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The role of macrofungi in environmentalconservationEef Arnolds aa Biological Station, Centre for Soil Ecology, Kampsweg 27, 9418 PD,Wijster, The NetherlandsPublished online: 14 Sep 2009.

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Giorn. Bot. Ital., 12G: 779-795, 1992

The role of macrofungi in environmental conservation -'.

EEF ARNOLDS Biological Station, Centre for Soil Ecology. Kampsweg 27, 9318 PD Wijster, The Netherlands

ABSTRACT. - The motives for the conservation of fungi are briefly outlined. The data and methods needed for successful use of fungi in environmental conservation are mentioned. A survey is given of the main habitats of threatened fungi in Western and Central Europe, including data on the significance of their mycoflora, the causes of decrease and possible measures to improve the situation. The value of macrofungi as bio-indicators for environmental quality is discussed and demonstrated with three examples: the indi- cator value of wood-inhabiting fungi for undisturbed forest sites; of ectomycorrhizal fungi for air pollution stress in forests and of saprotrophic fungi for the duration of undisturbed grassland use. Lists of indicator species with different indicator values for air pollution stress and grassland use are provided.

Key words: Nature conservation, macrofungi, bioindicators.

INTRODUCTION

Concern about the quality of the environment has rapidly spread in only 30 years and is now a major political issue on a local, regional, national, supranational and world- wide scale. Environmental quality includes such different subjects as the quality of air, water and soil in connection with economic process and human health; quality and quan- tity of human resources and biodiversity in terms of landscapes, communities and spe- cies. In this paper I shall mainly focus on aspects of biodiversity.

Thus far fungi play only a subordinate role in studies on biodiversity and changing species composition, as well as in the selection and management of nature reserves, although fungi are essential components of ecosystems in view of their irreplaceable role in decomposition processes and in mutual, symbiotic relationship with green plants. Re- cent investigations have demonstrated that the mycoflora in some parts of Europe has undergone drastic changes during the last decades and that fungi are very valuable as bioindicators. In this paper I shall concisely treat our present knowledge on the most important habitats of threatened macrofungi and the value of fungi as bioindicators. In addition attention is paid to methods used in this context. The presented data are exclusively based on research in Europe north of the Alps and east of Russia since data of other parts of the world are hardly or not available and since my personal experience is confined to Northwest Europe.

* Comm.no 462 of the Biological Station, Wijster.

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MOTIVES FOR THE CONSERVATION OF FUNGI

Motives for the conservation of fungal species and populations are not fundamen- tally different from motives to conserve other organisms (WINTERHOFF, 1978b; WIN- TERHOFF & KRIEGLSTEINER, 1984). They include (after ARNOLDS, 1991): - Ecological importance: Fungi are essential components of biocoenoses by their func-

tions as decomposers of organic matter, parasites of other organisms and mutualistic symbionts in mycorrhizas. This is also true for ecosystems of economic value, in par- ticular forests.

- Value as indicator organisms: Fungi can be exellent bioindicators in view of their functions, niche-differentiation and species diversity. For instance, many species of ectomycorrhizal fungi are indicators for the degree and kind of air pollution (e.g. FELLNER, 1985, 1989; ARNOLDS, 1991b); wood-inhabiting fungi for intensity of fore- stry practice (RENVALL etal., 1991a, b), and grassland fungi for soil conditions, type of management, and degree of disturbance (ARNOLDS, 1982; NITARE, 1988).

- Economic importance: Fungi are important (potential) sources of food and medicines, and they can be used for selective delignification of wood and straw as well as the degradation of xenobiotics (e.g. BUMPHUS & AUST, 1987).

- Importance for science: Conservation of the gene pools of fungi is needed in order to extend our understanding of evolutionary processes and the resulting diversity in taxa, morphological structures, and ecological strategies.

- Value for recreation and education: Collecting of edible sporocarps of wild fungi is an important form of healthy recreation in many regions; the study of fungi is an hobby of a growing number of naturalists (KREISEL, 1960).

- Esthetic value: Sporocarps of many macrofungi are appreciated by many people as interesting and beautiful components of our environment and a source of joy and creative activities, for instance photography.

- Ethical motive: Mankind is responsible for the continued existence of the variety of life-forms (including fungi), developed during evolution.

In my opinion the latter motive on its own is sufficient justification of efforts in the field or fungal conservation.

NECESSARY DATA AND AVAILABLE hlETHODS

The possible use of macrofungi (as wel! as of the other organisms) in the scope of environmental conservation depends on the availability on the following data:

- Taxonomic knowledge: Ideally modern revisions of all groups of macrofungi should be available. The existence of comprehensive floras of large areas would be very use- ful, as would be concensus on species concepts and nomenclature. Although taxonomy of European macrofungi has made much progress in recent years, the situation is still far from ideal and many years behind the taxonomy of green plants. The knowledge on various taxonomic groups in unbalanced. A well-known group are for instance the Polypores whereas an agaric genus such as Hebeloma is in bad need of a serious monograph. Some regions, for instance the mediterranean area and southeastern Europe, are still insufficiently explored. Great discrepancies

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exist between species concepts of, for instance, the French school of ccsplittersn (e.g. M. Bon) and the Dutch school of ctlumpersn (e.g. BAS et al., 1988). In spite of these imperfections, taxonomic knowledge in many groups is sufficient to serve as a basis for ecological and distributional studies.

- Geographical distribution: For conservation work it is essential to have information on the geographical distribution and frequency of species, by preference in their en- tire area. In fact, systematic mapping of macrofungi has only recently been initiated in some European countries. The first more or less representative distribution atlas has been produced only last year in Western Germany (KRIEGLSTEINER, 1991). Other distributional data in books are usually based on more or - more often - less accurate impressions of mycologists in a certain area. There is still a lot of work to do in this field and European coordination of the efforts would be very useful.

