Remarks to the ecology of the boreo-montane polypore Amylocystis lapponica based on data from the Czech Republic and Poland

In the Czech republic, the rare polypore Amylocystis lapponica (romell) Singer continuously occurs in the Boubínský prales virgin forest (southern Bohemia: šumava mts) where it has been documented for more than 60 years. Similarly, in Poland it has been known only from the Puszcza Białowieska virgin forest (northeastern Poland) for more than 50 years. Generally, it is considered a species of boreal coniferous forests of northern europe (taiga) and montane coniferous forests in central and Southern europe. However, the data from the Czech republic, Slovakia and western ukraine show that it also grows in mixed montane forests composed mainly of Fagus sylvatica, Picea abies and Abies alba. In Poland, the locality is situated in a lowland forest consisting mostly of Carpinus betulus, Quercus robur and Tilia cordata. In central europe, A. lapponica occurs only in virgin forest refuges with the following habitat conditions: vegetation continuity (never cut), natural tree species composition, multiaged structure, rich presence of dead wood in various stages of decay, relatively large area of the virgin forest surrounded by near-natural forests, stable, cold and humid mesoand microclimate. consequently, A. lapponica may be considered an indicator of long-term vegetation continuity and stable habitat conditions.

virgin forest (Białowieża National Park, PL). The occurrence in the Czech republic was summarised by K o t l a b a (1984) and K o t l a b a et al. (1995), who used data from older publications by K o t l a b a and Po u z a r (1963) and P i l á t (1965), labels of herbarium collections and field notes. more recent collections from the Boubín area were published on-line by V l a s á k (2005). In Poland, the occurrence of A. lapponica was recently evaluated by P i ą t e k (2005).
From the conservational viewpoint, Amylocystis lapponica is considered a critically endangered species, included both in the red Book (K o t l a b a et al. 1995) and red List of fungi of the Czech republic (H o l e c , B e r a n 2006). Consequently, it is protected by law in the country (a n t o n í n , B i e b e r o v á 1995). In Poland, the species is also included in the national red List as a critically endangered species (Wo j e w o d a , Ł a w r y n o w i c z 2006) and also protected by law (Ł a w r y n o w i c z 2004; also see K a r a s i ń s k i 2007: http://grzyby.strefa.pl/chronione.html).
recently, D a h l b e r g and C r o n e b o r g (2003) summarised data on habitats of Amylocystis lapponica in europe. unfortunately, the habitats present in the Boubínský prales and Puszcza Białowieska virgin forests are not mentioned.
The aims of this paper are (1) to summarise older data and publish new data on the occurrence of Amylocystis lapponica in the Boubínský prales virgin forest, (2) to compare the habitats of A. lapponica in the Czech republic and Poland, and (3) to add data on habitat preferences of A. lapponica in these countries to the incomplete habitat characteristics published by D a h l b e r g and C r o n e b o r g (2003).

maTerIaL aND meTHoDS
The search for Amylocystis lapponica in the Boubínský prales virgin forest by J. Holec was not intensive and not focused only on this species. It was carried out as part of a study of mycobiota in the šumava mts in the years 1996-2006 (see e.g. H o l e c 2000; H o l e c , B e r a n 2007). Consequently, his find of A. lapponica is rather accidental. However, when combined with published and herbarium data and provided with a phytosociological analysis, it can be used for an ecological synthesis. This paper is a reply to a request from the agency for nature conservation and Landscape Protection of the Czech republic for monitoring protected species of fungi.
Habitat of Boubínský prales. mixed montane forest composed of Fagus sylvatica, Picea abies and Abies alba, with admixed Acer pseudoplatanus and Ulmus glabra, phytosociologically a mosaic of herb-rich beech forests, montane sycamore beech forests, acidophilous beech forests and montane spruce forests with Calamagrostis villosa (a l b r e c h t et al. 2003), with some trees aged 400-500 years. The core area (46.67 ha) has never been cut nor managed by foresters. It represents a remnant of original natural vegetation, with a high amount of dead and fallen decaying trunks. It has been protected as a nature reserve since 1858. Forest surrounding the core area is also natural. The total area of the reserve is 677 ha, its altitude ranges from 874 to 1362 m, the core area (enclosed by a palisade) from 930 to 1100 m. Coordinates of the centre of the core area are 48° 58´ 39´´ n, 13° 48´ 41´´ e.
Mycological characteristics of Boubínský prales. See K u b i č k a (1960,1973), partial data are present in dozens of publications on taxonomy, biodiversity and ecology of fungi both in Czech and foreign literature.
Records of Amylocystis lapponica in the Boubínský prales. For details see Table 1. on mt. Boubín, the species is known from several microlocalities. most of them are situated in the core area (enclosed by a palisade, see below) of the Boubínský prales National Nature reserve.
The exact geographic position of older finds by J. Herink and J. Kubička is not known, but it is highly probable that they originate from the core area of BPVF which was traditionally visited by mycologists at that time. However, finds of A. lapponica are known from more microlocalities in the mt. Boubín complex. The exact location of finds by V l a s á k (2005) is not clear from data published by him but it is probable that those indicated by a question mark in Table 1 are from BPVF (core area enclosed by a palisade plus surrounding parts of the National Nature reserve). However, Vlasák mentions also two finds from mt. červený vrch which is situated in the mt. Boubín complex but approximately 3.5 km S of the summit of mt. Boubín and outside of the Boubínský prales National Nature reserve. There are also nearnatural forests on slopes of mt. červený vrch which resemble those in the BPVF. The well-documented find by Holec published here (see below) is situated in the National Nature reserve but outside the core area. Good colour photographs of A. lapponica from Boubínský prales were published by Pa p o u š e k (2004: no. 237).
The This is a potential natural vegetation type, which would be present if the site were free of human influence. In fact, the site is a high-grown (approximately 100 years old) man-influenced forest with dominance of Picea abies and admixed Fagus sylvatica. It is not a virgin forest (trees are of the same age, dominance of Picea instead of Fagus, low amount of dead wood). However, natural forest almost untouched by man (virgin forest) is situated at 10 m distance (behind the palisade enclosing the core area of the Nature reserve).

