GRONER, U. (2010). Calicioid lichens and fungi in the Muota Valley, central Switzerland: high species diversity in a small area. Candollea 65: 377–391. In English, English and French abstracts.
An inventory of the calicioid lichens and fungi of the Bödmeren Forest region in the Muota Valley, central Switzerland, produced no less than 51 lichenized and non-lichenized taxa on bark and wood. This mainly subalpine spruce forest in the Prealps hosts the most diverse calicioid flora in Switzerland today and a similarly rich flora from a comparably small area is unknown from neighbouring countries. Calicium denigratum (Vain.) Tibell and Chaenothecopsis oregana Rikkinen are new to Switzerland. Calicium adaequatum Nyl., Chaenothecopsis tasmanica Tibell and Chaenothecopsis vainioana (Nádv.) Tibell are also reported for the first time from this region. Chaenotheca chrysocephala (Ach.) Th. Fr., Calicium viride Pers., Chaenotheca trichialis (Ach.) Th. Fr. and Microcalicium disseminatum (Ach.) Vain. are the most frequent species, the majority of the other taxa are uncommon or rare. The diversity of the calicioid (and other) species is due to several favourable conditions, such as the presence of old-growth forest stands and the occurrence of standing dead trees. More than 40% of the recorded lichenized calicioid taxa are threatened; the state of the poorly known non-lichenized calicioids is undetermined as they were not evaluated in the “Red List”.
Introduction
The knowledge about calicioid lichens and fungi (the “Caliciales”) in Switzerland went relatively unchanged for a long period of time. Only a few references are cited by Clerc (2004) for the lichenized genera, such as the pin-lichens Calicium Pers., Chaenotheca Th. Fr., etc., between about 1940 and 1980. In the 1990s, only two noteworthy contributions, the works of Camenzind & Wildi (1991) and Groner (1994), were published. Data were gathered, mainly of the lichenized calicioid species, during field work for the “Red List of epiphytic lichens” (Scheidegger & al., 2002). Shortly after that, the pertaining distribution maps were available on “swisslichens” (Stofer & al., 2008). Stimulated by the checklist of Clerc (2004) and by the periodically updated results on the cited web site, descriptions of new discoveries have steadily increased (Bürgi-Meyer, 2005; Dietrich & al., 2008; Dietrich & Bürgi-Meyer, 2008b; Zimmermann, 2009), as well as reports about additional localities of rare taxa (Bürgi-Meyer, 2007; Dietrich & Bürgi-Meyer, 2008a).
The region of the Bödmeren Forest in the Muota Valley in central Switzerland has been described as lichenologically very interesting and species-rich (Groner & Clerc, 1988; Groner, 1990, 1997). Data regarding calicioid lichen and fungus species, however, have only recently been published within the scope of the project “Urwaldcharakteristiken des Bödmerenwaldes” (Groner & Frei, 2005; STIFTUNG URWALDRESERVAT BÖDMEREN, 2005). This recent work on calicioid taxa, together with the wealth of the calicioid flora in the region, were inspiration for an overview of the genus Chaenothecopsis Vain. for the whole of Switzerland (Groner, 2006). The current paper is part of a long-term project, the lichen inventory of the wider Bödmeren Forest area. All calicioid taxa are treated in an annotated list, and the substrates, frequencies of the species and threats are discussed. Calicium denigratum (Vain.) Tibell, collected in 2007 and Chaenothecopsis oregana Rikkinen, collected in 2003, are reported here for the first time from Switzerland.
Study area
The Bödmeren Forest and its surroundings in the Muota Valley (municipality of Muotathal, canton of Schwyz) including the primeval forest reserve of ca. 70 ha, is probably the scientifically best documented region of the northern Prealps in Switzerland (Fig. 1). It has been described in numerous papers with geological, botanical and zoological content; most of these contributions were published in Bettschart (1982, 1990, 1994) and Lienert (2001). The area of Bödmeren-Twärenen-Silberen is part of the largest contiguous karst in the country. High annual precipitation and long periods of snow cover are characteristics of the regional climate. However, no substantial surface water is present because of the subterranean drainage. The forested karst includes the montane-high montane and the subalpine zone, and as might be expected given the geological conditions, the vegetation is predominantly calciphile. A spruce-dominated Abieti-Fagetum is developed at lower altitudes; the larger part, however, is covered by a locally dispersed Piceetum subalpinum and cattle pastures. The study area of ca. 14 km2 more or less corresponds to the area described in Groner (1990). It includes the forested terrain between the altitudes of 1100 m in the west and about 1800 m in the east, with the Bödmeren Forest in the centre. Its boundaries mostly follow the topographical conditions except for the western delimitation.