- Ecology. Data on the range of habitats and substrates and the way of habitat exploi- tation of species are essential for an ecological interpretation of mycofloristic data and for useful conservation measures. In addition, representative data on the quan- titative composition of mycocoenoses in the more important habitat types (myco- coenological data) are essential to study effect of natural and disturbed environmental processes. Accurate autecological data are still scanty. Autecological data in text books are often superficial and based on limited observations.

- Changes in frequency of fungi. Data on processes in the mycoflora are essential to determine possible threats to fungi. Such data should include natural fluctuations in undisturbed situations, as well as changes in more disturbed areas. Four methods have been used with success at present: (1) comparision of foray reports of different periods from a limited area (e.g. ARNOLDS, 1988a); (2) comparision of distribution maps of well-investigated species (e.g. ARNOLDS, 1985; PARENT & THOEN, 1986); (3) comparision of observations in selected plots or stands over the years (e.g. NITARE, 1988; ARNOLDS, 1991b); and (4) data on the supply of edible macrofungi to markets (DERBSCH & SCHMIIT, 1987).

Quantitative and semiquantitative data on changes in the macrofungal flora are scarce in comparison with data on phanerogams and larger animals. This is due to some fundamental and practical methodological problems. The former concern: (1) the de- pendence on the above ground appearance of reproductive structures, sporocarps, which abundance may not closely reflect the below ground occurrence of vegative structures which are difficult to study and attribute to relevant fungi - in addition some sporocarps remain below the soil surface, and such hypogeous fungi may be quantitatively impor- tant in some ecosystems (e.g. FOGEL, 1976); (2) the short lifespan of sporocarps which may therefore be missed when spot observations.are made (e.g. RICHARDSON, 1970); (3) the pronounced periodicity of most species (e.g. KRIEGLSTEINER, 1977; ARNOLDS, 1985); (4) fluctuations from year to year attributable to weather conditions (e.g. THOEN, 1976; AGERER, 1985); and (5) which are coincidental with successional changes in plant cover and environment (e.g. ARNOLDS, 1988b).

The numbers of sporocarps of a species may vary from year to year by factors of ' 10 to 100, while the intervals between the successive appearance of the same fungus on a certain spot may extend from several years to decades. Practical problems involved in long-term studies include: (1) the scarcity of reliable older, and often recent, data;

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(2) difficulties related to taxonomy and nomenclature. Together these problems neces- sitate careful attention to methods and the reliability of different interpretations (AR- NOLDS, 1985, 1988a; DERBSCH and SCHhiITT, 1987).

It is evident that the conclusions in the forthcoming sections are based on results of a limited number of studies. However, the results are in many cases so striking that some general conclusions are permitted.

THREATENED MACROFUNGI AND THEIR HABITATS

A fungal species is considered as threatened in a certain area when (1) its number of localities is substantially decreasing, (2) the size of its populations (i.c. numbers of sporocarps on its localities) has been considerably reduced, (3 ) it occurs in a threatened habitat or (4) when it is (and has been) restricted to small populations on few localities. The latter category is often distinguished as <(potentially threatened>> or <(vulnerable)> species (e.g. WINTERHOFF & KRIEGLSTEINER, 1984). The phenomena under (1) and (2) often coincide with each other. Only few accurate data are available on changes in dis- tribution of fungi (see above), so that indirect factors as mentioned under (3) play an important role in drawing up lists of threatened fungi. Such lists, usually called Red (data) lists, have been published in nine European countries in the period 1780-1791. In addition several regional lists were made (for survey, see ARNOLDS, 1991a).

In this paper I confine myself to a concise enumeration of the more important habitat types for macrofungi, which are threatened in some way in large parts of North, West and Central Europe. A habitat is regarded as threatened in a certain area when (1) its range is strongly decreasing, (2) its ecological qualities (species composition, structure, soil properties) are deteriorating or (3) it has a small distribution area in a vulnerable region. However, not all threatened habitats are inhabited by a diverse and/or charac- teristic mycoflora. For instance, salt marshes, other eutrophic wetlands and herb-rich arable fields have only a poorly developed mycoflora and therefore they are not dis- cussed here.

The most important, threatened habitats far macrofungi include:

1. Virgin and tiear-virginforests. These forests are characterized by the (almost) com- plete lack of human interference during at least some centuries, which resulted in a charac- teristic structure of vegetation and soil including tree trunks in d stages of decay, natural rejuvination of the tree layer, undisturbed soil development and a characteristic microrelief. Virgin forests may belong to any forest type distinguished on the basis of floristic composition and usually they are not characterized by a specific flora of green plants. Such forests are nowadays in Europe mainly found in Scandinavia and isolated mountain regions.

Accurate mycological data on virgin forests in comparison to exploited forests are not known. Virgin forests are in the first place characterized by the occurrence of a wide variety of wood-inhabiting species, in particular species restricted to large, dead tree trunks. Many species are (almost) confined to (near-) natural forest stands, for in- stance Fomitopsis roseu and Amylocystis lupponica (RENVALL et al., 1991). Red lists of countries with remains of original forests comprise a high proportion of wood-inhabiting fungi, e.g. 39% in Sweden (FLORAVKRDSKOMMIIT~N FOR SVAMPAR, 1971); 39% in Nor-

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way (H~JILAND, 1988) and 37% in Finland (RASSI & VAISANEN, 1987). In addition, some rare ectomycorrhizal fungi seem to be confined to undisturbed soils in very old forests, for instance the false truffle Chamonixia caespitosu and the agaric Hygrophotrrs inocybifor- nzis (FLORAV~~RDSKO~MMITT~N, 1991).

