The shrub and Black Locust communities of chosen parts of the Hron downs, the Slovak Republic

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Blažena Benčaťová1, Ján Koprda2, Tibor Benčať3 1

Department of Phytology, Faculty of Forestry, Technical University in Zvolen, T. G. Masaryka 24, SK-960 53 Zvolen, Slovak Republic, e-mail: [emailprotected] 2 Športová 86, 95152 Slepčany, Slovak Republic, e-mail: [emailprotected] 3 Department of Landscape Planning and Design, Faculty of Ecology and Environmental Sciences, Technical University in Zvolen, T. G. Masaryka 24, SK-960 53 Zvolen, Slovak Republic, e-mail: [emailprotected]

Abstract Benčaťová, B., Koprda, J., Benčať, T. 2013. The shrub and Black Locust communities of chosen parts of the Hron downs, the Slovak Republic. Folia oecol., 40: 157–162. The scrubland and Black Locust phytocoenoses belong to the substitute communities which constitute an important component in present cultural landscape. They arose and evolved according to certain rules. Their presence and arrangement is mainly dependent on the type of agricultural land. In the study area, around the Arborétum Mlyňany, these systems have become a permanent component of the vegetation. In our article there are given phytocenological and ecological characteristics of scrubland and Black Locust forest stands in the municipalities of Vieska nad Žitavou, Tesárske Mlyňany andSlepčany. Within the scrubland we determined association Ligustro-Prunetum R.Tx. 1952 with the ecological variations of Prunus spinosa and Vitis vinifera and within the Black Locust we determined association Chelidonio-Robinietum Jurko 1963, with ecological variant withHedera helix and with facias with Rubus caesius and Vinca minor and association Bromo sterilisRobinietum Jurko 1963 prov. For the allocation of communities were used numerical methods (JUICE, TWINSPAN), ecological analysis was conducted in the program JUICE. On the basis of the performed ecological analysis we can conclude that the communities are very similar in their ecological claims. Keywords the Arborétum Mlyňany, Black Locust, cultural landscape, phytocoenoses

Introduction The scrubland and Black Locust phytocoenoses belong to the substitute communities which constitute an important component in present cultural landscape. They arose and evolved according to certain rules. Their presence and arrangement is mainly depended on the type of agricultural land. In the study area, around the Arborétum Mlyňany, these systems have become a permanent component of the vegetation. In the past not much attention was paid to the study of both types of communities and even nowadays the syntaxonomy of these communities is not definitely worked out. Besides Jurko (1964) the study of shrubs in Slovakia was performed mainly by Kontriš (1966) who

described the shrub field communities of north-western part of Liptovská kotlina basin. In the recent years the theoretic questions of syntaxonomic position of the shrub communities were studied by Valachovič (2002, 2007). Koprda (2008) described in his diploma thesis the shrub communities of the part of Žitavská pahorkatina hills and the newest paper about hazel communities of the Veľká Fatra Mts was published by Kliment et Jarolímek (2011). Problem of classification of the Black Locust communities in Slovakia was examined by Ščepka (1982, 1985), Jurko (1963), Jurko et Kontriš (1982), in the recent period by Šimonovič et al. (2002), Benčaťová et Benčať (2005, 2008), Benčaťová et al.157, (2008), Koprda (2008). 157

The study area is located in the cadastres of villages Vieska nad Žitavou and Slepčany and because of the fact that the paper follows the papers by Benčaťová et Benčať (2005), Benčaťová et al. (2008), characteristic of the territory is described in the mentioned references.

Material and methods Phytocenological research was performed during the growing seasons of the years 2006–2007. Within the field research and vegetation synthesis was followed the Zürich-Montpellier School method with 7 degree scale abundance and dominance (Braun-Blanquet, 1964). Nomenclature of plants is given according to Marhold et Hindák (1998), nomenclature of syntaxa according to the actual vegetation units of Slovakia by Jarolímek et Šibík (2008). Phytocenological records were saved in the database program TURBO(VEG) (Hennekens, 2005). Output numeric matrix of the program with the phytocenological records were used as an input data for the next management in the program JUICE (Tichý, 2002) for the following purposes: differentiation of the syntaxonomic units with the program TWINSPAN (Hill, 1979), indirect unimodal gradient analysis DCA and ecological analysis of the communities.

Results and discussion The following syntaxonomic units were selected with the numeric classification methods in the study area: o The scrubland communities Studied scrubland communities syntaxonomically belong to Rhamno-Prunetea Rivas-Goday and Borja-Carbonell 1961 family and to the two alliances. Berberidion vulgaris Br.-Bl. 1950 alliance includes Ligustro-Prunetum R. Tx. 1952 association and Arctio-Sambucion nigrae Doing 1962 alli-

ance includes Anthrisco-Lycietum halimifolii Jurko 1964 association (Fig. 1). – Ligustro-Prunetum R. Tx. 1952 association The scrubland communities of the association are spread in the whole study area. The most significantly they are represented on the slight slopes, mainly with the western or south-western exposition. Common sign of most of the records of the community is their occurrence on the sites that were in the past intensively used mostly like pastures, alternatively like mown meadows or they are developed like narrow stripes of shrubs among vineyard areas. The sites are affected by intensive human acting and so they were supplied with sufficient supply of nutrients, mainly with nitrogen in the initial phase of their creation. Association is represented by the poorest scrubland community according to species diversity in the area. The tree layer is negligible, created mainly with the stronger individuals of the shrubs Prunus spinosa, Crataegus monogyna, or the fruit tree Cerasus avium. In the case of older shrubs in the studied area phytocenoses there are Quercus cerris and Robinia pseudoacacia that infiltrate to the shrubs from the surrounding black locust stands. The cover of the shrub layer is large, averagely it reaches the value 90%. In the layer there occur mainly three types of shrubs which create typical, nearly impenetrable structure – dominant Prunus spinosa species, with associated Crataegus monogyna and Rosa canina species. Herb layer is in the most of the records very poor, with the average cover 15%. Herbs are spread especially in the marginal parts of the communities where they are particularly represented with nitrophilous species with high demand for nutrients; Galium aparine, Geum urbanum, Glechoma hederacea, Anthriscus cerefolium, Urtica dioica and others. From the grass species there is constantly

Modified TWINSPAN Dendogram

Fig. 1. Dendrogram created from the three groups of records in the scrubland communities representing syntaxonomic units (1st association Anthrisco-Lycietum halimifolii, 2nd variant withPrunus spinosa, 3rd variant withVitis vinifera).

158

occurred Poa nemoralis species. We also recorded quite high occurrence of the juvenile stages of Ligustrum vulgare species, Prunus spinosa and Rosa canina woody species. On the basis of certain differences in the habitat conditions, in the floristic composition there were selected two variants within the association: phytocoenoses with predominant Prunus spinosa species as an ecological variant with Prunus spinosa of the association Ligustro‑Prunetum R. Tx. 1952. and phytocoenoses with the floristic composition affected with the higher addition of cultural plant species (Vitis vinifera), as an ecological variant of association Ligustro‑Prunetum R. Tx. 1952 association withVitis vinifera which suggests big anthropic influence in these localities. – Anthrisco-Lycietum halimifolii Jurko 1964 association The shrubs of the association occupy very small area in the locality as they are located at the top parts of moderate slopes sufficiently warmed by the sun. They create differently wide stripes (2–15 m) at the interface of fields and vineyards ensuring favourable habitat conditions and additional nutrient supplementation from the fertilization of agricultural land. The cover of the tree layer is negligible and we can find there not high specimens of Acer campestre and Juglans regia. Average cover of the shrub floor reaches 95 % and the layer is characteristic with mono-dominance of introduced Lycium barbarum species which creates dense and impenetrable stands. Prunus spinosa and Rosa canina species are characterised with high stability with lower cover. Herb layer is very sparse with the average cover 20%. Inside the shrubs, there only rarely occur herbaceous species, mostly thermophilous ruderal and nitrophilous species of therophytes – Ballota nigra, Anthriscus cerefolium, Fallopia convolvulus, Galium aparine, Geum urbanum, Urtica dioica,

Arum alpinum and before the foliage of the shrubs Veronica hederifolia and Lamium purpureum. From the grass species occurs in the layer mesophyte Poa nemoralis. o The Black Locust communities The studied Black Locust communities syntaxonomically belong to Robinietea Jurko ex Hadač and Sofron 1980 family and to the two alliances. Chelidonio-Robinion Hadač and Sofron 1980 alliance includes Chelidonio-Robinietum Jurko 1963 association and Balloto nigrae-Robinion Hadač and Sofron 1980 alliance includes Bromo sterilis-Robinietum Jurko 1963 association (Fig. 2). – Chelidonio-Robinietum Jurko 1963 association In the studied territory it is the second most spreading and with the number of species the richest Black Locust community. It occurs especially on the slopes with western exposition and slight tendency. Common feature of all plots is sufficiency of soil moisture and increased mineral content of the soil. The tree layer is created with the dominant Black Locust completed in some cases with the native oaks (Quercus cerris, Q. robur). Average cover of the layer is 75% whilst average cover of the shrub layer is 10–55%. Within the species composition there dominates Sambucus nigra, another dominant species are Ligustrum vulgare and Robinia pseudoacacia. The differential species which differentiate one association from the other are represented by introduced Mahonia aquifolium and Prunus cerasus species which penetrate to the stands from the scrubland communities. Mahonia aquifolium species is nowadays considered to be an invasive species and probably it got to the community from the neighbouring Arborétum Mlyňany. Physiognomy of the herb layer is largely identified with nitrophilous species with the dominant Chelidonium majus. The layer is also rich on Galium aparine, Allium vineale, Urtica dioica and others.

Modified TWINSPAN Dendogram

Fig. 2. Dendrogram created from the four groups of records within the Black Locust communities representing syntaxonomic units (1 Chelidonio-Robinietum association, 2 facia withRubus caesius, 3 facia withVinca minor, 4 Bromo sterilis-Robinietum association).

159

In the community there is noticeable influence of the time aspect and therefore there is conditional species occurrence on the basis of growing season as well. Early in spring there is visibly increased occurrence Ficaria bulbifera species and Veronica hederifolia species, but later there dominates Chelidonium majus species with the mixture of grasses, especially Bromus sterilis species. During the summer period the herb layer becomes dry. On the basis of certain differences in the phytocenological, ecological and habitat conditions and dominant representation of Hedera helix species, but also on the basis of the highest similarity with this association we selected ecological variant with Hedera helix with facias with Rubus caesius and Vinca minor within the association. – Bromo sterilis-Robinietum Jurko 1963 prov. association In the studied area it is the most spreading association but regarding the species number it is poorer than previous association. It occurs on the similar habitats regarding exposition and slope tendency but generally on the bright and drier places with sandy and mineral-poor soils. The tree layer is created with the dominant Black Locust which is, however, a bit lower in its growth. The layer is also rich on Acer campestre, Carpinus betulus, Quercus cerris and Q. robur species. The shrub layer has lower cover (1–50%) and again it is characterised with the domination of Sambucus nigra species, constantly also occurs Robinia pseudoacacia species and significantly is also represented Euonymus europaeus species. Within the herb layer there is dominant Bromus sterilis species, with the lower cover occur nitrophilous Stellaria holostea, Galium aparine, Cheli-

donium majus, Arum alpinum, Geum urbanum and Urtica dioica species. Compared to the previous association, this association especially differs with Anthriscus sylvestris, Arrhenatherum elatius, Ballota nigra, Geranium robertianum, Lamium purpureum and Viola hirta species. Also in this association there is noticeable seasonal character of the herb layer, significant spring aspect at the beginning of summer (after fading of Bromus sterilis species) changes and understorey becomes poor. Ecological analysis of the study communities For the purposes of ecological analysis was used the selection into the shrub and Black Locust communities, the communities were compared with each other regarding their demands on different factors of the environment and we found out the following facts. Demands of both community groups regarding their light requests are quite equal. The numerical values are in the range 5.5–6.4 and they can be evaluated as the half-shade-like or even half-light-like communities. With respect to the temperature they are thermophilic communities, relatively more thermophilic seem to be the shrubs. In the question of continentality both groups can be classified as oceanic or even sub-oceanic, i.e. containing the species that occur in most parts of Central Europe. Demands of both groups regarding their requests on soil moisture are equal, most of the phytocoenoses’ species inclines to the dry or even freshly wet soils. In the question of soil reaction there are no differences between the groups. Eco-index 7 characterizes the species of acidic to neutral soils. Regarding the content of nitrogen substances in the soil, both groups can be described as nitrophilous communities.

Black Locust Scrubland

Fig. 3. Comparison of the scrubland and Black Locust communities on the basis of average eco-indexes (L, light; T, temperature; C, continentality; M, moisture; R, soil reaction; N, soil nitrates).

160

Ecological analysis of the communities confirmed that the scrubland and also Black Locust communities are ecologically similar in the studied area and differentiate with each other only slightly in the individual ecological indicators (Fig. 3).

Conclusion In the paper we focused on geobotanical and ecological characteristics of the scrubland and Black Locust communities in the Arborétum Mlyňany surroundings which cover quite large areas on this territory. We confirmed the fact that besides aesthetic function the shrubs in the agricultural land also have all-round biological and economic importance. Acquired phytocoenological data represent only a fragment of vegetation diversity of the shrub and Black Locust stands on the territory of Central Požitavie. Despite we hope that the information will contribute to the knowledge about the real state of scrubland and Black Locust communities on the whole territory of Slovakia and we also believe that the paper will help to the deepening of public awareness about the studied area.

Acknowledgements The work was supported by projects of VEGA Slovakia No. 1/0551/11 and 2/0059/11.

References Benčaťová, B., Benčať, T. 2005. The black Locust Communities in the Northen Part of „Pohronská pahorkatina“ Hills. Thaiszia, 15: Suppl. 1, 191–195. Benčaťová, B., Benčať, T. 2008. Black Locust communities from Slovak Gate to Danube. Thaiszia, 18: Suppl. 1, 3–7. Benčaťová, B., Benčať, T., Koprda, J. 2008. Agátové spoločenstvá okolia Arboréta Mlyňany [Black Locust communities of the Arborétum Mlyňany surroundings]. In Dendrologické dni vArboréte Mlyňany SAV. Zborník referátov z medzinárodnej vedeckej konferencie, Arborétum Mlyňany SAV, 15. – 16. 10. 2008. Vieska nad Žitavou: Arborétum Mlyňany SAV, p. 167–174. Braun-Blanquet, J. 1964. Pflanzensoziologie. Grund züge der Vegetationkunde. Wien, New York: Springer. 865 p. Hadač, E., Sofron, J. 1980. Notes on syntaxonomy of cultural forest communities. Folia geobot. phytotax., 15: 245–258. Hennekens, S. M. 1995. TURBO(VEG). Software package for input, processing and presentation of phytosociological data. User’s quide. Instituut voor Bosen Natuur,Wageningen and Unit of vegetation Science, University of Lancaster, Lancaster.

Hill, M.O. 1979. TWINSPAN: a FORTRAN program for arranging multivariate data in an ordered twoway table by classification of the individuals and attributes. Ithaca, New York: Cornell University, Section of Ecology and Systematics. 90 p. Jarolímek, I. , Šibík, J. (eds). 2008. Diagnostic, constant and dominant species of the higher vegetation units of Slovakia. Bratislava: Veda. 332 p. Jurko, A. 1963. Zmena pôvodných lesných fytocenóz introdukciou agáta [Change of autochthonous forest phytocoenoses by introduction of Black Locust]. Českoslov. Ochr. Prír., 1: 56–75. Jurko, A. 1964. Feldheckengesellschaften und Uferweidengebüsche des Westenkarpatengebietes. Biol. Práce, 10/6. Bratislava: Veda, p. 5–102. Jurko, A., Kontriš, J. 1982. Fytocenologická aekologická charakteristika agátin vMalých Karpatoch [Phytocoenological and ecological characteristic of Black Locust communities in the Malé Karpaty Mts]. Biologia, Bratislava, 37 (1): 67–74. Kliment, J., Jarolímek, I. 2011. European hazel shrubs in the Veľká Fatra Mts: syntaxonomy and nomenclature. Haquetia, 12 (2): 149–170. Kontriš, J. 1966. Poľné spoločenstvá krovín severozápadnej časti Liptovskej kotliny [Field shrub communities of north-western part of Liptovská kotlina basin]. Biol. Práce, 12/ 9. Bratislava: Veda, p. 41–78. Koprda, J. 2008. Geobotanicko-ekologická charakteristika krovinných a agátových spoločenstiev v k. ú. obcí Slepčany aVieska nad Žitavou [Geobotanyecological characteristic of shrub and Black Locust communities in the cadastre areas of villages Slepčany and Vieska nad Žitavou]. Diploma work. Zvolen: Technical University in Zvolen, Faculty of Ecology and Environmental Sciences. 93 p. Marhold, K., Hindák, F. (eds) 1998. Zoznam nižších avyšších rastlín Slovenska [List of lower and higher plants of Slovakia]. Bratislava: Veda. 687 p. Ščepka, A. 1982. Spoločenstvá s agátom bielym (Robinia pseudoacacia L.) vjužnej časti Východoslovenskej nížiny [Communities with Black Locust in southern part of Východoslovenská nížina lowland]. Acta bot. slov., ser. A, 6: 172–181. Ščepka, A. 1985. Vegetačné pomery južnej časti Východoslovenskej nížiny [Growing conditions of the southern part of Východoslovenská nížina lowland]. Acta bot. slov., ser. A, 8: 141–151. Šimonovič, V., Šomšák I., Kollár, J., Kanka, R., Nikodémová, Z. 2002. Charakteristika spoločenstiev s agátom bielym na Borskej nížine [Characteristic of the communities with Black Locust in Borská nížina lowland]. Phytopedon (Bratislava), Suppl., 1: 211–216. Tichý, L. 2002. Juice, software for vegetation classification. J. Veg. Sci., 13: 451–453.

161

Valachovič, M. 2002. Trnkové alieskové kroviny [Blackthorn and hazel shrubs]. In Stanová, V., Valachovič, M. (eds). Katalóg biotopov Slovenska. Bratislava: Daphne – Inštitút aplikovanej ekológie, p. 36–37.

Valachovič, M. 2007. Klasifikácia spoločenstiev krovín na Slovensku – možný koncept riešenia [Classification of shrub communities in Slovakia – possible solution concept]. Bull. Slov. bot. spol., 29: 169–176. Received December 6, 2012 Accepted April 14, 2013

162

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Diversity in honey locust (Gleditsia triacanthos L.) seed traits across Danube basin

Peter Ferus*, Marek Barta, Jana Konôpková, Silvia Turčeková, Peter Maňka, Tomáš Bibeň Mlyňany Arboretum SAS, Vieska nad Žitavou 178, 951 52 Slepčany, Slovak Republic

Abstract Ferus, P., Barta, M., Konôpková, J., Turčeková, S., Maňka, P., Bibeň, T. 2013. Diversity in honey locust (Gleditsia triacanthos L.) seed traits across Danube basin. Folia oecol., 40: 163–169. Honey locust (Gleditsia triacanthos L.), in the past planted as ornamental, technical or forest tree, is presently considered as casually invasive tree in Danube basin. Since plant invasiveness is usually tightly associated with its reproduction biology, in this work we focused on characterization of seeds from honey locust populations across this area. Analysing seed coat colour, thousand seeds weight (TSW), seed projection area, seed thickness, percentage of germinated seeds and their germination energy, as well as portion of seeds infested by honey locust seed beetle (Megabruchidius tonkineus), consumed part of seeds and their germination ability in relation to seed characteristics, local temperature means and precipitation sums during vegetation period, we came to the following conclusions: seed coat colour diversity decreases with geographical latitude; TSW, seed projection area and thickness were negatively correlated to mean temperature and positively to precipitation sum; between percentage of naturally germinated seeds and TSW as well as seed thickness we found positive correlations; germination energy showed positive relation to mean temperature and a negative one to precipitation sum; and the same relations were observed for infested seeds percentage and consumed seed part. No infested seed was able to germinate. From these results we can conclude that in colder and wetter conditions higher seed germinability, and in warmer and drier conditions enhanced germination energy of seeds supports spreading of this tree species. However, honey locust seed beetle can significantly affect seed germinability in regions with warm and dry summers. Keywords Danube basin, honey locust (Gleditsia triacanthos L.), honey locust seed beetle (Megabruchidius tonkineus), invasiveness, seed traits,

Introduction Honey locust (Gleditsia triacanthos L., Caesalpinaceae) is aleguminous tree originating in the middle and eastern part of North America, which was in Southern Slovakia and Hungary widely planted in parks as ornamental species, round vineyards, gardens and fruit groves as thorn-hedge, along roads and fields as wind barrier, and as a component of floodplain forests (Chrtková and Jasičová, 1988; György, 2007; Haraszthy, 2001). Into Europe it was introduced in 1700, and its first plantation

in the area of present Slovakia was established in 1806 in Dolná Krupá castle (Benčať, 1982). However, presently this species is ranked as often escaping from culture (Gojdičová et al., 2002) or newly as naturalized in Slovakia (Medvecká et al., 2012) and casually invasive in Hungary (Balogh et al., 2004). It causes extreme complications in Argentinian Pampa grasslands and central natural forests, where a large effort in ecological research has been paid (Chaneton et al., 2004; de Viana and Speroni, 2003; Marco and Paez, 2000). Finally, in 1993 Autralian Queensland

*Corresponding author: e-mail: [emailprotected]; phone +421 37633 4211-138.

163

spent 400,000 $ for eradication campaign ´search and destroy´, stimulated by honey locust infestation of 1,000 ha of Brisbane valley (Csurhes, 2004). Benčať (1982) classified this tree as very tolerant to industrial and transport emissions; Schindelbeck and Riha (1988) described its relatively high resistance to low soil reaction (up to pH 5). More recent works focus on its extraordinary tolerance to high temperature (Graves et al., 1991; Graves and Wilkins, 1991; Godoy et al., 2011) and drought (Graves and Wilkins, 1991; Burton and Bazzaz, 1991; Burton and Bazzaz, 1995), very helpful for its expansion. Pyšek and Richardson (2007) summarizing results of 59 studies in 64 alien plant species ordered traits associated with their invasiveness. As expected, features connected to generative reproduction appeared among them. In honey locust, combined clonal and sexual reproduction, short juvenile period, high seed production and high seed germinability were described (Marco and Paez, 2000). However, there is no reference about changes in this kind of traits with climatic conditions in literature. Therefore, in this work we analysed honey locust seeds from populations across the Danube basin.

