Environmental Pollution 130 (2004) 1732 www.elsevier.

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Vegetation of the selected forest stands and land use in the Carpathian Mountains
Krystyna Grodzinskaa,*, Barbara Godzika, Witold Frcczekb, Ovidiu Badeac, a c d e Julius Oszlanyi , Daniela Postelnicu , Yuriy Shparykf
w, Institute of Botany, Polish Academy of Sciences, 31-512 Krako Lubicz Str. 46, Poland b Environmental Systems Research Institute, Redlands, CA 92373-2853, USA c Forest Research and Management Institute, 72904 Bucuresti, Stefanesti Str. 128/2, Romania d nikova Str. 3, Slovakia Institute of Landscape Ecology, Slovak Academy of Sciences, 814 99 Bratislava, Stefa e nia SRL, Str. Roma nr 8, 71219 Bucuresti, Romania Geosystems Roma f The Ukrainian Mountain Forestry Research Institute, Hrushevsky St. 31, 76 000 Ivano-Frankivsk, Ukraine Received 30 June 2003; accepted 17 October 2003
a

Capsule: Vegetation and land use maps of forested mountain areas in central Europe are presented.
Abstract Within the framework of the project Eects of forest health on biodiversity with emphasis on air pollution in the Carpathian Mountains 26 permanent study sites were established in the vicinity of the ozone monitoring sites. The study sites were located on the NWSE transect through the Western (12 sites), Eastern (11 sites) and Southern (3 sites) Carpathians in forest ecosystems typical of each area. Some of the forest monitoring sites were located in national parks, biosphere reserves and areas of protected landscape. Each permanent site of 0.7 ha area consisted of 5 small 500m2 circular plots, arranged in the form of a cross, i.e. four placed on the cardinal points (N, E, S, W) and one in the center. Phytosociological records were done twice during the 1998 growing season using the BraunBlanquets method. The study sites represented various types of forest: Picea abies stands (8), beech (Fagus sylvatica) stands (10), r (Abies alba) stands (2) and mixed beechr, sprucer and beechspruce stands (6). Age of most stands was 80100 years. Degree of crown damage varied greatly between sites, a percentage of damaged trees decrease in Carpathians from West to East. It corresponds well with the O3 level in these areas. Typical damage by O3 in herb layer species in several Carpathian sites were found. Land-use map for the entire Carpathian Mountains and two detailed land use maps for Tatras (Western Carpathians) and Retezat (Southern Carpathians) are presented. A little more than half of the Carpathian territory is forested. The most densely forested are Eastern Carpathians, while the most sparsely Western Carpathians. Arable lands occupy 22.6% of the Carpathians, pastures and meadows 6.2%, water bodies 1.9%, and build up areas several percent. In the highest elevation of the Carpathians alpine meadows (11.3%) and rocks (3.5%) are distributed.
Keywords: Forests; Species diversity; Forest health; Ozone; Carpathian Mountains

1. Introduction There is a growing concern among scientists, managers of natural resources and conservationists about diminishing diversity of biological resources (Wilson, 1988, 1989). Many investigations suggest that biodiversity may be declining throughout the world in response to anthropogenic stresses such as habitat

* Corresponding author. Tel.: 48-12-421-56-78; fax: 48-12-421-97-90. E-mail address: grodzin@ibpost.ib-pan.krakow.pl (K. Grodzinska). 0269-7491/$ - see front matter # 2003 Elsevier Ltd. All rights reserved. doi:10.1016/j.envpol.2003.10.031

modications and input of toxic chemicals to the air, water, and soil (Barker et al., 1991; Szaro and Johnston, 1996). During the past 50 years, large areas of forests in Eastern and Central Europe have come under the adverse inuence of air pollutants. This has brought about a serious eect on forest sustainability and yet to be determined eects on biodiversity. The eects of environmental change on biodiversity including the impact of air pollution have been recognized as a high priority thrust for future research (Szaro and Johnston, 1996). The Carpathians harbor some of the most beautiful natural areas in Europe and are a unique reservoir of

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many endemic plant and animal species. Numerous national parks and nature reserves have been established in the Carpathianssome of them are on the UNESCO list of specially protected areas. Forests in the Carpathian Mountains have been under human impacts for many centuries, however, most of those impacts have been quite local. In the past 4050 years the long-range transport of pollutants from various European industrial areas has resulted in the high deposition of sulfur, nitrogen and heavy metals in the Carpathian forests (Grodzinska and Szarek-ukaszewska, 1997). The industrial air pollutants have caused an extensive decline of the Carpathian forests (Godzik and Sienkiewicz, 1990; Grodzinska et al. 1995; Oszlanyi, 1997). The toxicity of some of these pollutants, especially of sulfur and nitrogen oxides (SO2 and NOx, respectively) can be enhanced by the presence of ozone (O3) (Guderian et al., 1989). Recently, the rapidly growing number of cars in Central and Eastern Europe resulted in an increase of NOx and hydrocarbons emissions, which lead to higher rate of photochemical-smog formation. The long-range transport of ozone (O3) from Western Europe and O3 generated locally are likely to result in elevating concentrations of this highly phytotoxic pollutant in Central and Eastern Europe, including the Carpathian Mountains. An increase in ozone concentrations is possibly a major factor in continued mountain forest decline of the region, despite the recent reduction in sulfur dioxide (SO2) and heavy metal concentrations. To prove this hypothesis, information from a network of O3 monitoring sites and vegetation survey plots are needed in the Central European mountain regions. The aims of two joint projects Evaluation of ozone air pollution and its phytotoxic potential in the Carpathian forests and Eects of forest health on biodiversity with emphasis on air pollution in the Carpathian Mountains which started in 1996 and 1997 were: (1) to determine levels of O3, SO2, NO2 air pollutants, and (2) to describe forest vegetation and health status of selected forest stands in the Carpathian Mountains. In this paper species composition, geographical, ecological, dendrological characteristics and health status of selected Carpathian forest stands as well as the land use patterns in the whole Carpathian range are described.

2. General characteristics of the Carpathian Mountains The Carpathians are the largest mountain range of Central Europe in terms of maximum elevation and the total span of the mountain chain (Fig. 1). Geographically, the Carpathians are divided into three sections: Western, Eastern, and Southern (Transylvanian Alps) (Warszynska, 1995; Volos cuk, 1999) (Fig. 2). The length of the Carpathian Mountains arch is about 1400

km while its width varies from about 170 km in the central sections of the Eastern and Western Carpathians to less than 80 km in the central part of the Southern Carpathians and the eastern part of the Western Carpathians. There are six of the outstanding sub-ranges with alpine relief, out of which the Tatras (its primary summit, Gerlachovsky Stit, is elevated 2654 m a.s.l.) shared by Slovakia and Poland, is the highest. The remaining ve ranges with extensive bare rocks areas are located in Romania: Fagaras (2544 m), Paring (2519 m), Retezat (2509 m), Bucegi (2505 m), and Rodnei (2303 m) Mountains. All these ranges are in the Southern Carpathians, except for the last one, which is in the Eastern Carpathians close to the Ukrainian border. Politically, the Carpathian mountain range is administered by ve countries. Of the total area of the Carpathians (158,000 km2), 52.9% is shared by Romania, 18.8% by Slovakia, 14.8% by Ukraine, 10.8% by Poland, and 2.7% by the Czech Republic. The Carpathian range is situated at the edge of the Atlantic and Continental climatic regions. The western climatic type with its anticyclone weather pattern dominates over most of these mountains. However, on the eastern slopes of the Eastern Carpathians, the continental climate prevails. In the foothills of the Western Carpathians the mean air temperature of July is 19  C and in the Southern Carpathians 22  C. In the southwestern part of the Carpathians, with every 100 m increase of altitude, a temperature decreases by 0.81  C (temperature gradient), and in the southeastern part of the Carpathians by 0.50  C. Annual precipitation amounts from approximately 500 mm at the southeastern foothills of the Southern Carpathians to over 2000 mm at the summits of the Tatra Mountains (Volos cuk, 1999). Altitudinal changes in climatic conditions and changes in species and vegetation units distribution allowed to distinguish several vegetation belts. Altitudinal limits of particular vegetation belts usually dier slightly from one Carpathian sub-range to another, depending on their massiveness, elevation and geographical location (Mirek and Pickos -Mirkowa, 1992) (Fig. 3). The sube c montane (foothills) belt ranges from 400 to 650 m a.s.l. Originally, mostly oakhornbeam forests and mixed oakpine forests covered this belt. The lower forest montane belt extends between 650 and 1250 m a.s.l. It is occupied by beech forests and in some ranges by beech r, r-spruce or mainly articial spruce forests. The upper forest montane belt ranges from 1250 to 1550 m a.s.l. and is dominated by spruce forests. The subalpine, alpine and subnival belts occupy the highest elevations in the Carpathian Mountains from 1550 to 2655 m a .s. l. High latitudinal dierences and large extent of Carpathian Mountains as well as favorable ecological conditions resulted in abundant variety of plants and animals in the post-glacial period, and rich biodiversity.