Material and methods
The current paper is based on numerous field observations and more than 100 specimens of calicioid species, collected on many excursions during the last 24 years. Part of the data — from the Bödmeren Forest s.str. — originates from the above mentioned project (Groner & Frei, 2005). The specimens were analyzed under dissecting binoculars and a light microscope. The keys of L. Tibell (in Poelt & Vezda, 1981), Tibell (1999a) and Wirth (1995) were used for determination. The work of Titov (2006) includes modern keys and descriptions of the non-lichenized genera. Reagent tests were performed on the thalli and ascomata of several lichenized taxa; such tests, mainly with K (10% KOH) and N (50% HNO3), are necessary for the separation of Chaenothecopsis taxa. Overall descriptions of genera and species are given in the cited literature, but some morphological, anatomical and chemical features, especially the criteria for separating similar species, are mentioned in the list. The species with stalks may have branched ascomata and/or divided (multiple) capitula (Fig. 14, 16). Size, colour or pruina may vary and differ from the descriptions given in the literature (Fig. 6, 15, 19). Pale hyphae on a stalk or capitulum (Fig. 19), or colourless paraphyses growing out of a mazaedium, may possibly be mistaken for pruina. Accumulations of mature spores are sometimes observed on capitula without a mazaedium (e.g. Mycocalicium subtile (Pers.) Szatala). L. Tibell determined or revised some of the cited specimens; the resinicolous Chaenothecopsis oregana was examined by J. Rikkinen and H. Tuovila.
All calicioid genera belong to Ascomycota. The nomenclature adopted here for the lichenized species (Table 1) follows Clerc (2004). Taxa not considered in that work, and the non-lichenized species, are named as in Tibell (1999a, b) and Titov (2006), respectively. Altitudes mentioned in the table correspond to the lowest and the highest collection locality. Multiple occurrences (i.e. on 2 to 5 tree trunks) at a test site in Groner & Frei (2005) are treated as one record (observed frequency, Table 1). Only modern information about species frequencies in Switzerland from Stofer & al. (2008) is cited. Data of Chaenothecopsis are taken from Groner (2006). The categories of threat (“Red List” category) “CR” (critically endangered), “EN” (endangered) and “VU” (vulnerable) are added for the lichenized taxa included in Scheidegger & al. (2002). Collection data will be available on Stofer & al. (2008). All specimens were collected by the author and are deposited in the author's herbarium. The photographs were taken of recently collected herbarium material, exceptions are mentioned in the figure legends; the cited numbers are specimen (herbarium) numbers.
Results
Comments on the species listed in Table 1.
Calicium abietinum
The documented collections in Switzerland (Stofer & al., 2008) are epiphytic; the lignum of conifers, the common habitat of C. abietinum, has obviously not been investigated so far. In the field confusion possible with C. pinastri (see below).
Calicium adaequatum
All Swiss observations of this species on twigs of sycamore, in contrast to the substratum information from northern Europe (Tibell, 1999a). Stalk and excipulum Iodine + violet to dark blue. The thalli of all examined specimens are not immersed as mentioned in Tibell (1999a), but instead have warty to almost squamulose, greenish grey areoles (Fig. 2). Photobiont trebouxioid.
Calicium adspersum
Collections on the Swiss Plateau on oak; in the upper montane and subalpine belts mainly of the northern Prealps, on conifers. Mazaedium with more or less faint yellow pruina. Thallus usually distinct, K+ red (see C. trabinellum).
Calicium denigratum
The collection locality of C. denigratum is in an open Piceetum subalpinum with scattered mountain pines; on dry wood of a standing dead trunk (12 cm diam.). The lichen covers an area of about 3 dm2, it is accompanied by Calicium trabinellum. Small thalli of Imshaugia aleurites (Ach.) S. L. F. Mey., Parmeliopsis ambigua (Wulf.) Nyl. and Buellia griseovirens (Sm.) Almb. present as well (collection date: October 3, 2007). Thallus of Calicium denigratum immersed; ascomata of the specimen mostly slender, without pruina, the stalks glossy black; capitula cup- to bell-shaped (Fig. 3, 4). Asci cylindrical, the spores uniseriately arranged, 1-septate; mature spores with irregular pattern of cracks, therefore the surface appearing areolate. The mentioned characteristics correspond with the description in Tibell (1999a). This Calicium species is distinct and hardly confusable with any other taxon.