Material and methods Honey locust (Gleditsia triacanthos L.) pods from different regions of Slovakia and Hungary (locations with GPS coordinates are listed in Table 1) were collected at the end of November 2011. From each locality three trees were sampled. Trees were determined using determination keys of Rehder (1990) and Krüssmann (1960). Table 1. Collecting sites of honey locust pods with GPS coordinates Locality

GPS coordinates

Vieska n/Ž. (SK)

N 48°19´00.6´´ E 18°22´05.4´´

Szirák (HU)

N 47°49´49.4´´ E 19°30´36.4´´

Gyöngyös (HU)

N 47°45´56.4´´ E 19°56´42.3´´

Debrecén (HU)

N 47°34´06.3´´ E 21°35´50.3´´

Mezőtúr (HU)

N 47°00´11.4´´ E 20°38´41.0´´

Békéscsaba (HU)

N 46°40´50.1´´ E 21°02´50.1´´

Szeged (HU)

N 46°22´38.3´´ E 20°02´25.5´´

Pods were stored at 3 °C till seeds were shucked (3 days). Then we let them dry at room temperature (21/17 °C during the day/night). In the course of four months lasting storage in these conditions, adults of honey locust seed beetles (Megabruchidius tonkineus Pic, 1904) have enclosed and flied out from seeds. Injured seeds were collected and analysed for weight reduction related to seed beetle life cycle (%) and germination ability (%). Thereafter, hundred healthy seeds from each tree sample were submitted to multiple analyses: o Seed coat colour o Thousand seeds weight (g) o Seed projection area (mm2) o Seed thickness (mm) o Germinated seeds (%) o Germination energy (mm d–1). Seed coat colour was determined visually. Thousand seeds weight (TSW) we calculated from weight of hundred seeds, dried at room temperature. Seed projection area was defined by scanning and area analysis using ImageJ software (ver. 1.46). For seed thickness measuring we utilized a slide calliper. For the latter two tests, seeds have been swelling in water for 1 week, continuously transferred into transparent plastic boxes lined with a water-soaked tissue and analysed for natural germinability. Thereafter, non-swelled seeds were scarified in 93% sulphuric acid for one hour, washed thoroughly in water (Asl et al., 2011), let swell for 24 h and transferred into plastic boxes, as described above. Germination energy was analysed after 4 days of germination as hypocotyl length growth rate. All these procedures were done at 25 °C in constant diffuse light. Hungarian Central Statistical Office provided us meteorological data (monthly temperature means (°C) and monthly precipitation sums (mm) for year 2011) for most of the analysed collecting sites in Hungary. Mlyňany Arboretum SAS (in Vieska nad Žitavou) dispose of its own meteorological station. From these data we calculated mean temperatures and precipitation sums for a period June–October 2011 (period of flowering, pod/seed establishment, pod/seed growth and ripening; see Table 2). Experimental data were submitted to statistical analysis of variance (Anova) using Statgraphics Plus v.4.0 software. LSD tests at the 95% confidence level were performed to thousand seeds weight (TSW), seed

Table 2. Average meteorological data for the period June–October 2011 in most of the collecting sites Locality

Nearest meteo station

Vieska n/Ž. (SK)

Average temperature [°C] 17.4

Precipitation sum [mm] 307

Gyöngyös (HU)

Budapest (HU)

19.5

154

Debrecén (HU)

18.4

249

Mezőtúr (HU)

Kecskemét (HU)

18.9

212

Szeged (HU)

19.1

116

164

projection area and thickness as well as germination parameters. Correlation analysis between respective parameters was accomplished using application MS Excell 2010.

Results Seed coat colour of seeds from two northern collecting sites in Hungary (Szirák and Gyöngyös) showed larger variability than samples from Debrecén, Mezőtúr and Békéscsaba (Table 3). Only trees from alley near Szeged produced seeds uniform in colour. In Vieska nad Žitavou we found green-brown and medium brown seeds. In weight of thousand seeds (TSW) LSD test distinguished three hom*ogenous groups of samples: 1. with TSW round 220 g (Vieska nad Žitavou, Szirák and Békescsaba), 2. round 175 g (Gyöngyös and Szeged) and 3. intermediate seeds of TSW about 200 g (Debrecén and Mezőtúr).There was no statistical difference in projection area of seeds from respective collection sites. By seed thickness, localities can be ordered

this way: 1. Debrecén, 2. Szirák, Gyöngyös, Szeged, 3. Vieska nad Žitavou, Mezőtúr and 4. Békéscsaba. Naturally germinating seeds represent only a negligible portion from the total number of analysed seeds (Table 4): in Vieska nad Žitavou and Gyöngyösup up to 0.5%, in Debrecén, Mezőtúr and Szeged round 1%, in Szirák 1.35%, and in Békescsaba the portion slightly exceeded 1.5%. One hour pre-treatment by concentra-ted sulphuric acid almost completely released the germination process of any seed sample. We found values ranging from 95 to 98.5%. Only in samples from Debrecén and Békéscsaba less than 95% germinated seeds was identified. Germination energy of most of the samples showed similar level (approximately 9 mm d–1). Extremes were observed only in Vieska nad Žitavou (8.09 mm d–1) and Mezőtúr (10.56 mm d–1). In samples from Szeged we revealed the highest (9.15%) seed infestation by honey locust seed beetle (Table 5). In those from Békéscsaba almost 7.7%, in Gyöngyös and Debrecéna little more than 4%, in Szirák and Mezőtúr near 2.5% and in Vieska nad Žitavou no infested seeds were found. However, although ranging

Table 3. Parametrization of seeds from respective collecting sites. Abbreviations: TSW, thousand seeds weight; gb, green- brown; mb, medium brown; db, dark brown; lb, light brown. Letters indicate a statistically significant difference at P = 0.05 Locality

TSW [g]

Seed projection area [mm2]

Seed thickness [mm]

*221.75 ± 20.51b

57.57 ± 3.75a

4.04 ± 0.26bc

gb, lb, mb

225.92 ± 29.27b

59.64 ± 7.84a

3.79 ± 0.26ab

gb, lb, mb

173.49 ± 14.90a

48.90 ± 6.97a

3.64 ± 0.26ab

519

gb, mb

193.61 ± 23.42ab

57.07 ± 9.20a

3.48 ± 0.37a

848

gb, lb

202.86 ± 26.37ab

51.91 ± 5.92a

4.05 ± 0.31bc

598

lb, mb

217.07 ± 1.06b

49.47 ± 3.87a

4.35 ± 0.24c

973

175.96 ± 3.99a

50.84 ± 0.84a

3.58 ± 0.08ab

Analyzed pods

Analyzed seeds

Seed coat colour

Vieska n/Ž. (SK)

562

gb, mb

Szirák (HU)

575

Gyöngyös (HU)

463

Debrecén (HU)

Mezőtúr (HU)

Békéscsaba (HU) Szeged (HU) *

Average ± SD.

Table 4. Germination characteristics of honey locust seeds from respective collecting sites in Slovakia and Hungary. Letters indicate a statistically significant difference at P = 0.05

Germinated seeds [%]

Locality

Germination energy [m d–1]

Non-pretreated

Pre-treated

Vieska n/Ž. (SK)

0.50

*96.94 ± 4.33a

8.09 ± 1.68a

Szirák (HU)

1.34

97.97 ± 2.02a

9.30 ± 0.30ab

Gyöngyös (HU)

0.33

97.29 ± 2.05a

9.23 ± 0.35ab

Debrecén (HU)

1.07

93.78 ± 7.54a

9.00 ± 1.21ab

Mezőtúr (HU)

1.00

95.29 ± 4.55a

Békescsaba (HU)

1.68

92.54 ± 11.13a

8.89 ± 0.39ab

Szeged (HU)

1.01

98.59 ± 1.22a

9.22 ± 0.30ab

10.56 ± 1.89b

Average ± SD.

165

from 24.5% (Vieska nad Žitavou) to 41.37% (Szeged) in average, no statistically significant difference in consumed part of endosperm across Danube basin was detected. And finally, no infested seed was able to germinate. Performing correlation analyses, we found that TSW was strongly determined both by seed projection area and seed thickness (r values were a little higher than 0.6; Table 6). However, there was a weak negative relation between the latter two seed characteristics (r = –0.188). We also observed a strong negative correlation between TSW and mean temperature for June–October period (r = –0.897) and a strong positive one between this parameter and precipitation sum for the same period (r = 0.917). Seed projection area was strongly correlated with mean temperature (r = –0.918) and precipitation sum (r = 0.885). For seeds thickness we only observed moderate correlations to these meteorological parameters (r = –0.459 and r = 0.507, respectively). Despite of no relation of the percentage

of non-pretreated germinated seeds to seed projection area and mean temperature, moderate positive correlations of this characteristics to TSW (r = 0.449) and seed thickness (r = 0.389) as well as a weak one to precipitation sum (r = –0.117), were revealed. Percentage of pretreated germinated seeds was strongly related to precipitation sum (r = –0.544), moderately related to seed thickness (r = –0.467) but only weakly to the rest of parameters. Germination energy showed a strong correlation with mean temperature (r = 0.633) but a moderate one with precipitation sum (r = –0.434). However, it was not correlated with seeds characteristics. Except of seed thickness (r = –0.105), percentage of seeds infested by honey locust seed beetle was strongly related to all seed/weather characteristics. In the case of seed part consumed by beetle(s) we observed a strong negative correlation to TSW (r = –0.728), seed projection area (r = –0.676) and precipitation sum (r = –0.742). It was moderately related (r = 0.486) to mean temperature and weakly to seed thickness (r = –0.236).

Table 5. Percentage of infested honey locust seeds, consumed part of seeds and germinated infested seeds as influenced by collection locality. Letters indicate a statistically significant difference at P = 0.05 Locality

Seeds infested by seed beetle [%]

Consumed part of seeds [%]

Germinated infested seeds [%]

Vieska n/Ž. (SK)

0.00

–

Szirák (HU)

2.61

*24.90 ± 5.73a

Gyöngyös (HU)

4.54

39.53 ± 4.45a

Debrecén (HU)

4.05

34.11 ± 3.99a

Mezőtúr (HU)

2.71

30.32 ± 21.79a

Békescsaba (HU)

7.69

37.76 ± 9.05a

Szeged (HU)

9.15

41.37 ± 1.49a

Average ± SD.

Table 6. Correlation coefficients (r) of relations between respective seed/weather parameters in honey locust populations for year 2011 TSW

Seed projection area

TSW

–

0.6***

Seed projection area

–

Seed thickness Non-pretreated germinated seeds

– 0.449**

–

Seed thickness

Mean June–October temperature

June–October precipitation sum

0.655***

–0.897***

0.917***

–0.918***

0.885***

–0.188* –

–0.459**

0.049

0.389**

0.056

–0.117*

0.507***

0.172*

–0.467**

0.24*

–0.544***

Pre-treated germinated seeds

–0.251*

Germination energy

–0.01

0.051

0.633***

–0.434**

Infested seeds

–0.512***

–0.659***

–0.105*

0.69***

–0.894***

Consumed seed part

–0.728***

–0.676***

–0.236*

0.486**

–0.742***

0.03

*** – strong (1 > r ≥ 0.5), ** – moderate (0.5 > r ≥ 0.3) and * – weak linear regression (0.3 > r ≥ 0.1).

166

Discussion Within following locations ordered by lowering geographic latitude we found decreasing diversity of seed coat colour (from green-brown, light brown and medium brown to medium brown). Schoeps (2002) defined that for this pigmentation chlorophylls, carotenes and xanthophylls are responsible. Since sampled trees were components of alleys, and since this species has polygamous character and it is fertilized by insects (Chrtková and Jasičová, 1988; Koblížek, 1995), gene flow between neighbouring individuals was expected. In the work of Schnabel and Hamrick (1995) we can read about 17–30% minimum estimates of pollen gene flow, depending on maternal trees distance (round 100 and 200 m). However, in general seed coat colour is a trait, which is highly influenced by environmental conditions (Souza and Marcos-Filho, 2001). Ertekin and Kirdar (2010) studied effect of different seed coat colour on other seed characteristics of honey locust. They observed higher hundred seeds weight, seed coat weight, endosperm weight and embryo weight as well as germination in yellow coloured seeds compared with light and dark brown seeds. Despite of different seed coat colour scale, we can see similar trend in TSW but not in the percentage of germinated seeds. According to Asiedu and Powell (1998), slow rates of imbibition, caused by seed shrinkage and greater seed coat adherence to cotyledons during maturation, are associated with pigmentation. As indicated by Geneve (2009), after dormancy release lens and micropyle function as a primary water gap for seed imbibition in honey locust as well as water locust (Gleditsia aquatica Marsh.). Barnabás et al. (2008) summarize that high temperature and water deficit can impair ovary and embryo sac development, cause pollen sterility as well as fruit/ seed abortion or reduce their growth by restricted allocation of storage materials in cereals. This is in agreement with a general view of Fenner (2010) and Martre et al. (2011) on seed morphogenesis. Therefore, it is not a surprise that we found negative correlations of TSW, seed projection area and seed thickness with mean temperature as well as positive correlations of these seed parameters with precipitation sum in respective locations of Danube basin for a period from June till October, when reproductive cycle of honey locust has been accomplished. However, even in the warmest and driest regions stress was not enough intense to endanger seed germinability (Gutterman, 1991). Borges et al. (2005) focused on seed maturation process in Caesalpinia echinata Lam., a species relative to honey locust, at one place but in two years differing in rainfall during reproduction period. At the full fruit ripeness, seeds differed only in thickness (lower in drier year 2002).

Marco and Paez (2000) present honey locust as a plant species conferring high potential to become invasive: fast growth, clonal and sexual reproduction, short juvenile period, high seed production and high seed germinability. This coincides with a general ranking of plant features associated with their invasiveness (Pyšek and Richardson, 2007). Compared with Acacia aroma Hook. et Arn., a native to Argentina, invasive honey locust disposes of higher seed production per plant, percentage of scarified seed germination and density of seedlings around the focal individuals (Ferreras and Galetto, 2010). Herrera and Laterra (2009) show in an ecological study from flooding Pampa grassland that addition of seeds of invasive species promoted seedling emergence, and this effect was higher for large than for small-seeded species. Similar results obtained Eisenhauer and Scheu (2008) stating that established grassland community and invader seed size significantly affected the number of invader plants, while invader biomass was only affected by the established community. Jakobsson and Eriksson (2000) also found that the relative recruitment in undisturbed sward increased with increased seed size, and both recruitment success and seedling size were positively related to seed size. Although it was less strong, we revealed a correlation between the percentage of naturally germinated seeds and their TSW and thickness, as well. It can be explained by rising seed coat impermeability for water, associated with increasing adversity of environmental conditions during later stages of seed maturation (Souza and Marcos-Filho, 2001), responsible also for reductions in seed weight. But since germination energy was positively correlated to mean temperature and negatively to precipitation sum, seedlings of larger vigour can be expected in warmer and drier conditions. It is interesting that dark brown seeds, more frequently produced in such conditions, need much less water for germination than seeds of more lightly toned seed coats (Ertekin and Kirdar, 2010). However, warmer and dried environment was associated with higher seed infestation by honey locust seed beetle (György, 2007; Majzlan, 2011; Jermy et al., 2002), which exclude them from the pool of potentially germinating seeds. So, this is not the case of relative Bruchidius dorsalis Fahraeus, considered as a crucial bio-agent providing germination of Gleditsia japonica Miq. seeds (Takakura, 2002). Thus, we can distinguish two different strategies supporting honey locust spreading: i) through higher germinability of larger seeds associated with lower temperatures and higher precipitation, and ii) through higher germination energy of smaller seeds connected with higher temperatures and lower precipitation. These results support the knowledge about high environmental plasticity of honey locust described by Godoy et al. (2011).

167

Acknowledgements This work was supported by research projects Vega 2/0156/11, Vega 2/0085/09, Vega 2/0076/09 and SKRO-0013-10. Special thanks to Dr. Pavol Eliáš for technical help.

References Asiedu, E.A., Powell, A.A. 1998. Comparisons of the storage potential of cultivars of cowpea (Vigna unguiculata) differing in seed coat pigmentation. Seed Sci. Technol., 26: 211–221. Asl, M. B., Sharivivash, R., Rahbari, A. 2011. Effect of different treatments on seed germination of honey locust. Modern Appl. Sci., 5: 200–204. Balogh, L., Dancza, I., Király, G. 2004. Actual list of neophytes in Hungary and their classification according to their success. In Mihály, B., BottaDukát, Z. (eds). Biological invasions in Hungary – invasive plants. Budapest: Természet BÚVÁR Alapítvány Kiadó, p. 61–92. Barnabás, B., Jäger, K., Fehér, A. 2008. The effect of drought and heat stress on reproductive processes in cereals. Pl. Cell Envir., 31: 11–38. Benčať, F. 1982. Atlas rozšírenia cudzokrajných drevín na Slovensku [Atlas of exotic tree species spreading in Slovakia]. Bratislava: Veda. 368 p. Borges, I.F., Neto, J.G., Bila, D.A.C., FigueiredoRibeiro R.L., Barbedo, C.J. 2005. Maturation of seeds of Caesalpinia echinata Lam. (brazilwood), an endangered leguminous tree from the Brazilian Atlantic forest. Brazil. Arch. Biol. Technol., 48: 851–861. Burton, P.J., Bazzaz, F.A. 1991. Tree seedling emergence on interactive temperature and moisture gradients and in patches of old-field vegetation. Am. J. Bot., 78: 131–149. Burton, P. J., Bazzaz, F.A. 1995. Ecophysiological responses to tree seedlings invading different patches of old-field vegetation. J. Ecol., 83: 92–112. Chaneton, E.J., Mazia, C.N., Machera, M., Uchitel, A., Ghersa, C.M. 2004. Establishment of honey locust (Gleditsia triacanthos) in burned Pampean grasslands. Weed Technol., 18: 1325–1329. Chrtková, A., Jasičová, M. 1988. Gleditsia L. In Bertová, L. (ed.) Flóra Slovenska IV/4 [Flora of Slovakia IV/4]. Bratislava: Veda, p. 11–12. Csurhes, S. M. 2004. Successful control of honey locust trees (Gleditsia triacanthos L.) in Queensland. In Sindel, B.M., Johnson, S.B. (eds). Fourteenth Australian Weeds Conference. 6–9 September 2004, Wagga Wagga, NSW. Sydney: Weed Society of New South Wales, p. 673–675. De Viana, M.L., Speroni, F.C. 2003. Invasion of Gleditsia triacanthos L. (Fabaceae) in San Lorenzo 168

mountain forest (Northwest Argentina). In Child, L., Brock, J.H., Brundu, G., Prach, K., Pyšek, P., Wade, P.M., Williamson, M. (eds). Plant invasions: ecological threats and management solutions. Loughborough, 6th International conference on the ecology and management of alien plant invasions. Leiden: Buckhuys, p. 277–287. Eisenhauer, N., Scheu, S. 2008. Invasibility of experimental grassland communities: the role of earthworms, plant functional group identity and seed size. Oikos, 117: 1026–1036. Ertekin, M., Kirdar, E. 2010. Effects of seed coat colour on seed characteristics of honeylocust (Gleditsia triacanthos). Afr. J. Agric. Res., 5: 2434–2438. Fenner, M. 2010. Environmental influences on seed size and composition. Hort. Rev., 13: 193–213. Ferreras, A. E., Galetto, L. 2010. From seed production to seedling establishment: Important steps in an invasive process. Acta oecol. – Int. J. Ecol., 36: 211–218. Geneve R.L. 2009. Physical seed dormancy in selected Caesalpinioid legumes from Eastern North America. Prop. Ornam. Pl., 9: 129–134. Godoy, O., de Lemos, J. P., Valladares, F. 2011. Invasive species can handle higher leaf temperature under water stress than Mediterranean natives. Envir. Exp. Bot., 71: 207–214. Gojdičová, E., Cvachová, A., Karasová, E. 2002. Zoznam nepôvodných, inváznych aexpanzívnych cievnatých rastlín Slovenska [List of alochthonous, invasive and expansive vascular plant species of Slovakia]. Ochr. Prír., 21: 59–79. Graves, W.R., Joly, R.J., Dana, M.N. 1991.Water-use and growth of honey locust and tree-of-heaven at high root-zone temperature. Hort. Sci., 26: 1309– 1312. Graves, W.R., Wilkins, L.C. 1991. Growth of honey locust seedlings during high root-zone temperature and osmotic-stress. Hortscience, 26: 1312–1315. Gutterman, Y. 1992. Environmental conditions during seed maturation affecting seed germination. Acta Horticulturae, 314. Leuven: International Society for Horticultural Science, p. 179–187. György, Z. 2007. To the biology of the honey locust seed beetle, Megabruchidius tonkineus (Pic, 1904) (Coleoptera: Chrysomelidae: Bruchinae). Folia Ent. Hung., 68: 89–96. Haraszthy, L. 2001. The floodplain forests in Hungary. In Klimo, E., Hager, H. (eds). The floodplain forests in Europe: current situation and perspectives. Leiden: Brill, p. 17–24. Herrera, L.P., Laterra, P. 2009. Do seed and microsite limitation interact with seed size in determination invasion patterns in flooding Pampa grassland? Pl. Ecol., 201: 457–469.

Jakobsson, A., Eriksson, O. 2000. A comparative study of seed number, seed size, seedling size and recruitment in grassland plants. Oikos, 88: 494–502. Jermy, T., Szentesi, A., Anton, K.W. 2002. Megabruchidius tonkineus (Pic, 1904) (Coleoptera: Bruchidae) first found in Hungary. Folia Ent. Hung., 63: 43-45. Koblížek, J. 1995. Gleditsia L. In Slavík, B. (ed.). Květena České republiky 4 [Flora of the Czech Republic 4]. Praha: Academia, p. 326. Krüssmann, G. 1960. Handbuch der Laubgehölze. Berlin-Hamburg: Paul Parey. 495 p. Majzlan, O. 2011. Faunistické príspevky zo Slovenska Coleoptera 6 [Faunistic contributions from Slovakia, Coleoptera 6]. Naturae Tutela, 15: 103–107. Marco, D.E., Paez, S.A. 2000. Invasion of Gleditsia triacanhos in Lithraea ternifolia Montane forests of central Argentina. Env. Manag., 26: 409–419. Martre, P., Bertin, N., Salon, C., Génard, M. 2011. Modellling the size and composition of fruit, grain and seed by process-based simulation models. New Phytol., 191: 601–618. Medvecká, J., Kliment, J., Májeková, J., Halada, Ľ., Zaliberová, M., Gojdičová, E., Feráková, V., Jarolímek, I. 2012. Inventory of the alien flora of Slovakia. Preslia, 84: 257–309.