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Fig. 1. Location of the Carpathian Mountains in Europe.

In this area more than 3200 species of higher (vascular) plants grow representing more than a quarter of the European ora. Of special importance are Carpathian endemic species, which constitute approx. 12% of the total ora. There are many endemic and relic fauna species representing most of systematic groups of Europe (Volos cuk, 1999).

3. Data and methods 3.1. Land use The spatial denition of the Carpathian Mountains and the delineation of digital data chosen to illustrate the reported research were performed using the methods and technology of Geographic Information Systems (GIS). The GIS analyses were completed and the maps were created using ArcMap/ArcGISTM 8.3 software produced by Environmental Systems Research Institute, Redlands, California. Analysis of the land use information layer with a special emphasis on forest category was considered the principal criterion-determining outline of the mountain range. From the land use data of the neighborhood of the Carpathians, forest stands larger than 100 ha were extracted. The dense cluster of the forested areas was strongly spatially concentrated along the axis of the

Carpathians. All these forests forming the monotonous forested zone were considered as the Carpathian framework. The collateral criteria were derived from analyses of the underlying geologic structure and elevation data. Conceptually, in a manner similar to the grouping of the signicant forest stands, the elevation data were used. The elevation data included into the polygon of the monotonous area contained all geomorphic forms of the maximum altitude higher than 600 m a. s. l. Finally, the precise shape of the outline of the Carpathian Mountains was determined by including all the qualied features of all three information layers within a polygon dening the Carpathian Mountains study area. The shape of the outline polygon was smoothed for cartographic reasons (Bytnerowicz et al., 2002a,b). The digital land use data applied in this project was adapted from the Co-Ordination of Information on the Environment (CORINE), Landover Project funded by the European Union and completed in 1998. The aim of the CORINE project was to accurately and uniformly map the land use of Europe based on computer-assisted photo interpretation of satellite data (Landsat TM and SPOT) by using ancillary data (topographical and thematic maps, statistics, etc.). The resulting land use data for Europe consisted of 44 dierent land use types. The surface area of the smallest mapped unit was 25 ha (an area of 500500 m). For linear elements, such as rivers, the minimum width of a mapped geographic object was

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Fig. 2. Geographical divisions of the Carpathian Mountains.

100 m. The interpretation and map compilation results were scanned and integrated into a GIS to create a digital map comparable to a topographic map of 1:1 000 000 in terms of its spatial accuracy. Unfortunately, the territory of Ukraine was not covered by the CORINE project. The relevant land use data for the Ukrainian section of the Carpathian Mountains were obtained from the Ukrainian sources and merged with the data for the other four Carpathian countries (Romania, Slovakia, Poland, and the Czech Republic). The acquired Ukrainian data were signicantly less detailed in terms of both, spatial resolution and the number of land use types, as it was compiled of only 10 land use categories. However, for the purpose of this project, which focused mostly on main forest types, the Ukrainian data were applicable. In the next step, the combined land use classication of the CORINE and the Ukrainian data was rasterized into 100 m resolution raster and reclassied into the same 10 categories as determined by the Ukrainian data. 3.2. Vegetation Twenty-six permanent vegetation study sites were established in the vicinity of the ozone monitoring sites

in the Carpathian Mountains (Fig. 4) throughout the Czech Republic, Slovakia, Poland, Ukraine and Romania. Twelve of the sites were located in the Western Carpathians, 11 in the Eastern Carpathians and three in the Southern Carpathians; most of them in national parks, biosphere reserves and areas of protected land scape (Volos cuk, 1999). The study sites were situated at elevations ranging from 520 to 1360 m a.s.l., with most of them between 700 and 1100 m a.s.l. (Table 1). The study sites diered in exposure, inclination of slopes and bedrock type. Each permanent study site was 0.7 ha in size. According to the criteria established by the European Intensive Monitoring Network for forest ecosystems, each site consisted of ve 500 m2 circular plots (Fig. 5). In each plot all trees (> 7 cm in dbh) were marked permanently (number on a trunk) and their height and diameter at a breast height (dbh) were measured. Phytosociological records were done in each plot twice over the growing season (spring, summer 1998), using the BraunBlanquets method (Szafer and Zarzycki, 1972). The phytosociological material comprised 260 records (10 records during the growing season for each of the 26 permanent plots). Each permanent plot (total area of 2500 m2) is represented by one phytosociological record (Table 2).

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Fig. 3. Vegetation belts in the Carpathians ranges (Mirek and Pickos -Mirkowa, 1992). Western Carpathians: 1Nizke Tatry; 2Babia Gora; 3 e c Tatras; 4Gorce. Eastern Carpathians: 5Bieszczady; 6Cernohora; 7Cyvcyny Mts.; 8Rodnei; 9Calimani. Southern Carpathians: 10 Siriu; 11Bucegi; 12Sebesuku; 13Retezat; 14Valdeasa. Vegetation belts: asubmontane; blower forest montane; cupper forest montane; dsubalpine; ealpine; fsubnival.

Cover of each species in 10 small plots (500 m2) was determined using a six-grade scale (for explanation please see Table 2) (Szafer and Zarzycki, 1972). Species were listed according to their frequency of occurrence in the phytosociological material. Constancy of species was determined using a ve-grade scale (for explanation please see Table 2) (Szafer and Zarzycki, 1972). Sporadic species were listed below in Table 2. The nomenclature of vascular plant species follows Mirek et al. (1995) and the moss species Ochyra and Szmajda (1978).

4. Results 4.1. Land use in the Carpathian Mountains Due to the lack of the cartographic legibility, it was not feasible to present the detailed, 10-class land use classication map of the Carpathians at the scale applicable for this publication. Because of that, the authors decided to include the generalized map of the land use data with only four classes but showing the entire territory of the Carpathians (Fig. 6). However, in

order to show richness of the 10 class land use data as well as the dissimilarity in land use between the regions, two additional maps in a much larger scale were compiled for the vicinities of the Tatras and Retezat Mountains. These two sections of the detailed land use map are presented as Figs. 8 and 9. The signicant dierences in climate and geography of this 1400 km long, arch shaped mountain range result in a complex pattern of the land use classes distribution. For instance, the elevation of the timberline, which inuences positioning of ecological zones varies depending of the aspect of the mountain range, e.g. it reaches higher elevations on the southern (Slovak) side of the mountains than on the northern (Polish). The dierences between timberline elevation on the northernmost section of the Carpathians and the southernmost one are as large as 200300 m. The alpine meadows located above the timberline cover 11.3% of the entire mountain area, and occur along the ranges major axis. The highest zone of the Carpathian Mountains are the outcrops of bare rocks that make 3.5% of their total area. Territories designated as an agricultural land use category (mainly arable land) are located at the lowest

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Fig. 4. Location of study sites in the Carpathian Mountains (see also Table 1).