Calicium glaucellum
Short-stalked herbarium specimens of “Calicium glaucellum” with a distinct thallus, dated before 1999, should be checked; these probably belong to C. montanum. The similar C. parvum Tibell, currently not recorded in the region, has clavate asci and diffractaic acid as the main lichen substance. C. glaucellum contains sekikaic acid.
Calicium lenticulare
At present, all observations of C. lenticulare in Switzerland have been on bark; additional records on wood are to be expected. Stalk and exciple I+ dark blue.
Calicium montanum
A recently described species (Tibell, 1999b), it grows on bark of old and/or dead branches of Picea and other conifers, and on wood. Pruina sometimes difficult to see. Thallus loosely granular to compact-warted, with divaricatic acid, and therefore easily separated from C. glaucellum with short stalks (see remarks under C. glaucellum).
Calicium pinastri
The specimen collected on July 19, 1988, with short, nonpruinose ascomata and obconical capitula (Fig. 5), had provisionally been determined as C. glaucellum. Asci cylindrical; spores 9–13.5 × 5–6.5 μm, ornamented with flat warts and shallow cracks. Calicium pinastri, published together with C. montanum (Tibell, 1999b), has recently been reported from Toregg (canton of Lucerne; Dietrich & Bürgi-Meyer, 2008b). Confusion possible with short-stalked C. abietinum, which has somewhat larger spores with a minutely warted surface. C. parvum usually has a whitish pruina (see C. glaucellum).
Calicium salicinum
Widely distributed on wood of conifers (e.g. Tibell, 1999a; Wirth, 1995), but in Switzerland almost exclusively corticolous (see remark under C. abietinum). Thallus immersed or thin, K+ yellowish. Lower side of capitulum not always brown, but more or less black.
Calicium trabinellum
Swiss records of this lichen are mostly corticolous, just like C. abietinum and C. salicinum. Thallus immersed; yellow pruina on lower side of the head.
Calicium viride
Widespread in Switzerland, lacking only in the lowest parts of the country. Thallus very rarely immersed; a single collection with unusually yellow to brown stalks.
Chaenotheca brachypoda
A species mainly on the bases of conifers and deciduous trees. Additional localities for this lichen are to be expected in the Prealps and Alps.
Chaenotheca brunneola
Chaenotheca brunneola is usually lignicolous, but all Swiss records to date are from bark (Stofer & al., 2008). Distinguished from more or less similar species by the absence of pruina, the immersed thallus, and the photobiont, which is trebouxioid (Dictyochloropsis).
Chaenotheca chrysocephala
Several specimens observed with the parasitic/parasymbiotic Chaenothecopsis consociata (Fig. 6).
Chaenotheca ferruginea
Predominantly in rather dry situations on acid bark and wood. Thallus K-, yellowish pigmented or rust-coloured parts K+ dark red.
Chaenotheca furfuracea
A corticolous and terricolous lichen species, found almost exclusively in rain-protected, shaded habitats.
Chaenotheca gracilenta
Ascomata partly parasitized by Chaenothecopsis cf. savonica (Räsänen) Tibell (Fig. 13). Ecological requirements about the same as Chaenotheca furfuracea.
Chaenotheca hispidula
One of the two known locations in the study area was destroyed in November 2007 (see Discussion). Thallus immersed, photobiont Trentepohlia.
Chaenotheca laevigata
Another Acer with this lichen was discovered after the loss of the first tree (see remark under C. hispidula, and Discussion). Chaenotheca laevigata has unmistakable spores and its thallus is warted-areolate, like that of C. trichialis.
Chaenotheca phaeocephala
In a variety of rather well-lit situations; apparently less sensitive than most other calicioid lichens regarding humidity.