Pyšek, P., Richardson, D.M. 2007. Traits associated with invasiveness in alien plants: Where we do stand? In Nentwig, W. (ed.) Biological Invasions. Ecological Studies, vol. 193. Berlin: Springer, p. 97–125. Rehder, A. 1990. Manual of cultivated trees and shrubs. Portland: Dioscorides Press. 996 p. Schindelbeck, R.R., Riha, S.J. 1988. Soil acidity, and the growth, biomass partitioning and leaf mineral composition of honey locust (Gleditsia triacanthos L.) seedlings. Tree Physiol., 4: 361–369. Schnabel, A., Hamrick, J.L. 1995. Understanding the population genetic structure of Gleditsia triacanthos L.: the scale and pattern of pollen gene flow. Evolution, 49: 921–931. Schoeps, B. 2002.Pigment composition and location in honey locust (Gleditsia triacanthos) seeds before and after desiccation. Tree Physiol., 22: 285–290. Souza, F.H.D., Marcos-Filho, J. 2001.The seed coat as amodulator of seed-environment relationships in Fabaceae. Revta Bras. Bot. Sao Paulo, 24: 365–375. Takakura, K. 2002. The specialist seed predator Bruchidius dorsalis (Coleoptera: Bruchidae) plays acrucial role in the seed germination of its host plant, Gleditsia japonica (Leguminosae). Funct. Ecol., 16: 252–257.

Received December 6, 2012 Accepted March 22, 2013

169

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Diversity of flora in historical parks on example of Sokolow Podlaski Region in Poland

Beata Fornal-Pieniak Department of Environmental Protection, Faculty of Horticulture, Biotechnology and Landscape Architecture, Warsaw University of Life Sciences – SGGW, Nowoursynowska Str.161, 02-787 Warsaw, Poland, e-mail: [emailprotected]

Abstract Fornal-Pieniak, B. 2013. Diversity of flora in historical parks on example of Sokolow Podlaski Region in Poland. Folia oecol., 40: 170–175. The paper is focused on diversity of flora in historical parks on example of Sokolow Podlaski Region. The research was done in 20 historical parks. Parks were established in XVIII–XIX centuries (landscape historic style) on oak-hornbeam habitat. Nowadays these parks are without anthropic pressure since II World War. The methodology included two research stages: field research and indoor studies. Research assumed flora and syntaxonomic analysis. The field research was done in years 2010–2012, including 70 phytosociological records on the area 500 m2. Plant species are represented by natural, semi-natural and synantropical vegetation. There was observed impact for migration species from surroundings areas in study areas. Keywords flora, historical parks, oak-hornbeam habitat, Sokolow Podlaski Region

Introduction Agricultural landscape is as a ‘sea’ with ‘green islands’ (Buček et al., 1996). Manor parks are sometimes the last sites for existing forest plants in agricultural landscape (Olaczek, 1972). Many plants and animal species typical for agricultural landscape are rare and on the verge of extinction (Robinson and Sutherland, 2002; Hermy and Stieperaere, 1981). Studies concerning historical parks mostly include issues related to a dendrological inventory, the history of the manor parks’ ownerships, objects’ condition and parks’ cultural values and functions alongside with a proposal of their protection and restoration. There are no many studies about flora in these objects. The aim of the study is diversity of flora in historical parks on example of Sokolow Podlaski Region.

zowiecko-Poleski section according to Matuszkiewicz (1993). The research was done in 20 historical parks in Sokolow Podlaski Region (Fig. 1). Parks were established in XVIII and XIX centuries (landscape historic style) on oak-hornbeam site.

Material and methods Sokolow Podlaski Region (137.18 km² surface) is located on east part of Poland. This area belongs to Ma170

Fig 1. Location of study area.

The methodology included two research stages: field research and indoor studies. The field research was done in years 2009–2011. 70 phytosociological records on the area 500 m2 (Braun-Blanquet, 1964) were done on afforested areas of parks. Plant species were grouped by phytosociological system following Matuszkiewicz (2001). The next stage of work was vegetation evalua-

tion of manor parks. Vegetation evaluation included 11 criteria as number of tree species in tree layer, origin of trees, types of plantings, tree-covered areas (%), health of plantings, number of shrub species, origin of shrubs, shrub-covered areas (%), number of herb species, origin of herb species, herb-covered areas (%) and bonitation from 0 to 3 points (Table 2). Parks were grouped

Table 1. Vegetation evaluation of historical parks Criteria

Bonitation points Tree layer

Number of tree species

Origin of trees (dominated in park)

Types of plantings

Tree-covered areas [%]

Health of plantings

Above 6 tree species

4–5 tree species

1–3 tree species

n – native species

c – cultivator species (planted trees)

ex–exotic species (introduced artificial)

Avenues, group of trees, individual trees

Group of trees, individual trees

Only group of trees

Above 50%

25%–50%

1%–24%

Good (mostly without canopy losses, tree hollows, diseases)

Medium (sometimes with canopy losses, tree hollows, diseases)

Bad (many canopy losses, tree hollows, diseases)

Shrub layer Number of shrub species

Origin of shrubs (dominated in park)

Shrub-covered areas [%]

Above 6 plant species

4–5 plant species

1–3 plant species

n – native species

c – cultivator species (planted shrubs)

ex–exotic species (introduced artificial)

Above 50%

25%–50%

1%–24%

1 Herb layer

Number of herb species

Origin of herb species (dominated in park)

Herb-covered areas [%]

Above 6 plant species

4–5 pant species

1–3 plant species

n – native species (apophytes, spontanophytes)

c – alien species (antropophytes)

Above 50%

25%–50%

0%–24%

171

into fours groups: parks with high vegetation values (from 25 to 33 points), parks with medium vegetation values (from 18 to 24 points ), parks with low vegetation values (from 8 to 17 points) and parks with very low vegetation values (from 0 to 7 points) (Table 1). Table 2. Plant species in tree layer in manor parks in Sokolow Podlaski Region

Table 3. Plant species in shrub layer in historical parks in Sokolow Podlaski Region Latin name

Syntaxonomic class

Abies alba Mill.

Vaccinio-Piceetea

Acer campestre L.

Querco-fa*getea

Acer platanoides L.

Querco-fa*getea

Acer pseudoplatanus L.

Querco-fa*getea

Latin name

Syntaxonomic class

Aesculus hippocastanum L.

Companion species

Acer campestre L.

Querco-fa*getea

Betula pendula Roth

Epilobietea angustifolii

Acer platanoides L.

Querco-fa*getea

Carpinus betulus L.

Querco-fa*getea

Acer pseudoplatanus L.

Querco-fa*getea

Corylus avellana L.

Querco-fa*getea

Acer pseudoplatanus Atropurpureum L..

Companion species

Crataegus monogyna Jacq.

Rhamno-Prunetea

Euonymus verrucosa Scop.

Querco-fa*getea

Aesculus hippocastanum L.

Companion species

fa*gus sylvatica L.

Querco-fa*getea

Alnus glutinosa L.

Salicetea purpureae

Fraxinus excelsior L.

Querco-fa*getea

Betula pendula Roth.

Companion species

Padus avium Mill.

Querco-fa*getea

Carpinus betulus L.

Querco-fa*getea

Philadelphus coronarius L.

Companion species

Corylus avellana L..

Querco-fa*getea

Populus alba L.

Salicetea purpureae

Euonymus verrucosa Scop.

Querco-fa*getea

Quercus robur L.

Companion species

fa*gus sylvatica L.

Querco-fa*getea

Ribes rubrum L.

Companion species

Fraxinus excelsior L.

Querco-fa*getea

Robinia pseudoacacia L.

Companion species

Malus domestica Borkh.

Companion specie

Rubus idaeus L.

Epilobietea angustifolii

Padus avium Mill.

Querco-fa*getea

Sambucus nigra L.

Epilobietea angustifolii

Populus alba L.

Salicetea purpureae

Sorbus aucuparia L.

Companion species

Prunus avium (L.) Moench.

Querco-fa*getea

Syringa vulgaris L.

Companion species

Prunus domestica L.

Companion species

Taxus baccata L.

Companion species

Prunus mahaleb (L.) Mill

Companion species

Thuja occidentalis L.

Companion species

Quercus robur L.

Companion species

Tilia cordata Mill.

Querco-fa*getea

Quercus rubra L.

Companion species

Ulmus laevis Pall.

Querco-fa*getea

Robinia pseudoacacia L.

Companion species

Sambucus nigra L.

Epilobietea angustifolii

Tilia cordata Mill.

Querco-fa*getea

Ulmus laevis Pall.

Querco-fa*getea

Results and discussion Plant species recognized in manor parks were represented by 7 plant communities of eutrophic forest community (Querco-fa*getea), riparian forest and brush of river valley (Salicetea purpureae) coniferous forest communities (Vaccinio-Piceetea), cut-over communities (Epilobietea angustifolii), bush communities (Rhamno-Prunetea), meadow and pasture communities (Molinio-Arrhenatheretea), margin communities, ruderal communities (Artemisietea vulgaris) and companion plant species (Tables 2–4). There have occurred plant species from all 3 syntaxonimic classes in tree layer. In shrub, there were distinguished plant species from 5 plant communities and 6 plant communities in herb layer.

172

Table 4. Plant species in herb layer in historical parks in Sokolow Podlaski Region

Latin name

Syntaxonomic class

Acer platanoides L.

Querco-fa*getea

Acer pseudoplatanus L.

Querco-fa*getea

Aegopodium podagraria L.

Querco-fa*getea

Aesculus hippocastanum L.

Companion species

Ajuga reptans L.

Companion species

Allium ursinum L.

Querco-fa*getea

Anemone nemerosa L.

Querco-fa*getea

Anemone ranunculoides L.

Querco-fa*getea

Asarum europaeum L.

Querco-fa*getea

Carex pilosa Scop.

Querco-fa*getea

Carex umbrosa L.

Querco-fa*getea

Carpinus betulus L.

Querco-fa*getea

Cerastium sylvaticum Waldst. & Kit

Artemisietea vulgaris

Chelidonium majus L.

Artemisietea vulgaris

Convallarja maialis L

Companion species

Corydalis cava L.

Querco-fa*getea

Corylus avellana L.

Querco-fa*getea

Dactylis glomerata L.

Molinio-Arrhenatheretea

Dactylis polygama Horv

Querco-fa*getea

Euonymus verrucosa Scop

Querco-fa*getea

fa*gus sylvatica L.

Querco-fa*getea

Ficaria verna Huds.

Querco-fa*getea

Fragaria vesca L.

Epilobietea angustifolii

Gagea lutea L.

Querco-fa*getea

Galeobdolon luteum Huds.

Querco-fa*getea

Galium schultesii L.

Querco-fa*getea

Galium sylvaticum L.

Artemisietea vulgaris

Geum urbanumL.

Artemisietea vulgaris

Glechoma hederacea L.

Artemisietea vulgaris

Most of the species were represented by the community from eutrophic forest community (Quercofa*getea) in all layers (tree layer – 55%, shrub layer – 40% and herb layer – 55%). Percentage of cover plant species from Rhamno-Prunetea and Vaccinio-Piceetea classes was not very high in all parks (Figs 2–4). There were distinguished many native plant species such as: Acer pseudoplatanus, Carpinus betulus, fa*gus sylvatica, Euonymus verrucosa, Corylus avellana, Galeobdolon luteum. Plant species of eutrophic forest community like: Carpinus betulus, Milium effusum which are typical for oak-hornbeam habitat. Tilia cordata, Fraxinus excelsior, Anemone nemerosa, Gagea lutea, Galeobdolon luteum, Corydalis cava, dominated in all parks. Plant species from Epilobietea angustifolii class are represented by: Sambucus nigra, Betula pendula and Rubus idaeus. Percentage of cover plant species from Rhamno-Prunetea and the other synthaxonomic classes was not very high in all parks. Fig. 2. Percentage cover of plant species in different syntaxonomic unities in tree layer in historical parks.

Hedera helix L.

Companion species

Impatiens noli-tangere L.

Querco-fa*getea

Impatiens parviflora L.

Artemisietea vulgaris

Lamium album L.

Artemisietea vulgaris

Lamium maculatum L.

Artemisietea vulgaris

50%

Lamium purpureum L.

Artemisietea vulgaris

40%

Lathyrus vernus (L.) Bernh.

Querco-fa*getea

Luzula pilosa (L.) Willd.

Companion species

Lysimachia nummularia L.

Molinio-Arrhenatheretea

20%

Maianthemum bifolium (L.) Schmidt

Companion species

10%

Milium effusum L.

Querco-fa*getea

Oxalis acetosella L.

Companion species

Plantago major L.

Molinio-Arrhenatheretea

Poa nemoralis L.

Querco-fa*getea

Polygonatum multiflorum L. Querco-fa*getea Prunella vulgaris L.

Molinio-Arrhenatheretea

Prunus avium L.

Querco-fa*getea

Pulmonaria officinalis L.

Querco-fa*getea

Quercus robur L.

Companion species

Robinia pseudoacacia L.

Companion species

Rubus caesius L.

Rhamno-Prunetea

Sambucus nigra L.

Epilobietea angustifolii

Scilla bifolia L.

Querco-fa*getea

Sorbus aucuparia L.

Companion species

Stachys sylvatica L.

Querco-fa*getea

Stellaria holostea L.

Querco-fa*getea

Taraxacum officinale L.

Molinio-Arrhenatheretea

Tilia cordata L.

Querco-fa*getea

Trientalis europaea L.

Vaccinio-Piceetea

Urtica dioica L.

Artemisietea vulgaris

Vinca minor L.

Querco-fa*getea

Viola mirabilis L.

Companion species

60%

30%

0% Querco-fa*getea

Companion species

Salicetea purpureae

Epilobietea angustifolii

Fig. 2. Percentage cover of plant species in different syntaxonomic unities in tree layer in historical parks.

Vegetation evaluation included eleven criteria: number of plant species in tree layer, origin of trees, types of plantings, tree-covered areas, health of plantings, number of plant species in shrub layer, origin of shrubs, shrub covered areas, number of plant species in herb layer, origin of herbs, herb covered areas (Table 5). High diversity of native plant species in tree, shrub and herb layer was observed in the park objects. Individual trees and groups of trees were typical plantings on study parks. Eight parks with high vegetation values and twelve parks with medium vegetation values were distinguished. Flora of manor parks is still modified by human and nature processes (Sikorski and Wysocki, 2003). Woody plant species were noticed in parks by many scientists e.g. Dzwonko and Loster (2001), Fornal-Pieniak and Wysocki (2006, 2009). Plants from Quercofa*getea occur in Sandomierska Basin park’s herb layer (Fornal-Pieniak, 2007) and Sokolow Podlaski Region. There were also observed plant species from MolinioArrhenatheretea and Trifolio-Geranietea sanguinei. 173

Fig 3. Percentage cover of plant species in different syntaxonomic unities in shrub layer in manor parks. 45% 40% 35% 30% 25% 20% 15% 10% 5% 0% Querco-fa*getea

Companion species

Epilobietea angustifolii

Rhamno-Prunetea

Salicetea purpureae

Fig 3. Percentage cover of plant species in different syntaxonomic unities in shrub layer in historical parks. Fig 4. Percentage cover of plant species in different syntaxonomic unities in herb layer in historical parks

70% 60% 50% 40% 30% 20% 10% 0% Querco-fa*getea

Artemisietea vulgaris

Companion species

MolinioArrhenatheretea

Epilobietea angustifolii

RhamnoPrunetea

VaccinioPiceetea

Fig 4. Percentage cover of plant species in different syntaxonomic unities in herb layer in historical parks.

Table 5. Planting evaluation of manor parks in Sokolow Podlaski Region Criteria / number of parks

A Tree layer Number of plant species

Origin of trees (dominated)

Types of plantings

Tree-covered areas

Health of plantings

Sum

Number of plant species

Origin of shrubs (dominated)

Shrub covered areas

Sum

B Shrub layer

C Herb layer Number of plant species

Origin of herbs (dominated)

Herb covered areas

Sum

Sum A + B + C

33–25 points: parks with high vegetation values; 24–18 points: parks with medium vegetation values; 17–8 points: parks with low vegetation values; 7–0 points: parks with very low vegetation values.

174

Nowadays we have very little information about condition of vegetation in historical parks in Poland. Many dendrological field researches were done but without list of plant species in herb layer. This pilot study is to show the vegetation evaluation diversity of historical parks in Sokolow Podlaski Region in Poland. Acknowledgement This research project would not have been possible without the support of prof. Czesław Wysocki and prof. Jan Supuka. References Buček, A., Lacina, J., Míchal, I. 1996. An ecological network in the Czech Republic. Brno: Veronica. 44 p. Braun-Blanquet, J. 1964. Pflanzensoziologie. Wien, New York: Springer Verlag. 865 p. Dzwonko Z., Loster S. 2001. The indicator plant species of ancient forests and their role for environment protection and mapping vegetation. Geogr. Stud., 178: 119–132. Fornal-Pieniak, B. 2007. Szata roślinna parków wiejskich Kotliny Sandomierskiej [Flora of rural parks on example of Sandomierska]. In Doktorant a rozwój nauk rolniczych. Wielokierunkowość badań w rolnictwie. Tom 1. Zeszyty Naukowe Akademii Rolniczej im. Hugona Kołłątaja w Krakowie. Sesja naukowa, zeszyt 92. Kraków: Wydawnictwo Akademii Rolniczej, p. 223–231.

Fornal-Pieniak, B., Wysocki, Cz. 2006. Struktura szaty roślinnej parków wiejskich na przykładzie Krainy Kotlina Sandomierska [Structure of vegetation in manor parks on example of Sandomierska Basin]. Acta Sci. Pol. Silv. Colendar. Rat. Ind. Lignar., 5 (2): 31–45. Fornal-Pieniak, B., Wysocki, Cz. 2009. Diversity of ancient forest plant species in country parks. Ann. Warsaw Univ. Life Sc. – SGGW, Hort. Landsc. Archit., 30: 201–205. Hermy, M., Stieperaere, H. 1981. An indirect gradient analysis the ecological relationships between ancient and recent riverine woodlands to the south Bruges (Flanders, Belgium). Vegetatio, 44: 43–49. Matuszkiewicz, J.M. 1993. Krajobrazy roślinne i regiony geobotaniczne Polski [Vegetation landscapes and geobotanical regions]. Prace geograficzne IGiPZ PAN, 158. Wroclaw: Zakład Narodowy im. Ossolińskich. 107 p. Matuszkiewicz, W. 2001. Przewodnik do oznaczania zbiorowisk roślinnych Polski [Guidebook of plant communities]. Warszawa: PWN. 537 p. Olaczek, R. 1972. Rural parks as a refugium for native flora of forest. Protec. Home Nature, 20 (2): 5–22. Robinson, R.A., Sutherland, W. 2002. Post-war changes in arable farming and biodiversity in Great Britain. J. appl. Ecol. 39: 157–176. Sikorski P., Wysocki, Cz. 2003. Nature of structure and plant species changing in shade trees in rural parks on example of West Mazurion Region. Acta Sci. Pol. Form. Circ., 2 (1): 71–86.

Received December 6, 2012 Accepted May 13, 2013

175

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Differentiation of some interspecific hybrids of firs (Abies sp.) according to the length of primary branches and number of their secondary branches

Martin Galgóci1, Peter Maňka1, Andrej Kormuťák2, Vladimír Čamek2, Dušan Gömöry3 Mlyňany Arboretum SAS, Vieska nad Žitavou 178, 951 52 Slepčany, Slovak Republic, e-mail: [emailprotected], [emailprotected] 2 Institute of Plant Genetics and Biotechnology SAS, Akademická 2, 950 07 Nitra, Slovak Republic, e-mail: [emailprotected], [emailprotected] 3 Faculty of Forestry, Technical University in Zvolen, Masarykova 24, 960 53 Zvolen, Slovak Republic, e-mail: [emailprotected] 1

Abstract Galgóci, M., Maňka, P., Kormuťák, A., Čamek, V., Gömöry, D. 2013. Differentiation of some interspecific hybrids of firs (Abies sp.) according to the length of primary branches and number of their secondary branches. Folia oecol., 40: 176–180. During 2011, the length of primary branches was measured in individual seedlings of firs representing 15 crossing variants. Measured branches were divided into separate groups according to the number of secondary twigs. Our data indicate the possibility for differentiation between the hybrid combinations based on length of their primary branches. The interspecific combinations A. pinsapo × A. alba and A. alba × A. pinsapo were more similar to mother species in this trait rather than to paternal parent. A given combination of different age differed primarily by the number of primary branches with ahigher number of secondary branches in older seedlings. Comparison involving both primary and secondary branches appears to be more efficient in discriminating between hybrid combinations than comparison primary based on secondary branches alone. Keywords Abies, branches, differentiation, hybrids, somatic heterosis

Introduction The genus Abies Mill. with its 49 species and about 126 known interspecific hybrids demonstrates the importance and potential opportunities for interspecific hybridization in forest tree breeding (Greguss, 1995; Kormuťák, 1994, 2004). Generally, it is concluded that extensive crossability between representatives of the genus Abies is a result of their high genetic relatedness due to specific pattern of speciation (Greguss, 1995). Klaehn and Winienski (1963) consider this to be result of geographic rather than genetic isolation. Recent studies also point to the fact that crossability in the genus is the result of geographical isolation, not genetic isolation (Kormuťák, 2004; Kormuťák et al., 2012; Kormuťák et al., 2013). 176

A high level of genetic diversity in interspecific hybrids is the result of heterozygous gene loci which leads to various forms of heterosis in hybrids (Rohmeder and Schönbach 1959; Mergen et al., 1964). Increased resistance to pests and diseases (Müller, 1989; Rohmeder and Schönbach, 1959) along with increased vitality and adaptability of the hybrids to changing conditions of the environment is generally referred to as adaptive heterosis (Greguss, 1995; Ausenac, 2002). It may also involve a higher proportion of surviving individuals, especially in relation to either immission load of the environment (Čítková, 1988; Rišková, 1982; Evans and Műller, 1972) or increasing annual temperature (Ausenac, 2002). Reproduction or seed heterosis is the ability of the offspring to produce increased amount of viable seeds

compared to the parental species. This applies in seed orchards and natural sites where adverse environmental conditions reduce the number of viable seeds (Uljukina and Derjuţkin, 1981; Greguss, 1995; Kormuťák, 1994, 2004). Somatic heterosis of interspecific fir hybrids is the most commonly observed phenomenon which is directly related to the enhanced production potential of forest trees (Hawlwey and de Hayes, 1985a, 1985b; Kormuťák, 1994, 2004; Rohmender and Eisenhut, 1961; Klaehn and Winienski, 1963; Mergen et al., 1964). The papers dealing with somatic heterosis in hybrid firs have been focused preferentially on height growth neglecting other growth characteristics of the hybrids. Considerable variation of morphological traits in fir hybrids makes a problem with their taxonomic identification (Greguss, 1995; Kormuťák, 1994, 2004). In present work we have tried to create a system of taxonomic differentiation of selected interspecific fir hybrids based on morphometric features of their primary branches.