Virgin forests are threatened by more intensive exploitation by forestry, in the most extreme form clear-cutting. In some countries, e.g. Germany and the Netherlands, new strict forest reserves are being established in exploited forests in order to restore natur- al processes on a local scale. Mycologists in the two countries have been involved in the description of the mycocoenoses at the start of spontaneous forest development (WIN- TERIIOFF, 1989; VEERKAMP, 1992).

2. Forests with a natural coinposition of the tree layer. Native forests are character- ized by certain combinations of indigenous trees and shrubs, dependent on the local soil conditions and microclimate. These forest communities are also characterized by specific combinations of macrofungi. Most mycocoenological studies were carried out in native forest communities, for instance in Belgium (DARIMONT, 1973), the Nether- lands (e.g. JANSEN, 1984), Germany (e.g. KRIEGLSTEINER, 1977), Poland (e.g. LISIEWS- KA, 1974) and Czechoslovakia (ShlARDA, 1972). In many areas mixed native forests are more and more replaced by plantations of one species, often exotic trees or trees with a different natural habitat. Examples in Western Europe are species-rich oak-hoarnbeam forests (Carpinion) on calcareous soils which have been replaced by spruce plantations and oak-birch forests on acid sand (Querco-Betuletum) which have been replaced by Scotch pine. Such measures mean a complete change in species composition and struc- ture of the forests and consequently drastic changes in the mycoflora. It remains to be seen to what extent such changes are reversible.

The introduction of exotics may in fact first increase the mycological diversity on a local or regional scale, but is detrimental in a wider context, since few commercial trees replace a diversity of indigeneous trees in a wide range of habitats. The mycoflora of such plantations is almost constantly a strongly impoverished variant of that in stands in the original area of the planted trees. For instance, among the many specific symbi- onts of Larix in the Central-European mountains only six species are descending into plantations in the Netherlands, and only one of them, Sirillis grevillei, is widespread (although strongly decreasing in recent years) (ARNOLDS, 1989a).

This process is also extremely important in (sub)tropical areas all over the world, where diverse indigeneous forests are <(replaced)> by large-scale plantations of a few spe- cies of Pinus and Eucalyptus..

3. Oligotrophic forests. Forests on soils which are very poor in nutrients, are charac- terized by a canopy of slow growing trees (usually belonging to a single species), a herb layer poor in species and often a well-developed moss layer. However, they are of out- standing mycological significance on account of a very rich and diverse mycorrhizal flora (e.g. JANSEN, 1984; WOLDECKE & WOLDECKE, 1990). In addition, some lignicolous and litter decomposing species are characteristic of these habitats. These forests are still widespread in large areas, for instance Scandinavia, but they are strongly threatened in more densely populated and industrialized regions, for instance the Netherlands (AR- NOLD~, 1989a, 1991b), Eastern Germany (KREISEL, 1980) and Western Germany (WOL- DECKE & WOLDECKE, 1990).

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The decline of ectomycorrhizal species in these forests is well-documented for the association of the Dicrano-Quercetum, forests of Qtiercus robtrr on dry, acid sand dunes with a poorly developed organic top soil, in the Netherlands. Three plots of this vegeta- tion types were studied with the aid of mycocoenological methods during the period 1972-1990. Some results are summarized in Table 1. Similar results were obtained in pine forests on sand dunes (Cladonio-Pinetum). The mycorrhizal diversity may be im- proved by removal of the accumulated litter layer.

TABLE 1

Niirnben of species and sporocaps of the three main ecologicaigroztps of nzacrojlmgi in three plots (1000 n9 each) in oak forests on sand dunes (oicrano-Qiiercetiinz) in Drenthe, the Netherlands, hrring three pm.ods. The niimber of species is expressed as the average ntimber ilt three plots per period. The number of sporocarps is expressed as the sum ofthe average rnaxirnrinz abiindances of all species of a groirp in three plots per period.

av. number of species av. number of sporocarps

Period of study 1972-1973 1976-1979 1988-1990 1972-1973 1976-1979 1988-1990

Numbers of species: Saprotrophic species on soil 18 30 22 2367 899 1123 Wood-inhabiting species 16 19 29 248 335 2140 Ectomycorrhizal species 41 33 14 4781 1109 662 All macrofungi 75 82 66 7396 2343 3925

In percents: Saprotrophic species on soil 24 37 35 32 39 29

Ectomycorrhizal species 55 40 21 65 47 17 Wood-inhabiting species 22 23 44 4 14 54

4. Boggy forests. Forests and scrub on wet soils, usually dominated by species of Salix, Altiiis or Bettila, have in most areas always been local and confined to the valleys of rivers and streams and to depressions. They are characterized by a very diverse mycoflora, including a large number af rare ectomycorrhizal symbionts and wood- inhabiting fungi (e.g. BUJAKIEWICZ, 1989).

These forest types are threatened in many places by canalization of rivers and streams, drainage of surrounding agricultural areas, eutrophication of surface - and soil - water and plantations.of exotic poplar hybrids. The numbers of characteristic ectomycor- rhizal fungi decrease dramatically in desiccated and nutrient-enriched alder- and willow swamps (Arnolds, unpubl. observations), but detailed studies of the effects of different disturbances on the mycoflora have not’ yet been carried out.

5. Peat bogs, moors arid swamps. Bogs are restricted to areas with a constant, high groundwater level. Open bogs have a rather poor, but highly characteristic mycoflora, well-documented in some mycofloristical and mycocoenological studies (e.g. LANGE, 1948; FAVRE, 1948; EINfrELLINCER, 1976). The surface of bogs and other marshlands has been strongly reduced in all of Europe by digging of peat, deep drainage and supply of nutrient- enriched water. They belong to the most vulnerable habitats. The few relics in north- western Europe are usually strongly affected by drainage of surrounding agricultural areas.