Table 1 Characteristics of permanent plots in the Carpathian Mountains Locality 1. Bily Kriz 2. Javorina 3. Male Karpaty 4. Mala Fatra 5. Kozie Chrbty 6. Polana 7. Stoliky 8. Brenna 9. Babia Gora NP 10. Tatry NP 11. Pieniny NP 12. Magura NP 13. Morske Oko 14. Bieszczady NP 15. Uzhokskij Pass 16. Kuzij 17. Synevir 18. Yablunitsa 19. Krivopilja 20. Vizhnitsa 21. Obcina Mare 22. Rarau 23. Magura Odobesti 24. Fundata 25. Retezat 26. Stana de Vale Latitude 49 490 2000 48 510 2000 49 250 5700 49 020 4600 49 020 4600 48 380 1700 48 530 8800 49 400 0700 49 350 1700 49 170 0700 49 250 3700 49 320 0000 48 540 5000 49 060 3800 49 000 1500 48 010 3700 48 370 2400 48 170 5000 48 130 0000 48 090 4000 47 440 1700 47 280 2300 45 510 4900 45 250 4600 45 180 3800 46 410 4100 Longitude 18 320 4500 17 400 0500 17 180 0600 19 0400 5300 19 550 0500 19 320 2800 20 300 3000 18 560 3700 19 350 0000 20 070 0000 20 210 3200 21 300 0000 22 120 0100 22 350 4000 22 520 5700 24 070 2000 23 410 5000 24 260 4600 24 390 5400 25 200 5500 25 380 3700 25 320 3300 26 560 4400 25 160 1400 22 470 5400 22 380 2100 Altitude (m a.s.l.) 910 590 660 880 840 855 875 600 1100 1080 640 520 640 880 910 850 1050 960 1020 750 1100 980 960 1360 1250 1300 Exposure S NW NW SW NW NE SE W NW E S SW E N E SW SW SE W SW NE N E SE N W Inclination ( ) 7 15 1540 1545 550 7 10 15 25 1015 1530 310 25 520 38 1520 25 20 1520 20 25 10 1020 2540 530 20

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Fig. 5. Schematic of an intensive permanent observation plots with ve subplots.

Fig. 6. Land use map for the Carpathian Mountains.

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elevations of the mountains. This land use class makes as much as 22.6% of the Carpathians. The pastures class and meadows covers 6.2% and is usually situated at small hilltops just above arable areas, or in large forest clearings when these are placed in valleys. The spots of build-up areas make 3.1% and are scattered typically at low elevations in the neighborhood of the agricultural regions. Water bodies, usually reservoirs, but also numerous, small-size, natural mountain lakes make total of about 1.9% of the Carpathians. A little more than half of the Carpathian Mountains territory is forested. The distributions of forest in general, as well as the individual types of forests are uneven within the Carpathians. The most densely forested are the Eastern Carpathians (both in the Ukraine and Romania), while the most sparsely forested are the Western Carpathians, especially on the Polish side of the mountains and at the westernmost part of the entire range. Hardly any forest can be found at low elevations in the entire range due to agricultural and residential development. Forests categorized into the coniferous, mixed and deciduous types, with the applied land use classication over 18.3, 10.3, and 22.5% of the Carpathian territory, respectively (Fig. 7). The three distinguished classes of forest are also unevenly dispersed. The coniferous forest covers the highest forested elevations and creates a huge

block of forested area on the northeastern side of the Ukrainian Carpathians. In the Southern Carpathians, due to a warmer climate, the deciduous trees are predominant except for the highest elevations, where the coniferous forests are characteristically set astride of the main mountain range. The deciduous forest is the major forest type in most of the Romanian Carpathian, except for the highest altitudes where the mixed and coniferous are most common. Noticeably, large continuous section of a pure broadleaf forest is located in the central section of the Carpathian arch, particularly in its Ukrainian segment, but also in eastern Slovakia, Polish Bieszczady Mountains, and Romanian Maramures and Radui Mountains. The two maps presented at a larger scale are the enlarged sample maps of the land use in the Carpathians with 10 distinguished categories. Both areas are good examples of land use and vegetation zonality caused by elevation change and harsher climate. One of these detailed maps shows the land use pattern in the Tatra Mountains range and its vicinity (Fig. 8, site 10), while the other, the Retezat Mountains and its surroundings (Fig. 9, site 25). The maps are of the same scale, their map extents are of the same size and they were generated in the same manner. Similarly, both mountain ranges are of almost the same height. Hence, the information on these two maps is comparable.

Fig. 7. Forest types in the Carpathian Mountains.

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Fig. 8. Land use map for the Tatra Mountains region and surrounding areas (Western Carpathians).

Fig. 9. Land use map for the Retezat region and surrounding areas (Southern Carpathians).

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Table 2 Vegetation of forest stands in Carpathian Mountainsa

Part of Carpathian Mts. Table no. of record Date Elevation a.s.l. (m) Exposition Inclination (o) Cover of tree layer A (%) Cover of shrub layer B (%) Cover of herb layer C (%) Cover of moss layer D (%) Area of record (m2) Trees (A) Fagus sylvatica Western 1 2 3 4 5 6 7 8 9 10 11 Eastern 12 13 14 15 16 17 18 19 20 21 22 Southern 23 24 25 26

May/JuneAugust/September 1998 910 S 7 3560 1 95 15 590 660 880 840 855 875 600 1100 NE SE 7 10 95 540 1 110 70 100 <1 315 2500 1080 640 520 640 880 910 850 1050 960 1020 750 1100 980 960 1360 1250 1300 Constancy records 126 W 20 5095 510 2085 5

NW N SW NW 15 1540 1545 550 70 75 85 90 <5 5 25 545 5595 1075 3090 14 5 210 1050

W NW E S SW E N E 15 25 1015 2530 310 25 520 38 70 5070 6080 80 70 100 5090 6085 85 3040 550 30 30 1 1020 520 90 90 90 100 80 540 6070 1550 70 6075 30 <5 <1 <5 13

SW SW SE W SW NE N E SE N 1520 25 20 1520 20 25 10 1020 2540 530 5580 6080 7085 7595 95 80 60 6590 5080 5075 1520 1015 1020 510 15 2560 1560 1025 1540 520 540 1060 2585 2070 1065 13 25 25 25 37 110 <2 120 2550 2500 2500

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin

Abies alba

Picea abies

Acer pseudoplatanus

Sorbus aucuparia

A B C A B C A B C A B C A B C

+ + + + . + 34 + 1 . . . . . 1

45 + 2 . . +1 . . . . . + . . .

35 14 . . . . . . . . . + . . .

34 14 1 3 + + 12 +1 + . + 2 . . +

2 + 1 14 . 1 35 . + . + + . + 1

4 1 1 1 . + 2 . + . 2 + . . +

. . . 1 + + 13 . 1 + + + . 4 2 5 + 4 . . + . . . . . 1 + + +

. + . . . + 34 . 2 . . . . 3 1

. + . + + 2 4 3 2 . + . . + +

. . . 45 . 1 + . + . + + . + +

25 + + 34 + 1 . . . . . + . . +

5 + 1 . . . . . . 1 + 1 . . .

35 2 1 . . . . . . 2 . + . . .

34 35 14 13 14 1 24 2 + 13 13 + . . .

25 34 34 + . + . . . 3 . + . . .

3 45 24 15 12 13 5 2 23 . . . . . .

14 . 12 23 . 12 15 . 12 . . . . 12 .

13 . . +3 . + 45 . + . . . . . .

4 . 1 34 1 2 + . . . . . . 12 .

23 . . . . . 45 . . . . . . . .

24 . . 3 . . 24 . . + . . . . .

35 + . . . . . . . 2 . . . .

5 . . . . . . . . . . . . . .

. . . + . . 5 . . . . . + . .

24 . . + . . 3 . . . . . . + .