Material and methods In summer 2011, the length of primary branches in seedlings of some interspecific combinations of firs and in control variants of the parental species from open pollination and controlled cross pollination were measured. We used hybrids of the following species: Abies alba Mill., Abies pinsapo Boiss., Abies numidica de Lannoy ex Carrière., Abies nordmanniana (Steven) Spach, Abies procera Rehder, Abies holophylla Maxim. We have analyzed 4-year old seedlings of A. pinsapo × A. numidica, A. pinsapo × A. alba, A. alba × A. pinsapo, A. alba – controlled crossing, A. alba – open pollination and A. pinsapo – open pollination along with 6-year old seedlings of A. nordmanniana × A. numidica, A. nordmanniana × A. procera, A. nordmanniana × A. alba, A. alba × A. numidica, A. alba – open pollination, A. nordmanniana – open pollination and 7-year old seedlings of A. nordmanniana × A. holophylla, A. nordmanniana × A. alba and A. nordmanniana – open pollination. Seedlings were grown in nurseries of Mlyňany Arboretum SAS in spaced 20 × 20 cm. They were also

Branch 1 degree

Branch 2 degree

Fig. 1 Branches of the first and second degree.

177

grown in standard light conditions without shading. Length measurement was carried out with the help of a ruler Fig. 1. Measured branches of the first degree were divided into groups according to the number of their secondary branches. These groups were statistically evaluated and tested by nested anova (SAS, 1988).

Results and discussion There were applied various approaches in characterization of intra- and interspecific hybrids of firs involving height growth parameters, morphology and anatomy of needles (Larsen, 1934; Klaehn and Winieski, 1962; Critchfield, 1988), pollen viability and seed quality (Galgóci, 2010), photosynthesing pigments, isoenzymes, terpenes and DNA (Gaudlitz, 1983; Zavarin et al., 1977; Wagner et al., 1987; Dong and Wagner, 1994). The length of branches asmorphological trait has not been used so far. Using Duncan grouping we were able to differentiate between some crossing variants according to the primary branches length and number of their secondary twigs (Table 1). It is of interest to mention that testing of individual groups of primary branches in 6- and 7-year old seedlings has not affected the descending order of the variants given in Table 1. Even achange of the order has not any statistical impact on differentiation of the crossing variants according to the type of their branches. It follows, therefore, that it does not matter which length parameter is used as acriterion for differentiation. In 6- and 7-year old seedlings both primary and secondary branches length characteristics may be used. It was not possible to discriminate between 7-year old seedlings of the crossing variants on the basis of length characteristics of the first three groups of the primary branches with 0–2 secondary twigs. Efficient in this respect were only the primary branches with three secondary twigs which had clearly differentiated the crossing variant A. nordmanniana × A. alba. In the category of primary branches with four secondary twigs, we have distinguished crossing variants A. nordmanniana × A. holophylla and A. nordmanniana – open pollination. Similar discrimination was also possible between 6-year old seedlings. The only exceptions in this respect were the variants A. alba – open pollination and A. nordmanniana × A. procera which had not complied discrimination criteria mentioned above (Table 1). Comparison of our data with those published by Galgóci et al. (2011) which refer to the height growth characteristics of the 6- and 7-year old progenies of the same crossing variants revealed ahigh degree of correlation with respect to the 7-year old seedlings only. At the 6-year old seedlings level the authors detected statistically significant differences between the pair of crossings A. nordmanniana × A. numidica – A.nordmanniana × A. alba and A. nordmanniana × A. numidica A. nordmanniana – open pollination. Quite different situation was observed at 178

the stage of 4-year old seedlings. At each group of the analyzed primary branches with corresponding number of twigs, the order of crosses has changed in this case at statistically significant level starting from the longest primary branch to the shortest one. The findings in the 5 groups scale of primary branches were those which had enabled to differentiate the 4 year old seedlings of all the analyzed crossing variants based on their primary branches. Comparison of the length characteristics in two different age categories of A. nordmanniana – open pollination seedlings have not revealed statistically significant differences in average lengths of the primary branches in 6- and 7-year old progenies. Both these age categories were represented by the seedlings with primary branch containing maximum 2 twigs. In the crossing variant A. nordmanniana × A. alba statistical differences between 6- and 7- year old seedlings exist only in the group of primary branches without and/or with one secondary twig. Seven year old seedlings are involved in the group of primary branch with 3 and 4 secondary twigs. Six year old seedlings have not occurred in this group. Among 6- and 4-year old seedlings of A. alba – open pollination and controlled crosses of A. alba maternal species statistically significant differences were revealed between seedlings of the same age but not between seedlings of different age categories. We have not observed such phenomenon in other groups of primary branches. Mutual comparison of the growth characteristics in the crossing variants A. pinsapo – open pollination 2007, A. alba – open pollination 2007, A. pinsapo × A. alba and A. alba × A. pinsapo resulted in the conclusion that variants A. pinsapo – open pollination and A. pinsapo × A. alba differ significantly in several groups of the primary branches from the variants A. alba – open pollination and A. alba × A. pinsapo. As arule, the hybrids have exhibited thetendency to be similar in this respect to mother tree. The phenomenon may be ascribed to the matrilinear inheritance of growth characteristics. A typical feature of crossing variants with arid species A. numidica and A. pinsapo involved the parental specimen in the formation of secondary twigs in relatively young plants as well as in plants with short primary branches. It is supposed that this feature of the hybrids is related to the ability of the parental species to invade new localities.

Acknowledgement This study was supported by the VEGA Grant Agency, projects no. VEGA 2/0076/09, and 2/0110/13. References Aussenac, G. 2002. Ecology and ecophysiology of circum- Mediterranean firs in the context of climate change. Ann. Forest Sci., 8: 823–832.

0.7

3.3

3.5

14.9

0.5–0.6

2.58

1.47

42.0

37.6

–

4.4

4.1

14.2

0.6–0.7

2.56

1.46

43.0

39.5

0.6

2.9

3.1

19.7

0.0–0.1

2.67

0.93

65.2

51.9

3.1

10.2

11.4

32.9

0.1–0.2

2.41

1.20

50.2

48.6

0.6

1.0

1.3

27.6

0.2–0.3

2.46

1.51

38.6

38.2

–

0.4

0.3

18.1

0.3–0.4

2.54

1.23

51.6

46.5

0.5

4.6

4.8

27.6

0.4–0.5

2.51

1.19

52.6

49.1

0.6

2.9

3.2

29.2

ρs, particle density; ρd, bulk density; P, porosity; Pk, capillary pores; Ps, semi-capillary pores; Pn, non-capillary pores; VA, air filled porosity; Θp, available water capacity.

low 45% for loam tend to inhibit root penetration. High bulk density and low porosity may adversely affect soil biological properties and lead to decreasing of microbial biomass due to oxygen deficiency in the compacted soils (Tan et al., 2005). Obi (1989) noted that alluvial soils do not exhibit any definite pattern with regard to porosity. Where sandy deposits dominate, macro-pores would expectedly dominate. On the other hand, where the deposited material is of high clay and organic matter content, water-logging may be expected to cause pore instability with resultant tendency towards the formation of smaller pores. Nevertheless, according to Bedrna et al. (1989), values of total pore space (P) do not give any indication of pore size distribution. Optimal pore distributions are as follows: 1/3 macro-pores (Pn) where aeration and water drainage take place and 2/3 meso (Ps) and micro-pores (Pk) for water retention and capillary elevation. When considering optimal pores distribution, very low amount of macro-pores of total porosity was found in studied profiles (in soil profile 1: 14–21%, in soil profile 2: 6–14% and in soil profile 3: 1–16%). Air porosity was also very low, mainly in soil profiles 2 and 3 (Table 2). High content of capillary water and low aeration caused reduction conditions in lower horizons what resulted to the development of described redoximorphic feature.

240

Values of basic chemical parameters are written in Table 3. Total soil organic carbon (CT) decreased with depth in all examined soil profiles, with values ranging from 28.4 to 40.1 g kg–1 for A horizons and 9.1–17.6 g kg–1 for subsoils. Exchangeable soil reaction (pHKCl) ranged from slightly acidic to slightly alkaline. Increased content of carbonates in deeper parts of soil profiles corresponded to higher values of pHKCl. Compared with soil profiles 1 and 2 dug on the left and right bank of river Žitava, soil profile 3 dug near the confluence of rivers Nitra and Žitava contained significantly higher amount of carbonates. This was possibly due to different chemical composition of alluvial sediments deposited by the river Nitra versus Žitava. According to Čurlík and Šefčík (2006) around 80% of the territory which crosses river Nitra contains 5.72% carbonates in humus horizon, whereas the river Žitava crosses territory with 5.72% carbonates only on 65% and the rest of territory contains only 0.16% carbonates. Analogously, concentration of carbonates in soil forming substrates was higher in soils of river Nitra basin compared to soil of the river Žitava basin. Moreover, Szombathová et al. (2007) reported that Eutric Fluvisol in NR Žitavský wetland (48º09’ N, and 18º19’ E, 40 km from NR Alúvium Žitavy) did not contain carbonates, and they were presented only in CGo horizon of Mollic

Fluvisol. On the contrary, Tobiašová (2010) found in Eutric Fluvisol in floodplain of the river Nitra 1–5% of carbonates. Additional reason of higher amount of carbonates in studied soil profile 3 could be the content of shell fragments in lower parts of profile which contributes to the rise of carbonate concentration. Consequently, in soil profile 3 the lowest values of hydrolytic acidity (H) were found, since acidic protons were well neutralized by carbonates (Table 3). In general, the cation exchange capacity (CEC) is related to the type of sediments of predominantly loam to clay-loam character. In the surface layers, the value is also affected by the content of humus. Sum of bases (S) ranged from 86 to 205 mmol kg–1 and CEC from 88 to 212 mmol kg–1. Comparing the sum of bases it is evident, that the lowest values were found in soil profile 3, mainly in Fvc/Gl and in Glp horizons. Presumably, in this profile the majority of base cations, mainly Ca2+ (Noskovič et al., 2010), were bound with CO32– anions to solid particles or they were part of the shell fragments. Therefore, the sum of bases and consequently also the cation exchange capacity was low, in spite the clay content was the highest just in soil profile 3 (Tables 1, 3). On the contrary, the values of CEC and also S proportionally increased with increasing of clay content in soil profile 1, where the lowest concentration of carbonates was determined and shell fragments were not occurred (Tables 1, 3). In all soil profiles, the sorption complex was saturated with base cations, mostly ranging between 97–98 %. Studied soils exhibited high buffering capacity related to carbonates content, middle CEC capacity and the high degree of base saturation. Results obtained in this study showed that soil properties in soil profile 3 (dug in the locality, where the river Žitava flows into the river Nitra) distinctly differed from soil properties in profiles 1 and 2 dug at a greater distance (2.3 and 1.7 km) from the estuary of

the river Žitava. Differences were mainly determined in soil texture, content of carbonates and pH values. Soil properties in NR Alúvium Žitavy were distinctly influenced by different sediments deposited by the river Nitra compared to Žitava and ground water level.

Acknowledgement The paper was published thanks to G.P. No. 1/0513/12 and 1/0544/13 of the Scientific grant Agency of Ministry of Education, Science, Research and Sport of the Slovak Republic.

References Bedrna, Z., Fulajtár E., Zrubec F., Juráni B. 1989. Pôdne režimy [Soil regimes]. Bratislava: Veda. 224 p. Brady, N.G., Weil R.R. 1999. The nature and properties of soils. 12. ed. New Jersey: Prentice-Hall, Inc. Simons & Schuster A Viacon Company. 881 p. Čurlík, J., Šefčík, P. 1999. Geochemický atlas Slovenskej republiky [Geochemical atlas of the Slovak Republic]. Bratislava: Ministerstvo životného prostredia SR. 99 p. Fiala, K., Kobza, J., Matúšková, Ľ., Brečková, V., Makovníková, J., Barančíková, G., Búrik, V., Litavec, T., Houšková, B., Chromaničová, A., Váradiová, D., Pechová, B. 1999. Záväzné metódy rozborov pôd. Čiastkový monitorovací systém – Pôda [Obligatory methods of soil analysis. Partial Monitoring system – Soil]. Bratislava: VÚPOP. 142 p. Fulajtár, E. 2006. Fyzikálne vlastnosti pôdy [Physical properties of soil]. Bratislava: Výskumný ústav pôdoznalectva aochrany pôdy. 142 p.

Table 3. Selected ecological properties of soils Soil profile

Horizon

1 Calcaric Fluvisol

40.1

Fvc

15.8

7.4

176.6

9.1

3.5

204.5

2 Calcaric Fluvisol 3 Calcaric Fluvisol

[g kg–1]

Fvc/Gl

CEC

[mmol kg–1] 27.4

132.6

CO32–

BS [%]

160

pH KCl

0.8

5.73

184

0.9

6.19

208

3.4

6.37

28.4

7.1

204.9

212

1.4

6.58

Fvc

17.6

4.7

171.3

176

2.0

6.92

Fvc/Gl

12.5

3.8

156.2

160

3.6

6.97

33.2

3.1

164.9

168

4.4

6.92

Fvc/Gl

12.5

2.8

109.2

112

13.6

7.24

Glp

11.4

2.4

85.6

17.6

7.43

CT, total soil organic carbon; H, hydrolytic acidity; BS, base saturation; S, sum of bases (Na+, K+, Ca2+, Mg2+); CEC, cationic exchange capacity.

241

Kadlec, R.H., Knight R.L. 1996. Treatment wetlands. Bocca Ratoon: CRC Press. 893 p. Kukla, J., Kuklová, M. 2009. Soil-ecological properties and plant communities in the Nature Reserve Chynoriansky luh. Folia oecol., 36: 86–93. Lapin, M. 2012. Dlhodobý režim teploty vzduchu v Hurbanove a úhrnov atmosférických zrážok na Slovensku [Long term regime of temperature in Hurbanovo and sum of atmospheric precipitation in Slovakia] [cit. 2013-01-11]. http://www.dmc.fmph.uniba.sk/public_html/climate/THurbanovo.htm Lelkes G., Vasiová E., Kiss J. 2006. Program hospodárskeho a sociálneho rozvoja obce Martovce 2006 [Program of economic and social development of the village Martovce 2006]. 96 p. [cit. 2010-11-12]. http://www.rrakn.sk/images/downloads/132-SKmartovce_2010_verzia%201.pdf Lukniš, M., Bako J., Berta J., Ferianc O., FeriancováMasárová Z., Fusán O., Futák J., Hejný S., Jurko A., Korbel L., Kurpelová M., Mičian Ľ., Michalko J., Petrovič Š., Schmidt Z., Šimo E., Vlček F., Zaťko M., 1972. Slovensko 2. Príroda [Slovakia 2. Nature] Bratislava: Obzor. 917 p. Noskovič, J., Babošová, M., Kvetanová, Ľ. 2010. Hodnotenie koncentrácie dvojmocných bázických katiónov vo vode Prírodnej Rezervácie Alúvium Žitavy [Evaluation of bivalent basic cations in the water of the Nature Reserve Alúvium Žitava]. Acta hydrol. slov., 11: 97–107.

Noskovič, J., Palatická, A., Porhajašová, J., Kováčik, P. 2009. Concentration of phosphate phosphorus and total phosphorus in the water in different biotopes of the Nature Reserve Alúvium Žitavy. Folia oecol., 36 :125–133. Obi, M.E. 1989. Some physical properties of wetland soils with reference to the tropics. In Internal report IC/89/354. Trieste: International Centre for Theoretical Physics, p. 26. Orlov, V., Grishina, I. 1981. Praktikum po chimiji gumusa [Guide of humus chemistry]. Moscow: Moscow University Publishing. 124 p. Szombathová N., Noskovič J., Babošová M. 2007. Selected chemical properties of soil in the Nature Reserve Žitavský wetland. Folia oecol., 34: 61–65. Šeffer, J. 1996. Mokrade pre život [Wetlands for life]. Bratislava: Nadácia DAPHNE. 32 p. Tan, X., Chang, S., Kabzems, R. 2005. Effects of soil compaction and forest floor removal on soil microbial properties and N transformations in boreal forest long-term soil productivity study. Forest Ecol. Mgmt, 217: 158-170. Tobiašová, E. 2010. Pôdna organická hmota ako indikátor kvality ekosystémov [Soil organic matter as an indicator of ecosystems quality]. Nitra: Slovenská poľnohospodárska univerzita. 107 p. WRB. 2006. World reference base for soil resources 2006. Rome: FAO. 128 p. Zaujec, A., Chlpík, J., Nádašský, J., Szombathová, N., Tobiašová, E. 2009. Pedológia azáklady geológie [Soil science and base of geology]. Nitra: Slovenská poľnohospodárska univerzita. 399 p.

Received January 28, 2013 Accepted March 6, 2013

242

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Ecophysiological research of European beech (fa*gus sylvatica L.) in high-productive mixed forests of the Poľana Mts in Central Slovakia Tibor Priwitzer, Vladimír Čaboun National Forest Centre, T. G. Masaryka 22, 960 92 Zvolen, Slovak Republic, e-mail: [emailprotected], [emailprotected]

Abstract Priwitzer, T., Čaboun, V. 2013. Ecophysiological research of European beech (fa*gus sylvatica L.) in highproductive mixed forests of the Poľana Mts in Central Slovakia. Folia oecol., 40: 243–250. This paper presents the results of ecophysiological research of European beech (fa*gus sylvatica L.) in highproductive mixed forests of the Slovak Poľana Mountain. This research was performed in the research and demonstration object Poľana - Hukavský grúň. The radiation, temperature and humidity regimes, as well as daily dynamic of photosynthetic activity and electric resistance of cambial tissue are presented within the whole beech crown profile. The impact of meteorological conditions on selected physiological processes was studied. The results confirmed close correlation between a diameter of trees d1,3 and biofield, as well as between a biofield and cambial tissue’s electric resistance. The considerable differences in CO2 uptake within individual beech crown layers were determined. Keywords beech, cambial tissue, electric resistance, gas exchange

Introduction

Material and methods

The site conditions can be considered to be the determining complex of indices and factors from the point of view of quantity and quality of physiological processes in forest trees. Knowledge on time and spatial dynamic of individual characteristics within the whole forest ecosystem is necessary for quantification of impact of meteorological characteristics on photosynthesis and production processes, damage of foliage (frost, radiation, immissions), water and energy regime of tree species, etc. This paper presents the results of ecophysiological research of European beech (fa*gus sylvatica L.) – the tree species with the largest distribution in the Slovak forests. The main attention is paid to the impact study of site conditions on selected physiological processes (mainly photosynthetic activity and electric resistance of cambial tissue). This contribution is based on the results gained within the framework of ecological and ecophysiological research being realized in the research and demonstration object Poľana – Hukavský grúň.