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It is evident that the characteristic macrofungi of bogs are disappearing together with their habitat. However, this process have not been documented so far.

6. Heathlands. Heathlands are characterized by a dwarf-shurb layer dominated by Ericaceae, producing slowly decaying litter and having a special type of ericoid mycor- rhiza (REID, 1983). Most heathlands in western Europe are not natural, but have origi- nated after a process of forest degradation, caused by tree cutting, grazing and sod cutting. These semi-natural heathlands are inhabited by a rather poor saprotrophic mycoflora with some characteristic species (e.g. LANGE, 1948; ARNOLDS, 1981).

The areas of heathlands have been decimated in the past decades, mainly due to conversion into fertilized agricultural grasslands and fields and to afforestation, often with monocultures of exotic conifers. In the remaining areas, usually in nature reserves, heathlands are increasingly becoming dominated by grasses (Moliniu cuenrku, Deschumpsiu flexzrosu), bracken (Pteriditrz uqtrilimtn) or spontaneous forest development. This is due to lack of continuation of the appropriate management (grazing, sod cutting, burning) and in many regions also to increased nitrogen deposition (e.g. AERTS, 1989). The mycofl ora of grass-dominated former heathlands is only fragmentary developed (ARNOLDS, 1981) and after forest development the characteristic assemblage of saprophytes is replaced by fungi on leaf litter.

7. Sminutrrrul grassknds. Seminatural grasslands are regarded here as grass-dominated communities with a spontaneous floristical composition (not sown or planted), which are for their existence dependent on continuous, relatively limited human influence: grazing in low densities, annual removal of the sward by hay-making or burning. Pesti- cides, fertilizers and stable manure are not or hardly used. A wide variety of such grass- lands can be distinguished, which are adapted to different soil types and types of human influence, for instance limestone grasslands (Mesobromion erecti), grasslands in coastal dunes (Galio-Koelerion), hayfields along streams (Calthion palustris and Junco-Molinion).

All grassland types have a characteristic mycoflora (ARNOLDS, 1981). In the Nether- lands the number of characteristic grassland fungi outnumbers the number of grassland plants (ARNOLDS & DE VRIES, 1989). The richest in species and sporocarps are perma- nent, mesic to dry pastures on various nutrient-poor soils. They are known as Hygrocybe grasslands on the basis of the abundance of species of this and related genera (ARNOLDS, 1980; RALD, 1985). Other, well-represented genera in such grasslands are Entoloma, Demoloma, Geoglosstiin, Clavuriu and Clavtrlinopsis. It is striking that the species diver- sity of macrofungi increases with the duration of the permanent grassland management without additional fertilizer application. The rarest and most endangered species are restricted to undisturbed grasslands of at least 100 years old. The characteristic fungi are very sensitive to nitrogen fertilizers (ARNOLDS, 1989b).

All types of seminatural grasslands in Western and Central Europe are strongly threatened due to the EEC policy to concentrate grassland farming on a relatively small, very intensively used area, including grazing in very high densities and a high addition- al input of nutrients. So-called marginal grasslands are increasingly improved, abandoned, or planted with forests. In addition, nitrogen deposition has resulted in recent years in shifts in species composition towards dominance of a few tall grasses, for instance Bruchypodium pinnutztm in limestone grasslands (BOBBINK, 1989). Grasslands on steep slopes are particularly endangered, for instance most limestone grasslands. NITARE (1988)

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described the loss of seminatural grasslands in some parts of Sweden with its conse- quences for the mycoflora, in particular Geoglossaceae. In the most intensively cultivat- ed parts of Europe Hygrocybe grasslands have become very rare, for instance in Denmark (BOERTMANN, 1985; RALD, 1986) and the Netherlands (ARNOLDS in BAS et nl., 1988).

The only methods to maintain the precious mycoflora of old pastures are either the creation of large nature reserves where the management-regimes of grazing or hay- cutting are continued, or financial support for farmers who are prepared to continue traditional, low-input grassland use.

8. Sand dztnes. Sand dunes are in Europe mainly found along the sea coasts, but also locally along big rivers (e.g. upper Rhine valley) and in Pleistocene areas as a result of over-exploitation by farmers in the past. Although the environment seems not to be very appropriate for saprotrophic organisms, sand dunes are remarkably rich in charac- teristic fungi, in particular gasteromycetes, e.g. species of Geastrrrm, Bovista, Discixeda and Trrlostotmz. However, also a number of saprotrophic agarics are characteristic (e.g. ANDERSON, 1950; WINTERHOW, 1975). Sand dunes with spontaneous pine and oak trees are in many cases very rich in rare ectomycorrhizal fungi, e.g. Gyroporzrs cyaiiesceris and Tricholonza focale.

Sand dune areas are strongly threatened all over Europe due to expansion of towns (in particular tourist resorts), over-recreation, winning of sand and afforestation (e.g. WINTERHOFF, 1978a).

THE SIGNIFICANCE OF MACROFUNGI AS BIOINDICATORS

Fungi, like all other organisms, have their characteristic ecological range and in that respect each organism can be considered as a bioindicator of certain environmental conditions. A trivial fungus of disturbed habitats is in fact as indicative as a rarity de- pendent on very complex conditions. I shall focus here mainly on those bioindicators among fungi that indicate qualities of threatened habitats. As outlined in a previous section, the methods used to establish presence and abundance of macrofungi are much more complicated than those for green plants and many animals well. Therefore I shall treat at first a few general characteristics of fungi, which make fungi worthwile to use as bioindicators, next to plants and animals, before turning to some specific cases as examples.

