Publication foreign language title: Ecology
Author: Kollár, Jozef; Balkovic, Juraj; Mazúrová, Anna; Simonovic, Vojtech
Date published: July 1, 2011
Journal code: EKBR
As indicated by the name itself, this contribution is linked to a series of articles by Somsák, Kubíek ( 1 994, 1 995, 2000) which focused on phytocoenological and production characteristics in the original forest ecosystems and also the changes induced by Scotch-pine monocultures. These were massively applied throughout the entire region of the Borská nízina lowland during recent centuries. This article is closely related to the work of Somsák, Kubicek (1994) which examined Pino-Qurcetum association oak-pine forests. These occupy xeric sand dunes traditionally considered to be a major, or even exclusive, original forest type here (e.g. Krippel, 1965; Ruzicka, 1960). These authors also reported on derived Pleurozio schreberi-Pinetum association of Scotch-pine monocultures. However, in our opinion, the question of potential vegetation in sand dunes, and also in this entire region, is more complex than elucidated so far, and it still has not been satisfactorily answered. Earlier authors, including Ruzicka (1960), noted that some acidophilous oak forests may exist here, but certainly not to a wide extent. On the other hand, adjacent region of sand dunes in the Czech Republic list the occurrence of only acidophilous oak forest (Festuco ovinae-Quercetum roboris Smardal961 association of Geniste germanicae-Quercion NeuhäusletNeuhäuslov á-N ? ? o t ? á 1967 alliance). Other authors in Western Europe including PhUippi (1970) do not consider Scotch-pine to be a climax tree in their respective sandy regions. Following this idea, we recorded stands consisting entirely of oaks in this Borská nízina lowland region which, together with secondary pine forests, form the main subject of this study. We have not named this syntaxon here because its classification has so far remained unclear. However, syntaxonomical and phytocoenological issues are the subject of our next article already under preparation.
The study area is contained in the Borská nízina lowland in western Slovakia. Herein, we focused on its subunit portion - named Bor - which has served as a military training area since its declaration in 1928. The area is composed of eolie siliceous sands dominated by Podzols and Arenosols, and these are very poor in nutrients. Annual average temperature is 9-9.6 °C, but during the vegetation period it is 14-15 °C, while annual precipitation is in the range of 550-650 mm, with prevailing évapotranspiration.
Phytocoenological relevés in the field and classification of the forest communities were performed by the ZurichMontpellier school, plant names follow Marhold, Hindák (1998). Soil names are according to the IUSS Working Group WRB (2006). Estimation of the herb layer above ground biomass was carried out on the selected sample plots applying the method of indirect sampling (Kubiiek, Brechtl, 1970) modified for non-recurrent sampling (Kubicek, Jurko, 1975; Kubiiek, Simonovic, 1975; Kubiiek, Somsák, 1982; Kubíéek, 1983).
Results and discussion
Characteristics of oak forests
Oak forests, which we consider to be original here, are best-preserved in military training area in the sites adjacent to the firing ranges. They occupy stabilized sand dunes where is no impact of underground water. Most of mature soils belong to Brunie Arenosols (Dystric), which are typical for relatively deep cambic-like B horizon (20-60 cm thick), moderate organic carbon contents in A horizon (Cor -2%) and acidic soil reaction (pH-H20 ~ 4.6 and 5.0 in A and B horizons respectively). Vertical stratification of Brunie Arenosols (Fig. la) indicates that these soils have hot been significantly touched by forest management measures for a long time.
The tree layer is dominated by Quercus petraea agg. while other species form an admixture. The shrub layer formed by thermophilous species is usually poorly developed, while the herb layer is rich in species. The Festuca ovina agg. is a typical dominant species, and this is especially accompanied by light-demanding forest and ecotone species (Betonica officinalis, Trifolium alpestre, Teucrium chamaedrys, Vincetoxicum hirundinaria), species of shaUowless productive soils (Hylotelephium maximum, Poa compressa, Silčne nutans, Steris visearía), grassland species (Achillea millefolium, Agrostis capillari*, Hypericum perforatum, Poa angustifolia), plus others. Despite the sandy substrate, obligate psamophilous species are quite rare here, with the presence of Peucedanum oreoselinum most constant. The cover of the moss layer is not very high, with Pleurozium schreberi, Plagiomnium affine and Polytrichum formosum representing the most common and abundant species. A more detailed floristic composition is given in Table 1.