Chaenotheca sphaerocephala (Fig. 7)
Reported as new to Switzerland by Dietrich & al. (2008) from Krienser Hochwald (canton of Lucerne); that collection was also on a single tree. Easily confused in its habitat on tree bases and in root cavities with C. stemonea. The photobiont, a trebouxioid alga, is distinguishable only under a microscope (compare C. stemonea). The similar C. hygrophila Tibell is characterized by singly-produced asci (non-catenulate) and a well-developed excipulum, in contrast to both C. sphaerocephala and C. stemonea. The lichen products of these three species, however, are more or less identical (Tibell, 1999a).
Chaenotheca stemonea
Always on the lower part of the trunk and on the base. Thallus with Stichococcus. Confusion possible with Chaenotheca sphaerocephala (see that taxon).
Chaenotheca subroscida
Easily distinguished from other species with yellow pruina by the whitish granular to coralloid thallus.
Chaenotheca trichialis
On wood, the thallus is scarcely developed or immersed, much like C. xyloxena. In contrast to that species, the hyphae of the exciple of C. trichialis are parallel to the stalk surface (periclinally arranged; Fig. 8). Several times seen with the host-specific parasite/parasymbiont Chaenothecopsis epithallina.
Chaenotheca xyloxena
Thallus immersed. Outer hyphae of the exciple more or less perpendicular to the stalk surface (anticlinally arranged; Fig. 9). In the Bödmeren Forest area found a few times with Chaenothecopsis pusiola as a parasite/parasymbiont (Fig. 10).
Chaenothecopsis consociate
Capitula occasionally somewhat pruinose (Fig. 6), these forms can be confused with C. tasmanica. However, C. consociata with a K+ red turning greenish reaction.
Chaenothecopsis debilis (Fig. 11)
Only two other Swiss collections, both dated before 1900 (cantons of Basel-Stadt, Basel-Land). In Groner (2006) presumably the first published records from Switzerland. Characteristic reactions of the pigments with K and N (sometimes weak or ephemeral).
Chaenothecopsis epithallina
Ascomata K- (in contrast to C. pusiola). The affected areoles of Chaenotheca trichialis are often discoloured and disintegrating.
Chaenothecopsis hospitans (Fig. 12)
One collection outside of the study area, at 710 m; on Fraxinus L. Chaenothecopsis hospitans grows on saxicolous Haematomma ochroleucum (Neck.) J. R. Laundon as well; however, only four corticolous collections are known in the country. The first mentioned Swiss records probably in Groner (2006). Ascomata sessile or very short-stalked; yellowish or reddish pigments K+ distinctly red.
Chaenothecopsis nana
Ascomata K-. C. nana has spores similar to those of Mycocalicium subtile, but distinctly shorter asci than that species; compare also Chaenothecopsis savonica.
Chaenothecopsis oregano
A single collection on resin and resin-soaked bark of a fir tree in the upper montane zone (coll. November 12, 2003). Recalls C. rubescens Vain. because of the non-septate spores and the K+ purplish reaction of the exciple. However, the substratum and ecology do not match with any known European species. The analysis of its DNA has shown that this specimen belongs to C. oregana (J. Rikkinen, pers. comm.); the species has been described rather recently (Rikkinen, 2003). This is the first record of a resinicolous Chaenothecopsis taxon in Switzerland.
Chaenothecopsis parasitaster
Observed several times in the Bödmeren Forest area by C. Scheidegger (pers. comm.). Chaenothecopsis parasitaster is very similar to C. pusilla and has by some authors been included as a synonym (e.g. Tibell, 1999a; Groner, 2006). However, recent molecular results indicate that C. parasitaster is a distinct species (L. Tibell, pers. comm.).
Chaenothecopsis pusilla
Rather variable with regards to size, colour and pruina. This probably represents a complex of unresolved species (Tibell, 1999a).
Chaenothecopsis pusiola (Fig. 10)
All specimens with greenish pigments (not yellowish to reddish as in Tibell, 1999a), reacting K+ dark red.
Chaenothecopsis savonica
Probably juvenile (without spores) on Chaenotheca trichialis and C. gracilenta (Fig. 13); possibly also saprobic on bark. Asci short, just like Chaenothecopsis nana, but the spores not fusiform. Greenish pigments K+ yellowish brown.
Chaenothecopsis tasmanica
Two other collections since Groner (2006); the only known locality in the country at present. Also on crustose lichens and non-lichenized algal colonies (Tibell, 1998; Selva & Tibell, 1999). Capitula bluish grey or bluish pruinose (Fig. 14); ascomata K+ yellowish to brownish. May be confused in the field with pruinose C. consociata.