1. Fungi have fundamentally different functions in ecosystems as plants and animals. They are completely dependent on other organisms to meet their carbon needs and are not mobile like animals. Saprotrophic fungi may provide information on the dominant processes of litter and wood decomposition. Ectomycorrhizal sporocarps are indicators of this vital symbiotic relationship between tree roots and fungi and may provide infor- mation on the nutrient status of forest soils and on tree vitality. Parasitic fungi may be indicators of both natural processes in forests and tree vitality.

2. Fungi contribute highly to the biodiversity. In almost any forest community fungi outnumber green plants. In forest communities on nutrient-poor soils the number of green plants is very restricted, but the number of fungi is very high. For instance, AR- NOLDS & DE VRIES (1989) reported for coniferous forests in the Netherlands 439 charac- teristic fungi and 7 phanerogams, for-. deciduous forests 1324 fungi against 213

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phanerogams. In such forests the mycoflora may be the most important reason for na- ture conservation or even the only possibility to make a classification. Also outside forests the numbers of fungi are considerable. The number of characteristic grassland fungi in the NETHERLANDS (365) is higher than that of green plants (345) (ARNOLDS & DE VRIES, 1989).

3. Fungi are representative of the actual environmental conditions in a certain area. Species of phanerogams and animals may maintain themselves a long time in a certain area as relics, although the environment is in fact not appropriate anymore. On the other hand, species of plants and animals may lack in a suited habitat due to difficulties to reach that area by diaspores, for instance on account of large distances to other popula- tions or to geographical barriers between populations. Such phenomena are not known in fungi: the spores are easily distributed and it seems that they react very quick to environmental changes, for instance clear cutting and fertilizer treatment (ARNOLDS, 1989b).

Three interesting groups of indicator species among fungi are concisely treated her as examples.

1. Wood inhubiting ftrngi us indicators for uirgin forests. Virgin and near-virgin forests are not so much characterized by a special composition of green plant communities, but by structural characteristics, especially the presence of vast quantities of decaying trunks in all stages of degradation. Dead trees are ideal substrates for a great number of wood- rot fungi. RENVALL et ul. (1991a, b) demonstrated that part of these species are (almost) confined to virgin forests, apparently on account of the permanent presence of suffi- cient substrates. In other forests with scattered dead trunks these species are rarely found. RENVALL et ul. (1.c.) considered as characteristic species of spruce-dominated virgin forests in northern Finland for example Arnylocystis lupponicu and Foinitopsis roseu, of pine- dominated forests Antrodiu ulbobrrrnneu and Postiu lowei. HBILAND (1991) is develop- ing a method for the evaluation of virgin forests in Norway for nature conservation, using polypores as indicators. He distinguishes four groups of indicators: (1) qualitative indicators, whose sole presence indicate virgin forest (40 species); (2) quantitative indi- cators, which indicate virgin forest when they occur in large quantities (66 species); (3) indifferent species, occurring in both virgin forests and forests under strong human in- fluence (36 species); (4) negative indicators, which occur mainly in forests influenced by man (13 species).

2. Ectornycorrhizul jkngi as. indicators for air pollution. Forests on soils, poor in nutrients, are inhabited by a diverse and abundant flora of ectomycorrhizal fungi, liv- ing in association with the roots of the trees. Examples were described in a previous section. I t has been demonstrated that air pollution leads to a strong reduction of ec- tomycorrhizal fungi, being deleterious for most of the species and stimulating only few species. Species can be classified according to their sensitivity for air pollution (Table 2) and consequently used for the evalution of the impact of air pollution on forest ecosystems. However, some precaution should be kept in mind. Firstly, the indicator values concern only the sporocarp formation of the species. The sensitivity of mycorrhi- zas and extramatrical mycelia is still unsufficiently known, although it is probable that a close relationship exists between the influence of air pollutants on vegetative and reproductive organs. Secondly, air pollution is a complex mixture of different chemicals

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(in the Netherlands mainly ozone, SO,, NH, and NO,, but also traces of pesticides, heavy metals, etcetera), producing various direct and indirect effects on plants and soils. Since the exact working mechanism on ectomycorrhizal fungi is not yet known (see Ar- nolds in this volume: Ecological research on ectomycorrhizal fungi, an introduction), the indicator values can not yet be specified for different components of air pollution. In the Netherlands nitrogen deposition is probably the most important cause of decrease of ectomycorrhizal fungi. In other regions of Europe different air pollution climates occur and therefore the mentioned indicator values are possibly only of regional sig- nificance.

FELLNER (1985, 1988, 1989) considered ectomycorrhizal fungi also as valuable in- dicators for air pollution effects in forests and as a result for the vitality of forest stands in Czechoslovakia. He distinguished three phases of impoverishment within assemblages of ectomycorrhizal fungi: (1) inhibition of sporocarp productivity; (2) loss of species diver- sity with the proportion of mycorrhizal species in the total count of macromycetes drop- ping to ca. 35%; (3) partial to total destruction of ectomycorrhizal fungi which then contribute only l0-20% of the total number of macromycete fungi. The third phase is correlated with symptoms of (strongly) reduced vitality of the trees. Consequently, ectomycorrhizal fungi may be extremely valuable as predictive indicators of forest decline, In the Netherlands also a positive correlation was found between the numbers of ec- tomycorrhizal fungi and the vitality of the stands (TERNORSHUIZEN, 19901, but this rela- tionship is less obvious as in Czechoslovakia and seems to be more dependent on local factors, such as soil type. More research is needed before ectomycorrhizal fungi can be generally used as indicators for forest vitality.