Characteristics of secondary Scotch-pine forests
These stands enjoy the same the conditions as the oak forests; the only difference is that the application of the Pinus sylvestris Scotch pine adversely affects the ecological conditions. These Scotch-pine plantations are on different soils, with young pine stands being planted on mechanically adjusted sites of Protic to Haplic Arenosols (Dystric), whereas older Scotchpine stands grow on Albic Podzols with acidic leaching. AU Albic Podzols are acidic in both the A and B horizons of pH-H20 ~ 4.1 and 4.6 respectively (Fig. Ib).
The tree layer comprises mono-dominant Pinus sylvestris, with other species only accessory. The shrub layer is almost negligible and the herb layer is generaUy poor in species with low cover. Dependent on type, its physiognomy is determined by species including Festuca ovina agg., Calluna vulgaris or Calamagrostis epigejos. The moss layer is distinctly weU-developed and it dominates the herb layer. The typically dominant species here is Pleurozium schreberi and this is accompanied by Dicranum polysetum and Pseudoscleropodium purum. Additional species, including Hypnum cupressiforme, Leucobryum glaucum and Polytrichum formosum, form an admixture.
Comparison of the floristic composition of the natural oak forest with pine monocultures
Our earlier data was used to compare floristic composition (Rollar, 2004; Mazúrová, 2006). Pine monocultures were sampled on sites located near oak forest areas with similar natural conditions, and the results are registered in Table 1. Changes in floristic composition were established as follows: pine monocultures triggered secondary regressive succession, with many herb layer species retreating or even disappearing in this process. Firstly, the following species tied to light-demanding forests, ecotone and grasslands became impacted; Achillea millefolium agg., Anthericum ramosum, Betonica officinalis, Carex caryophyllea, Polygonatum odoratum, Silčne nutans, Veronica chamaedrys, Vincetoxicum hirundinaria and others. Secondly, there was an obvious increase in the cover of some bryophytes, which colonize accumulated pine needle fitter. These bryophytes included Pleurozium schreberi, Dicranumpolysetum and Pseudoscleropodium purum. These results are compatible with those of Somsák, Kubicek (1994), with the course and intensity of these processes being dependent on time and forestry management.
Comparison of herb and moss layer production between natural oak forest and pine monocultures
The data on biomass production is shown in Table 2, and here herb and moss layer production reflect changes in floristic composition. It was established that total herb layer production of pine monocultures at 472.8 kg.ha"' is 3.8 times lower than that of the oak forest with 1 ,8 1 4.5 kg.ha'. (The latter above ground mass has a multiplication factor of 3.5 and the below ground mass has 4.25). However, the moss layer production increased dramatically by 6.75 times, moving from 283.5 kg.ha"' to 1 ,910.8 kg.ha"1 . This is so productive that the sum of herb and moss layer production is slightly higher than pine monocultures at 2,383.6 kg.ha"' when compared to oak forest production of 2,098 kg.ha"1. Therefore, mosses provide most biomass production in pine monocultures, while herb layers achieve this in oak forests. Both these types have the same production dominance by Festuca ovina agg. In pine monocultures, Calluna vulgaris and Calamagrostis epigejos also contribute significantly while Peucedanum oreoselinum plays a co-dominant role in oak forests. Other herb layer species are much less productive Overall, the pine forest Pleurozium schreberi moss species is the most dominant productive species with 1,792.3 kg.ha1.