IV IV III IV II IV IV II III II II III II II

Shrubs (B) Corylus avellana Sambucus racemosa

B C B C

. . . .

. . . .

. . . .

+ + . .

+ . . .

. . + +

. . + .

. . . .

. . . .

. . . .

1 + + .

+ + . .

. . . +

. . . .

3 . 23 .

. . . .

+ . . .

. . . .

. . . .

. . 2 .

. . . .

. . . .

. . . .

. . . .

. . 1 .

. . . .

II I II I

Herbs (C) Oxalis acetosella Athyrium lix-femina Galeobdolon luteum Galium odoratum Mercurialis perennis Mycelis muralis Dryopteris lix-mas Rubus idaeus Polygonatum verticillatum Senecio fuchsii

+ + . . . . . + . .

12 3 . 2 . + . . . 1

1 . . 2 4 2 . . . +

2 + + 2 3 1 + + 3 2

24 + + . 1 + 2 12 + 1

2 12 12 2 1 + 1 1 + 1

1 1 2 1 1 1 1 25 + .

+ + + . . . + + . .

1 2 . . . . . 2 + .

2 +2 + . . . . 12 1

34 12 23 2 2 1 2 14 + 13

12 2 + . . . 2 . . 3

. 1 2 1 12 + + . . .

2 2 12 3 1 . 2 . + +

1 1 1 1 + + + 1 + 1

. 2 + 1 2 . + . + .

23 2 1 + . 1 . 2 + .

23 1 1 2 1 1 + + 2 2

24 1 + + + + + +

. 1 + 2 1 + 1 . . .

13 . . 1 . . + . . .

12 . + 1 . 2 . . . +

12 2 13 12 . . 1 . .

. . + 12 12 + . . 1 +

24 . . . + . 2 . . .

23 2 2 . . 1 . 12 . 1

V V IV IV IV IV III III III III

(continued on next page)

Table 2 (continued) Part of Carpathian Mts. Gymnocarpium dryopteris Rubus hirtus Urtica dioica Dentaria bulbifera Dentaria glandulosa Dryopteris dilatata Geranium robertianum Maianthemum bifolium Paris quadrifolia Salvia glutinosa Vaccinium myrtillus Actaea spicata Euphorbia amygdaloides Fragaria vesca Anemone nemorosa Calamagrostis arundinacea Impatiens noli-tangere Luzula luzuloides Prenanthes purpurea Viola reichenbachiana Luzula sylvatica Stellaria nemorum Carex sylvatica Deschampsia exuosa Glechoma hirsuta Hieracium murorum Milium eusum Sanicula europaea Veronica montana Gentiana asclepiadea Poa nemoralis Circaea lutetiana Cystopteris fragilis Galium schultesii Luzula pilosa Phyteuma spicatum Ranunculus lanuginosus Bryophytes (D) Dicranum scoparium Hypnum cupressiforme Polytrichum formosum Dicranum montanum Plagiothecium curvifolium Plagiothecium laetum Brachythecium salebrosum Brachythecium rutabulum Brachythecium velutinum Western . + . . . + . +2 . . 23 . . . . 2 . . . . . . . 35 . . . . . . . . . . . . . . 1 . . . . + . . + . . . . . . 1 . . 1 . . 1 . . + . + . . . + . . . . . . . 1 13 1 . 2 . . + . + . + . 2 . . + + . . . + . + . 1 + . 1 1 . + . . . + . + + 1 . + 2 + 1 + + 1 1 . . . . 2 2 . . . + . 2 . . . . + . . + . 1 + 2 . + . . + + 3 1 . 2 + 1 1 . + . . 1 . + . + . . 1 + + + 1 + . + 1 . + + + 1 1 2 . + 1 + + + . + . + 2 . + . + 1 . 1 + . . . . 1 + . . + . . . . + . + . + 2 . + . 1 . + 12 + . 1 + + . . . . 24 + . + . 1 . . . 2 24 + . 1 . + . + . . . + . . . + + 1 . + . 1 . . . + . + . 3 . + . . . . . . . . . . . 1 . + . . 23 . 2 . . 45 . . . . + . . + . 1 + . 1 . . . . . + . . . . . . . 1 . . . . 23 . 12 . . 34 . . . . 3 . 12 1 . + . . 2 . + . . + 1 . . . . + + . + . + + 2 1 + + + 12 . 1 + + + + 23 . . + . . + . + + + + . . + + . + + + + Eastern + 34 . . + 3 . 1 + . + . . . + + 1 . . . . + . . + . + . + . + . . . + . . . 2 + 1 2 . + . . + . + + . + . + . . + . . . . + . . . . . . . . . . . . + 23 + 1 3 1 . . + . + + + . 1 . + + 1 . . 1 + . + . . + + . . 1 . + . + + + 4 . . . . . 1 + + + . . . + . + . + . . . + . + 12 + . 1 . . + . . + . + . + 1 12 . . . + + . + . . 1 . . . . . . . + . . . . + . . . . . . . . . + 13 . . + . . . + . 1 . . . + 1 + . 1 . 1 + . . . . 1 . . . . . 1 . . . . + . . . 1 . + . + + . + + 1 + . . . . . 1 1 . . . . 1 . 2 . . + . 1 . + . . . 1 + . . . . . . + . . 1 . . . . . + . + . . . + . + . . . + . 1 . . . . . . . . . + . + . . + + . . + 1 . + . . . . . . . . . . . . . . . . . . + + . . + 1 . . . . . . . . . . + . + + + . . 1 . . . . . . . . . + . . Southern . 1 12 . . . . . + . . . 12 + . . . 12 . . . . . . . . 12 . . . . . . . . . . . . 2 + . . + . . + . . 1 + . . . 1 . . . . . . . + . . . . 12 . + . . . . . . . + . . . . . . 2 . . . . . . + . . 3 + . . . . . . . 1 . . . . . . . . 2 . 2 2 . . . . . 1 . . . 24 . . 12 . . 1 . . 12 . . . . . . . . . . . . . III III III III III III III III III III III III III III III II II II II II II II II II II II II II II II II II II II II II II

+ .. . . 2 + . + . . + 2 . . . 1 . 1 2 . . . 1 . . . . . . . . 1 . . .

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin

+ . + + . + . . .

. . 1 . . . . . .

. + + . . . 1 . .

2 1 + . . . 1 . .

2 + 1 . + . + + .

+ + + + . . . . . + . . 1 . 1 . + 1

2 . . + + + + . .

2 . 4 + + + . . .

+ . 4 + + + . . .

. + 4 + . + + . +

. + . + + + . + +

+ + + . + . . + 1

+ + . . . + + + .

. . + . . . . + .

. + . . . . . . +

+ + . . . + . . .

. . + . . . . . .

+ + + . . . . . .

1 . . . . . . . .

. . . . . . . . .

. . . . . . . . .

. . . . . . . . .

. . . . . . . . .

. . . . . . . . .

. . . . . . . . .