Description of research plot All measurements were done in the research and demonstration object (RDO) Poľana - Hukavský grúň which is a part of the Biosphere Reserve – Protected Landscape Area Poľana. The highest attention of research activity has been paid to the Permanent Research Plot – 0 (PRP 0) of which more detailed description, as well as the whole spectrum of problems being solved were published by Čaboun et al. (1996). Basic geographic, pedological, meteorological and typological characteristics of the RDO and PRP 0 are presented in (Table 1). Measurement of meteorological characteristics and ozone concentration All meteorological characteristics within the PRP 0 were continuously measured and recorded to the data logger (DELTA T). The following meteorological characteristics were measured: air temperature (Ta) and rel-

243

Table 1. Selected characteristics of the RDO Poľana – Hukavský grúň and the PRP 0 Characteristics of RDO

Characteristics of PRP – 0 (area 0.55 ha)

Orographic unit

Poľana

Elevation

850–860 m

District

Detva

Slope

5–15 %

Forest Enterprise

Kriváň

Stand age

90–120

Forest Administration

Poľana

Number of trees

342

Elevation

820–915 m

Tree species

Beech

70.0 %

Exposure

North-east

Spruce

19.6 %

Slope

5–35 %

Sycamore maple

4.6 %

Geological substrate

Volcanic

Fir

3.5 %

Forest Management

Beech forests (411)

Ash

2.0 %

Type Group fertile

Fertile fir beech

Aspen

0.3 %

Number of PRP

Forests (511)

Canopy

Forest type

nitrophilous fir beech forest (5302)

Soil type

Ando-eutric Cambisol

ative air humidity (RH) (0.3 m, 6 m, 29, 34, 46 m above the soil surface), soil temperature (Ts) (depth 0, 5 and 10 cm), global radiation (Q) (34 m), photosynthetically active radiation (PhAR) (37 m, 32 m, 29 m, 19 m above soil surface), precipitation (34 m), wind direction (46.5 m), wind speed (46.5 m, 38.8 m, 37m). More detailed technical parameters of meteorological measurements were published by Čaboun et al. (1996). A daily dynamics of Ta, RH was determined for three height levels (34, 29, 6 m). The amount of incident PhAR upon the forest stands was calculated from the values of Q according to the relations presented by Rovňáková (1986). The values obtained by this method were consequently quantified for individual height levels on the basis of direct measurements (cloudless days with typical daily course of PhAR). In addition to meteorological characteristics, the concentrations of atmospheric ozone (O3) were continuously measured above the stand. The analyzer ML 8810 (based on UV photometry) was used. The ozone concentrations were calculated according to Lambert– Bersch rule. More detailed methodology of atmospheric quality measurement in PRP 0 is presented by Čaboun et al. (1996). Determination of photosynthetic activity of beech foliage – ecophysiological measurements were realized on representative co-dominant beech (diameter d1,3 = 49 cm and height 35 m.). Its crown was divided into three separate crown layers (upper 32 m, central 29 m, lower 19 m from soil surface) on the basis of the previous measurements of leaf area, anatomical leaf structure, density of stomata, chlorophyll content (Priwitzer et al., 1996), as well as the incident PhAR. For determination of photosynthetic activity was used the portable photosynthetic system Li-6200 (Licor, Nebraska, USA). The detailed description of in244

strument and method of work with it was published by Priwitzer (1993, 1998). All measurements were performed directly in the crown space using the tower. The measurements were usually performed from June to September during favourable weather (days without precipitation) on physiologically mature leaves (leaves in phenophase of adult leaf) and ambient CO2 concentration (330–350 ppm). Daily dynamic of photosynthetic activity was determined by two methods as follows: o Measurement of CO2 exchange in hourly intervals for upper part of crown (sun type of leaves). More detailed methodology was published by Priwitzer et al. (1996). o Calculation using light response curves (relation of photosynthesis and PhAR intensity) determined on the basis of direct measurement, as well as conversed for individual crown layers according to the methodology presented by Marek et al. (1989), Marek et al. (1992) and Pirochtová and Marek (1991). Measurement of electric resistance of cambial tissue – within the framework of observing the relative tree vitality was used the TREE VITALITY METER – TVM 01. It is a portable electronic instrument for determination of health condition of standing trees, and occasionally damage of wood mass, operating on principle of electronic measurement of cambial layer resistance and wood tissue respectively. Air temperature is being scanned by thermal sound. The 24-hour dynamic, as well as seasonal resistance dynamic were finding out. More detailed methodology of measurement of forest trees’ cambial resistance was published by Čaboun (1994, 1997). Measurement of tree species biofield – a magnitude of tree species biofield was measured in sense of the methodology published by Čaboun (1993).

Results As an example of beech ecophysiology we present the measurement performed on warm summer day with allday occurrence of cumuliform cloudiness. Radiation regime – the curve of global radiation showed typical daily course with the values around 900 Wm–2 between 10 a.m. and 2 p.m. (daily max. 950 Wm–2 around the midday). Daily course of ozone concentration has slightly upward character with maximal values (58–63 ppb) between 2 p.m. and 4 p.m. At that time, the maximal values of air temperature were measured (Fig.1). The daily dynamic of PhAR above the stand has a similar character as for global radiation. Changes in intensity and amount of PhAR in crown space are showed in the Fig 1. The individual daily course differs significantly in certain parts of crown and during the day. While PhAR in the upper part of crown showed very similar daily dynamic like above the crowns (max. – 1,397 µE m–2 s–1 around the midday), in central part of crown (29 m) the max. daily value of PhAR (1,085 µE m–2 s–1) was recorded between 7 a.m. and 8 a.m. Fairly

balanced intensity of PhAR characterized the lower part of the crown during the whole day. An increase of PhAR was recorded only in very short periods of time, while maximal daily values (520 µE m–2 s–1) were found out between 8 a.m. and 9 a.m. As regards a decrease of PhAR amount, due to its penetration through the crown (Fig. 1), it can be observed for selected part of the day, that 95% of PhAR from the amount measured above the crown was found out in the upper part of crown at 8 a.m., 61% at midday, and 39% at 6 p.m. Within the central crown layer the values of PhAR reached 50, 42 and 8% of values measured above the crown resp. 11%, 5% and 2% in the lowest part of crown. Similarly there was found out that in the upper part of crown 70% of PhAR values were in the interval of 500–200 µE m–2 s–1, while in the lower layer 70% of values in the interval of 0–50 µE m–2 s–1 during the day. Temperature and humidity regime – daily course of air temperature and air humidity above the stand, as well as in the individual layers of the stand is shown in the Figure 2. Air temperature (Ta) in all height layers has gradually increased since early morning and it reached maximal daily values (from 18.7 to 20.4 °C)

1.4 1.2 1 0.8 0.6 0.4 0.2 0

Fig. 1. Daily course of global radiation (Q) and ozone (O3) concentration above the forest stand (34 m) and PhAR measured in 37, 32, 29, 19 m above the forest floor.

245

at around 3 p.m. The highest values of Ta during the day were measured in the height of 34 m above the surface of soil. It is a consequence of quicker warming of air layer in the area immediately above the crowns and followed radiation of crowns’ active surface within the infrared range of spectrum. Air humidity (RH) was fairly constant (80%) during the period between 6 a.m. and 10 a.m. in the whole vertical profile of the stand. After that time we recorded its decrease while minimal daily values (48–57%) it reached in all height layers at around 3 p.m. The lowest RH values were measured in 37 m, between 10 a.m. and 6 p.m. Photosynthetic activity of individual parts of the crown – daily dynamic of CO2 (AN) uptake determined for individual parts of the crown is shown in the Figure 3. When we look at the course of measured values (Fig. 3 – right), we find out the considerable differences in CO2 uptake in individual crown layers. While assimilation apparatus reached maximal daily values AN (8.09 µmol CO2 m–2 s–1) at around 11 a.m. in the upper part of the crown (sun type of leaves), in lower part of crowns (shade type of leaves) it was between 8 a.m. and 9 a.m. In the central crown layer (occurrence of both types of leaves), the maximal daily values AN (4.37 µmol CO2 m–2 s–1) were found out at around the midday. A considerable depression of CO2

uptake was recorded mainly in the central (at around 2 p.m.) and lower (between 11 a.m. and 1 p.m.) part of the crown. After a decrease of CO2 uptake we recorded more considerable increase of AN value (crown centre at around 3 p.m., lower part at around 2 p.m.) in the both crown layers. Significant decrease of CO2 uptake in all layers of the crown occurred after 4 p.m. When comparing the maximal values AN we can find out that central part of the crown reaches 49% and in lower crown part only 28% of AN of the upper crown part. When assessing the daily dynamic of CO2 uptake (sun leaves) obtained on the basis of direct measurement we recorded certain differences in comparison with simulated daily dynamic. While in direct measurement the daily maximum was between 6 a.m. and 8 a.m., and after that there occurred the consequent whole day decrease of values AN, in case of simulated determination of AN the daily maximum was between 10 a.m. and 11 a.m., and values of CO2 uptake were fairly constant during the bigger part of the day (7 a.m.–3 p.m.). Electric resistance of cambial tissue – daily course of electric resistance of cambial tissue in the investigated beech is given in the Figure 4. From the values measured in various cardinal points we can see that values of electric resistance of cambial tissue have similar daily dynamic, however, the values of resistance measured in northern direction are higher than in three other direc-

Fig. 2 Daily course of air humidity and air temperature, measured in height of 34, 29, 6 m a bove the forest floor.

246

Fig. 3. Daily dynamic of CO2 uptake (AN) simulated for upper (32 m), middle (29 m) and lower (19 m) part and measured in upper (32 m) part of beech crown.

tions. The increased resistance of tissues in the north side has been significantly expressed also in average values. The difference between average values of electric resistance of cambial tissue calculated from three and four measured values can be seen in the Figure 4. The highest dependence between electric resistance of cambial tissue and air temperature was discovered. Biofield of beech – the daily dynamic of biofield’s value has not been expressed. The following values of biofield: 56 cm, 47 cm, 38 cm, 29 cm, 20 cm and 11 cm from the stem of the tree were recorded. On the basis of measured values we can determine a regular 9-centimetre interval, it means a wave course of biophysical component of observed beech’s biofield.

Discussion The daily dynamic of physiological processes can, according to Schulze and Hall (1982), provide the basic information on responses and adaptation of plants to the nature conditions of the environment. At the same

time it is a reflection of effect of outside factors on individual physiological processes, as well as it can provide a great number of information about behaviour of tree species in their natural environment. Kozlowski et al. (1991) present that photosynthetic rate frequently varies between the tree species and their provenances, between the sun and shade type of leaves, during the day, as well as during the growing season. These variations are the result of interactions between the characteristics of plants such as leaf age, its structure and position, development of canopy, behaviour of stomata, amount and activity of Rubisco, as well as the factors of the environment such as light intensity, temperature, water supply, atmospheric CO2 concentration, air pollutants and soil conditions. Larcher (2003) presents that the gas exchange rate, investigated directly under the conditions of the forest stand, is aresult of mutual effect of number of internal factors and factors of the environment of which the specific effect can be recognized in avery difficult way. From the whole group of factors, one of them usually limits photosynthetic rate, while others support it further. We can see from the daily dy-

247

10 9.8 9.6 9.4 9.2 9 8.8 8.6 8.4 8.2 8 7.8 7.6 7.4 7.2

Fig. 4. Daily course of electric resistance of cambial tissue measured from north, east west and south direction and average values from 3 direction (dotted) and 4 direction (hatched) (E,W,S, N).

namic of selected physiological processes a close connection between individual processes and meteorological characteristics. The daily dynamic of photosynthetic activity of beech leaves stated by simulation is decisively being determined by light conditions in the individual parts of tree crown. The reason is the fact that intensity of PhAR was considered in advance to be the main factor influencing the CO2 uptake. The influence of other factors has not been considered since the measurements of dependence of PhAR – AN were done at leaf 248

temperature 20 °C and relative atmospheric humidity 60%. In addition, we can see from the daily dynamic of meteorological characteristics that Ta and RH have not reached the values which influence more considerably the CO2 uptake in beech leaves (Schulze, 1970). As for the Ta, Schulze (1970) presented that max. values of AN in mature beech leaves were measured at air temperature between 18–20 °C. A favourable RH (80%), sufficient intensity of PhAR, as well as low ozone concentration can be considered a reason for the maximal daily values of AN in

upper part of crown in the morning. The consequent whole-day decrease of CO2 uptake, as well as the occurrence of considerable depression of photosynthesis between 11 a.m. and 3 p.m. could be caused by a high intensity of PhAR, reduction of RH, increase of leaf temperature (Čaboun et al., 1996), and by an increase of ozone concentration during this part of the day. For instance Masarovičová and Štefančík (1990), Xu and Shen (1996) present the high intensity of PhAR, low RH and high Ta as causes of midday depression of photosynthesis. The most well–known dependence of values of cambial tree tissue’s electric resistance, frequently denoted as relative vitality, is a dependence on diameter of measured tree. The dependence of values of cambial tissue’s electric resistance on their diameter, was found out in observing the daily or seasonal dynamic of electric resistance within all cases. On the basis of our current and previous research results (Čaboun et al., 1993; Čaboun, 1994, 1997) we can state that the less diameter tree species has, the higher annual variability of values of cambial tissue’ electric resistance is. Thus, tree species with lower diameter react more sensitively on the environment’s influences. Similarly, daily dynamic in trees with bigger diameter is not so marked than in trees with low diameter. Considerably it has been expressed in all cases and had an opposite course than atmospheric temperature. We have found out a considerable correlation in all our measurements between temperature and electric resistance of cambial tissue. The closest correlation has been found out between electric resistance of cambial tree tissue and average maximal temperature calculated from maximal temperatures of three days before measuring the resistance. From the above mentioned follows that the resistance of cambial tissue is influenced more by three-day, mainly maximal temperature, resp. by the weather (where temperature plays a dominant role especially in beech) than moment temperature during measurement. On the basis of previous results we can state that since the yearly or seasonal dynamic of electric resistance is considerably higher than daily dynamic, the date of measurement is essentially more important than day’s time. We have found out, within our long-term measurements in a great sample of tree species, very close correlation between a diameter of trees d1,3 and biofield (Čaboun, 1993; Čaboun et al., 1996). We have also found out a close correlation between biofield and electric resistance since it correlates very closely with diameter of measured tree species as well. In spite of certain variability of a magnitude of tree species biofield, considerable trend of biofield change during the year has not been expressed. From the measurements we can see that correlation between the magnitude of biofield and temperature of atmosphere does not exist as well. The differences in magnitude of biofield during the year and also during the day, however, are not so high

that we could speak about the seasonal or daily biofield dynamic but only about average values of biofield individual levels. However, besides diameter and kind of tree species also site conditions and intraecosystem relations have an influence on a magnitude of tree species biofield. On the basis of our previous observations we can state that biofield of trees is not the same for each measuring man. When we proceed with the definition of allelopathy (Čaboun, 1990); each organism can react differently on biochemical and biophysical effect of other organism.

Acknowledgements This work was supported by the Slovak Research and Development Agency under the contracts No. APVV0608-10 and No. APVV-0111-10.

References Čaboun, V. 1990. Alelopatia v lesných ekosystémoch [Alelopathy in forest ecosystems]. Bratislava: Veda, SAV. 120 s. Čaboun, V. 1993. Biopole lesných drevín [Biofield of forest tree species]. Lesn. Čas. – For. J., 39: 415–425. Čaboun, V. 1994. Sledovanie relatívnej vitality drevín elektrickou odporovou metódou [Research of tree species relative vitality by electric resistence method]. Acta Fac. Ecol. Zvolen, 1: 53–75. Čaboun, V. 1997. Relative vitality of forest trees on research area Hukavský grúň in Biosphere reserve Poľana. Ekológia (Bratislava), 6: 33–47. Čaboun, V., Minďáš, J., Priwitzer, T., Štrba, S., Hladká, D., Šablatúrová, E., Tužinský, L, Škvarenina, J., Kukla, J., Meszároš, I., Molnár, Ľ., Zaušková, J., Konôpka, M., Ferjenčík, L., Slivková, E. 1996. Výsledky ekologického a ekofyziologického výskumu lesných ekosystémov na Výskumno – demonštračnom objekte Poľana Hukavský grúň [Results of ecological and ecophysiological research of forest ecosystems in the research and demonstration object Poľana – Hukavský grúň]. Lesnícke informácie, č. 1. Zvolen: Lesnícky výskumný ústav. 82 p. Kozlowski, T.T., Kramer, P.J., Pallardy, S.G. 1991. The physiological ecology of woody plants. New York: Academic Press. 657 p. Larcher, W. 2003. Physiological plant ecology. Berlin, Heidelberg: Springer-Verlag. 513 p. Marek, M., Masarovičová, E., Kratochvílová, I., Eliáš, P., Janouš, D. 1989. Stand microclimate and physiological activity of tree leaves in oak-hornbeamforest. II. Leaf photosyntetic activity. Trees, 4: 243–240. 249

Marek, M., Pirochtová, M., Marková, I. 1992. Production activity of mountain cultivated Norway spruce stands under the impact of air pollution. II. Vertical distribution of photosynthetic activity in the stand canopy. Ekológia (Bratislava), 11: 121–132. Masarovičová, E., Štefančík, L. 1990. Some ecophysiological features in sun and shade leaves of tall beech trees. Biol. Plant., 32: 374–387. Pirochtová, M., Marek, M. 1991. Metoda matematického vyhodnocování fotosyntetické aktivity lesních dřevin [Evaluation of photosynthetic activity of forest trees by mathematical method]. Lesnictví, 37: 399–408. Priwitzer, T. 1993. Prenosný fotosyntetický systém Li 6200 a jeho využitie v lesníckom ekologickom výskume [Portable photosynthetic system Li 6200 – using for forest ecological research]. In Kamenský, M. (ed.). Lesníctvo a výskum v meniacich sa ekologických a ekonomických podmienkach v Slovenskej republike. Zborník z jubilejnej konferencie LVÚ. 2. sekcia. Zvolen: Lesnícky výskumný ústav, p. 216–221. Priwitzer, T. 1998. Denná dynamikavýmeny CO2 aH2O slnných listov buka lesného (fa*gus sylvatica L.) pri rôznych meteorologických situáciách [Daily dynamics of CO2 and H2O exchange in sun beech leaves during different meteorological conditions]. In Škvarenina, J., Minďáš, J., Střelcová, K. Atmos-

férická depozícia a ekofyziologické procesy v ekosystémoch. Zborník referátov z medzinárodného pracovného seminára, Poľana 12.–13. jún 1996. Zvolen: Technická univerzita, p.201–208. Priwitzer, T., Šablatúrová, E., Hladká, D. 1996. Vybrané fyziologické a biochemické charakteristiky asimilačného aparátu buka [Selected physiological and biochemical characteristics of beech foliage]. Lesn. Čas. – For. J., 42: 371–380. Rovňáková, A. 1986. Fotosynteticky aktívne žiarenie [Photosyntetically active radiation]. Diploma work. Bratislava: Comenius University in Bratislava, Faculty of Mathematics and Physics. 57 s. Schulze, E.D. 1970. Der CO2-Gaswechsel der Buche (fa*gus sylvatica L.) in Abhängigkeit von den Klimafaktoren im Freiland. Flora, 159: 177–232. Schulze, E.D., Hall, A.E. 1982. Stomatal responses water loss and CO2-asimilation rate of plants in contrasting environments. In Lange, O.L., Nobel, P.S., Osmond, C.B., Ziegler, H. (eds). Physiological plant ecology II. Encyclopedia of plant physiology, new series, vol. 12B. Berlin, Heidelberg, New York: Springer-Verlag, p. 181–230. Xu, D.Q., Shen, Y.K. 1996. Midday depression of photosynthesis. In Pessarakli, M., (ed.). Handbook of photosynthesis. New York: Marcel Dekker, p. 451–459.

Received December 6, 2012 Accepted April 8, 2013

250

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Somatic embryogenesis: method for vegetative reproduction of conifers Terézia Salaj, Lenka Fráterová, Martin Cárach, Ján Salaj Institute of Plant Genetics and Biotechnology, Slovak Academy of Sciences, Akademická 2, P. O. Box 39 A, 950 07 Nitra, Slovak Republic, e-mail: [emailprotected], [emailprotected], [emailprotected], [emailprotected]

Abstract Salaj, T., Fráterová, L, Cárach, M., Salaj, J. 2013. Somatic embryogenesis: method for vegetative reproduction of conifers. Folia oecol., 40: 251–255. For Pinus nigra Arn. somatic embryogenesis has been initiated from immature zygotic embryos enclosed in megagametophytes. The initiated embryogenic tissues contain bipolar structures – somatic embryos consisted of meristematic embryonal part and long vacuolised suspensor cells. The embryogenic tissues/ cultures are usually maintained on solid or liquid nutrient media. For long-term storage, recently the method of cryopreservation has been used to replace the time consuming regular transfers to nutrient media. The initiated cell lines represent individual genotypes and the structure of somatic embryos as well as their maturation is cell line dependent. The maturation of early somatic embryos occurs on media containing abscisic acid and osmotica. The process of somatic embryogenesis is completed by plantlet (somatic seedlings) regeneration. Keywords cryopreservation, in vitro, micropropagation, pine, somatic embryogenesis

Introduction Somatic embryogenesis refers to the process in which somatic or non-sexual cells are induced to form bipolar embryos through aseries of developmental steps similar in those occurring during in vivo embryogenesis (Stasolla et al., 2002). The initiated bipolar structures are capable of development producing cotyledonary stage somatic embryos that in appropriate conditions germinate and their development is completed by whole plants (somatic seedlings) regeneration. Owing to the fact through the developmental process of somatic embryogenesis large number of plants can be obtained in relatively short period of time, the method became an attractive tool for clonal propagation (Klimaszewska and Cyr, 2002). For conifers somatic embryogenesis has been initiated for different species belonging to genera Picea, Pinus, Abies, Pseudotsuga and plantlet (somatic seedlings) regeneration has been achieved, but some prob-

lems still remain. For Pinus species the relatively low initiation frequencies represent serious problem and recently efforts have been made to optimize the initiation process. For Pinus radiata Hargreaves et al. (2009) obtained on average 55% initiation rates although the initiation was depending on families and collection time of explants. The initiation process can be enhanced through seed family screening, zygotic embryo staging as well as media adjustment (Montalbán et al., 2012). The development of early bipolar somatic embryos present in embryogenic tissues can be stimulated by using abscisic acid (ABA) combined with non-penetrating osmoticum as polyethylene glycol (Svobodová et al., 1999; Vooková and Kormuťák, 2009) or carbohydrates maltose and sucrose (Salajová et al., 1999; Lelu-Walter et al., 2008). The advantage of embryogenic tissues is their ability to regenerate after cryopreservation – storage in liquid nitrogen at –196 °C. In our laboratory, somatic embryogenesis for Pinus nigra Arn. has been repeatedly initiated as well

251

as plantlets/somatic seedlings regeneration has been achieved. The aim of presented paper is to evaluate somatic embryogenesis for the mentioned species.

Material and methods Plant material In our experiments megagametophytes containing immature embryos have been used as explants. The green cones of Pinus nigra Arn. have been collected at the beginning of June (usually between 1 and 15). The cones were stored at 4 °C for several days, after washed in tap water and the immature seeds were dissected. Surface sterilization of seeds was done by 10% H2O2 for 10 min., and then four washings in sterile distilled water followed. Finally, the megagametophytes were excised and placed on the culture medium. Culture media For the initiation as well as maturation of somatic embryos in most of cases DCR medium (Gupta and Durzan, 1985) has been used. Other media as LV (Litvay et al., 1981) or LP (Quoirin and Lepoivre, 1977) were also tested. The media were supplemented with enzymatic caseinhydrolysate (500 mg l–1), glutamine (50 mg l–1) as well as myo-inositol (200 mg l–1) and solidified with 0.3% gelrite (duch*eva). Plant growth regulators as 2,4-dichlorophenoxyacetic acid (2,4-D, 2 mg l–1) and 6-benzyladenine (BA, 0.5 mg l–1) have been incorporated into the nutrient media as well. Sucrose (2%) was used as carbohydrate source. Maturation occurred on DCR medium containing abscisic acid (25 mg l–1) and 6–9% maltose or high concentration of Phytagel (1%). After appearing of cotyledonary somatic embryos the tissues were transferred to media without ABA. The germination medium contained activated charcoal (1%). The cultivation of explants occurred in dark at 23 °C (except the culture of regenerated somatic seedlings that were transferred to light (110 µM s–1/day). Cryopreservation of embryogenic tissues For cryopreservation of embryogenic tissues the slowfreezing method has been used. In the experiments altogether 46 cell lines were included. On the 8th day of growth cycle 3.0 g of tissues was re-suspended in 9 ml of DCR medium containing 180 g l–1 sucrose. After 1 hour incubation in this medium gradually 15% DMSO was added to reach the final concentration 7.5%. Subsequently 1.8 ml of suspension was pipetted into cryovials and placed into the Mr. Frosty container. The samples were incubated in deep freezer (–80 °C) until the temperature in controlled cryovial reached –40 °C. Finally,

252

the cryovials were plunged into liquid nitrogen and kept there for 1 hour to 1 year. Thawing of tissues occurred at 40 °C in water bath. Following, the tissues were cultured on DCR medium as mentioned above in dark at 23 °C. Pretreated but not cryopreserved tissues were considered as control 1 (C1). Visual observations have been done in 3–4 days intervals. Growth analysis of tissues occurred three months after cryopreservation. Microscopic observations The structure of somatic embryos was investigated by light microscopy (Axioplan 2, Zeiss) using squash preparations and 2% acetocarmine staining.

