Acta Bot. Need. 37(2), June 1988, p. 251-263

Macromycetes in alpine snow-bed communities —

mycocoenological investigations

B. Senn-Irlet

Systematisch-Geobotanisches Institut der Universitdt Bern, Altenbergrain31, CH-3013 Bern,

Switzerland

SUMMARY

For a period of 3 or 5 years, six plots in the alpine zone of Switzerland,

mainly in the Bernese Oberland, have been observed for macromycetes.

In snow-bed communities, 88 species could be found, 25 on siliceous

ground and 63 on calcareous ground. Fungi with a wide ecological

and distribution dominate in range a large area acidic snow-bed

communities(Salicetum herbaceae(SAH)) and fungi with distinctly

specified ecological claims dominatein calciphilous snow-bed

communities(Salicetum retuso-reticulate (SAR)). In both vegetation

types mycorrhizae-formers play a most important role. The differences

in the between for flora the two snow-bedcommunities are greater the

macromycetes than for the phanerogams. Both vegetation types are

characterized by macromycetes.

Key-words: , alpine zone, discomycetes, mycocoenology,

snow-bed communities.

INTRODUCTION

Snow-bed vegetation often passes for a paragon of a high alpine vegetation type, yet this vegetation type takes up only a small surface of the entire alpine plant cover. In fact, grass heaths dominate the alpine zone of the central European Alps. Snow-bed

in communities occur habitats with extreme ecological conditions, normally at the bottom of northern with thick a exposed slope a snow cover during winter, or at higher altitude in with The short regions relatively high precipitation. very vegetation period, the main feature of snow-bed lasts 3 months in favourable and communities, years 8 weeks in unfavourable ones (Walter & Breckle 1986). Among the very few plants found that can develop under these rough climatic conditions are the dwarf willows. The rest of the species are tiny, nestled to the soil. Liverworts and mosses play an important role in such vegetation types.

Macromycetes are a very substantial part of these alpine communities. It was Favre

(1955) who first focused attention on the mycoflora of such extreme habitats. More-

he discovered of of over, a high diversity mycorrhizae-forming fungi dwarf shrubs.

The aim of this investigation was to assess snow-bed communities of siliceous and

and of limestone areas both quantitatively qualitatively. This paper is part a more com- prehensive study on the ecology, sociology and of alpine macromycetes

(Senn-Irlet 1986).

251 252 B. SENN-IRLET

Fig. 1. Map of Switzerland showing the investigation areas.

MATERIALS AND METHODS

The investigations were carried out in four areas of the alpine mountains of Switzerland

2 (Fig. 1). At each site, plots ofat least 30 m were selected in vegetations consideredto be of

described with releves and homogeneous composition. Vegetation was Braun-Blanquet

symbols. Mosses and lichens were collected and identifiedin the laboratory.

Mycocoenological analyses

In accordance with the proposals ofBarkman (1976) and Arnolds (1982), permanentplots

of 2 established and observed for of 3 30-100 m were a period (four plots) or 5 years (two

and information about plots) to achieve detailed qualitative quantitative macromycete

and removed from abundance, frequency and biomass. The carpophores were counted

the plot then dry weight was measured by weighing some mature carpophores. The

calculated: following parameters were

(1) Density (£>) ofcarpophores per year:

z:c x 100

DC = y ,

s

where x = number of visits, C=number of carpophores, s = surface of the plot, y = number ofobservation years.

Total numberof (2) carpophores per plot:

tDC = Y.DC r

(3) Average density (aD) of carpophores per year;

tDC

aDC = •

y ALPINE SNOW-BED MACROMYCETES 253

(4) Maximum density (mD) of carpophores per visit:

cwlOO

— mDCv(iy) ——,

5

where = number cm maximum of carpophores established on one visit during the observation period.

RESULTS

Vegetation

Snow-bed vegetation on limestonediffersstrikingly from that on siliceous ground. Based on the composition ofthe phanerogams, all releves can be ascribed to already well-known plant communities (Braun-Blanquet 1949). Yet, plot 33 shows an interesting species

The unusual of composition. occurrence Arabis coerulea and Salix reticulata in plot 33, in with is caused an area predominantly acidophilous flora, by glacially transported dolomiticmaterial in moraine. the is still a However, community interpreted as a SAH.