Chaenothecopsis vainioana
Specimen with a few small ascomata; stalks pale brownish, not black or brownish black (Fig. 15). Grows as a parasite or parasymbiont on Trentepohlia, lichens containing Trentepohlia and apparently also with Trebouxia (Titov, 2006). Currently no other Swiss records.
Chaenothecopsis viridialba
Probably also parasitic or parasymbiotic on Chaenotheca chrysocephala. Ascomata without the reddish pigments (i.e. with pale stalks) give a ± yellowish brown, not green, reaction.
Chaenothecopsis viridireagens (Fig. 16)
In the study area on Chaenotheca xyloxena and C. stemonea, among others. The stalks just below the head are often distinctly reddish brown.
Cyphelium inquinans
On one occasion together with C. karelicum. Thallus K+ yellowish (to reddish brown), very similar to C. karelicum! The (juvenile) spores possess a distinct ornament of more or less longitudinally arranged ridges and irregular cracks (Fig. 17).
Cyphelium karelicum
Alpine distribution like C. inquinans, but more frequent. Thallus K± yellowish, not Chas mentioned in Wirth (1995) and Tibell (1999a). According to these authors, C. karelicum has smaller apothecia than C. inquinans. However, the two species are not distinguishable without microscopical examination; young spores of C. karelicum are smooth, while mature spores bear deep cracks; their ornamentation is thus coarsely areolate (Fig. 18).
Cyphelium lucidum
Easily recognized by the yellow thallus and the distinct yellow pruina on the margin of the apothecia.
Cyphelium tigillare
On the wood of dead branches of Picea. In the dryer parts of the Alps most probably widely distributed.
Microcalicium disseminatum
Ascomata rarely with distinct short stalks. Found on Chaenotheca chrysocephala several times with conspicuous pycnidia (Fig. 6).
Mycocalicium subtile
Size, form and colour of the ascomata are all rather variable (Fig. 19). Easily confused in the field with species of Chaenothecopsis, but distinguished microscopically by its non-septate, broadly fusiform spores and long asci (always >30 μm).
Phaeocalicium compressulum
Probably everywhere in stands of Alnus in the area. Spores dark brown; excipular hyphae periclinally arranged. The outer layer of the exciple of the similar P. betulinum (Nyl.) Tibell consists of isodiametric to irregular cells; in addition, the spores are pale brown (Tibell, 1999a).
Sphaerophorus globosus
Sphaerophorus is the only calicioid genus in the Bödmeren Forest region with shrubby thallus. A protected species (Naturund Heimatschutz-Verordnung; NHV). Medulla I+ dark blue.
Sphaerophorus melanocarpus
Several sterile thalli found on a single tree. A remarkable oceanic element of the lichen flora, currently known only from Gurnigel (canton of Berne) and from Alpthal (canton of Schwyz). Protected species (NHV). Medulla I-.
Sphinctrina anglica
Four out of seven observations were on Protoparmelia hypotremella van Herk, Spier & V. Wirth. The host species of the other Swiss records are, unfortunately, not mentioned in Stofer & al. (2008).
Stenocybe major
This fungus usually prefers old Abies (Wirth, 1995; Titov, 2006); in Switzerland possibly much overlooked.
Thelomma ocellatum
A single collection from a wooden fence post; the lichen was mostly sterile with only a few scattered apothecia. Resembles sterile morphs of Trapeliopsis flexuosa (Fr.) Coppins & P. James. Therefore, several old reports and specimens should be verified. Medulla I+ dark blue.
Table 1.
List of the calicioid lichens and fungi of the Bödmeren Forest area. The rarer cases under “Host” and “Substrate” are set in brackets.
continued
Frequencies are given on a rough relative scale (absolute numbers in brackets).
Fig. 4.
Calicium denigratum (Vain.) Tibell, most ascomata pressed flat (Schaerer s.n. (G [G00057263]). Scale = 1 mm.
Fig. 5.
Calicium pinastri Tibell and pinkish discoloured areoles of Hypocenomyce sorophora (Vain.) P. James & Poelt (Groner 610, herb. Groner; old specimen). Scale = 1 mm.
Fig. 6.
Chaenothecopsis consociata (Nádv.) A. F. W. Schmidt and pycnidia of Microcalicium disseminatum (Ach.) Vain. on Chaenotheca chrysocephala (Ach.) Th. Fr. (Groner 1612, herb. Groner). Scale = 1 mm.