Results and discussion Initiation of embryogenic tissues The production of embryogenic tissues has been observed approximately 3 to 5 weeks after placing the explants to the culture medium. The tissues were protruded from micropylar end of the megagametophyte (Fig. 1a), and reaching the size about 5 mm in diameter, they were separated from the explants and cultured individually as cell lines. The embryogenic tissues are of white color, translucent and relatively rapidly growing (Fig. 1b). Microscopic observations revealed the presence of bipolar structures – somatic embryos as the most important components of the tissues (Fig. 1c). The initiation frequencies reached values from 1.53% to 24.11% and changed from year to year. Plant growth regulation treatment as well as basal media formulations affected the initiation process. The highest initiation frequencies were obtained on medium DCR. The most important factor as we experienced is the developmental stage of zygotic embryos used as starting explants. For Pinus nigra the very early developmental stages – immature precotyledonary embryos gave the best initiation frequencies. As the maturation of zygotic embryos progressed the initiation frequencies dropped and finally the explants failed to produce embryogenic tissues. Although the bipolar structures are present in embryogenic tissues their structural organization is changing in dependence on the cell line. Relatively low initiation frequencies are characteristic for Pinus species and many attempts were done in order to achieve improvement. For Pinus nigra the developmental stage of original zygotic embryos is decisive and this phenomenon was also confirmed for Pinus radiata (Hargreaves et al., 2009; Montalbán et al., 2012), Pinus pinaster (Miguel et al., 2004), Pinus sylvestris (KeinonenMettälä et al., 1996).

Fig. 1a. Initiation of embryogenic tissues (ET) on megagametophyte explants (MG).

maturation medium numerous precotyledonary somatic embryos appeared on the surface of embryogenic tissues. The suspensor was still present and connected the developing embryos to the tissue. Cotyledonary somatic embryos appeared around the eighth week of maturation (Fig. 1d). Their quantity was strongly cell line dependent. For germination somatic embryos at least with four cotyledons were selected. The germination occurred on ABA free medium and resulted in somatic seedling formation (Fig. 1e).The obtained plantlets were transferred to soil and survived five to six months. During somatic embryo development (maturation) structural changes were visible in the developing somatic embryos. The most conspicuous features were differentiation of root meristem and later the procambium formation. The maturation of conifer somatic embryos is genotype dependent (Keinonen- Mettälä et al., 1996; Salajová et. al., 1999) and is influenced by composition of the maturation medium (Carneros et al., 2009; Montalbán et al., 2010).

Fig. 1b. Proliferating embryogenic tissue 8 days after transfer to fresh medium.

Fig. 1d. Cotyledonary somatic embryos (arrows) developed on the maturation medium.

Fig. 1c. Bipolar somatic embryos composed of meristematic embryonal cells (E) and long vacuolised suspensor (S).

Maturation of somatic embryos The early bipolar structures are capable of development and plantlets (somatic seedlings) production. Transfer of tissues from proliferation medium containing 2, 4-D and BA to maturation medium with ABA and maltose resulted in the development of early somatic embryos. Approximately around the fifth week of culture on

Fig. 1e. Plantlets (somatic seedlings) regenerated from somatic embryos.

253

Cryopreservation of embryogenic tissues Cryopreservation of embryogenic tissues enables their storage for long-term period. Out of 46 cell lines cryopreserved 35 survived the storage in liquid nitrogen. The regeneration of tissues after thawing (Fig. 1f) was dependent on cell line although the duration of storage in liquid nitrogen had no significant effect on the growth and behaviour of tissues. Our examinations also showed no correlation exists between the maturation ability and cryotolerance of cell lines. The use of cryopreservation method for long-term storage of conifer embryogenic tissues has been demonstrated for several species (Aronen et al., 1999; Vondráková et al., 2010).

Fig. 1f. Re-growth of embryogenic tissue (ET) after cryopreservation.

The obtained results for Pinus nigra give evidence that it is possible to initiate embryogenic cell lines from immature zygotic embryos, but to obtain higher initiation frequencies the method/approach needs refinement. Although the maturation of somatic embryos is cell line dependent, cotyledonary somatic embryos were produced and somatic seedling regeneration occurred as well. Cryopreservation of embryogenic tissues using slow-freezing was also successful for the majority of tested cell lines.

Acknowledgement The study was supported by the Slovak Grant Agency VEGA, proj. No. 2/0144/11.

References Aronen, T.S., Krajnaková, J., Häggman, H., Ryynänen, L.A. 1999. Genetic fidelity of cryopreserved em254

bryogenic cultures of open pollinated Abies cephalonica. Pl. Sci., 142: 163–172. Carneros, E., Celestino, C., Klimaszewska, K. 2009. Plant regeneration in stone pine (Pinus pinea L.) by somatic embryogenesis. Pl. Cell Tiss. Org. Cult., 98: 165–178. Gupta, P.K., Durzan, D.J. 1985. Shoot multiplication from mature trees of Douglas fir (Pseudotsuga menziesii) and sugar pine (Pinus lambertiana). Pl. Cell Rep., 4: 177–179. Hargreaves, C.L., Reeves, C. B., Find, J.I., Gough, K., Josekutty, P., Skudder, D., Van Der Maas, S.A., Sigley, M.R., Menzies, M.I., Low, C.B., Mullin, T.J. 2009. Improving initiation, genotype capture, and family representation in somatic embryogenesis of Pinus radiata by acombination of zygotic embryo maturity, media and explant preparation. Can. J. Forest Res., 39: 1566–1574. Keinonen-Mettälä, K., Jalonen, P., Eurola, P., von Arnold, S. 1996. Somatic embryogenesis of Pinus sylvestris. Scand. J. Forest Res., 11: 242–250. Klimaszewska, K., Cyr, D.R. 2002. Conifer somatic embryogenesis: I. Development. Dendrobiology, 48: 31–39. Lelu-Walter, M.A., Bernier-Cardou, M., Klimaszewska, K. 2008. Clonal production from self- and cross pollinated seed families of Pinus sylvestris (L.) through somatic embryogenesis. Pl. Cell Tiss. Org. Cult., 92: 31–45. Litvay, L.D., Johnson, M.A., Verma, D., Einspahr, D., Weyrauch, K. 1981. Conifer suspension culture medium development using analytical data from developing seeds. IPC Techn. Pap. Ser. Inst. Pap. Chem. (Appleton, WI), 115: 1–17. Miguel, C., Concalves, S., Tereso, M., Maroco, J., Oliveira, M. 2004. Somatic embryogenesis from 20 open-pollinated families of Portuguese plus trees of maritime pine. Pl. Cell Tiss. Org. Cult., 76: 121–130. Montalbán, I.A., De Diego, N., Moncaleán, P. 2010. Bottlenecks in Pinus radiata somatic embryogenesis: improving maturation and germination. Trees, 24: 1061–1071. Montálban, I. A., De Diego, N., Moncaleán, P. 2012. Enhancing initiation and proliferation in radiata pine (Pinus radiata D. Don) somatic embryogenesis through seed family screening, zygotic embryo staging and media adjustment. Acta Physiol. Plant., 34: 451–460. Quoirin, M., Lepoivre, P. 1977. Études des milieux adaptés aux cultures in vitro de Prunus. Acta Hort., 78: 437–442. Salajova, T., Salaj, J., Kormutak, A. Initiation of embryogenic tissues and plantlet regeneration from somatic embryos of Pinus nigra Arn. Pl. Sci., 145: 33-40. Stasolla, C., Kong, L., Yeung, E.C., Thorpe, T.A. 2002. Maturation of somatic embryos in conifers:

morphogenesis, physiology, biochemistry, and molecular biology. In Vitro Cell. Dev. Biol. – Pl., 38: 93–105. Svobodová, H., Albrechtová, J., ku*mstýrovÁ, H., Lipavská, H., Vágner, M., Vondráková, Z. 1999 Somatic embryogenesis in Norway spruce: anatomical study of embryo development and influence of polyethylene glycol on maturation process. Pl. Physiol. Biochem., 37: 209–221.

Vookova, B., Kormuťak, A. 2009. The improved plantlet regeneration from open-pollinated families of trees of Dobroc primeval forest and adjoining managed stand via somatic embryogenesis. Biologia, Bratislava, 64: 1136–1140. Vondráková, Z., Cvikrová, M., Eliášová, K., Martincová, O., Vágner, M. 2010. Cryotolerance in Norway spruce and its association with growth rates, anatomical features and polyamines of embryogenic cultures. Tree Physiol., 30: 1335–1348.

Received December 6, 2012 Accepted April 15, 2013

255

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Selection and breeding of stress-tolerant woody ornamentals for urban plantings

Gábor Schmidt Department of Floriculture and Dendrology, Faculty of Horticultural Science, Corvinus University of Budapest, 1118 Budapest, Villányi út 29-43, Hungary e-mail: [emailprotected]

Abstract Schmidt, G. 2013. Selection and breeding of stress-tolerant woody ornamentals for urban plantings. Folia oecol., 40: 256–260. Because of this geographic position, climate and soils, Hungary lends itself for selection of woody plants which tolerate environmental stresses. Selection and breeding of the woody ornamentals for extreme urban conditions started in the early 1950s at the former University of Horticulture and Food Industry (at present: the Faculty of Horticulture of Corvinus University), Budapest. The first results were 8 Sorbus, 3 Tilia and 2 other cultivars, and selected clones from Fraxinus, Cornus, Juniperus and others. In the recent 20 years, many new hardy cultivars and named clones are brought up, the most important of which are as follows: Ailanthus altissima (Mill.) Swingle cv. Purple Dragon, Acer campestre L. cv. Zentai Upright`, Celtis occidentalis L. cv. Straight Stem, Crataegus pinnatifida Bunge cv. Tahi, Hedera helix L. 9 cultivars, Platanus × hispanica Münchh. cv. Budapest, Prunus padus L. cv. Aurora, Prunus × davidopersica cv. Rubin (P. L. cv. Piroschka), Prunus tenella Batsch. cv. Pink Carpet, Pyrus nivalis Jascq. cv. Kartália, Salix matsudana Koidz. cv. Golden Spiral, Syringa josikaea J. Jacq. ex Rchb. cv. Smaragd, Tilia tomentosa Moench. cv. Zenta Silver, Tilia × euchlora K. Koch. cv. Saint Stephan. Keywords Ailanthus, new cultivars, Prunus, stress-tolerance, Tilia, urban trees

Introduction Because of this geographic position, Hungary lends itself for selection of woody plants which tolerate environmental stresses: the summer is warm with temperatures reaching a maximum of 30–35 oC, and the winter is cold and irregular with temperatures falling sometimes –26–30 oC. These extremities are multiplied by the poor sandy and salinity soils of the Great Hungarian Plains, the dry limestones and dolomites of low hills and by the dry warm and polluted atmosphere of cities and towns. Selection of woody ornamentals for such conditions started in the early 1950s by the Department of Horticulture and Dendrology of the former University of Horticulture and Food Industry (at present: the Faculty of Horticulture of Corvinus University, Budapest). The first results were 8 Sorbus, 3 Tilia and 2 other

256

cultivars, and selected clones from Fraxinus, Cornus, Juniperus and others (Sipos, 1964; Read and Schmidt, 1987). In the recent 20 years, many new hardy cultivars and named clones are brought up (Lukács et al., 2009; Orlóci et al., 2009; Schmidt and Sütöriné, 2011). The aim of the present paper is to give a short description of (and experiences with) the new selections of this period.

Material and methods The breeding work was carried out in two basic ways: 1. on-site selection of adult specimens in urban or roadside plantings; 2. selection of young seedlings from mass-propagation in the nursery.

In the case of on-site selection of adults specimens in urban or roadside plantings, expeditions were organized to the larger cities and towns for the search of such adult specimens which in the given urban environment showed higher tolerance towards urban injuries than the other ones (their neighbours standing in the same alley or group) and also showed some increased ornamental or technological value, like better crown shape, shiny and healthy or colourful leaves, good structure of branch system, etc. In the case of selection of young seedlings from mass-propagation in the nursery, this sort of breeding started with mass-propagation by seed of such species or individuals which were known as genetically stresstolerant in Hungary (and having sometimes also additional values like red leaves, or disease – resistance etc.) Later from the seedling lot (minimum 400 liners, but usually much more) those were picked out, which showed better qualities than the other ones. In both cases, at the first step minimum 10–12 specimens were selected (picked out) and propagated vegetatively (by budding or by cuttings) in an amount of minimum 50 cuttings or grafts per clone. The second step was the nursery trial of the clones (speed of growth, healthiness of leaves, straightness of stem of trees or bushiness of shrubs) and, of course, repeated bonitations were made on their ornamental value. The third step was to plant and try them out in urban conditions, usually in Budapest. Finally, those clones which proved to be the best both in the nursery and in urban plantings were propagated again in the nursery. They were given cultivar names and submitted to official cultivar – recognition to the respective Institution and Community for testing and approval (see later at chapter New cultivars for urban planting.)

aging. The dark purple winged fruits are born in abundant clusters and retain their intensive colour from July through August and early September. Foliage is shiny green with red petioles and leaflet nerves, the shoots are purplish brown. (Breeder: G. Schmidt, 1996.) Celtis occidentalis L. cv. Straight Stem. The Hackberry Tree Celtis occidentalis L. is perhaps the hardiest urban tree in Hungary, which equally tolerates the poor urban soils, polluted atmosphere and also the negative effects of human vandalism like heavy injuries of the trunk, cutting the branches and the roots, etc. The only (but great) disadvantage of the traditionally used type is the irregular growth (curved trunk) and the overhanging branches which create problems both in the nursery and in the street plantings. The new cultivar Straight Stem has the high tolerance of the traditional species without its disadvantage in habit and growth: The stem of this cultivar is growing straight (and fast) in the nursery so (in contrast to the “traditional” type) it does not need staking (Fig. 1); later (on the final place) along urban streets. The crown becomes upright oval. The branches are not overganging at all, so they do not disturb the traffic. (Breeder: G. Schmidt 2006.)

Results Description of the recent cultivars and selections New cultivars in Hungary are inspected and tried for several years by the National Institute for Agricultural Quality Control. If they meet the necessary criteria, including the DUS-requirements, an official recognition and certificate is awarded by the Hungarian Cultivar Qualification Council (HCQC). In the recent 20 years, many new hardy cultivars and named clones are brought up and recognized by HCQC, the most important of which are as follows: Ailanthus altissima (Mill.) Swingle cv. Purple Dragon. The heaven tree, Ailanthus altissima (Mill.) Swingle, tolerates drought and bad soils extremely well in Hungary and grows like weed in polluted urban environment. The cultivar Purple Dragon is a female form found in Budapest. It has a straight leader, fast growth and a regular rounded crown becoming flattened with

Fig. 1. Celtis occidentalis L. cv. Straight Stem.

Hedera helix L. Hungarian cultivars. The English Ivy Hedera helix L. is native to practically all woodlands in Hungary (Bényei-Himmer, 1994a). It is a multifunctional plant in landscaping. The juvenile form

257

is suitable for a groundcover (both in shadow and on the sunshine), for covering walls of the buildings and fences and climbs easily on pergolas or on trunk of trees as well. The adult form makes a fine and hardy rounded bush, or, if pruned, can be planted as a low evergreen semi-low hedge. The cultivars of the Faculty of Horticultural Sciences originate from different parts of Hungary and fall into two groups: 1. the spreading climbing (juvenile) cultivars, and 2. the bushy (adult) ivies. They were bred by Mrs Bényei-Himmer M. at the Department of Botany (Bényei-Himmer, 1994b). Spreading – climbing (juvenile) cultivars are: H. helix L. cv. Börzsöny. A fast spreading cultivar, with thick growth. Leaves are elongated triangular in form (f. sagittifolia), the leaf blade leader them, dark green with well marked nerves (Fig. 2). Makes an excellent groundcover. (Breeder: Bényei-Himmer M. 2000).

Fig. 3. Hedera helix L. cv. Krokó.

Fig. 2. Hedera helix L. cv. Börzsöny.

H. helix L. cv. Zebegény. Middle-strong growth, very good tendency for branching. Leaves are markedly five-lobed (f. pedata) with vivid green colour and silvery leaf-nerves. Suitable as a ground-cover (especially in small gardens) or for balcony-boxes. (Breeder: Bényei-Himmer M. 2000). H. helix L. cv. Krokó. A slow to medium-strong growing spreading form, with slightly lobed and widely silvery-nerved leaves which give the plant a spectacular (“crocodile-like” appearance, Fig. 3). Excellent as a grand cover for small gardens or in balcony-boxes. (Breeder: Bényei-Himmer M., G. Botlik 2004). H. helix L. cv. Negro. A medium-strong growing spreading form, whose leaves are very dark green (almost black). An interesting ground cover, makes a good contrast if planted in one group with silvery or with golden-coloured cultivars. (Breeder: Bényei-Himmer M. 2004). 258

Bushy (adult) cultivars are: H. helix L. cv. Soroksár. A wide-rounded low or medium-sized bush, with vivid green shiny leaves. Black fruits from February till the end of March. (Breeder: Bényei-Himmer M. 2000). H. helix L. cv. Blue Star. A medium-sized bush, with shape and size like that of the former cultivar, but the fruits are shiny blue, appearing in abundant loose cymes. (Breeder: Bényei-Himmer M. 2000). H. helix L. cv. Marble. An upright bush of smaller (later medium) size. Dark green leaves with undulate margins and marked light-green veins are giving a marbled effect. It has abundance of yellowish flowers in September–October and black fruits (in compact cymes) during winter. (Breeder: Bényei-Himmer M. 2000). H. helix L. cv. Csocsoszan. A rounded bush whose leaves are not lanceolate but wide obovate with crenate leaf-margins resembling a Japanese fan. Not so winter-hardy as the former cultivars. (Breeder: BényeiHimmer M. 2004). Prunus × davidiopersica cv. Rubin (P. × d. cv. Piroschka). A small tree with flattened crown. Ultimate height is 4–5 m, diameter 6–8 m. Leaves are dark rubyred during the intensive shoot growth and are turning dull greenish red when the growth stops. This change of

colour (with new and new flushes of growth) is repeating 2–3 times in one vegetation. Large white flowers with a small pink eye bloom in late March–early April immediately after bud-brake. The cultivars are resistant to mildew and to Taphrina deformans (Berk.) Tul. Tolerant to drought and early frost. (Breeders: G. Schmidt and F. Incze. 1996). Prunus padus L. cv. Aurora. The “Bird Cherry” Prunus padus L. is a medium-sized bushy tree widely distributed on the Northern Hemisphere including Hungary. The first red-leaved Prunus padus cultivar Coloratus was introduced to Hungary some 30 years ago. This cultivar did not distribute in Hungary because of its poor tolerance to continental climate and the limy soils (the cultivar was selected in Sweden, under the humid climate of Scandinavia). The Hungarian cultivar Aurora is developing leaves which are much darker red under our conditions, than those of cv. Coloratus and, in contrast to the mentioned Scandinavian cultivar do not burn neither become necrotic (yellow) in the hot summer. It brings abundance of dark pink flowers (Fig. 4) blooming in upright panicles during mid- or late April. (Breeder: G. Schmidt, 2005, further selected clones still in process of trials).

young. Later the side-shoots turn upright and are only slightly thorny. Flowers are white, blooms 2 weeks later than Pyrus communis L. The fruit is a 3 cm wide, yellowish green pear. No pests. Suitable in parks, small streets. Drought tolerant. (Breeders: I. Tóth, and A. Terpó 1995). Salix matsudana Koidz. cv. Golden Spiral. A fast growing corkscrew-willow, probably a spontaneous hybrid between S. matsudana Koidz. ‘Tortuosa’ and S. alba L. ‘Tristis’. It was found as a chance seedling near Velencei lake. Shoots, twigs and branches are much contorted, light yellow in the summer turning rich golden orange in the winter (Fig. 5) (Breeder: G. Schmidt. 1993).

Fig. 5. Salix matsudana Koidz. cv. Golden Spiral.

Fig. 4. Prunus padus L. cv. Aurora.

Pyrus nivalis Jacq. cv. Kartália. A 4–6 m high, slow growing small tree with wide columnar form. Side shoots are short, squat and grow horizontally when

Syringa josikaea J. Jacq. ex Rchb. cv. Smaragd. A 2–3 m high, strong growing, compact shrub with stiff, upright branches. Leaves are 6–10 cm wide, elliptic, dark emerald green, leathery through the whole summer. Dark lilac-coloured flowers, bloom in upright compact panicles, in mid- or late May (2 weeks after the common lilac (S. vulgaris L. cultivars). Cv. Smaragd is not susceptible to mites, and tolerates half-shade. Use: alone or in groups in parks and in home gardens. (Breeder: G. Schmidt. 1993). 259

Tilia tomentosa Moench. cv. Zenta Silver. A strongly upright-growing tree, making a straight leader in the nursery and reaching an ultimate height of 15–20 m in parks. The crown is wide columnar when young, becoming compact conical with aging. Branches are upright. Shoots are greyish green, downy. New leaves are light dull green above, strongly silvery tomentose beneath. Highly scented flowers bloom in late Juneearly July. Grows fast in the nursery and tolerates urban climate well. Suitable for streets and parks. (Breeders: B. Nagy and G. Schmidt. 1996). Tilia × euchlora K. Koch. cv. Saint Stephan. A strongly growing tree, making a straight stem and continuous leader in the nursery and reaching an ultimate height of 15–20 m. Crown is narrow-ovate, with pointed top (Fig. 6). Branches are upright. Shoots are brownish-green, glabrous. Leaves are leathery, shiny green above, dull green and glabrous beneath. Slightly fragrant flowers bloom in early July and almost no fruits later. Tolerates urban climate well. (Breeders: E. Jámbor-Benczúr, Z. Ifjú., I. Tóth and G. Schmidt 2000).