The three plots of SAR show several species typical of dwarf shrubs and of calcareous heaths. These vegetation types can be foundin the neighbourhood, thus they formcontact communities.Above plot 41 the Rhododendro-Vaccinietum,another type of dwarf shrub heath, is present. Adjacent to plots 42 and 43 Caricetum ferrugineae has developed a

heath moist grass community. These neighbouring vegetation types influencethe floristic composition of the SAR plots.

in Mosses play an important role the structure of snow-beds. In SAH there are nearly

beside whereas exclusively acrocarpeous mosses some tiny hepatics, in SAR pleuro-

dominate. the richness of lichens is in carpeous mosses Moreover, striking SAR. In relation to their cryptogam composition the two snow-bed communities differ greatly; there are only a few species in common. For the mycological study it is interesting that only five species ofhigher plants may form ectomycorrhizas, i.e. Salix herbacea, S. retusa,

S. reticulata, Dryas octopetala and Polygonum viviparum.

Mycoflora

Two examples of the original mycocoenological data are shown in Tables 1 and 2. The

differences between these siliceous the great two plots, one on ground other on calcareous ground, are significant.

The plot in the SAH is very poor in fungal species; however, some compensateby their abundance. Many of the SAH macrofungi have a wide ecological range, a large distri- bution area, and are found in a wide altitudinalrange. The mycoflora is considered to be rather trivial. Psilocybe chionophila, a species close to P. montana, and Galerina chionophila are exceptions. They are known only from arctic-alpine snow-beds.

The of in is minimal: 2 productivity macrofungi plot 11 1 g dried substance per 100 m

l An often of small bodies late in the year~ . singular production fruiting very season, due to harsh climatic conditions, explains the typical low macrofungi productivity of SAH communities. Indeed, carpophores with a cap size between 0-5 cm and 2 cm are standard, rarely some medium-sized carpophores (e.g. Cortinariusfavrei) occur.

The plot in SA R reveals quite a differentpicture. Overall 37 species have been foundin 5 years. Apart from some fungi with a wider ecological range, several species have to be con- sidered found as endemic exclusively in the alpine zone ofCentralEurope and Scandinavia.

bound Obviously they are to special plant species. For instance, Marasmius epidryas only

dead of Other occurs on lignified parts Dryas octopetala (Rosaceae). fungi as Inocybe 254 B. SENN-IRLET

mgmg 6060 (3y) 2944 804 1298 258 524 d.w.

6 mDC 84 53 31 59

97 tDC 134134 6-26-2 106 311313197

2 35 1 32 aDC 2-6 116 4545 35

) 3 3 2 9494 — 53 — 38 2 1985 m

(100 1984 0 0 — — — — — 50 5 6 3 DC 6 53 59 1983 253253 131

3 9/9 3030 1 — 17 — 12

1985 22/8 00 — — — — —

25/7 00 — — — — —

20/8 00 — — — — — 1984

3/8 00 — — — — —

4 1400006666 27 2 17 — 19 23/9 . herbaceae 2 m

1 24/8 1515 1515 1 32 1983 1983 — — exp.; Salicetum 11/8 0 0 — N

0 0 — — ——— — — 27/7 m.ii.M.,

3 6 1 3 2 —— —— — + community 17/9 1982 — — — 2315 21/7 0 0

02030029 6 3 — + snow-bed 16/9 1981 Zinggenstock, 4/8 0 0 — — — — — acidic of

an slope of mycocoenosis cinnamomeoluteacirmamomeolutea Grimsel-Oberaar, species carpophores chionophilachionophila vittaeformisvitlaeformis tetraspora conferendum The II:

1. no. no. no. Table Total Total Psilocybe Galerina Laccaria Entoloma Dermocybe Plot ALPINE SNOW-BED MACROMYCETES 255

egenula, I. luteipes and Cortinarius favrei form mycorrhizae with dwarf willows (Salix

other seel. Glaciales), Dryas octopetala and presumably with some alpine phanerogams.

also be bound substrates. Debaud showed that Saprophytic fungi may to special (1983)

litter extracts of Dryas octopetala stimulated the growth of several calciphilous Clitocybe

species while an extract of Salix herbacea had the opposite elfect at least on Clitocybe

lateritia. the of SAR have found that in Among remaining species we many are very rare

such Galerina and albidum. In Europe, as acerosa, pseudocerina Gerronema plot

42 Marasmiusandrosaceus must be regarded as an alien species; it grows on the needles of

Picea abies that have been blown by wind into the plot. In fact the presence ofMarasmius

androsaceus indicatesthe proximity of subalpine spruce forest.