III III III II II II II II II

(continued on next page) 27

28

Table 2 (continued) Part of Carpathian Mts. Eurhynchium striatum Herzogiella seligeri Plagiothecium denticulatum Rhizomnium punctatum Tetraphis pellucida Number of vascular plant species incl. sporadic species Number of bryophytes incl. sporadic species Total number of species Western . . . . . 45 10 55 . . . . . 24 3 27 . . . . . 47 12 59 . . + . . 74 19 93 1 . 12 . . 91 22 113 1 + . . 1 . 1 1 . . 53 11 64 85 8 93 . + . . + 22 12 34 . . . . + 27 14 41 . . . . + 35 15 50 . + . + + 81 24 105 Eastern . + + + + 38 14 52 + + + + . 28 16 44 . . + + . 54 21 75 . . . . . 53 2 55 . + . . . 32 8 40 . . . . . 31 8 39 . + . . . 49 4 53 . . . . + 29 6 35 . . . . . 25 5 30 . . . . . 20 20 . . . . . 25 2 27 Southern . . . . . 40 1 41 . . . . . 30 1 31 . . . . . 22 3 25 . . . . . 27 1 28 II II II II II

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin

Sporadic species: Trees: Acer platanoides A 3:3; 4: 2; 6:1; 23: 1; B 6:1; 13; 23; C 3: 1; 5; 6; 13: 1; Betula pendula A 7; 20; 23; 25; B 7: 1; C 1; Carpinus betulus C 12; Fraxinus excelsior A 6:1; 13:1; 16:2-3; B 6:3; 11; 13; 16; C 2:1; 3; 4; 6:2; 13; 16:2-4; Larix decidua A 5; B 7; 25:1; Pinus sylvestris B 7:1; 16; Populus tremula A 6; 20; 23; B 6; 20; 23; C 6; Salix caprea B 7; Sorbus aria A 4; B 17: 4; C 4; Tilia cordata B 11; Ulmus glabra A 13:1-2; 16: 3; B 16; Shrubs: Crataegus sp. 12; Frangula alnus 16:3-4; Lonicera nigra 5; 11; 15:2; 18: 2; 25: 2; C 5:1; Lonicera xylosteum B 5:1; 11:2; C 4; 5: 1; 15: 1; Ribes alpinum C 11; Ribes petraeum B 5; C 5; Ribes uva crospa B 7; 11; C 3; 4; 5; 11; Rosa canina C 7; Rosa pendulina B 5; C 5; Rosa sp. 24; Sambucus nigra B 2; 6; 11:1-2; 13; 14; C 11; 12; 13; 14; Spiraea ulmifolia 10; Viburnum opulus B 11; C 5:1; 11; Herbs: Achillea millefolium 23:1; Aconitum moldavicum 4; Adoxa moschatellina 7; 14; 24:1; Aegopodium podagraria 7:3; 11: 2; 15; 16; 18; Ajuga genevensis 23; Alliaria petiolata 3; 6; 7; 11; 24; Alopecurus pratensis 23; Anemone ranunculoides 6:1; 22; Anthriscus sylvestris 15; Aposoeris foetida 15; Arabis alpina 3; Arctium lappa 6; Asarum europaeum 4: 1; 5: 1; 7: 1; 11: 1; Asplenium trichomanes 4; 5; 11; Asplenium viride 4; 5; Astragalus glycyphyllos 7; Astrantia major 4; Athyrium distentifolium 9; Barbarea vulgaris 7; Blechnum spicant 8; 9; Brachypodium sylvaticum 3: 2; 4:1; 11: 1; Bromus benekenii 4:1; 5:1; Bromus ramosus 6; Calamagrostis epigeios 7:2; Calamagrostis varia 4; Calamagrostis villosa 1:2; 5: 1; Campanula patula subsp. abietina 21:1; 24; 25; Campanula persicifolia 5; 7; 24; 26; Campanula trachelium 4:1; 5; 11; Cardamine amara 11; 14; Cardamine impatiens 7; 10; 24; Cardamine pratensis 14; Cardaminopsis arenosa 18; Cardaminopsis halleri 4; 5; Carex alba 4:4; 11; Carex digitata 4; 5; 11; 13; 20; Carex pilosa 20:1; Carex pilulifera 8; Carex remota 14; Carex spicata 23; Chaerophyllum hirsutum 5; 11: 1; 14; Chamaenerion angustifolium 7; 26: 1; Cheiranthus cheiri 23; Chrysosplenium alternifolium 7; 11; 14; 15; Cicerbita alpina 15; 17; 18: 1; Cimicifuga europaea 5; Circaea alpina 4; 11: 2; 14:1; 17; 18: 1; Cirisium erisithales 4; 5:1; Clematis alpina 3; 5:1; 7; Clinopodium vulgare 4; 5; 7; Convallaria majalis 5; Corallorhiza trida 5; Corydalis solida 6; 16: 1; Cruciata glabra 4; 23; Cuscuta epithymum 7; Cytisus austriacus 23; Dactylis glomerata 23:1-3; Dactylorhiza fuchsii 15; Dactylorhiza maculata 5; Daphne mezereum 3; 4; 5; 18; Dentaria enneaphyllos 3; Deschampsia caespitosa 5; 7; 26:1-2; Digitalis grandiora 7; 11; Doronicum austriacum 15; 17; 18: 1; 19; Dryopteris carthusiana 8: 1; 9; 10; 11; 12; Dryopteris disjuncta 22:1; Epilobium montanum 3; 5; 7; 11; 24: 1; Epilobium roseum 18; Epipactis helleborine 2; 4; 9; 10; Epipactis sp. 5; Equisetum sylvaticum 15; Equisetum arvense 11; 12; Euphorbia carniolica 15; 18; 19; Fagopyrum tataricum 7; Festuca altissima 12; 24; 26:1-2; Festuca drymeja 23:2; Festuca gigantea 2; 3; 11; Filipendula ulmaria 15: 1; Galeopsis speciosa 7; 13; Galeopsis tetrahit 7: 1; 12; 14; Galium aparine 7:1; Galium intermedium 18: 1; 20:1; Galium mollugo 7; Galium rotundifolium 8: 1; 10; Genista tinctoria 23:1; Geranium phaeum 16; 18; Geum urbanum 24; Glechoma hederacea 24:2; Gnaphalium sylvaticum 7; Gymnocarpium robertianum 17: 1; Heracleum sphondylium 4; 5; Hieracium pilosella 23; Hieracium sp. 18; 19; Homogyne alpina 9:1-3; 10:1-3; 25: 1; Hordelymus europaeus 3; 4; Huperzia selago 9; 10; 14; 19; 25; Hypericum hirsutum 6; 11; Hypericum maculatum 7; Hypericum perforatum 7; 10; 26:1-2; Impatiens parviora 2; 3; Isopyrum thalictroides 6; 14; 16: 1; 24: 1; Lamium maculatum 11; 23:1; Lapsana communis 7; 11; Lathyrus vernus 4:1; 5; 11; Leontodon hispidus subsp. hastilis 8; Leucoium vernum 26:2-4; Lilium martagon 4; 5; Linaria vulgaris 7; Lithospermum purpurocaeruleum 21; Lunaria rediviva 16: 1; Luzula luzulina 5; Luzula sp. 24; Lycopodium annotinum 9; 10; Lycopodium clavatum 22; Lysimachia nemorum 12; 14; Lysimachia nummularia 7; 15: 2; Melampyrum sylvaticum 5: 2; Melandrium rubrum 7; 11; Melica nutans 4:1; 7:1; 11: 1; Melica uniora 3; Moehringia muscosa 4; Moehringia trinervia 3; 11; 13; Moneses uniora 22; Myosotis palustris 14; 15; Myosotis sylvatica 3; 7; 11; Orobanche ava 5; Orthilia secunda 20: Parietaria ocinalis 3:4; Petasites hybridus 15; 16: 3; 18: 1; Petasites sp. 14; Petasitus albus 6; 10;11; Phegopteris connectilis 8:1; 10; Pimpinella major 4:1; Platanthera bifolia 4; Polygonatum multiorum 11; Polygonatum odoratum 4; Polypodium vulgare 5:2; Polystichum aculeatum 4; 5; 14; Potentilla argentea 23; Potentilla recta 23: 2; Primula elatior 5; 6; 14; Primula vulgaris 15; Prunella vulgaris 23; Pteridium aquilinum 12: 3; Pulmonaria obscura 15; Pulmonaria ocinalis 4; 5; 7; 23; Pulmonaria rubra 25:1-2; Pyrola minor 15; Ranunculus acris 4; Ranunculus carpaticus 21; 22; Ranunculus cassubicus 25; Ranunculus platanifolius 5; Ranunculus repens 5; 7:2; 14; 23; Rubus caesius 18; 19; Rubus saxatilis 3; 5; Rumex crispus 23; Rumex obtusifolius 6; Rumex sanguineus 3:2; Rumex sp. 26:1; Sambucus ebulus 23:2; Scrophularia nodosa 2; 7; 18; 23: 1; Senecio nemorensis 7: 1; 14; 16: 1; 17: 2; Senecio sylvaticus 3; Sesleria calcaria 4: 1; Soldanella carpatica 12:1-2; Soldanella montana subsp. hungarica 5; 25:1; Solidago virgaurea 4; 5; 7; 10; 14; Stachys sylvatica 6; 11: 1; 16; 18; Stellaria holostea 11; 20; Stellaria media 14; 21; 26:1-2; Stellaria sp. 24:1; Streptopus amplexifolius 9; 12; 15; 17; Symphytum cordatum 12; 14: 2; 15; 18: 2; 19; Symphytum popovii 19; Symphytum tuberosum 21; Tanacetum corymbosum 5; Taraxacum ocinale 3; 5; 7; Telekia speciosa 18; Thalictrum aquilegifolium 4; 5; Thymus serpyllum 23:1-2; Tussilago farfara 3; 4; 5:1; Vaccinium vitis-idaea 5; Valeriana tripteris 4: 2; 5:1; 11; Veratrum album 15; Veronica chamaedrys 2; 7: 2; 23:3; Veronica ocinalis 7; Vicia sativa 23:2; Vicia sylvatica 5; Viola alba 3; Viola arvensis 7; Viola mirabilis 18; 19; Viola riviniana 5; Bryophytes: Amblystegium serpens 14; Anomodon attenatus 11; 14; 16; Atrichum angustatum 5; Atrichum undulatum 2; 11; 12; 13; 19; Bazania trilobata 18; Blepharostoma trichophyllum 9; 10; Brachythecium geheebi 4; Brachythecium populeum 14; Brachythecium reexum 14; Brachythecium starkei 3; 10; Bryum capillare 11; Bryum accidum 14; Bryum sp. 1; 2:2; Calapogeia azurea 9; Calapogeia integristipula 8; Cephalozia bicuspidata 9; Ceratodon purpureus 3; 8; Chiloscyphus pallescens 13; Cirriphyllum piliferum 11; Climatium dendroides 6; Ctenidium molluscum 4:1; 5; 11; Dicranella heteromalla 1; 4; 8; 12; Dicranodontium denudatum 14; 17; 18; Dicranum tauricum 8; Ditrichum heteromallum 3; 4; Entodon orthocarpus 3; Eurhynchium angustirete 11:2; 17; Eurhynchium swartzii 6; Eurhynchium zetterstedtii 5:1; Fissidens cristatus 4; Hemallia trichomanoides 16; Heterocladium sp. 16; Homalothecium sericeum 5:1; Hylocomium splendens 5; 20; 23; 25:2; Hypnum pallescens 17; Hypnum vaucherii 4; 5; Isothecium alopecuroides 11; 14; Isothecium myurum 16; 19; Lepidozia reptans 1; 5:1; 10; 12; Leucodon sciuroides 3; 4; Lophocolea heterophylla 9; 10; 13; Metzgeria furcata 5; 14; Mnium hornum 12; Neckera complanata 5; Neckera crispa 3; 4; Oxystegus tenuirostris 14; Paraleucobryum longifolium 14; Pedinophyllum interruptum 4; Plagiochila asplenioides 4; 5: 2; 11; 20: 1; Plagiomnium ane 5: 2; 6: 1; 7: 2; 10; 11; Plagiomnium cuspidatum 14; 17; Plagiomnium denticulatum 11; Plagiomnium rostratum 10; 11; 12; Plagiomnium undulatum 5; 9; 11:2; 20: 1; Plagiothecium nemorale 11; 14; Plagiothecium roseanum 4; Plagiothecium sp. 1; Plagiothecium undulatum 9; Pleurozium schreberi 10; 19; 20: 1; 22; 25: 1; Pohlia nutans 1; 4; 8; 12; 14; Polytrichum commune 10; 14; 22: 1; 25:2; 26:2; Polytrichum juniperinum 7; Ptilidium ciliare 9; Pylaisia polyantha 4; Rhacomitrium heterostichum 5; Rhytidiadelphus loreus 9; 10; Rhytidiadelphus triquetrus 22:1; Sanionia uncinata 9; Solenostoma tristis 3; Sphagnum cymbrifolium 24:2; Syntrichia subulata 3; Thuidium tamariscifolium 11; 16; 17; Tortella tortuosa 3; 4; 5: 1; 11; Tritomaria quinquedentata 10; a Scale of coverage [13]: 575100%; 45075%; 32550%; 2 <25%; 1abundant but coverage low; few numerous. Scale of constancy [13]: Vpresent in 80100% of records; IV6080%; III4060%; II2040%; Isporadic species <20%.

