PD+ red reaction, and grows over moss.
Specimens: Björk 9393, 10812.
Local Status: Rare.
References: Printzen & Tønsberg (1999).
Biatora flavopunctata (Tønsberg) Hinteregger & Printzen
Thallus medium-sized, forming patches up to 3 cm wide, creamy green-yellow, smooth or slightly crackled, with more or less evenly spaced roundish soralia that remain discrete; soredia 10-20 (‑25) µ, apothecia occasional, 0.2-0.5 mm wide; disc and rim pale yellow-tan, rim receding with maturity and disc becoming domed; upper hymenium lacking granules or any distinct pigment layer; hypothecium colourless; asci 8-spored; spores colourless, ellipsoid, mostly 1-celled mixed with a few 2-celled. 8-10 × 2.5-4 µm.
Reactions: K+ yellow.
Contents: Atranorin, usnic & isousnic acids, small amount of stictic and cryptostictic acids and an unknown terpenoid.
Habitat: On smooth bark of shrubs within the winter snowpack. Lower to (especially) upper forested elevations.
Similar Species: Lecanora farinaria has poorly delimited soralia that become confluent with age.
Specimens: Tønsberg 14415 (BG).
Local Status: Common.
Biatora globulosa (Flörke) Fr.
Thallus immersed and inconspicuous, sometimes surficial, but thin, whitish; apothecia 0.2-0.5 mm wide, dark grey-brown or blackish, strongly convex, with a thin, quickly receding rim; upper hymenium greenish, K+ deeper green, N+ red; hypothecium colourless; paraphyses swollen at the tips to 3-6 µ wide; spores narrowly oblong, 0- to (mostly) 1-septate, 7-12 (‑16) × 2-4 µ.
Reactions: Spot tests all negative.
Contents: No known lichen substances.
Habitat: On wood, especially snags. All forested elevations.
Similar Species: Some Micarea species can be very similar, but very few have a greenish, K+ green upper hymenium. They differ also in having a Micarea-type ascus in which the IKI+ blue apical thickening has a darker blue tube ascending through its entire height.
Specimens: Björk 8965; Goward 01-209.
Local Status: Uncommon.
Biatora hypophaea Printzen & Tønsberg
Thallus smooth or areolate, or sometimes immersed, light green or grey-green; apothecia medium to dark grey, with a paler rim that is thin and recedes with maturity; exciple formed of cylindrical hyphae arrayed in a fan-like pattern as seen in cross-section; hymenium medium to dark streaky grey-green or blue, sometimes purplish; hypothecium medium to dark brown or grey-brown; spores 8 per ascus, mostly simple, but with at least a few 1-septate spores intermixed.
Reactions: Thallus PD+ orange-red.
Contents: Argopsin, ± norargopsin.
Habitat: On conifer branches in humid forest, less often on broad-leaf trees or shrubs. Lower to middle elevations.
Similar Species: No other local epiphytic crustose lichen combines dark grey apothecia, a thin, paler rim, and occurrence in humid forests. Rarely Lecania naegelii produces grey apothecia, but is easily distinguished by its 3-septate spores.
Specimens: Björk 12324, 13976.
Local Status: Common.
Biatora meiocarpa (Nyl.) Arnold var. tacomensis (Printzen & Tønsberg) Printzen & Tønsberg
Thallus thinly areolate, inconspicuous, light grey-green, of areoles 0.3-0.6 mm wide; apothecia convex, sometimes lumpy, constricted at the base, 0.25-0.45 mm wide, pale tan to medium orange-brown; exciple in the rim formed of elliptical cells up to 6 µ wide; hymenium colourless or with a diffuse pale yellowish pigment; hypothecium colourless; paraphyses often swollen to 4 µ at the tip; spores (8.5‑) 10-15 (‑23.5) × (3‑) 3.5-4 (‑5) µ.
Reactions: All spot tests negative.
Contents: 2,4,7-trichloro-3-O-methylnorlichexanthone with minor amounts of 5,7-dichloro-3-O-methylnorlichexanthone, 3-O-methylasemone, and 5,7-dichloronorlichexanthone.
Habitat: On smooth bark, especially of ericaceous shrubs. All forested elevations.
Similar Species: Biatora subduplex has paraphyses scarcely swollen at the tips, often larger spores, narrower excipular hyphae, and lacks xanthones. Lecidea albohyalina has narrower spores, but is otherwise very similar.
Specimens: Björk 12887, 13968.
Local Status: Common.
Biatora pallens (Kullhem) Printzen
Thallus pale green-gray or whitish, thin or slightly lumpy; apothecia 0.15-0.3 mm wide, pruinose, whitish or pale pinkish yellow, flat or low-convex, with a thin but persistent rim; exciple formed of hyphae with rounded or elliptical cells; hypothecium colourless; spores narrowly elliptic, 1- to (mostly) 3-septate, 9-18 × 2-3 µ.
Reactions: All thallus spot tests negative, but the apothecia may react KC+ yellow.
Contents: Usnic acid (especially in the apothecia), and zeorin.
Habitat: Mostly on bark of twigs and branches above the winter snowpack. Lower to middle elevations.