“ 1 The productivity of fungal biomass in SAR is 10 so it is 10 times higher thanin g year ,

SAH. Beside many tiny fruiting bodies some single big carpophores occur, such as Russula

delica which reached normal size. the lower altitude the (plot 42), cap Due to vegetation

of the SAR lasts than that in the SAH This be period investigated plots longer plots. may

the main for such differences in the the the reason great productivity. In SAH plots snow

melts only during July; in 1981 and 1984 there was thick snow cover even in the first week

ofAugust, whilethe SAR plots were free ofsnow from mid-July. In SAH the fructification

period starts in the second half of August (Table 1) and is often interrupted by snowfalls

in September. Only in years with exceptionally high September temperatures can the mycoflora of SAH develop fully. This was the case in autumn 1985.On 30 September 1985 plot 33 had eight species, with a totalof 33 fruiting bodies; thisis the best record ofall visits

to a SAH plot.

with a wide and distribution dominate in Fungi ecological range a large area SAH, while fungi with specified ecological demands dominate in SAR. This is true not only in the case of plot 11 versus 42 but also when comparing all plots (Table 3). Comparing the values given for the maximum number of carpophores established on one visit

(mDCy (3y)) it is obvious that in SAH only few species occur but with rather high

whilein SAR but with low densities densities, many species occur always (Table 4).

The annualfrequency ofcarpophores

While and in SAH less the in SAR species pattern seasonality are more or regular, fungi appear very irregular over the years (see Table 2): two-thirds of all species occurred only

in 5 once years. That means that either the plot size was too small or a much longer observation period is needed to obtain a more consistent picture of the species composition. The size of the sample plot is a very great problem. Barkman (1968) clearly demonstrateda minimumarea for macrofungi butas alpine plant communitiesoften form small patches one location seems to be far below the minimum area for fungi. The three plots within SAR cover almost the entire surface of SAR vegetation type in all localities studied so a larger plot size could not have been chosen. So for calciphilous snow-bed communities a much longer observation period is needed and the number of plots must be enlarged, however, for acidic snow-beds a 3-year investigation period seems to be sufficient for a representative inventory of the mycoflora.

Mycocoenology, a preliminary list ofcharacteristic species

A list including the abundance of all fungi observed in snow-bed communities is given in

Table 3. After observation of in an period 3 or 5 years (e.g. plot 42) the numberof fungal species exceeds that of mosses and lichens, and in SAR communities even that of higher plants. 256 B. SENN-IRLET

mg 33 166 190 510 216216 91 33 391 136 d.w. (3y)(3y) 28924 — — —

mDC 3-3 10 10 4343 — 6-6 3-33-3 — 23 3-3

tDC 3-3 10 10 4343 — 6-6 3-33-3 — 20 3-3

1 7 aDC 78 M11 3-3 3-3 14 — 2-3 1-010 — 7-7 1-010

7 3 13 20 ) 19851985 130 3-3 — — — — — — 2 m

6 (100 1984 40 10 — — — — — — — DC DC 3 3 5 19831983 63 — 10 4343 — — —

4 1 26/926/9 11 — — — — — — — —

2 6 1985 3/93/9 10 20 l1 — — — — — —

27/7 1 8 27/7 — 3 6 3 24/824/8 12 1984 1/81/8 0 0 — — — — — — — —

1 retuso-reticulatae 8/98/9 2 2 — — — — — — —

3 3 1| 18/818/8 17 1313 1983 0 0 0 — Salicetum 6/86/8

0 0 — 21/721/7 0 — — — — — — — —

5 1982 26/726/7 83680032061 20 — 4 1 — — — — —

6 0 community + 3 3 10/910/9 — — —— — — — — — — — —

3 0 3/93/9 — — — + — — — + .— — — -——— — — — 0 snow-bed 26/826/8 15 + + — — — — — — + + 1981 7 0 ++ 0 18/818/8 — — — — — + -(-+ + + + _l_+ + +