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin

29

The dierences in the land use pattern between the two comparable environments are signicant. In the Tatras region, the areas just below timberline are predominantly covered with coniferous forest while the lower elevations are typically occupied by agriculture. In the Retezat area, where even at the highest elevations the relief is not as dramatic as in the Tatras, alpine meadows and deciduous forests are the dominating land use classes. In the Tatras (Fig. 8), the areas located above about 2200 m a.s.l. frequently represent the dramatic alpine type of relief. In this subnival zone many herbaceous plant species, mosses and lichens can be found. However, from the land use classication point of view, these areas are described as the bare rocks land use category. The alpine meadows are located below and around the zones of the bare rock class. These meadows of numerous grass and herb species at their upper transition zone are typically intermixed with groups of bare rocks and at their lower transition zone with alpine dwarf pine (Pinus mugo) stands. With the decreasing elevation, clusters of dense Pinus mugo become more common and locally prevailing. Since these two categories are typically situated next to each other, they were blended together and represented on the maps as the alpine meadows class. Below the timberline there are large stands of coniferous forest, usually dominated by Norway spruce (Picea abies). As the elevation decreases, Picea abies is more and more frequently interchanged with white r (Abies alba). While describing mountain vegetation, normally, the zones of upper and lower coniferous forests are considered. However, the applied land use classication did not dierentiate between the two. Instead, the categories of coniferous, mixed, and broad-leaved forests were used. Consequently, in the Tatra Mountains, the coniferous category on the maps represents both upper and lower coniferous forest zones. Below the forested areas, in the vicinity of the Tatras, the arable land intermixed with pasture is the characteristic type of land use. Since the density of population in this area is the highest within the entire Carpathian Mountain range, the build-up areas make a signicant part of the land use. The Retezat Biological Reserve area (Fig. 9) is another good representation of the vegetation zonality resulting from plant adaptation to an altitude. A signicant dierence between the Tatras and Retezat is that in Retezat, the upper limits of all the relevant land use categories are elevated by 200300 m relative to the Tatras. Another noteworthy dierence is that in Retezat, the coniferous forest in some locations is not present on the southern slopes of the mountains. On these slopes, the mixed, and occasionally even the deciduous forests reach up to the category of alpine meadows. Conrming the general trend observed in the Car-