Fig. 6. Tilia × euchlora K. Koch. cv. Saint Stephan.

Conclusion In the recent 20 years 11 new urban trees and shrubs have been bred at the Department of Floriculture and Dendrology and 8 Hedera helix L. cultivars at the Department of Botany of the Faculty of Horticulture, Corvinus University, Budapest. All of them are available in the leading Hungarian tree nurseries, registered at the 260

Central Agricultural Office and also are kept records by the civil organization “Commission for Hungarian Ornamental Cultivars”. The home page of this organization (http://www.magyarfajtak.hu) contains a more detailed description of the above-listed cultivars too, illustrated by digital photos.

Acknowledgement The Author would like to express his thanks to the organizers of the excellent Conference in the Mlyňany Arboretum (Sept. 18–19., 2012) and also to Stephan Bakay for completing the Slovakian abstract of this paper.

References Bényei-Himmer, M. 1994a. A borostyán mint őshonos növény Magyarországon (Variability of Common Ivy as a native plant to Hungary). Dissertation for Candidate of Sciences degree. Budapest: University of Horticulture and Food Industry, Faculty of Horticulture, p. 14–16. Bényei-Himmer, M. 1994b. Hedera fajok és fajták Magyarországon (borostyánhatározó) [Identification ad description of Hedera species and cultivars in Hungary]. Kert. Élelm.-Ip. Egy. Közlem., 54: 60–70. Lukács, Z., Orlóci, L., Schmidt G., Csikor J., Honfi P., Sütöriné Diószegi, M. 2009. A magyar kertészeti dendrológiai nemesítés felmérése [A survey of Hungarian dendrological breeding]. In Hagyomány és haladás a növénynemesítésben. Budapest: MTA, Növénynemesítési Bizottsága, p. 307–311. Orlóci, L., Lukács, Z., Schmidt, G. 2009. Kontinentális klímán kipróbált, Magyarországon tesztelt díszfák, díszcserjék [Ornamental trees and shrubs tried on continental climate and tested in Hungary]. In Hagyomány és haladás a növénynemesítésben. Budapest: MTA, Növénynemesítési Bizottsága, p. 406–411. Read, P.E., Schmidt, G. 1987. Stress tolerant plants for urban landscape – the Nebraska-Hungary cooperative experience. Acta hort., 496: 401–407. Schmidt, G., Sütöriné Diószegi, M. 2011. Magyar nemesítésű díszfák-díszcserjék gyűjteményeinek fejlesztése a BCE Budai Arborétumában [Collections of Hungarian woody ornamental cultivars in the Buda Arboretum of Corvinus University of Budapest]. Kertgazdaság, 43 (4): 3–12. Sipos, E. 1964. A hazai Sorbus fajok értékelési metodikája [Evaluation of indigenous Sorbus species]. Publ. High School Hort. Viticult., 37 (1): 255–265.

Received December 6, 2012 Accepted April 8, 2013

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Influence of vegetation on microclimate in the urban environment Monika Strelková1, Zuzana Hečková1, Zdenka Rózová1, Anna Tirpáková1, Dagmar Markechová1 1

Department of Ecology and Environmental Science, Faculty of Natural Sciences, Constantine the Philosopher University in Nitra, Tr. A. Hlinku 1, 949 74 Nitra, Slovak Republic, e-mail: [emailprotected], [emailprotected], [emailprotected], [emailprotected], [emailprotected]

Abstract Strelková, M., Hečková, Z., Rózová, Z., Tirpáková, A., Markechová, D. 2013. Influence of vegetation on microclimate in the urban environment. Folia oecol., 40: 261–265. There are many factors in the urban environment influencing its microclimate conditions. Vegetation is one of the main components participating in this process. Our study compares microclimatic factors (the air temperature and the air humidity) of two sites with different ratio between built-up area and greenery. The measurements have been realized in the chosen areas of the built-up area of Nitra town in the spring months of 2012. The aim of the research was to compare the air temperature and the air humidity depending on the percentage of the vegetation cover in the urban environment of the built-up area of Nitra town. Keywords air humidity, air temperature, microclimate, vegetation

Introduction Greenery in the urban environment is irreplaceable. The cities are being characterized by the presence of different surfaces changing the microclimatic characteristics of the environment. The presence of different surfaces is particularly reflected in the influence of the air temperature and the relative air humidity conditions of the human environment. Greenery is a major component of the nature being involved in the treatment of this condition in the cities. It is characterized by several functions that make us a great deal in such disturbed environment as the urban settlements are. The microclimatic function of the greenery is the basic one. Microclimatic function consists of cooling of the urban environment by greenery during the warm months and avoiding of the large temperature fluctuations during the day and night. All the trees and bushes regulate the air humidity of the atmosphere. Their space capacity and biomass assimilation adapt the climate, air temperature, solar radiation and air flow (Supuka et al., 1991; Rózová and Mikulová, 2009).

The microclimatic function is being understood as the ability of greenery to influence the air humidity, shade provision, reduction of temperature fluctuations, etc., by its transpiration function. For example, the adult birch in growing season can evaporate up to 7,000 l of water, the urban parks reduce the air temperature by an average of 1 °C if compared to the temperature in the streets. The green areas increase the air humidity (the average value is 5 to 7 per cent) (Hudeková et al., 2007). The aim of this work was to evaluate the development of the microclimatic characteristics of the areas with different ratio of built-up area and greenery in the spring of the year 2012. The relative air humidity in % and the air temperature in °C, the main indicators of the microclimatic conditions, have been the measured characteristics.

Material and methods The research was carried out in the selected parts of Nitra town. There were selected two model areas: Site

261

Excel 2010 program and evaluated by two-factor analysis of variance by Statistical Program.

Results and discussion The measurements were realized in spring during the months of March, April and May in 2012. Change in the development of the air temperature was observed in the selected months in the monitored areas. We assumed that the air temperature would be higher in the areas with the higher ratio of the built-up area to the greenery. Such assumption was confirmed in March (Fig. 1). The air temperature during the monitored week was higher at Site 2 than at Site 1 with the higher ratio of the greenery coverage. If sub-sites were considered, the biggest differences in temperature were recorded at sub-site with the hard surface and the smallest differences were at the contact interface of the grass-vegetation. In April (Fig. 2) the air temperatures became the equal at both sites. A significant difference is visible at the sub-sites with the grass coverage. The minimal differences were observed at the sub-sites with the hard surface and the contact areas with grass and vegetation. The measurements in May (Fig. 3) showed differences in the air temperature values at the selected sites. The greatest difference in temperature was observed at the sub-site grass – vegetation. The higher temperatures in May were measured at Site 1, in March at Site 2. Our assumption that the air temperature is higher at site with

2 m from the building

Hard surfacegrass

Hard surface

Vegetationhard surface

Vegetation

Grassvegetation

Grass

Air temperature [°C]

1/Chrenová III. – parking place on Akademická Street, area with 0–50% of greenery and Site 2/Chrenová I. – place behind the Student Dormitory Nitra, area with 51–100% of greenery. The ratio of the built-up areas and the green areas on the same size surface has been the main criteria of the selection process. The squares of 50 × 50 m have been determined within the area. Chosen squares had to include parts with different types of cover (hard surface, grassland and vegetation section). There were created two categories, 0–50% and 51–100% of greenery of the whole area. The research was carried out during the spring months (March, April and May) in 2012. Anemometer with multifunctional climate probe, VELOCICALC ® (9565) – TSI, has been used for the measurements. The relative air humidity in % and the air temperature in °C were measured simultaneously. Data logger recorded data at weekly intervals (7 days), beginning on the second Monday of month at 7 a.m., 2 p.m. and 9 p.m. with the periodic repetition. The measurements have been realized in both localities in the middle of the selected areas with the vegetation, grass and hard surface and at the contact interfaces of these parts: grass – vegetation, vegetation – hard surface, hard surface – grass and at a distance of 2 meters from the building. The measurements have been performed at 2 m above the ground level. Twenty entries were recorded at the specific time in each area for the purpose of the statistic evaluation. Collected data have been processed to the tables in

Site 1 Site 2

Sub-site Fig. 1. Comparison of mean air temperatures measured during one week at sub-sites of sites 1 and 2 in March 2012. Site*Sub-site; LS Means. Current effect: F (6.280) = 0.01259, p = 0.99999. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

262

2 m from the building

Hard surfacegrass

Hard surface

Vegetationhard surface

Vegetation

Grassvegetation

Grass

Air temperature [°C]

Site 1 Site 2

Sub-site Fig. 2. Comparison of mean air temperatures measured during one week at sub-sites of sites 1 and 2 in April 2012. Site*Sub-site; LS Means. Current effect: F (6.266) = 0.01815, p = 0.99997. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

full foliage of trees occurs in May. The growing season of trees in Slovakia lasts from April 1 to September 30. The relative air humidity and the air temperature were measured at the same time and at the same places. The research was performed in spring of 2012. The higher values of the air humidity were measured in all

2 m from the building

Hard surfacegrass

Hard surface

Vegetationhard surface

Vegetation

Grassvegetation

Grass

Air temperature [°C]

the higher ratio of the built-up area to the countryside has not been confirmed. This situation might be caused by the consequence of the fact that the measurements were taken at the beginning of the growing season of the trees. The vegetation in our latitude is without leaves in March. The leaves start to grow during April and the

Site 1 Site 2

Sub-site Fig. 3. Comparison of mean air temperatures measured during one week at sub-sites of Sites 1 and 2 in May 2012. Site*Sub-site; LS Means. Current effect: F (6.280) = 0.00496, p = 1.0000. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

263

was the biggest difference in values of the air humidity between the monitored sites. The difference in the following month, in April, was smaller one. In May, the values of the humidity became equal at sites 1 and 2.

2 m from the building

Hard surfacegrass

Hard surface

Vegetation-hard surface

Vegetation

Grassvegetation

Grass

Air humidity [%]

months at Site 1, where the greenery dominates over the built-up area. Figures 4–6 show the changes in the air humidity at the sub-site with the hard surface. In March, there

Site 1 Site 2

Sub-site

2 m from the building

Hard surfacegrass

Hard surface

Vegetationhard surface

Vegetation

Grassvegetation

Grass

Air humidity [%]

Fig. 4. Comparison of values of mean relative air humidity measured during one week at sub-sites of Sites 1 and 2 in March 2012. Site*Sub-site; LS Means. Current effect: F (6.280) = 0.03431, p = 0.99983. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

Site 1 Site 2

Sub-site Fig. 5. Comparison of values of mean relative air humidity measured during one week at sub-sites of Sites 1 and 2 in April 2012. Site*Sub-site; LS Means. Current effect: F (6.266) = 0.00456, p = 1.0000. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

264

2 m from the building

Hard surfacegrass

Hard surface

Vegetationhard surface

Vegetation

Grassvegetation

Grass

Air humidity [%]

Site 1 Site 2

Sub-site Fig. 6. Comparison of values of mean relative air humidity measured during one week at sub-sites of Sites 1 and 2 in May 2012. Site*Sub-site; LS Means. Current effect: F (6.280) = 0.00243, p = 1.0000. Effect hypothesis decomposition. Vertical bars denote 0.95 confidence intervals.

This situation is caused by the equalizing of the temperature daily routine. The air temperature is higher and its fluctuations are reduced during the day. We did not take into consideration the precipitation when evaluating data and results. In the future, the precipitation regime of Nitra town needs to be considered in comparison with the microclimatic characteristics.

Acknowledgements This study is the result of the project implementation: Environmental Aspects of the Urbanized Environment, ITMS: 26220220110, supported by the Research & Development Operational Programme funded by the ERDF, scientific projects: Slovak Ministry of Education, project VEGA No. 1/0042/12 and Constantine the Philosopher University in Nitra, project UGA No. VII/35/2012.

References Hudeková, Z., Krajcsovics, L., Martin, P., Pauditšová, E., Reháčková, T., Hudek, V. (eds). 2007. Ekologická stopa, klimatické zmeny amestá [The ecological footprint, climatic changes and towns]. Bratislava: Regionálne environmentálne centrum. 52 p. Rózová, Z., Mikulová, E. 2009. Vegetačné úpravy vkrajine [Vegetation arrangement in the country]. Prírodovedec, č. 365. Nitra: Fakulta prírodných vied Univerzity Konštantína Filozofa vNitre. 155 p. Supuka, J., Benčať, F., Bublinec, E., Gáper, J., Hrubík, P., Juhásová, G., Maglocký, Š., Vreštiak, P., Králová, K. 1991. Ekologické princípy tvorby aochrany zelene [Ecological principles of creation and protection of greenery]. Bratislava: Veda. 308 p.

Received December 6, 2012 Accepted March 8, 2013 265

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Development, changes and assessment of tree alleys in town streets

Ján Supuka Department of Garden and Landscape Architecture, Faculty of Horticulture and Landscape Engineering, Slovak University of Agriculture in Nitra, Tulipánová street No. 7, 949 01 Nitra, Slovak Republic, e-mail: [emailprotected] Abstract Supuka, J. 2013. Development, changes and assessment of tree alleys in town streets. Folia oecol., 40: 266–271 Street tree alleys were started to plant as apart of great reconstruction of European and world cities after industrial revolution from the beginning of 19th century. Important participation at first street alleys plantation have had decoration associations organised at town’s of Slovakia territory on 2nd half of 19th century. Street tree alleys and river embankments have wide spectrum of functions useful for man and improving of environmental conditions. In a new terminology we are talking on services those should be classified as follow: supporting, provisioning, regulating and cultural services. Street trees have high value at designing of the pedestrian zones and new squares. In Nitra town conditions we have assessed 26 most important street tree alleys. For this purpose was elaborated a new methodical approach, where besides basic biometrical dates the following tree characteristics were assessed: potential of environmental adaptability, potential of biology-ecological value, potential of culture value, potential of disturbing and undesirable influences. Each group of characteristics was valuated in three following levels of significance (1) low, (2) medium, (3) high. There were assessed 22 woody plant species and cultivars mapped at Nitra town streets. Key words street trees, changes, Nitra town, service assessment

Introduction The street free alleys represent important component in vegetation structure of town settlements (Supuka et al., 2008). Tree alleys were in the past used as a component part of designed landscape, planted especially along with roads and as internal composition element of historical gardens and parks or prolongation element from those parks into open country. Street alleys as town design element were mainly established after industrial revolution during urban reconstruction of larger cities for instance Paris, where spacious street boulevards and alleys were constructed in time after beating up Bastila in 1779. Since that time establishment of new public parks has been dated (Kalusok, 2004; Kupka, 2010). Moreover, such reconstructions were realised at other European cities also including Bratislava which was succeeded by creation of new esplanades, reconstruction of former market squares to the public spaces. Since period of the 2nd 266

half of 18th century, establishment the first public park – Aupark has been dated in central European countries in Bratislava (in 1775, nowadays as Sad Janka Kráľa in Petržalka). There are registered plantations of street trees and alleys such as locust-tree, linden, maple, and plane. At tree plantings was importantly participated city’s decorative association (Tomaško, 1967). Activities related to Nitra town reconstruction, mainly after revolution during 1948–1949 years, were aimed to the construction of new buildings of state and public governance, theatre and other objects of culture, recreation, amusem*nt and military services (barracks). The new streets were paved, installed public lighting and planted trees at streets and squares under active participation of Nitra decorative association, which was established in 1888s by 123 founding members (Fusek and Zemene, 1998). Activity of decorative associations passed over progressive development in Slovakia and at present time, the care of cities green areas including street trees passed under leadership of the city authority

profession departments. Street tree alleys, at river embankments and bank side roads have wide spectrum of functions, those in a new terminology are classified as services for man and improving of environmental conditions. Basically they might be divided into four groups as are supporting, provisioning, regulating and cultural services (Reyers et al., 2009; Yong, 2010). Particular classification and quantification of functional importance and effectiveness of green spaces including street trees have been published by Supuka et al. (1991). Accent is given on environmental quality improving (climate, physical, chemical, micro-organism air quality, and water regime), ecological aspects of urban spaces (biodiversity, ecological stability, ecological and green nets), social and cultural ones (trees as urban design elements, aesthetic, culture, education, recreation effect). Moreover most actual research articles at international level have presented functional and service tree’s values in urban environment. Green spaces and trees in USA cities have absorbed annually 711 thousand tons of allochtonous components on average, especially glasshouse gases such as ozone, carbon, nitrogen and sulphur oxides (Nowak et al., 2006). Climate improving like shade, refrigeration, air humidity up to 10% effectiveness was published in other article (Armson et al., 2012; Tomaško, 1996). The negative influences of woody plants in the relation to human body where a lot of woody species have caused allergy sickness were published as well (Zlínska, 1996). Assessment of the landscape gardening values and healthy conditions of poplar-trees in Nitra streets and green spaces has been published by Verešová (1999), tree alleys of 9 streets of Nitra town were evaluated from the phytopatology point of view (Tkáčová, 2003). The cities are characterised as spaces with expressive environmental changes, which limits selection of trees for street alleys. For this reason a lot of authors and research projects have aimed in issue solution and definition of criterions on advisable tree selection for urban environment (Quigley, 2004; Saebo et al., 2005; Supuka, 2005; Vreštiak, 1994 and others). The aim of the paper is to assess framework development in establishment and changes of streets in Nitra town, to evaluate current state of selected street spaces from point of species composition, ecological importance, environmental adaptability and cultural-aesthetic values. The specific street’s tree assessment method was defined and applied in Nitra town conditions.

Material and methods Historical development of street tree alleys establishment in relation to urban structure reconstruction at Nitra town was assessed by using of published historical documents (Fusek and Zemene, 1998) and as a result of own field study. Current state of selected 26 most inter-

ested street alleys were assessed complexly by using of published particular criterions (Benčať, 1982; Machovec et al., 2000; Juhásová et al., 1991; Supuka, 2005; Verešová, 1999; Vreštiak, 1994; Zlínska, 1996). Throughout syntheses particular marks and characteristics we have defined following categories and assessing criteria for street trees alleys in Nitra town: a) Characteristic dates aa) Locality, street, square ab) Name of woody plant and cultivar ac) Average tree alley high ad) Average tree alley age ae) Average landscape gardening value b) Potential of environmental adaptability to ba) Urban soils bb) Urban climate bc) Soil salinity bd) Complex of immission impact c) Potential of biology-ecological values ca) Potential for ecological networks and ways cb) Topical and trophycal potential for biodiversity support cc) Potential of phytoncidy activity and microbiol ogy regulation cd) Potential of pest and disease impact ce) Potential of tree pruning tolerance and regene ration ability d) Potential of culture values da) Potential of aesthetic effect and perception db) Potential of species rareness and gene-pool value e) Potential of negative influences ea) Potential of phyto-allergy eb) Potential of invasive demonstration ec) Potential of litter fall contamination. Each potential within particular categories might be valuated in three degrees: (1) low, (2) medium, (3) high.