“

6 0 12/812/8 367 + + — + + + — — —

calciphilous 3 0 6/86/8 + + + — — — — — — — —

a of mycocoenosis rusticellus epidryas androsaceusandrosaceus albidum marginatulum cyanulum sp. concava sarcitulum cf unicolor pseudocerina acerosa The species carpophores luteipes galopus petiginosa

2. no. no. Table Total "fotal Entoloma Cortinarius Inocybe Marasmius Clitocybe Marasmius Mycena Gerronema Entoloma CortinariusCortinarius Inocybe Galerina Galerina Arrhenia Hebeloma ALPINE SNOW-BED MACROMYCETES 257

62727 198 863863 150 120120 143143 70 600 1680 2310 6 9659 1020 9900

2020 10 3-33.3 3-3 17 13 3-3 17 20 13 6-6 3-3 3-3

20 6-6 3-33.3 3-3 10 13 3-3 17 20 6-6 6-6 3-3 3-3

0 0 0 0 6-6 3-3 10i-o 10 5-5 4-3 10 5-6 6-6 4-3 2-3 10 10 1 1 1 1

7 3 20 7 7 17 20 7 3 3 20 — — — 13

3 3 3 3 3 — 10 13 3 — —

3-3 — — — 6 — 22

1 1 — 2 — 2 3 2 ______5 3 1— ______1i 3 4 1 -— — —— —— —— _ _—— 1I

1 2 5 +

. -_—_— 2 1— m

• — ___—— —— 30 +——— _——— __—— _——— +_—— —— _ — —— _—— _——— _——— __——— —— —— ——— —— _—— exp.; N + + + + m.ii.M., — — — —— 2060 —Daube, sp. Platte Nolanea 2 brevipes rufostriatus rigidus favrei Schynige favreana subgen. tenellum citrinomarginata aa vittaeformis favrei nitidiuscula gibbagibba sp. 42: egenula sp. geophylla arenosaarenosa urania purpur sp. sp.sp. delica no. Cortinarius Melanoleuca Mycenella InocybeInocybe Entoloma Clavaria:Clavaria Entoloma Inocybe Entoloma Mycena ClitocybeClitocybe Mycena Inocybe Cortinarius Cortinarius InocybeInocybe:InocybeInocybe: Geopora Mycena Galerina Russula Clitocybe Plot 258 B. SENN-IRLET

9-39-3 5-6 2-3 1 1 5-6 1 1 SAR — — — — — — — — — — — — — — — 4 4 2 mDCv

6 7-3 3-3 3-3 2-3 6 1-6 7-3 0-6 4 4-3 3-3 3-3 2-3 0-6 2-6 1 4-3 1 3-6 2-62-6 2 1 1 0-6 4-3 1 1 4-3 1 SAH 3838 7373 3030 4 15 18 2 22

Switzerland er* THh TPbpTPbp TPhm TPbp THh TPhm TPhm TPhmTPhm TPhm TPhm VC VC TPhmTPhm TPhm TPhm THh THh TPhmTPhm TPhm TH1 THh TPhm TH1TH1 TPbs TPhm

in

5 3 43 13 23 — — — — — — — — — — — — — — — + 3 — — — — — — 1+

1 0-4 1 4 7 2 — communities — — — — — — — — — — — — — — — 1 — 7 — SAR 42 16 36

— — — — 6 5 5 — — 1 — 1 — — — — — — — — — — — — — — — 41 II11 28 — — — snow-bed — from — -— — — — — — — — 1 4 4 2 — 3 22 2 4 1 4 33 10 11 — 13 — — — — — — — — — — 16 + — — — — densities — — 0-5 0-5 7 1 0-5 4 5 3 2 — — — — 4 4 — — — — — 3 SAH 22 12 16 1313 52 47 4 4