Sampled sites are characterized by the following phytocoenological relevés:
1. Oak forest
22. 7. 2004, Záhorie military training area, geographical coordinates (WGS 84): 48°3G8.57" N, 17°13'36.55" E, relevé area: 400 m2, sand dune top and its upper slopes, covers: E3 70%, E2 2%, E1 95, E0 10%, height of trees: 10 m, number of specie: 53, soil: Brunie Arenosol (Dystric)
E3: Quercus petraea agg. 4, Quercus robur r
E^sub 1^: Festuca ovina agg. 2, Peucedanum oreoselinum 1, Betonica officinalis +, Brachypodium sylvaticum +, Calamagrostis epigejos +, Campanula persici/olia +, Campanula rotundifolia agg. +, Carex caryophyllea +, Clinopodium vulgare +, Lembotropis nigricans +, Dianthus carthusianorum agg. +, Tithymalus cyparissias +, Fallopia convolvulus +, Agrostis capillaris +, Fragaria vesca +, Galium mollugo agg. +, Galium verum +, Pilosella officinarum +, Hypericum perforatum +, Koeleria pyramidata +, Luzula campestris +, Steris visearía +, Peucedanum cervaria +, Anthericum ramosum +, Poa angustifolia +, Polygonatum odoratum +, Anthoxanthum odoratum +, Asperula tinctoria +, Hylotelephium maximum +, Silčne nutans +, Teucrium chamaedrys +, Thymus serpyllum +, Trifolium alpestre +, Trifolium montanum +, Veronica chamaedrys +, Veronica officinalis +, Vicia hirsuta +, Vincetoxicum hirundinaria +, Galeopsis tetrahit r, Solidago virgaurea r, Achillea millefolium agg. r, Allium senescens subsp. montanum r, Pseudolysimachion spicatum r, Geranium sanguineum r, Moehringia trinervia t, Scabiosa columbaria r, Scorzonera purpurea r, Frángula alnus r, Viola rupestris r
E^sub 0^: Pleurozium schreberi 1, Plagiomnium undulatum +
2. Secondary pine forest
14. 06. 2001 and July 2005, Záhorie military training area, geographical coordinates (WGS 84): 48°33'6.09" N, 17°16'0.66" E, slope: 3°, aspect: SE, covers: E3 65%, E1 40%, E0 85%, height of trees 18 m, number of species: 35, soil: Albic Podzol
E3: Pinus sylvestris 4, Quercus petraea agg. +
E : Calluna vulgaris 3, Festuca ovina agg. 1, Agrostis capillaris +, Campanula rotundifolia agg. +, Carex ericetorum +, Lembotropis nigricans +, Danthonia decumbens +, Tithymalus cyparissias +, Anthericum ramosum +, Fragaria vesca +, Hieracium murorum +, Pilosella officinarum +, Hypericum perforatum +, Luzula campestris +, Peucedanum oreoselinum +, Anthoxanthum odoratum +, Pinus sylvestris +, Calamagrostis epigejos +, Quercus petraea agg. +, Acetosella vulgaris +, Silčne nutans +, Thymus serpyllum +, Viola rupestris +, Steris visearía r, Pulsatilla pratensis r, Solidago virgaurea r, Cerastium arvense r, Asperula tinetoria r
E0: Pleurozium schreberi 3, Dicranum polysetum 2, Pohlia nutans +, Polytrichum piliferum +, Cladonia macilenta subsp. macilenta +, Cladonia furcata subsp. furcata +, Hypnum cupressiforme +
In Table 2, we have attached part of Somsák and Kubicek's (1994) results for comparison with our production data. In principle, it has been established that these two data sets correspond quite well. This is especially true and relevant for high moss layer production, and also for the low ratio between herb and moss layer production in secondary pine forests.
Translated by the authors
English corrected by R. Marshall
This contribution was financially supported by VEGA grant No. 2/0059/11.
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JOZEF KOLLÁR1, JURAJ BALKOVlC2, ANNA MAZÚROVÁ3, VOJTECH SlMONOVIC1
1 Institute of Landscape Ecology, Slovak Academy of Sciences, Stefánikova 3, 814 99 Bratislava, P.O. Box 254,
Slovak Republic; e-mail: email@example.com
2 Department of Soil Sciences, Faculty of Natural Sciences, Comenius University, Mlynská dolina, Bratislava,
3 Kapitána Rasu 35, 841 Ol Bratislava, Slovak Republic