DIScuSSIon
Occurrence in the Czech Republic. To the present date, Boubínský prales virgin forest is the only locality of Amylocystis lapponica in the Czech republic. The species has not been found in other virgin forests of the country (e.g. Žofínský prales in south-eastern Bohemia, mionší, Salajka and razula in north-eastern moravia) in spite of a long and intensive study of polypores in the Czech republic (e.g., P i l á t 1936-1942; K o t l a b a 1984; K o t l a b a , Po u z a r , Va m p o l a in H o l e c , B e r a n 2006).
The data document continuous occurrence of Amylocystis lapponica on Mt. Boubín for more than 60 years. The vitality of the local population is illustrated by the fact that the species is known from several microlocalities. The following habitat conditions (which are unique in the Central european landscape) seem to be decisive: vegetation continuity (never cut), natural tree species composition, multi-aged structure, rich presence of dead wood in various stages of decay, relatively large area of the virgin forest surrounded by near-natural forest (the importance of larger reserves for maintaining sensitive species was stressed by S i i t o n e n et al. 2005), and a stable, cold and humid meso-and microclimate. as it was illustrated in another paper (H o l e c , B e r a n 2007), these "virgin forest conditions" of BPVF enable the occurrence of other fungi preferring natural forests (e.g., Phellinus pouzarii Kotlaba, Comparison of Czech and Polish habitats. The following habitats of Amylocystis lapponica in the Puszcza Białowieska virgin forest are given by P i ą t e k (2005): Vaccinio uliginosi-Pinetum Kleist 1929, Tilio cordatae-Carpinetum betuli Tracz. 1962, and the grouping association Querco-Carpinetum medioeuropaeum Tüxen 1936. In all cases, the species was found on fallen trunks or branches of Picea abies. Puszcza Białowieska virgin forest is a lowland forest (mean altitude 170 m), where spruce grows in wet depressions, the microclimatological conditions are colder than in the surrounding.
In the Czech republic, the locality is surrounded by the natural montane sprucefir beech forests of the very large coniferous complex of the šumava mts (montane spruce forests in upper, and spruce cultural forests in lower position, dozens of square kilometres). In contrast, the Polish locality is situated in a lowland deciduous forest complex with a higher proportion of pine and spruce forests in wet depressions near the bog. Therefore both localities are especially similar in their historical management (minimal influence by man) rather than habitat.
Remarks to habitats and occurrence in Europe. D a h l b e r g and C r o n e b o r g (2003) compiled the following data on habitats of Amylocystis lapponica in europe: western taiga (Natura 2000 priority habitat 9010) and Fennoscandian herb-rich forests with Picea abies (9050) in nordic countries and estonia, acidophilous Picea forest of the montane to the alpine levels (9410) in Slovenia and alpine Larix decidua and/or Pinus cembra forests (9420) in Italy. It is clear that habitats known from the Czech republic and Poland are not included. These differ e.g. by a high percentage of broad-leaved trees (esp. Fagus sylvatica, Carpinus betulus, Quercus robur, and Tilia cordata) forming the habitats Asperulo-Fagetum beech forests (9130, Boubínský prales) and Galio-Carpinetum oak hornbeam forests (9170, Puszcza Białowieska), respectively.
The number of localities in various european countries is very different. While hundreds of localities in Sweden and Finland and dozens in Norway (D a h l b e r g , C r o n e b o r g 2003) are known, in countries of Central and Southern europe the species is extremely rare and mostly occupies only one locality per country. It clearly shows that localities in central and Southern europe represent refuges of A. lap-ponica outside the zone of boreal forests of northern europe. These refuges are formed by azonal soil or topoclimatic conditions in the temperate forest zone.
In northern europe, Amylocystis lapponica is considered as a good indicator of old-growth and species-rich spruce forests which are in special need of conservation  (1999) showed in Swedish spruce forests that if indicator species are to be used for evaluation of their biodiversity, these should be chosen from several groups of organisms (e.g. vascular plants, bryophytes, epiphytic lichens and wood-inhabiting fungi). This is certainly true but at least in Central europe, A. lapponica indicates high continuity of forest vegetation which has never been cut. Supporting data from the Czech republic and Poland are summarised here. Data from Slovakia (K o t l a b a , Po u z a r 1963; P i l á t 1965; K o t l a b a 1984; Va r j ú 1994; š k u b l a 2003) and eastern Carpathians in western ukraine (P i l á t 1936-1942, 1940; for current names of his localities, see H o l e c 2002) are almost identical. Amylocystis lapponica occurs there exclusively in virgin forests (central Slovakia: Dobročský prales, voucher specimens in Prm and herb. Vlasák; eastern Carpathians: altogether 19 collections from several localities, voucher specimens in Prm). These forests are composed mainly of Fagus sylvatica, Abies alba and Picea abies and have the same "virgin forest character" as was specified in the first chapter of the Discussion.