1 2 5 1 carpophore 11 13 35 — — — — — — — — — — — — — — — — — — — 32 4545 - annual average Mos. their Quel.Quel. Mos. Noord, and (Ort.) Mer.)Mer.) (Ort.) Gill. Hend. Sing. (Britz.)(Britz.) gracilis exex Lam.Lam. Senn-Irl. (Fr.) ex (Fr.) Orton f. Sing. Sing. (Bull,(Bull, Favre macromycetes Bruchet Favre Favre ReidReid Mos. Noord. Fr.Fr. occultopigmentatum itiailia cinnamomeoluteacinnamomeolutea of velutipes hinnuleus favrei type observations conferendum chionophila tetraspora chionophila minus sericeum semiovata tetraspora uvidus sp. later favrei vittaeformis chionophila sp. later favrei taxa rhacodes norvegica calamistrata cf List giacomi sp.

3. no. no. no. Galerina Omphalina HebelomaHeheloma Cortinarius Cortinarius Cortinarius Entoloma Discomycetes Galerina DermocybeDermocyhe Table Vegetation Plot Total Total Entoloma PsilocybePsilocybe Laccaria Galerina EntolomaEntoloma Inocybe Inocybe Anellaria ConocybeRussula Inocybe Lactarius Entoloma Entoloma ClitocybeClitocybe Inocybe Galerina ALPINE SNOW-BED MACROMYCETES 259

0-3 0-3 0-3 0-3 0-3 0-3 0-30-3 1-3 3-6 4-6 3-33-3 7-3 2-3 2 3-3 5-3 1 1 0-3 0-3 0-3 — 2 7 2 2 1 SARSAR 11 13 mDCv

SAHSAH — — — — — — — — — — — — — — — — — — — — — — — — — —

er*er* TPhm TPhm THh TPhm TPhm TPhm TPhm THh THh TPhm THh THh THhTHh TPhm LR TPhm TPhm THh THI TPhmTPhm TPbs TPbs TPhm THI TPhm THh THI

1 5 1 1 1 4 5 2 6 43 13 2323 — — — — — — — — — — — — — — + — + — +— — +— — — -— — — — — —— — — — — — — — — 1— 1—— 1 3 — 1 1 1 3 — 8 2 4 — — — — — — — — — — — — — — — — — — 3— — SARSAR 42 16 3636 42 —

0-3 0-3 0-30-3 0-6 0-3 0-6 0-3 0-3 0-3 0-6 0-8 2 0-3 0-6 1 1 1 0-6 0-6 0-3 0-3 0-3 + 1 2 1 2 1 — — — — 41 11 2828 17 12

33 10 II11 — — — — — — — — — — — — — — — — — — — — — — — — — —

— — — — — — — — — — — — — — — — — — — — — — — — — — SAHSAH 22 12 16

5 11 13 — — — — — — — — — — — — — — — — — — — — — — — — — —

Bruch. Fr. Sing. Fr.) Kum. Kum. Sing. Karst. & ex (Favre) Sing. Fr. Fr.) Sing. Lam.Lam. Quel. FavreFavre (Fr.) Kuhn. Smith Favre (Fr. Favre. Kuhn. Favre (Murr.) 1. (Fr.) 1. (Bull;(Bull: (Sow.iFr.)(Sow.:Fr.) sensu s. Vel. (Fr.) sensu (Fr.) reclinis Fayod (Fr.:Fr.) caryophyllea oreobius tenebricus anomalusanomalus poliopus epidryas mucronella Kill. marginatulum albidumalhidum type observations conicaconica aff.ajf. taxa scabripes linkii dryophila acerosaacerosa pseudocerina concava (Continued) taxa monochroa dulcamara cf fastigiata geraniodora lutescens pumila nana geophylla urania sp. no.no. no. 3. no. Thelephora Cortinarius Cortinarius Cortinarius HyerocybeHygrocybe Gerronema MarasmiusMarasmius Hygrocybe Hebeloma Gerronema Inocybegeophylla Table VegetationVegetation Plot Total TotalTotal Thelephora Inocybe Hydropus Inocybe Cortinarius Cortinarius Cortinarius Entoloma Inocybe Entoloma Bovista Gerronema Inocybe Inocybe Laccaria Hygrocybe Collybia Russula Arrhenia Galerina Gerronema Inocybe ClitocybeClitocybe Mycena 260 B. SENN-IRLET

on

5-6 6-6 3-3 6-6 2-3 1-81-8 1 1 1 5 1 1 1 1 1 — — — — — 10 1 — 14 — 1 — 5 — — lignicolous 10 11 SAR 1 =