FELLNER (1985), from observations in the Giant mountains, Czechoslovakia, sug- gested that in subalpine Pinus mugo scrub the following ectomycorrhizal fungi were rela- tively tolerant to air pollution: Russiila ochroletica, R. emetica, Hygrophonis olivaceoalbus, H. bypothqus and Laccaria laccatu; in beech forests Lactaritis subdukis, Xerocomus pniinatzis and Amanita submembranacea. CUDLIN et al. (1987) identified the following species as

being tolerant in spruce forests in the Ore Mountains (Czechoslovakia): Nygrophorus olivaceoalbus, Laccaria laccata, Lactarius nqh, Paxillus involiitiis and Riissiila ochroleuca. Hygrophorus olivaceoalbus, Xerocomus pniinatiis and Amanita submembranacea are very rare in Netherlands, so that their possible indicator value is unknown. Most of the other species, mentioned by these authors, are indicated as constant or increasing species in the Netherlands (Table 2). However, Lactarius nfus and Russiila emetica show a ten- dency to decrease in the Netherlands and Hygrophonrs hypothejzis is listed among the sensitive species. These differences are probably due to differences in pollution climate between the areas.

Ectomycorrhizal fungi may also be used as indicator species for other environmen- tal factors. VESTERHOLT (199 1) demonstrated the significance of species of Cortinarim subgenus Phi'egmacium as indicator species for valuable old deciduous forests on calcareous clay in Denmark.

3. Soil-inhabiting saprotrophs as indicators for old, tindistzirbed grasslands. Old, un- disturbed grasslands are often well-characterised by communities rich in phanerogams and bryophytes, including rare species. However, this is not always the case. Some grass-

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TABLE 2

Examples of ectomycorrhizal firngi with different indicator values for air pollution stress (combination of NH,, NO, and S02) in the Netherlands, based on the presence of sporocarps.

1. Very sensitive species, at present extinct or having lost over 90% of their localities.

Bankera firligineoalba Boletopsis leucomelaena Boletus calopus Canthare flus tubaeformis Cortinarius a Iboviolaceus Cortinarius bolaris Cortinarius bovinus Cortinarius caninus Cortinarius castaneus Cortinarius mucosus Cortinarius multiformis Cortinarius violaceus Craterellus cornucopioides Dermocybe cinnabarina Gomphidius ghtinosus Gyroponrs cyanescens Hydnellum airrantiacum Hydnellirm caenrleirm Hydnellum compactum

Hydnellum concrescens Hydnellum femigineirm Hydnellum peckii Hydnellum spongiosipes Hygrophorus eburneus Hygrophorus leucophaeus Hygrophorus nemoreus Hygrophorus russula Inocybe sambucina Lactarius piperatus Leccinirm testaceoscabmm Pseudocratellirs sinuosrrs Ramaria botrytis Ramaria formosa Rozites caperata Rtissula decolorans Rtrssula olivacea Rtissula rosacea Rirsszrla virescens

Sarcodon imbricatus Sarcodon joeides Sarcodon scabrosus Sarcodon undenuoodii Suillus grevi I lei Suillus variegatics Tricholoma albobnrnnerrm Tricholoma argyraceum Tricholoma aurantium Tricholoma airratrim Tricholonza columbetta Tricho lonza focale Tricholoma imbricatum Tricholoma portentosum Tricholoma saponaceirm Tricho loma sca@turatirm Tricho loma sciodes

2. Sensitive species, having lost over 50% of their localities and numbers of sporocarps strongly reduced.

Amanita aspera Amanita inaurata Amanita potphyria Amanita virosa Boletinus cavipes Boletus queletii Boletus satanas Chantarellirs cibarius Chroogomphus rutilm Coltricia perennis Cortinarius elatior Cortinarius hinnu leus Cortinarius pholideus Cortinarius toruus Democybe crocea

Dermocybe semisangirinea Elaphomyces grandatus Elaphomyces mtrricatirs Gyroporus castaneus Hydnum repandum Hydnum ru fescens Hygrophoncs airreus Hygrophonrs hypothejus Inocybe xanthomelas Lactarius chrysonheus Lactarius deliciosus Lytarius vellereus Lactarius vietics Leccinum airrantiacum Phyllopoms rhodoxanthirs

Rtrssirla adusta Rrrssrrla coenrlea Russula foetens Russula luteotacta Russula paludosa Riissula sangirinea Russula sardonia RuiiuIa solaris Ruiitth xerampelina s. strict0 Siiillus bovinus Suillus luteus Tricholoma irstale Tylopiltrs fe l leus

3. Rather sensitive species, having lost a considerable part of their localities and/or showing a dis- tinct decrease in abundance of sporocarps on their localities.

Anzanita citrina Amanita fulva Anianita gemmata Amanita pantherina Amanita phalloides Amanita spissa Boletus edulis

Boletus erythropus Boletus luridus Clitopilus prunu lus Cortinarius anomalus Cortinarius delibutus Cortinarius paleaceus Cortinarius trivialis

Gomphidius roseus Hebeloma cylindrosporum Hebeloma sinapizans Inocybe mixtilis Inocybe napipes Inocybe ovaticystis Inocybe praetervisa

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lnocybe unrbrina Lactarius b fennius Luctarius camphoratrrs Luctarius deterrimus Luctarius glyciosmirs Lactariris befuris Luctarius pirbescens Lactariris 1741s Lactariiis serijluus Leccinum qirerciniim

Rhizopogon futeofus Rrissula aeruginea Russrrla a fbonigra Riissiila atroptilpcrrea Russula chamaefeontina Russu fa cyanoxantha Riissiifa defica Rrrssrr fa emetica var. sy fuestris