Fig. 8.
Exciple of Chaenotheca trichialis (Ach.) Th. Fr. with periclinal hyphae. Squash preparation with KOH. Scale = 30 μm.
Fig. 9.
Exciple of Chaenotheca xyloxena Nádv. with ± anticlinal outer hyphae. Squash preparation with KOH. Scale = 30 μm.
Fig. 10.
Chaenothecopsis pusiola (Ach.) Vain. on Chaenotheca xyloxena Nádv. (Groner 2848, herb. Groner). Scale = 1 mm.
Fig. 12.
Chaenothecopsis hospitans (Th. Fr.) Tibell on Lecanora albella (Pers.) Ach. (Groner 486, herb. Groner ; old specimen). Scale = 1 mm.
Fig. 13.
Chaenothecopsis cf. savonica (Räsänen) Tibell, young ascomata and pycnidia, on Chaenotheca gracilenta (Ach.) Mattsson & Middelb. (Groner 2587, herb. Groner). Scale = 1 mm.
Fig. 14.
Chaenothecopsis tasmanica Tibell, capitula subdivided, on Chaenotheca chrysocephala (Ach.) Th. Fr. (Groner 2843, herb. Groner). Scale = 1 mm.
Fig. 15.
Chaenothecopsis vainioana (Nádv.) Tibell, stalk pale brown, on Lecanactis abietina (Ach.) Körb. with white pruinose pycnidia (Groner 810, herb. Groner ; old specimen). Scale = 1 mm.
Fig. 16.
Chaenothecopsis viridireagens (Nádv.) A. F. W. Schmidt with branched ascomata, on indet. crustose lichen ; with Chrysothrix candelaris (L.) J. R. Laundon (Groner 2943, herb. Groner). Scale = 1 mm.
Fig. 17.
Ascospores of Cyphelium inquinans (Sm.) Trevis.; ornament of mostly longitudinally arranged ridges and irregular cracks. Scale = 15 μm.
Discussion
Calicium denigratum
According to Clerc (2004), neither observations nor collections of C. denigratum have ever been published in Switzerland. However, there are certainly specimens of this taxon deposited in various herbaria, with outdated nomenclature or named using old systematic concepts. I have examined one of L. E. Schaerer's specimens from the upper Emmental, canton of Berne (Fig. 4), which is actually C. denigratum (Schaerer s.n. (G [G00057263]). The label reads: “Calicium nigrum v. pusillum Schaer. Ad truncos putridos in valle Buembach. Jul. 1825” (the year not clearly legible). As stated by Tibell (1999a), the species occurs in Scandinavia, in continental Europe, and in Siberia. Both the habitat and substrate of C. denigratum in the Bödmeren Forest agree well with the description in Tibell (1999a). On the other hand, the usual companion species Chaenothecopsis fennica (Laurila) Tibell mentioned for Scandinavia has not yet been recorded in Switzerland.
Species diversity
The Bödmeren Forest area in the Muota Valley has by far the most diverse flora of calicioid lichens and fungi in Switzerland, with 51 lichenized and non-lichenized species currently listed (Table 1). However, the calicioids in other Swiss regions are, with few exceptions, poorly known at best (see comparison in Camenzind-Wildi & al., 1996). About 68 calicioid taxa are known today in the country — of which a considerable 75% are present in the Bödmeren Forest region. Likewise, the species diversity of the investigated area of only 14 km2 is remarkably high, compared with the numbers of calicioid taxa of neighbouring countries, although not all counts are up to date. Wirth (1995), several years ago, listed ca. 58 calicioid lichens and fungi for the German Land Baden-Württemberg, Hafellner & Türk (2001) mentioned 54 species for Austria, and Nimis & Martellos (2008) noted 65 taxa for Italy. No comparably rich calicioid flora is known today from a similarly small area in these countries; the study area is probably exceptional in central Europe.