= mDCv TH1 LR SAH _ — — — — — — — — — — — — — — — — — — — — — — — — — — humus, bryophytes, of fungus on = fungus er* THh THh TPhm TPhm TPhsTPhs THh TPhm TPhm THh THh TPhm TPbsTPbs THh TPbs THh TPhm TH1 TPhm TPhm THhTHh TH1TH1 TPhm THh— THh THhTHh— TPhm THh THh fungus, saprophytic

7 1 7 1 1 43 13 23 1010 + — — — — — — — — — — — — — — — — — — II11 + + + — — — — — — — — — — — — — — — — = — — — — — mycorrhizal TPhs

0-2 + + + 7 7 1 + 1 + 1 + 3 + + + + + + 1 + + 7 5 7 I 3 — — — — — SAR 42 16 36 14 42 = TPHm bryophytes, _ — — — — — — — — — — — — — — — — — — — — — — — — — — 41 II11 28 on (1982): fungus

33 10 11 _ — — — — — — — — — — — — — — — — — — — — — — — — — — Arnolds

to parasitic

= — — — — — — — — — — — — — — — — — — — — — — — — — — SAH 22 12 16 according TPHp

5 list 11 13 — — — — — — — — — — — — — — — — — — — — — — — — — — leaves, the in and indicated stems on is Am. fungus fungus Rom.) Fr. Fr.) each dung. & Sacc. Noord. Kum. Sacc. Gill. of ex saprophytic on (L.:Fr.) Korf Favre Kum. Gill. Sing. Fr.) Favre (Kuhn. = Hk, (Lasch (Britz.)(Britz.) function fungus (Favre) Fr. (Fr.:Fr.) Favre (Fuck.) (Fr.)(Fr.) (Pers.: Favre TPhs = Fr. (Pers.:Fr.) VC androsaceus rufostriatus (yellow)(yellow) brevipes rusticellus stridula litter, type observations tenellum sarcitulum favreana rigidus cyanulum citrinomargiata sericellum ecological citrinomargiata sp. (Continued) taxa gibba sp. arenosa sp. unicolor nitidiuscula roots, egenula delica sp,sp. pura luteipes petiginosa galopus nauseosa corium no. no. The on 3. no. Inocybepetiginosa Entoloma Entoloma Melanoleuca MelanoleucaMelanoleuca *er: Table Vegetation Plot Total Total Entoloma Entoloma Mycenella Inocybe Marasmius Geopora Cortinarius Cortinarius ClitocybeEntoloma RussulaRussula Clavaria Mycena GalerinaGalerina Clitocybe Inocybe Mycena Inocybe Inocybe Mycena Cortinarius Entoloma Conocybe Russula Helvella decomposing fungus ALPINE SNOW-BED MACROMYCETES 261

Table 4. Species composition of snow-bed communities

SAH* In common SAR* Sum (SAH + SAR)

Phanerogams 37 11 41 78

Mosses and lichens 17 3 52 69

Macromycetes 25 9 63 88

= *SAH Salicetum herbaceae, SAR = Salicetum retuso-reticulatae.

Table 5. Species in each category ofdiagnostic taxa

Salicetum herbaceae Salicetum retuso-reticulatae

Characteristic species

Psilocybe chionophila LaccariaLaccariapumila

Laccaria tetraspora Galerina pseudocerina

Clitocybe concavaconcava

Entoloma tenellum

Constant species

Galerina chionophilachionophila Russula nana

Inocybe calamistrata Arrhenia acerosa

Marasmius epidryasepidryas

Mycenellafavreana Inocybe egenula Entolomasarcitulumsarcitulum

Hebelomamarginatulum

Collybia dryophila Dermocybe cinnamomeolutea Dermocybe cinnamomeolutea CortinariusCorlinariusfavrei CortinariusCorlinariusfavrei