Russu fa felfea

Russufa hagifis Riissufa graveofens Russir fa nigricans Rirssrrfa odorata Riissirfa vesca Thefephora terrestris Tricho foma jlavo brunnerrm Tricho foma sir fphrirerrm Xerocomus sirbtomentotrs

4. Indifferent species, remaining approximateIy constant in numbers of localities and sporocarps.

Amanita muscaria Amanita rubescens Amanita vaginata ss. str. Cortinariiis saniosrrs Hebeloma helodes Hebeloma latifo fium Hebeloma fongicaudum Hebeloma mesopbaeum lnocybe asterospora Inocybe brevispora lnocybe cookei Inocybe du fcarnara Inocybe fastigiata Inocybe hirieffa lnocybe facera

Inocybe faniiginefla lnocybe petiginosa Inocybe sqiraniata Laccaria amethystea Laccaria faccata Laccana tortifis Lactaritis controversus Luctarius necator Lactariris quietus Lactarius subdufcis Leccitairm scabrum Leccinrttn variecofor Naucoria bobemica Paxillus involutiis Russula amoeno fens

Riissufa emetica uar. betufarum

Riissufa flaua Russufa mairei Riissrr fa nitida Russrda paraziirea RrissriIa pectinatoides Scferodema are0 fatirm Sclerodema verrucosum Trichofoma poprrlinartn Xerocomus badiris Xeroconrus cbrysenteron Xerocomus porosponrs Xerocomus nibelfus

5. Increasing species, showing an increase in numbers of localities or sporocarps. Laccaria bico for Laccaria proxinra

Lactaritis hepaiiciis Lactarizts theioga firs

Riissu fa ochro feuca Sc feroderma citrintrni

lands with very high mycological significance in the Netherlands have a rather trivial species composition of green plants, for instance the grasslands on the Cannerberg and along the Drongelens Kanaal (ARNOLDS, 1980). On the other hand, some grasslands with many rare phanerogams are poor in fungi, for example, some limestone grasslands which have been recently created in the southern Netherlands on former arable fields (e.g. Wrakelberg, Putberg). The determining factor for these differences is apparently the duration of undisturbed grassland use. Consequently fungi can be used as indicators of old grasslands.

Also NITARE (1988) emphasized the importance of continued grassland use for mac- rofungi in Sweden. He developed a simple model for the evaluation of Swedish grass- lands on the basis of the numbers of species in several characteristic taxonomic groups (Fig. 1). ARNOLDS (1980) and RALD (1985, 1986) used the number of species of Hygrophoraceae as criterion for the mycological value of grasslands in the Netherlands and Denmark, respectively.

Like Mycorrhizal fungi, the grassland inhabiting fungi can be arranged in species groups with a certain indicator value concerning the significance of their habitat for

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30

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L4 Y g 10

8

3 a

HYCROCYBE + CAhiAROPAYLLOPSlS

EKTOLOMZI

DERUOLOMA

LOW IUTAER HIGH VERY E l G E HIGH

VALUE FOR NATURE CONSERVATION

Fig. 1 - A scheme for assigning the value for nature conservation to seminatural grasslands in Sweden, based on the number of species of grasslands fungi (modified from Nitare, 1988).

TABLE 3 Indicator values of 225 selected macrofrngi for the degree of disturbance ofgrassland communities on mesic to dry soils in northwestern Enrope. Species, which occur also or mainly outside grasslands are marked with *. The mentioned indicator value is restricted to grasslands only.

1. Species of very old, stable, unfertilized grasslands, very sensitive to changes in management (in par- ticular use of fertilizers), mostly very rare or strongly decreasing in grasslands in W. Europe.

Camarophyllopsis schrrberi Dermoloma josserandii Demoloma pragensis Dermoloma pseu- docirneifolitim Entoloma anatinrim Entoloma atromarginatum Entoloma bloxamii

(= madidunz) Entoloma caenileum

Entoloma caenileofloccosunz '

Entoloma carneogriseum Entoloma chloropolium Entoloma cruentatum Entoloma exile Entoloma griseocyaneum Entoloma kerveniii Entoloma lampropus Entoloma lividocyanulum Entoloma porphyrophaeirnz

Entoloma pninirloides Entoloma roseiinz Geoglossum barlae Hygrocybe aurantiosplendens Hygrocybe chlorophana Hygrocybe cystidiata Hygrocybe flavescens Hygrocybe fivipes

(= kzcmus ss. auct.) Hygrocybe intermedia

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Hygrocybe nitiosa Hygrocybe nitrata Hygrocybe obrussea

Hygrocybe ovina (= quieta ss. auct.)

Hygrocybe perplexa Hygrocybe unguinosa hlicroglossum niidipes

Hygrocybe punicea Poronia punctata (on dung) Hygrocybe radiata Tbueminidium atropurpureum Hygrocybe spadicea Tricbog lossurn variabile

(= sciophana ss. auct.)

2. Species of old, unfertilized grasslands, sensitive to changes in management (in particular use of fer- tilizers), mostly decreasing in grasslands in W. Europe.