The first published records in Switzerland of Calicium adaequatum (Groner, 1994), of Chaenothecopsis tasmanica and C. vainioana were from the Bödmeren Forest region (Groner, 2006). The same presumably applies to C. debilis, C. hospitans and C. pusiola, also listed in the cited work. Calicium denigratum and Chaenothecopsis oregana, as described above, are additional calicioid taxa new to the country; moreover, this is probably the first report of C. oregana in central Europe. Other remarkable species found as part of this survey include the recently reported Calicium pinastri (Dietrich & Bürgi-Meyer, 2008b) and Chaenotheca sphaerocephala (Dietrich & al., 2008), the latter recorded here for the second time in the country. The study area hosts 14 of the 16 currently known species of Chaenotheca in Switzerland, as well as 11 of the 14 Calicium-species, and 14 of the 18 Chaenothecopsis- taxa. These numbers definitively confirm the species-richness of the area, as stated previously in an inventory of the epiphytic macro-lichens (Groner, 1990). The diversity of the calicioid flora is, just like that of the macro-lichens, favoured by several ecological factors combined in the Bödmeren Forest area: Stands with elevated air humidity as a result of high precipitation; numerous suitable micro-habitats on bark and wood; many highly inaccessible spots without any logging; carefully managed stands without documented large clear-cuttings in the past; stands of trees demonstrating long ecological continuity, with many old trees and substantial quantities of dead wood (primeval forest relics ; STIFTUNG URWALDRESERVAT BÖDMEREN, 2005). The Bödmeren Forest area almost perfectly matches the ecological requirements of the majority of the calicioid taxa.
Frequencies, Substrata, Distribution
The most frequent species in the Bödmeren Forest s.str., as well as in the rest of the study area, are Chaenotheca chrysocephala, Calicium viride and Chaenotheca trichialis (Table 1). C. chrysocephala was noted on two thirds of the 300 spruce trees studied by Groner & Frei (2005), whereas the two other taxa were present on somewhat more than half of the trees. Microcalicium disseminatum was found on 28% of the trees; all other calicioid lichens and fungi were less frequent or rare. As expected, the number of species on bark and wood of conifers is high, while the number on deciduous trees is rather low. However, some rare calicioids such as Chaenotheca gracilenta, C. hispidula and C. laevigata, and the substrate-specific Calicium adaequatum and Phaeocalicium compressulum occur only on deciduous trees or shrubs. An important substrate is dead wood: 40% of all species (parasites included) grow on wood and on bark, eight taxa are exclusively lignicolous. In general, the number on stumps is low, and the records on logs are very rare. The presence of snags (standing dead trees) is therefore of crucial importance for diversity and frequency of the calicioid taxa. Calicioid species on rocks or on saxicolous lichens have until now not been recorded in the study area.
The calicioid lichens and fungi in the Bödmeren Forest region are not evenly distributed, but clearly delimited areas with concentrations of calicioid species (“hot spots”) apparently do not exist, as demonstrated in the central part of the study area (Fig. 20, 10 relevés are added in the southern part compared to Fig. 3.7-2 of Groner & Frei, 2005). The recorded localities of a taxon are either more or less dispersed or, in several cases, too scanty for an interpretation. Most species occur between ca. 1250 and 1650 m throughout the forested parts, obviously more frequent and with higher species numbers in stands offering suitable habitats and appropriate climatic conditions.
Indicator species
The occurrence and the absence of epiphytic lichens, namely of rare species and/or species with higher ecological requirements, allow conclusions to be drawn about the state and evolution of a particular forest. The method of using selected taxa as indicator species, and the calculation of indices to assess the ecological continuity of forests, goes back to the work of Rose (1976). In an investigation by Tibell (1992), several crustose lichens and calicioid species proved suitable as indicator organisms for old-growth forests, i.e. as indicator species for forests with long continuity. Selva (2002) suggests the use of calicioid taxa alone for this purpose.
An investigation with this approach in the Bödmeren Forest s.str. was only partly successful (Groner & Frei, 2005); therefore, several relevés have been added later. On this basis, the distributional and frequency data roughly match the other evaluated characteristics of old-growth forest (Liechti & Burger, 2005). The test sites with the highest numbers of calicioid lichens and fungi were recorded in the forest parts 5 and 6 (Fig. 20), which contain not only the oldest trees of the investigated area (400–500 years), but in addition showed high tree age averages on several plots (250–300 years; Horat & al., 2005). Most of parts 5 and 6, together with part 2 (the forest reserve), are regarded as primeval or close to primeval forest by Liechti & Burger (2005). The low species numbers on some plots here are mainly due to unfavourable climatic conditions (Groner & Frei, 2005). A few rather species-rich test sites were found in areas 1 and 4 (Fig. 20), where logging was common during the last 100 years (Liechti & Roth, 2005; Liechti & Burger, 2005). Apart from calicioid taxa, other old-growth forest indicator species were recorded as well (e.g. Usnea longissima Ach.; Groner & Frei, 2005). Obviously, the availability of suitable ecological niches, as observed also in these harvested parts of the forest, is very important. The calicioid lichens und fungi, and epiphytic lichens in general, of the Bödmeren Forest can be used as indicators of long ecological continuity only when combined with other indicators, such as tree age, occurrence and quantity of dead wood, etc. The study area with old trees and calicioid species in abundance apparently is not a perfect example for the potential of the calicioid taxa as indicators, but the results set a good standard for future studies in prealpine forests.