Dominant species (weight(weight dominance) Psilocybe chionophila Russula delica CortinariusCorlinariusfavrei CortinariusCorlinariusfavrei Entolomasarcitulumsarcitulum

Thelephoracaryophylleacaryophyllea

Despite the lack of similar investigations in other regions of the Alps, and despite the low numberof test plots, an attempt is made to characterize the two snow-bed communi- ties with macromycetes. Due to methodological problems mentionedabove, the results are preliminary. Based on Arnold’s (1982) report, three categories of diagnostic taxa can be formed (Table 5). (1) Taxa with a high frequency in only one plant community are considered characteristic defined with to be species. They are a higher frequency or a

than in other studied. that considerably higher carpophore density vegetation types Note this is identical the in of concept not to character species the sense Braun-Blanquet (1949).

(2) Constant taxa are those occurring in almost all plots of a certain syntaxon. (3) Domi- nant are those species with the highest carpophore density or productivity. They deter- mine to a large extent the appearanceof mycocoenoses. 262 B. SENN-IRLET

Fig. 2. Life forms of the higher plants and of the macromycetes in (a) Salicetum herbaceae and (b) Salicetum retuso-reticulatae . The life forms ofthe phanerogamsareindicated according to Landolt (1977): h = hemicrypto-

= = = t = herbaceous = i phytes, z woody chamaephytes, c chamaephytes, g geophytes, therophytes, evergreen nanophanerophytes.For the macromycetes see Table 3.

Life forms

The SAR and SAH differ considerably on the phanerogam species level and slightly less on the respective life form level. This counts even more for the cryptogams, especially the fungi studied. Hemicryptophytes and mycorrhizae-formers dominate(Fig. 2).

The higher diversity may be explained by slightly better climatic conditions and a greater variation in microrelief.

Humicolous abundant and in SAR: this is well fungi are very numerous explained by

humus of soil of the the high content the uppermost these plots. Despite numerous species and the of high coverage bryophytes, parasitic macromycetes on mosses are lacking, and

debris Mosses and saprophytic macromycetes on distinguishable organic are scarce. hepatics in SAR are more frequently attacked by bryophilous lichens such as Lecidea hypnorum, Bacidia sabuletorumand Rinodina mniaraea than by agarics, whereas in SAH agarics fill in this ecological niche. The fact that some lichenized fungi do attack mosses ALPINE SNOW-BED MACROMYCETES 263

has recently been shown by Mayrhofer & Poelt (1985). The few coprophilous fungi in

SAH reflect the whereas the SAR for that plots grazing by cattle, plots are too steep

purpose.

DISCUSSION

Literature about alpine macromycetes is still scanty. Mainly through the articles of

Kiihner (1972) and Lamoure (1972) the ‘combes a neige’ became well known for alpine

fungi. However, as Favre (1955) has pointed out, the snow-bed communities on siliceous

rather in observed manifold grounds are poor macromycetes. He a mycoflora only in

localities with a richer phanerogamic flora. So far Eynard (1977) has published the most

detailedstudy aboutthe ecology of agarics inacidic snow-bed communities. Altogether he

counted in 12 plots double the species I found in four plots. In these SAH communities

in Parc National de la Vanoise, constant species (i.e. species that occurred in most

plots) are Psilocybe chionophila, Dermocybe cinnamomeolutea, Laccaria laccata, Russula

norvegica, Hebeloma marginatulum and Cortinariusfavrei; observations that agree with

the present study.

Bon & Gehu (1973) mentioned 13 species for acidic snow-bed communitiesand 10 for

calciphilous communities. Out of these species I observed only three species in each

vegetation type. Kiihner (1972) stated that Galerinapseudocerina is a very characteristic

of snow-bed communities on observation confirmed in this species limestone; an paper.

ACKNOWLEDGEMENTS

I am indebted to Professor Dr H. Clemengon for support in taxonomy. I thank Dr R.

Watling for identifying Conocybe tetraspora, Dr Th. W. Kuyper for identifying some

Inocybe, Chr. Scheidegger and E. Ruoss for identifying some lichens, C. Scherrer for

technical assistance, and Dr K. Ammann for critically reading the manuscript.

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