pathians, the deciduous trees cover considerably larger percentage of area in the vicinity of the Retezat Mountains than in the neighborhood of the Tatra Mountains. Due to the less dramatic relief of the Retezat Mountains as well as the mentioned elevation shift of the vegetation zonality, the majority of its most elevated areas is occupied by alpine meadows and dwarf pines, instead of bare rocks, which outcrops exclusively along the ranges of summits. Because the population density at this part of Romania is considerably lower than in the environs of the Tatras, the agricultural and inhabited areas are concentrated in the valleys apart from the Retezat National Park leaving the large forested areas of the mountains relatively undisturbed. The above described elevation shift of oral zonality is caused by climatic dierences between the two regions, resulting from latitudinal distance of about 450 km between the Retezat and the Tatras. Therefore, the temperatures in the Retezat area are signicantly higher all year round and the precipitation is noticeably lesser. The anthropogenic pressure created by the high population density in the vicinity of the Tatras Mountains explains the high percentage of their area occupied by the agricultural usage. Consequently, the lower number of inhabitants around the Retezat area explains why the agricultural land use class is by far less signicant there. 4.2. Vegetation of the studied forest stands 4.2.1. Species composition Common beech (Fagus sylvatica), white r (Abies alba) and Norway spruce(Picea abies) are main tree species in the studied sites in the Carpathians. Sycamore maple (Acer pseudoplatanus) and mountain ash (Sorbus aucuparia) are less frequent. European larch (Larix decidua), maple (Acer platanoides), ash (Fraxinus excelsior), elm (Ulmus glabra), birch (Betula pendula), aspen (Populus tremula), and rowan (Sorbus aria) occur as a sporadic species. Forest stands are represented by pure beech and spruce stands and by mixed (beechspruce, beechr, beechrspruce) stands. While the shrub layer is not abundant, the herb layer is rich in species, especially in the sites on a calcium carbonate-rich substratum (e.g. records 4, 5, and 11; Table 2). The number of vascular plant species registered in all records was 269, ranging from 20 to 91 species in individual records (Table 2). More than half of the records comprised 2040 species each. Among the 269 vascular plant species, as much as 216 were sporadic and only 28 frequent components of the phytosociological records in the surveyed plots. Most of the recorded species represent plants of mesotrophic habitats, typical of broadleaved forests (e.g. Galeobdolon luteum, Dentaria bulbifera, D. glandulosa, Euphorbia amygdaloides or Mercurialis perennis). Species of poorer habitats, typical of coniferous forests (e.g. Vaccinium myrtillus, Maianthe-

30

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin

mum bifolium or Polygonatum verticillatum), occurred less frequently. Dierent geographical elements are represented among the plants growing on the surveyed plots. Dentaria glandulosa and Symphytum cordatum represent the Western-Carpathian subendemics; Soldanella carpatica, Galium rotundifolium and Luzula luzulina represent a Western-Carpathian element (records 5, 8, and 10). Veratrum album, Aposoeris foetida, Telekia speciosa and Festuca drymeja are limited to the Eastern Carpathians (records 15, 18, 23); and Campanula patula subsp. abietina is characteristic of the Eastern and Southern Carpathians. Many mountain species occur in the surveyed forest plots. Subalpine species are represented by Doronicum austriacum, Cicerbita alpina, Soldanella montana and Veratrum album, which have been noted almost exclusively in the records from the eastern part of the Carpathian arch (records 15, 17, and 18). Numerous were species of mountain forests (among others Dentaria glandulosa, Luzula sylvatica, Polygonatum verticillatum, Prenanthes purpurea, and Symphytum cordatum) and all-mountain species (among others Calamagrostis villosa, Gentiana asclepiadea, Homogyne alpina, Huperzia selago, and Valeriana tripteris). The invasion of Asiatic Impatiens parviora (records 2 and 3), a foreign species to our ora, has been observed in the investigated forests, and particularly in the Western Carpathians. Also worthy of mentioning is a frequent occurrence of Urtica dioica, a strongly nitrophilous species in the phytosociological records from the Western Carpathians. Presence of this species indicates a high level of soil-available nitrogen. Numerous moss species (88) were found in the examined material. Number of moss species ranged from 1 to 24 species in individual records. Just as for the vascular plants, sporadic species are the most numerous group (74 species). 4.2.2. Ecological and dendrological characteristics of forest stands Among 26 sites located in the Carpathians, there were eight (Picea abies) stands, 10 beech (Fagus sylvatica) stands, two r (Abies alba) stands and six mixed beech r, sprucer and beechspruce (Table 3). Most of forest stands developed on either a calcium-carbonatedecient Carpathian Flysh substratum (e.g. Brenna, Babia Gora, Yablunitsa, Krivopilja) or on a crystalline granite substratum (Tatry). The r stands occurred on a limestone substratum (e.g. Pieniny, Mala Fatra), while the beechr (e.g. Magura) and beech stands (e.g. Bieszczady, Kuzij, Uzhoksy Pass) on a calcium carbonate rich Flysch substratum. Participation of beech on forest stands of the Carpathian arch increases from the west to the east. Characteristics of the investigated stands are presented in Table 3. Age of most stands was 80100 years,

younger (4575 year old) forest stand occurred in Eastern Carpathians (e.g. Obcina Mare, Krivopilja). Older forest stands (160170 years old) were found also in the Eastern Carpathians (Kuzij, Synevir). In most cases a height of trees reached 2530 m, and a diameter (dbh) 3040 cm. In two stands in Eastern Carpathian (Romania) trees were lower (1617 m) and thinner (1819 cm). As a consequence of a various age of forest stands, tree volume and number of trees varied from low (144 m3/ ha) to high (229245 m3/ha). Stand quality varied between I and III class, with most stands included into class II. 4.2.3. Forest health status in the Carpathian Mountains A degree of tree crown damage varies greatly between the investigated sites. Trees from the western part of the Western Carpathians show the greatest degree of defoliation. In the sites located in the Czech Republic and Slovakia, a degree of damage to beech crowns varies from slight (510%) (Mala Fatra, Kozie Chrbty) to high (3950%) (Bily Kriz, Male Karpaty). For spruce (Picea abies), the defoliation reaches 37% in certain sites, and for r (Abies alba) even 50%. A percentage of damaged trees decreases in the Carpathians from the west to the east.

5. Discussion Geographical, ecological, oristical, and dendrological characteristics of the Carpathian forests, as described in the present paper on the basis of data from 26 permanent plots distributed all over the Carpathian arch, are consistent with data given for these mountains by other authors (among others, Korpel, 1987; Jaworski and Karczmarski, 1991; Jaworski et al., 1991; Szwagrzyk et al., 1995; Fabijanowski and Jaworski, 1995; Saniga and Vesely, 1998). In general, the Carpathian forests show a small or average (moderate) degree of damage. In the Carpathians the proportion of damaged trees, representing defoliation classes 24, is now between 12.2% and 12.8%. Somewhat bigger damage has been found in coniferous trees (Abies alba, Picea abies) as compared with deciduous ones (Fagus sylvatica). The most damaged forests occur in the Western Carpathians, while the least in the Southern Carpathians (Badea et al., 2002a,b). Data concerning the degree and spatial distribution of damage to stands in the 26 Carpathian permanent study plots, presented in this work, are consistent with the results obtained by Badea et al. (2002a,b). Many factors can be responsible for damage to the Carpathian forests. The most important factor is undoubtedly human activity, both direct (type of forest management) and indirect (industrial development accompanied by gaseous and dust emissions).

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin Table 3 Characteristics of forest stands (permanent plots) in the Carpathian Mountainsa Locality Western Brenna Babia Gora Tatry Pieniny Magura Eastern Bieszczady Uzhoksky Pass Kuzij Synevir Yablunitsa Krivopilja Vishnitsa Obcina Mare Rarau Magura Odobesti Southern Fundata Retezat Stana de Vale
a

31

Main tree species Spruce Spruce Spruce Fir Beech/r

Mean height (m) 37 24 27 31 23

Mean diameter (cm) 47 31 34 36 29

Tree volume of site (m3) 245 127 134 195 110

Tree volume 1 ha (m3) 982 509 536 780 440

Crop density 1.1 0.8 0.8 0.7 0.6

Stand quality I II II I II

Mean age of trees 120 80 100 100 80

No of trees site 69 126 96 109 142

No of trees 1 ha 276 504 384 436 568

Beech Beech Beech Spruce/r Spruce Spruce Beech/r Spruce Beech/spruce Beech

24 25 31 34 31 25 26 17 27 16

35 43 43 54 43 28 35 19 32 18

164 155 133 229 126 143 111 59 130 36

656 622 532 917 504 571 445 234 520 144

1 0.9 0.9 1 0.7 0.9 0.8 0.7 0.8 0.7

II Middle Middle Middle Middle Middle Middle III II III

110 105 160 170 90 65 75 45 80 50

80 . . . . . . 292 127 588

320 263 324 364 272 948 480 1669 634 2352

Beech Spruce Beech

21 25 27

38 35 50

52 113 106

297 451 265

0.7 0.8 0.4

III II II

127 80 120

196 127 52

582 634 519

Six stands (ve in Western and one in Eastern Carpathians) located in Slovakia were not included to the table due to incomplete information.