Threatened taxa
One third of the 32 calicioid lichen species present in the Bödmeren Forest region appear to be threatened (Table 1): 6 species are classified in “Red List” category VU, 4 in EN, and one in CR (Scheidegger & al., 2002). Unfortunately, not all recorded species are considered in the “Red List”, the state of the recently discovered and of the lignicolous taxa is not determinable. The percentage of endangered and vulnerable species is therefore presumably higher. A low estimate indicates that at least 42% of the calicioid lichens — currently 11 out of 26 classified taxa — are threatened; 19% are endangered or critically endangered. A “Red List” for the non-lichenized calicioid fungi does not exist, so the 19 parasitic, parasymbiotic (or commensalistic) and saprobic species recorded in this study were not considered. Nothing much is known about the distribution and rarity of these taxa, with the exception of Chaenothecopsis (Groner, 2006). The non-lichenized genera, traditionally studied by lichenologists, need to be included in an updated “Red List”.
Experience has shown that rare species often occur on a single tree or in a confined, small area; the exception in the Bödmeren Forest is Usnea longissima, growing on quite many spruce trees (Groner, 1990). On the other hand, examples for only once- or twice-recorded calicioid taxa are numerous: Calicium denigratum, Chaenotheca brunneola, C. sphaerocephala, etc. The risk of extirpation of such isolated populations is very high, as demonstrated in the case of C. hispidula-laevigata. One of only two known host trees was cut in autumn 2007. The sycamore in question, which supported C. hispidula, C. laevigata and Chaenothecopsis hospitans among other calicioids, had obviously been removed for a temporarily installed cable used for the transport of logs. The protection and conservation of rare species are difficult under these economically controlled circumstances, especially in the case of small, often inconspicuous taxa. However, a conservation strategy must be broadly based, not on the rare calicioid species alone, because this region harbours not only rare and threatened epiphytic lichen taxa, but also several rare species on rotten wood, on soil and on rock with different ecological requirements.
Lichenological research in the Bödmeren Forest area will continue after this inventory of the calicioid lichens and fungi, there are probably additional species to be discovered also from this fascinating group. The study area is obviously valuable from a conservation perspective, with numerous and rare taxa — this remarkable species diversity (not only of the calicioids) must be preserved. However, the pressure on the Bdmeren Forest ecosystem has considerably increased during the last years due to increasing human activities, like tourism, recreation, sports etc. A certain natural protection though exists in the area: Leaving the road, or the trails, is not recommended in places with a rough, rocky terrain, with karst pits and crevices. The significant enlargement of the forest reserve, as agreed in 2009 by cantonal authorities and the land owner, is certainly an important measure towards protection and conservation of threatened species. An inventory like the present study should, of course, document existing species, rather than extirpated or extinct ones.
Acknowledgements
I am most grateful for help in this mainly privately financed lichen project: to Leif Tibell (Uppsala) for the revision of specimens and for valuable information; to Jouko Rikkinen and Hanna Tuovila (Helsinki) for the determination of the resinicolous specimen; to Alexander Titov (St. Petersburg) for his book about the Mycocalicioids; to Rosmarie Honegger (Zurich) for information and literature concerning lichen algae; to Bruno Suter (Muotathal) for support during fieldwork in the Bödmeren Forest; to Michael Dietrich (Kriens) for corrections and improvements of an earlier draft of the manuscript; to the members of Stiftungsrat Urwaldreservat Bödmeren for a contract within their primeval forest project; to Elizabeth Feldmeyer-Christe (Birmensdorf) for correcting my French summary; to Scott LaGreca (Pittsfield) for improvement of my English text.