Similar Species: The combination of pale, pruinose, biatorine apothecia, 3-septate spores, and occurrence on branches and twigs makes this species unique in higher elevation forests.
Specimens: Björk 8970, 8976.
Local Status: Occasional.
References: Foucard (2001).
Notes: Our material differs from European specimens in several (albeit minor) regards. More work is needed.
Biatora rufidula (Graewe) S. Ekman & Printzen
Thallus grey or whitish, usually intergrowing inconspicuously with other crust lichens; apothecia numerous, 0.3-0.6 (‑1) mm across; discs orange-red to red-brown, flat when young, becoming domed, rim concolourous or slightly paler, becoming obscure with maturity; hypothecium colourless; asci 8-spored; spores colourless, ellipsoid, 3-septate, 13-16 × 5-6 µ.
Reactions: All spot tests negative.
Contents: No known lichen substances.
Habitat: On branches and twigs of conifers above the winter snowpack, especially in humid sites. All forested elevations, but mostly at lower elevations.
Similar Species: Biatora nobilis is very similar, differing as indicated in the keys.
Specimens: Björk 10805; Goward 78-328.
Local Status: Common.
Biatora subduplex (Nyl.) Printzen
Thallus mostly inconspicuous but expansive, of drab pale tan or whitish grey colours, smooth or lumpy-areolate, lacking soredia; apothecia abundant, often clustered, 0.2-0.7 (‑1.4) mm wide, the rim apparently only when young; disc strongly convex, often becoming spherical and lumpy; hypothecium pale to light brown; spores oblong to elliptic-oblong, colourless, 1-septate or (less often) non-septate, (8‑) 9.5-15 (‑21) × (3‑) 3.5-5 (‑6) µ.
Reactions: All spot tests negative.
Contents: No known lichen substances.
Habitat: On smooth bark of shrubs, also over mosses at the base of trees, always within the winter snowpack. All forested elevations.
Similar Species: Biatora vernalis differs in having a darker, less expansive crust and larger spores. Biatora meiocarpa usually has smaller, paler apothecia.
Specimens: Björk 8701, 13222; Goward 97-1, 02-05.
Local Status: Common.
Notes: Morphologically diverse and perhaps representing two species corresponding, e.g., to the moss-dwelling versus smooth bark-dwelling habits. The former, at any rate, has consistently larger, more heavily pigmented apothecia.
Biatora vacciniicola (Tønsberg) Printzen
Syn. Lecidea vacciniicola Tønsberg
Thallus small to medium-sized, green or grey-green with scattered loose, light green or yellowish green soralia that are mostly not in defined soralia; apothecia rare, mostly 0.2-0.4 mm across; disc pale reddish-brown, convex to globose, indistinctly rimmed by a pale, semi-translucent proper rim that soon recedes; hymenium 50 µ high, upper portions lacking granules or a distinct pigment layer; hypothecium colourless; spores 8 per ascus, ellipsoid, non-septate or (rarely) 1-septate, 8-10 × 3-4 µ.
Reactions: C+ pink-red.
Contents: Gyrophoric acid.
Habitat: On smooth bark of shrubs within the winter snowpack. Middle to upper elevation forests and bogs.
Similar Species: Biatora chrysantha and B. chrysanthoides are similiar but do not grow on the bark of shrubs. Biatora efflorescens is C-.
Specimens: Björk 13180, 13781.
Local Status: Common.
Biatora vernalis (L.) Fr.
Thallus grey or olive-green, consisting of coarse granules mostly 80-150 µm across, lacking soredia; apothecia abundant, 0.4-0.6 (‑1.5) mm wide, the rim thin and quickly receding, the disc medium orange-brown to red-brown, strongly convex to globose; spores colourless, ellipsoid, non-septate (rarely 1-septate), (10‑) 13-20 (‑25.5) × (4‑) 4.5-6 (‑7) µ; hymenium 45-95 µ high, upper portions lacking granules or a distinct pigment layer; hypothecium colourless or light brown.
Reactions: All spot tests negative.
Contents: No lichen substances.
Habitat: Over mosses at the base of trees. Middle to upper forested elevations.
Similar Species: Biatora subduplex, has a paler thallus, smaller spores, and tends to form large patches over moss.
Specimens: Björk 8552; Goward 96-796.
Local Status: Rare.
Biatora sp. nov. “gold nuggets”
Thallus formed of rather loose, weakly corticate, mustard-yellow or yellow-green granules 150-300 µ with a cottony, white medulla, lacking a hypothecium; apothecia unknown.
Reactions: All spot tests negative.
Contents: Unknown.
Habitat: On bark of conifer branches and trunks in humid oldgrowth forest. Lower elevations.
Similar Species: Bacidia rubella and Agonimia sp. have granules of similar size, but these are more firmly corticate, and less yellowish. Biatora aureolepra Spribille & Tønsberg, which may eventually be found in the study area, is somewhat similar, but has paler pigmentation and a more continuous thallus.
Specimens: Björk 12303.
Local Status: Rare.
Notes: Known only from inland BC.