The Carpathian forests for some decades have been under the inuence of air pollutants transported by westerly winds from urban-industrial centers in Austria, Germany, Czech Republic, Poland, Slovakia, and Hungary (Bytnerowicz et al., 2002b). During the past 1020 years the emissions of sulfur dioxide (SO2) and nitrogen dioxide (NO2) have been much reduced in entire Europe. Concentrations of these gases in the Carpathians are presently below the toxic level for vegetation (Bytnerowicz et al., 2002a,b). Therefore the Carpathian forests presently are not directly endangered by these pollutants. However, the recent decrease in atmospheric SO2 and NO2 concentrations is accompanied by an increase of ozone (O3) concentrations. In the Carpathians, concentrations of tropospheric O3 frequently reach a toxic level for forest trees (Bytnerowicz et al., 2002a,b). The O3 concentrations change considerably along the Carpathian arch. The highest O3 concentrations are generally found in the Western Carpathians, while the lowest in the Southern Carpathians (Bytnerowicz et al., 2002a,b). A decreasing proportion of the damaged stands along the Carpathian arch from the west to the east and south corresponds therefore with the level of O3 noted in these areas. The concentrations of O3 dier also between seasons. The high concentrations of O3 occur usually in May (Bytnerowicz et al., 2002a,b). In that season they can pose a particular threat to young leaves and needles of

trees, which are sensitive to the action of ozone. The herb layer plants are more sensitive to O3 than trees, so damage caused by O3 to these plants may be higher (Bull, 1996; Bytnerowicz et al., 2002b). This supposition has been conrmed by the results of studies by Manning et al. (2002) carried out in many locations in the Carpathians. They have found conspicuous, typical damage by O3 in more than a dozen herb layer species and in some species of open areas (meadows, road sides). It may be expected that ambient concentrations of O3 may further increase in the Carpathians, particularly in the warmer and stronger insolated Southern Carpathians. Therefore, managers and policy makers should be prepared for increased threats to the Carpathian forests caused by this type of air pollution.

6. Final remarks Clearly, the characteristics of the Carpathian forests, based on data from 26 vegetation sites scattered over the 1400 km long-mountains arch are sketchy. Data collected during these studies conrm, however, the richness and oristic diversity of these forests. To be able to better describe the diversity of the Carpathian forest stands, their relation to diverse geological and soil substrata, climatic conditions, and to assess their health, the number of sites should be increased by 45 fold.

32

ska et al. / Environmental Pollution 130 (2004) 1732 K. Grodzin Carpathian Mountains in central Europe. Environmental Pollution 116, 325. Fabijanowski, J., Jaworski, A., 1995. Gospodarstwo les ne (Silvi culture). In: Warszynska, J. (Ed.), Karpaty PolskiePrzyroda, Czowiek i jego dziaalnos c. Uniwersytet Jagiellonski, Krakow, pp. 253263. Godzik, S., Sienkiewicz, J., 1990. Air pollution and forest health in Central Europe: Poland, Czechoslovakia and German Democratic Republic. In: Grodzinski, W., Cowling, E.B., Breymeyer, A.I. (Eds.), Ecological RisksPerspectives from Poland and the United States. National Academy Press, Washington DC, pp. 155170. Grodzinska, K., Szarek-ukaszewska, G., 1997. Polish mountain forests: past, present and future. Environmental Pollution 98, 369374. Grodzinska, K., Mirek, Z., Szarek, G., 1995. Environmental damage and restoration attempts in Polandan overview. In: Urbanska, K., Grodzinska, K. (Eds.), Restoration Ecology in Europe. Geobotanical Institute SFIT, Zurich, pp. 113130. Guderian, R., Klumpp, G., Klumpp, A., 1989. Eects of SO2, O3 and NOx singly and in combination on forest species. In: Ozturk, M.A. (Ed.), Plants and Pollutants in Developed and Developing Countries. Ege Univ., Izmir, Turkey, pp. 231268. Jaworski, A., Karczmarski, J., 1991. Struktura i dynamika drzewos tanow o charakterze pierwotnym w Pieninskim Parku Narodowym (na przykadzie czterech powierzchni dos wiadczalnych). Zesz. nauk. AR, Lenictwo 20, 4583. c Jaworski, A., Skrzyszewski, J., Swic tkowski, W., Karczmarski, J., a 1991. Budowa i struktura dolnoreglowych drzewostanow o charakterze pierwotnym na wybranych powierzchniach w Bieszczadach Zachodnich. Zesz. nauk. AR, Les nictwo 20, 1743. Korpel, S., 1987. Dynamica s truktury a vyvoj bukovych prirodnych lesov na Slovensku. Acta Facultatis Forestalis 29, 5983. Manning, W.J., Godzik, B., Musselman, R., 2002. Potential bioindicator plant species for ambient ozone in forested mountains areas of central Europe. Environmental Pollution 119, 283290. Mirek, Z., Pickos -Mirkowa, H., 1992. Plant cover of the Western e c Carpathians (S Poland). In: Zarzycki, K., Landolt, E., Wojcicki, J.J. (Eds.), Contributions to the Knowledge of Flora and Vegetation of Poland. Vero. Geobot. Inst. ETH 107, pp. 116150. Mirek, Z., Pickos -Mirkowa, H., Zajc c, A., Zajcc, M., 1995. Vascular e c a c a c plants of PolandA Checklist. PAN. W. Szafer Institute of Botany, Krakow. Ochyra, R., Szmajda, P., 1978. An annotated list of Polish mosses. Fragm. Flor. Geobot 24, 93145. Oszlanyi, J., 1997. Forest health and environmental pollution in Slovakia. Environmental Pollution 98, 389392. Saniga, M., Vesely, L., 1998. Dynamika zmeny s truktury, produkcie a regeneracie bukoveho prirodneho lesa v s tatnej prirodnej rezervacii Ras tun. Lesnictvi-Forestry 44, 116125. Szafer, W., Zarzycki, K. (Eds.), 1972. Szata rolinna Polski. T. 11614. PWN, Warszawa. Szaro, R.C., Johnston, D.W. (Eds.), 1996. Biodiversity in Managed Landscapes1780. Oxford University Press, New York. Szwagrzyk, J., Szewczyk, J., Bodziarczyk, J., 1995. Structure of forest . stand in the Zarnowka reserve of the Babia Gora National Park. Folia Forestalia Polonica ser. A, Forestry 37, 111123. Volos cuk, I., 1999. The National and Biosphere Reserves in Car pathiansThe Last Nature Paradises. ACANAP, Tatranska Lomnica, Slovak Republic. Warszynska, J. (Ed.), 1995. Karpaty polskie. Przyroda, czowiek i jego dziaalnos c. Uniwersytet Jagiellonski, Krakow. Wilson, E.O., 1988. The current status of biological diversity. In: Wilson, E.O., Peter, F.M. (Eds.), Biodiversity. National Academy Press, Washington, DC. Wilson, E.O., 1989. Threats to biodiversity. Sci. Am 261, 108117.

Future, more detailed studies carried out in various parts of the Carpathians should undoubtedly reveal many new data which would allow to develop a more synthetic understanding of the Carpathian vegetation status.

Acknowledgements The study was supported by the USDA Foreign Agricultural Services, Oce of International Cooperation projects: No. PL-FS-111 Evaluation of ozone air pollution and its phytotoxic potential in the Carpathian Forestsa cooperative study between the Czech Republic, Poland, Romania, Slovakia, Ukraine and the United States and Eects of forest health on biodiversity with emphasis on air pollution in the Carpathian Mountains. The authors thank the anonymous reviewers for their valuable comments and critical remarks.

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