Camarophy I Iopsis atropuncta* Camarophy llopsis foetens* Camaropbyllopsis micacea

(= pbaoxantha) Camarophy Ilopsis phaeopbylla Clavaria straminea Clavulinopsis fusiformis Dermoloma atrocinereum Dermoloma ciineifoliunt Entoloma ameides Entoloma asprellum Entoloma atrocoenileum Entoloma cocles Entoloma cotuinum Entoloma excentricrim Entoloma farinaspreIlum Entoloma fomzosum Entoloma hispidulum Entoloma buysmanii

Entoloma incanum Entoloma jiibatum Entoloma juniperinum* Entoloma kuebnerianunz Entoloma mougeotii Entoloma poliopus Entoloma pseudoturci Entoloma sodale Entoloma solstitiale Entoloma turci Entoloma xantbocbroum Geog lossirm cookeianum Geoglossum fa flax Geoglossum glutinosum Geog lossurn nigritum Hygrocybe aurantioviscida Hygrocybe calciphila Hygrocybe calyptraeformis Hygrocybe ceracea Hygrocybe coccinea

Hygrocybe co lemanniana Hygrocybe constrictospora Hygrocybe fornicata Hygrocybe glutinipes Hygrocybe lacmus

(= subviolacea) Hygrocybe laeta Hygrocybe luteolaeta

Hygrocybe marchii Hygrocybe pratensis Hygrocybe psittacina Hygrocybe reai Hygrocybe reidii Hygrocybe russocoriacea Hygrocy be subg lobispora Ramariopsis pu lchel la Rhodocybe caehta Rbodocybe parilis Tricboglossurn hirsirtum

(= vitellina 5s. auct.)

3. Species of rather old, unfertilized grasslands, rather sensitive to changes in management, mostly decreasing in W. Europe.

Agaricus cupreobrunneus Agaricus porphyrocephalus Agrocybe pusiola Clavaria acuta Clavaria vermicularis Clavulinopsis coniiculata Clavrrlinopsis helveola Clavulinopsis laeticolor Clavu linopsis Iuteo-alba Clitopilus scypboides

Entoloma caesiocinctum Entoloma cbalybaeuni Entoloma clandestinum Entoloma fernanhe* Entoloma incarnatofuscens

( = cretatus)

Entoloma in/& Entoloma pallens Entoloma papillaturn Entoloma psilopus* Entoloma sarcitu lum Entoloma sericellum Entoloma semlatunz Entoloma turbidum * Entoloma undatum Entoloma vinaceum* Entoloma xantbocau Ion * Galerina calyptrata * Galerina cepha lotricba * Galerina mniopbila* Galerina pumila * Galerina unicolor*

Gymnopilus flavus Hygrocybe conica Hygrocybe conicoides Hygrocybe insipida Hygrocybe konradii Hygrocybe miniata Hygrocybe persistens

(= acutoconica) Hygrocybe virginea

(= nivea ss. auct.) Lepiota alba Psilocybe liniformans (on dune)

Ramariopsis kunzei Ramariopsis tenuiramosa Rhodocybe popina lis

4. Species of rather young to old, not to mederately fertilized grasslands, to some extent sensitive to changes in management, with a tendency to decrease in grasslands in W. Europe.

Agrocybe pediades Bovista pusilla Anellaria semiovata (on dung) Calocybe carnea

Calocybe constricta ( = leucocepba la)

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Clitocybe vibecina* Co llybia butyracea * Collybia dryophila* Crinipel lis stipitarius * Cystoderma Amianthinrim * Cystodema jasonis* Entoloma conferendrim

(= staurosporum) Entoloma neglectttm

Entoloma sericeum Galerina atkinsoniana * Galerina Hypnorum* Galerina vittaeformis * Lepista iuscina Lepista personata Lycoperdon liuidum ( = spadicerim)

hfacrolepiota excoriata

hfacro Iepiota gracilenta Afycena cinerella* Alycena pelliculosa* Omphalina pyxidata* Puneolus campanukztus

Pfeurotus eryngii Psilocybe montana

(on dung)

5. Species of both unfertilized and fertilized, stable and disturbed grasslands (neutral species) mostly constant in grasslands in W. Europe.

Agaricus campester Bovista nigrescens Bovista plumbea Calvatia utrifomis Clitocybe dealbata Clitocybe riuulosa Conocybe magnicapitata Conocybe rickeniana Conocybe semiglobata Conocybe sienophylla Ga lerina heterocystis * Hemimycena dekzctab fir Hemimycena mairei Leucoagaricus pudicus Leucoagaricus giganteus

hlacrolepiota procera Aiarasmius graminurn hfarasmius oreodes hfelanoleuca exscissa

( = cinerascens) hlelanoleuca polio leuca *

(= maleleuca* 5s. auct.) hlycena avenacea hlycena )?lopes* hlycena f7avoalba hlycena Ieptocephala * Afycena pura * Afycena sepia Ompha lina microspema Omphalina obscurata

Paneolus acuminatus (= rickenii)

Paneolus ater Psathyrel la panaeoloides * Psilocybe inquilina Psilocybe semilanceata Rickenella fibula* Rickenella setipes* Stropharia coronillo* Stropharia inuncta Stropharia pseudocyanea Tubaria furfurarea* Vasceilirm pratenre

6. Species of disturbed, fertilized, often young grasslands, mostly increasing in grasslands in W. Europe.

Agaricus amensis* Agaricus macrospoms Agaricus subperonatus * Agmcybe amah Aleuria aurantia * Bolbitius vitellinus Clitocybe agrestis Clitocybe amarescens

Conocybe ambigiia Afacrolepiota rhacodes * Conocybe nibiginosa Panaeolina foenesecii Conocybe sifiginea Panaeolcts fimicokz Conocybe tenera Panaeolus subbalteatus Coprinus impatiens* Stropharia caerulea* Coprinus p Iicatilis 5s. kzto * Volvariella speciosa Lepista nu&* (= gloiocephakz) Lepista sordida

environmental conservation. A first, tentative classification was made of a selection of 225 species, indicative of the duration of undisturbed grassland use and sensitivity for disturbance in northwestern Europe (Table 3). The species are arranged in six groups, including a group of indicators for disturbed grasslands (<<negative indicator vaIue4 and a group of species without preference for disturbed or undisturbed grasslands. The lists include a few coprophytic species, which seem to be characteristic of grasslands, poor in nutrients.

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