Results and discussion The graphic vedutes from 1st half of 19 century shows Nitra as romantic town surrounded by gardens with domain of sacral and governance buildings and churches. To the town leaded road bridged over Nitrička (second branch of the Nitra river to the south of castle hill), lined by both side tree alleys. In that time street trees were rather ambition that reality, the trees were most often as river bank side tree line, in domain of poplars, willows, ashes and alders. Effective planting of street tree alleys and park areas was started after revolution years 1848– 1849s, which was closely associated with reconstruction of urban structure of settlements. The contribution is coursed only at those new buildings and reconstruction which relates to new tree planting. In 1870–1873s there were made terraced modifications of the square in 267

front of Piaristic church and placed 12 apostles’ sculptures and throughout circumference were 100 pieces of horse chestnuts tree planted, which are found there at present time. In period of 1878–1885, the 14 chapels of cross-route were built-up at Mariansky hill associated with linden and horse chestnuts tree plantings. Construction of region theatre on contemporary Svätopluk square in 1885 was accompanied by tree planting of Acer platanoides ´Globosum´ at square circ*mstance. In 1905 was reconstructed the New Elisabeth road from town barracks to gasworks, 8 m in width, paved surface and tree alleys planted on both sides. Road from Nitra’s capitol till Taufel garden at Chmelova valley, constructed in 1888 was lined by tree alleys plantings. After the Nitra river regulation on the turn out of 19th and 20th century there was established city park Sihoť and embankment of new canal was planted up by tree alleys (linden, horse chestnut, and poplar). Tree alley was planted along with road from down town to rail way station, at Palanok street under castle were planted coniferous trees. High merit at tree alley planting has had Nitra´s decorative association established in 1888 at Chmelova valley town zone that was renamed to Zobor´s decorative association in 1889 and on 24 October 1897 to Town’s decorative association (first chair MUDr. K. Tarnóci). At tree alley plantations, parks establishment and a forestation of Mariansky and Šibeničny hill there was very active forester Marcel Boroš. In 1917 association has expired because of the 1st world war. The historical documents and photos of Nitra town from period of 1920ies show tree alleys from the Kralik restaurant as far as theatre. The new dwelling zones of Nitra town have been built up after 2nd world war and parallely there were planted street alleys and new park was established. In sixties of the 20th century there was appointed town’s enterprise of gardening and technical service, which has had solicitude at street trees and green areas. Since 1990 all activities related to city greenery have been under profession auspices of city authority Department of main architect, section of urbanism and architecture. Reconstruction, new plantings and maintenance of street trees, park trees and green areas are served by profession private enterprises. Establishment of street tree alleys is directly associated to urban development of the town. Constitution of the 1st Czechoslovak republic creates extensive opportunity for social-economy development of new state human society including Nitra town. There were established new dwelling zones such as Číneš, under Calvary, Post colony, under Zobor. In the new built up streets were planted new streets alleys, especially linden, maple, ash, locust-tree, in natural tree shape or as cultivars mostly in globes or column ones. After the 2nd world war additional dwelling town zones were built up, e.g. Chrenová, Párovce, Čermáň, Klokočina, Diely, Zobor (further intensification). Those zones were es268

tablished on area of original villages or on new areas at former agriculture land and vineyards. Along with transport lines, small squares and public spaces were planted new tree alleys. Besides traditional trees, a new woody species and cultivars were planted, such as Acer platanoides ´Globosa´, Acer dasycarpum, Carpinus betulus ´Columnare´, Fraxinus excelsior ´Globosa´, Prunus cerasifera ´Atropurpurea´, P.c. ´Nigra´, Ribinia pseudoacacia ´Umbraculifera´, R. p. ´Bessoniana´, and others. To the newest planted woody species and cultivars in the streets belongs e.g. Acer platanoides ´Columnare´, Koelreuteria panniculata, Gingko biloba, Liquidambar styraciflua, Prunus fruticosa ´Globosa´, Sorbus aria ´Lutescens´ and others. Those new tree forms and cultivars were successively planted to alleys during reconstruction of Svätoplukovo square and Štefánikova street at present time as pedestrian zone. Those two public spaces have representative and culture aesthetic dominant functions therefore they are designed in high architectonical level with applying of modern trees in alleys. New woody plant species and attractive cultivars were also planted as a part of reconstruction of old tree alleys or within process of humanisation of the down town and housing estates. Very important and high effective was change of tree alleys in Naperville street, where original older trees of Populus × canadensis were replaced by Celtis occidentalis street trees, mainly for reason of human body allergy elimination. Smaller poplar’s alley (P. × canadensis) is planted at the Nitra river embankment in neighbouring of birch Grove Park and also throughout circumference of football and hockey stadium as protection green barriers. Particular survey of species composition of contemporary tree alleys at selected street spaces and squares of Nitra town is presented in Table 1, including the tree assessment according to described methods. At the Nitra territory there have been mapped 26 most important street tree alleys where 22 species and cultivars were identified and assessed. To the most traditional street trees belongs linden, which represents the oldest alley at Nábrežie mládeže embankment. To the latest tree elements belong e.g. Gingko biloba L. ´Fastigiata´ and Liquidambar styraciflua L. Originally, perspective of Prunus fruticosa ´Globosa´ species shows its weak point because attack by Erwinia amylovora fungus has been identified at older trees in Sládkovičova street. At assessment method of the street tree alleys we paid attention to the marks, which have been used very rare till now in the research and practical mapping. Most often valuation marks up to the present time were pest and diseases occurrence at the vegetation organs, tree vitality characteristics, tree stability and social values of trees as well. Those assessment characteristics and methodical approaches according to individual authors are described in complex publication of group authors (Hrubík et al., 2011).

269

Tilia platyphyllos L.

Prunus fruticosa Pall. ´Globosa´

Fraxinus excelsior L. ´Globosa´

Tilia platyphyllos L.

Square of Cyril a Metod

Farská Street

Kupecká Street

Square of Svätopluk

Štefánikova Street pedestrian zone

Kúpeľná Street

Sládkovičova Street

B. Němcovej Street

Kmeťkova Street

Tilia platyphyllos L. Tilia cordata Mill.

Prunus cerasifera Ehrh. ´Nigra´

Prunus serrulata Franch. ´Hisakura´

Slančíkovej Street, Chrenová

Fraňa Mojtu Street

Štefánikova Street, OD-TESCO

20 4 7

Acer platanoides L. ´Columnare´

Liquidambar styraciflua L.

Štúrova Street, OD-MLYNY

Čsl. Armády Street

Nepervillská Street

Mostná Street

Župné Square

Janka Kráľa Street

Populus alba L. ´Boleana´

Acer saccharinum L. (yearly prunned)

Acer platanoides L. ´Globosa´

Pyrus calleryana Decne. ´Chanticleer´

Carpinus betulus L. ´Columnare´

Catalpa bignonioides Walt. ´Nana´

Štefánikova Street, VÚB

Celtis occidentalis L.

Koelreuteria panniculata Laxm.

Nábrežie mládeže and Wilsonovo nábrežie Embankment

Tilia cordata Mill.

Sorbus aria (L.) Crantz. ´Lutescens´ 12

Prunus fruticosa Pall. ´Globosa´

Corylus avelana L. ´Globosa´

Tilia plataphyllos L.

Gingko biloba L. Fastigiata´

10 5

a) Characteristics

Acer platanoides L. ´Globosa´,

Aesculus hippocastanum L.

Tilia cordata Mill.

Fraxinus excelsior. L. ´Globosa´

Tilia platyphyllos L., Tilia cordata Mill.

Štúrova Street ZSS + SOU

Bernolákova Street

Crataegus monogyna L. ´Paul´s Scarlet´

Štúrova Street Dom služieb

Damborského Street

Tilia platyphyllos L.

7. Pešieho pluku Street

Robinia pseudoacacia L.´Bessoniana´

Name of woody plant

Štefánikova Street OD MLYNY

Locality, street, square

Table 1. List of inventoried street alley trees in Nitra town and their evaluation

b) Environmental adaptability

c) Biology-ecological values

d) Culture values

e) Negative influences

Achieved results of our research activity have showed positive trends in tree alley reconstruction on the on hand. On the other hand many tree alleys in Nitra town are destroyed, trees are too old, and unseemly that would be reason for more effective decision to remove them from street soon. In some streets we may see alleys with different tree species, which looks very diversified and out of composition rules, e.g. Štefánikova street, Župné square. Every year tree branches cutting at Acer saccharinum L. in J. Kráľ street creates compacted and unified street alley. Species of Populus genus, with regards to fast growing character and big tree crown should be useful to the future also as elements for protection barriers, as shade tree and also for windbreaks around large sport centres, but selected male gender trees only (Varga, 1994). In conclusion, our research results show positive and negative feedback. In our opinion Nitra town needs to elaborate complex master plan for reconstruction and quality improving of main and the most attractive streets regarding tree alleys.

Acknowledgement Paper was elaborated thanks to financial supporting by the grant project KEGA No. 020SPU-4/2011 and No. 019SPU-4/2011 on Ministry of Education, Science, Research and Sport of the Slovak Republic.

References Armson, D., Stringer, P., Ennos, A.R. 2012. The effect of tree shade and grass on surface and globe temperature in an urban area. Urban For. Urban Greening, 11: 245–255. Benčať, F. 1982. Atlas rozšírenia cudzokrajných drevín na Slovensku arajonizácia ich pestovania [Atlas of the distribution of exotic woody plants in Slovakia and zoning of their cultivation]. Bratislava: Veda. 360 p. Fusek, G., Zemene, M.R. 1998. Dejiny Nitry [A history of Nitra]. Nitra: Nitrianske tlačiarne. 398 p. Hrubík, P., Kollár, J., Rovná, K., Tkáčová, S., Mňahončáková, E. 2011. Kvalitatívna inventarizácia, klasifikácia a hodnotenie zdravotného stavu drevín pre účely záhradno-architektonickej a krajinárskej tvorby [Qualitative inventory, classification and the assessment of health stage of woody plants for the purposes of garden architectonical and landscape creation]. Nitra: Slovenská poľnohospodárska univerzita. 100 p. Juhásová, G., Hrubík, P., Gáper, J. 1991. Škodlivé činitele zelene [Harmful agents of greenery]. In Supuka, J., Benčať, J., Bublinec, E., Gáper, J., Hrubík, P., Juhásová, G., Maglocký, Š., Vreštiak, P. Eko270

logické princípy tvorby a ochrany zelene. Bratislava: Veda, p. 235–294. Kalusok, M. 2004. Záhradná architektura. Malá encyklopedie [Garden architecture. Small encyclopaedia]. Brno: Computer Press. 192 p. Kupka, J. 2010. Krajiny kultúrní a historické [Culture and historical landscapes]. Praha: ČVUT. 180 p. Machovec, J., Hrubík, P., Vreštiak, P. 2000. Sadovnícka dendrológia [Landscape gardening dendrology]. Nitra: Slovenská poľnohospodárska univerzita. 228 p. Nowak, D.J., Crane, D.E., Stevens, J.C. 2006. Air pollution removal by urban trees and shrubs in the United States. Urban For. Urban Greening, 4: 115–123. Quigley, M. F. 2004. Street trees and rural conspecificies: Will long-lived trees reach full size in urban conditions. Urban Ecosystems, 7: 29–39. Reyers, P. et al. 2009. Ecosystem services, land-cover change and stakeholders: Finding and sustainable food hold for semi-arid biodiversity hotspot. Ecol. and Soc., 14, p. 38–46. Saebo, A., Borzan, Z., Supuka, J. et al. 2005. The selection of woody plant materials for street trees, park trees and urban woodlands. In Urban forests and trees: Areference book – COST E12. Heidelberg: Springer Verlag, p. 257-280. Supuka, J. 2005. Význam a stratégia výberu drevín pre komponovanie vegetačných štruktúr miest a sídelného prostredia [Importance and selection strategy of woody plants for vegetation structure composition in towns and settlement environment]. In Bernadovičová, S., Juhásová, G. (eds). Dreviny vo verejnej zeleni. Zborník z konferencie s medzinárodnou účasťou, 10. – 11. 5. 2005, Bratislava. Zvolen: Ústav ekológie lesa SAV, p. 72–77. Supuka, J., Benčať, J., Bublinec, E., Gáper, J., Hrubík, P., Juhásová, G., Maglocký, Š., Vreštiak, P. 1991. Ekologické princípy tvorby aochrany zelene [Ecological principles of green spaces creation and protection]. Bratislava: Veda. 308 p. Supuka, J., Feriancová, Ľ., Tomaško, I., Štrba, B., Štěpánková, R., Rózová, Z., Oboňová, M., Moravčík, Ľ., Laurová, S., Kuczman, G., Kubišta, R. 2008. Vegetačné štruktúry v sídlach: parky a záhrady [Vegetation structures in settlements: parks and gardens]. Nitra: Slovenská poľnohospodárska univerzita, Fakulta záhradníctva akrajinného inžinierstva. 499 p. Tkáčová, S. 2003. Príklady poškodzovania uličných drevín v meste Nitra [The instances of the trees injuring in the streets of Nitra town]. In Bernardovičová, S. (ed.) Dreviny vo verejnej zeleni. Zborník z konferencie s medzinárodnou účasťou, 27. – 28. 5. 2003, Košice. Košice: Univerzita Pavla Jozefa Šafárika, p. 137–145. Tomaško, I.. 1996. Využitie introdukovaných drevín na vyrovnanie negatívneho dopadu globálnych

klimatických zmien v lesnom hospodárstve a pri úpravách krajiny [Use of introduced woody plants to balance negative impact of global climatic changes in forestry and in landscape arrangement]. In Škvarenina, J., Minďáš, J., Čaboun, V. (eds). Lesné ekosystémy a globálne klimaticko zmeny. Zborník referátov z pracovného seminára, Zvolen, 22. február 1995. Zvolen: Lesnícky výskumný ústav, p. 140–143. Tomaško, I.. 1967. Vedecké základy systému mestskej zelene rozpracované na príklade Bratislavy [Scientific basis of the city green spaces elaborated at Bratislava capital study area]. In Benčať, F. (ed.). Problémy dendrobiológie a sadovníctva. Slávnostný sborník prác Arboréta Mlyňany SAV k 75. výročiu jeho založenia. Bratislava: SAV, p. 323–454. Varga, L. 1994. Rýchlorastúce dreviny v intravilánoch obcí a miest [Fast growing woody plants for village and towns intravilane]. In Vreštiak, P. (ed.). Stromy

v uliciach miest. Nitra: Slovenská poľnohospodárska univerzita, p. 51–57. Verešová, M. 1999. Zhodnotenie rodu Populus L. vmestskom prostredí [Assessment of genus Populus L. in city environment]. Diploma thesis. Nitra: Slovak University for Agriculture in Nitra. 51 p. Vreštiak, P. 1994. Sortiment stromov pre uličné stromoradie [ Tree species for street alleys]. In Vreštiak, P. (ed.). Stromy v uliciach miest. Nitra: Slovenská poľnohospodárska univerzita, p. 6–15. Yong, R.F. 2010. Managing municipal green space for ecosystem services. Urban For. Urban Greening, 9: 313–321. Zlínska, J. 1996. Zeleň obytných súborov z hľadiska peľových alergií [Green spaces of housing estates from the pollen allergy point of view]. In Fotta, M. (ed.). Revitalizácia obytných súborov. Nitra: Slovenská poľnohospodárska univerzita, p. 59–66.

Received December 6, 2012 Accepted September 30, 2013

271

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Effect of delayed tending on development of beech (fa*gus sylvatica L.) pole stage stand Igor Štefančík1, 2 1

National Forest Centre-Forest Research Institute in Zvolen, T. G. Masaryka 22, SK-960 92 Zvolen, Slovak Republic, e-mail: [emailprotected] 2 Department of Silviculture, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences in Prague, Kamýcká 1176, CZ-165 21 Prague 6 – Suchdol, Czech Republic

Abstract Štefančík, I. 2013. Effect of delayed tending on development of beech (fa*gus sylvatica L.) pole stage stand. Folia oecol., 40: 272–281. The paper deals with assessment of the long-term experiment (45 years of investigation) in beech (fa*gus sylvatica L.) stand with delayed tending started at stand age of 60 years. The research was performed on four partial plots by different methods of their management: (i) plot with heavy thinning from below (C degree according to the German forest research institutes from 1902), (ii) plot with the free crown thinning (thinning interval of 5 years), (iii) plot with the free crown thinning (thinning interval of 10 years) and (iv) control plot (with no thinning). From qualitative point of view, the best results according to the number of target (crop) trees were found on plots tended by the free crown thinning (thinning interval of 5 years), and the worst on plots with heavy thinning from below and/or plot with no tending (control plot). Consequently, the results showed lower number of target (crop) trees in comparison with our assumption and/or the model developed for beech stands in the past. On the other hand, from quantitative point of view, the best results were achieved on plot tended by heavy thinning from below, followed by the plot with the free crown thinning (thinning interval of 5 years). Keywords beech, crop trees, quantitative production, stand structure, tending

Introduction Tending of each forest stand has a crucial importance for its development. As a rule, it takes more than half of rotation age of forest stand. For the management of beech stands originated from natural regeneration, is very important not only the method of their tending, but also the stand age, when to start with tending. The papers dealt with the history of beech stands tending in Slovakia (Štefančík, 1985) and other literature focused on problems of thinning in beech stand (Štefančík, 1984) concluded, that tending of beech stands had not so special tradition in comparison to France, Denmark or Germany. It should be stated, that tending was realized only according to foreign knowledge and poor experiences of internal forest practitio-

272

ners. The systematic research started at the end of the 50-ies, of the last century. The aim of the research at this time was mainly to find the first scientific results on the mentioned topic under the natural condition of Slovakia. Within the framework of thinning problems, the attention was especially paid to know, which kind of selective thinning method should be considered the most suitable for beech stands under our natural condition. Nevertheless, actual experiences by application of the methods developed abroad, and also the condition of beech premature stands in Slovakia, have been taken into account (Štefančík, 1984). Research was by the first time focused on the beech thickets, not systematically tended until then (Réh, 1968, 1969) and/or small pole stage stands or pole

stage stands (Šebík, 1969; Štefančík, 1974). Within the framework of the research, all principal silvicultureproduction questions of thinning started to be solved step by step. In the initial stage of the research it was especially the problem related to thinning type (thinning from below, crown thinning), method of selection (positive, negative) and structure of pure beech stands, as well as thinning intensity, i.e. intensity, frequency and thinning interval, later on. Effect of two degrees of thinning from below (B and C according to German forest research institutes from 1902) and two crown thinning methods (qualitative according to Schädelin) and the free crown thinning according to Štefančík (1984) started to be verified by the research. Since the 70-ies of last century, the results of longterm investigation have been published (Štefančík, 1974, 1984; Šebík and Polák 1990; Štefančík et al., 1991, 1996; Štefančík, 2007; Štefančík and Bolvanský, 2011). The outcomes showed better results by application of crown thinning in comparison with thinning from below. Especially, the free crown thinning (Štefančík, 1984, 2007) appeared to be suitable for tending of pure beech stands in Slovakia. Since 1958, the above-mentioned method has been applied in the thinning research of beech stands. Nowadays, after long-term verification it was successfully put into the practise. The aim of this paper was to ascertain the changes of selected parameters of quantitative and qualitative production in beech stands, tended by different thinning methods for a long time (45 years of investigation).

Material and methods The research was carried out on the series of permanent research plot (PRP) Cigánka, established in the stand located in compartment 50, forest district Muráň, forest enterprise Revúca. The given beech stand originated from a natural regeneration. The stand age on the PRPs at their establishment (in autumn 1966) was 60 years. The mentioned series of PRPs consists of four partial plots (C, H, H2, 0) with the area of each plot of 0.25 hectare. The basic mensurational characteristics are presented in Table 1. On the plot (marked as C) a heavy thinning from below (C degree according to German forest research institutes from 1902) was realized. On the second and the third plot (marked as H and H2), the method of the free crown thinning (according to Štefančík, 1984) was applied. The mentioned method is focused on individual tending of the trees of selective quality (promising and target trees). These trees are selected on the base of the three criteria (quality, dimension and spacing). Thinning interval on plot H is 5 years, and on plot H2 10 years.

Table 1. The basic characteristics of the given series of permanent research plots (PRPs) Cigánka Characteristic

PRP Cigánka

Establishment of PRP

Autumn 1966

Age of stand [years]

60 (in 1967)

Site index

Geomorphologic unit

Stolické vrchy

Exposition

Northwest

Altitude [m]

560

Inclination [degree]

Parent rock

Gneiss (biotitic)

Soil unit

Haplic Cambisol (Dystric)

Forest altitudinal zone

4th beech

Ecological rank

A (Acid)

Management complex of forest types

405 – acid beech woods

Forest type group

fa*getum pauper (Fp) higher tier

Forest type

4301 woodrush beech woods (higher tier)

Average annual temperature [°C]

5.5

Sum of average annual precipitation [mm year–1]

918

The plot marked as 0 is control (with no thinning). No planned silvicultural interventions were carried out up to establishment of PRPs. The first measurement was performed in 1967. Since establishment of PRPs, 10 biometrical measurements were realized on each partial plot with the interval of 5 years, including the intervention (only on treated plots) with thinning interval of 5 years (plot C and H) and/or 10 years (plot H2). On all plots, the standard biometrical measurement and evaluation of stem and crown of trees were carried out. Within the framework of the measurement, the quantitative parameters (breast height diameter, both height of tree and base of tree crown, crown width) were assessed according to silvicultural and commercial classification. They were focused on evaluation of each tree, and separately on the trees of selective quality (promising and target trees). Silvicultural classification consists of: a) Biosociological position of trees according to growth (tree) classes (Štefančík, 1984): 1. dominant tree 2. co-dominant tree 3. intermediate tree 4. suppressed tree – decreased 5. suppressed tree – dying out b) Degree of stem quality: 1. well-shaped and straight, best stem quality, with out burrs

273

2. average shaped – average stem quality, crooked only in upper third of the stem, low number of burrs 3. bad shaped – worse quality of the stem, high num ber of burrs, very crooked c) Degree of crown quality: According to the type (form of ramification): 1. crown with continuous axis of stem up to the top of tree; 2. bouquet (cluster); 3. broom; 4. forked. According to size: 1. oversized; 2. normal size; 3. small size, asymmetrical developed, but able to regenerate; 4. too small size, not able to regenerate. According to crown density (sufficiency of foliage): 1. good density with complete foliation, also inside of the crown; 2. sufficient density, with foliation in outside of the crown only; 3. sparse, foliation quite well; 4. very sparse, insufficient foliation. Within the framework of commercial classification, only lower part of the stem up to crown base was assessed, separately for lower and upper half of the stem, respectively. Quality classes: 1. very high (A), 2. average (B), low quality, but industrial wood (C), 4. fuelwood (D). The calculation of the results was performed by standard methods for tending evaluation and production-silviculture relations, utilized the software package of QC Expert and growth simulator Sibyla (Fabrika, 2005). To find out the statistical significance of the differences, the single-factor analysis of variance (anova) was used.

Results and discussion Diameter structure The diameter development of the investigated PRPs is characterized by the diameter frequency distribution (Figs 1 and 2), as well as by the values of mean diameter (dg) presented in Table 2. In the initial stage of the research, the course of curves of diameter frequency distribution was found similar to all partial plots (Fig. 1). It is a type of lefthand asymmetric distribution, typical for young stands, as well as for the middle age ones, which were untouched (neglected by tending) until then. This is also the case of PRP Cigánka, where tending started at the growth stage of pole stage stand (60 year old), which is considered to be too delayed for beech stands. On the base of numerous experimental experiments, it is recommended to start with tending already in the thickets (Réh, 1968, 1969; Jurča and Chroust, 1973; Korpeľ et al., 1991) and/or no later than in small pole stage stand (Štefančík, 1974). The highest values of the mean diameter (dg) were found on plot C, and the lowest on control plot (0). After 45 years under different thinning regime, the differences among the plots increased (statistical significant differences at the level α = 0.05 were found only between plot C and each other plots). The order of plots remained unchanged, when the highest values of dg were found on plot C (heavy thinning from below),

Fig. 1. Diameter frequency distribution on plots in the initial stage of the research in 1967.

274

Fig. 2. Diameter frequency distribution on investigated plots in 2012.

and the lowest on control plot (with no treatment), Table 2. Simultaneously, the diameter frequency distribution was more or less changed to double-peak course

(Fig. 2). The above-mentioned development response the thinning methods realised. On the plots tended by the free crown thinning (H, H2), the interventions

Table 2. Development of stand characteristics Plot

Stand

Age

V7b

Mean Diameter d1,3 [cm]

Height [m]

[year]

[pcs ha–1]

[m2 ha–1]

[m3 ha–1]

[dg]

[hg]

Total

2,940

34.784

337.444

12.3

19.1

Main

2,276

35.768

357.892

14.1

19.8

Total Main

2,004

35.956

392.920

15.1

21.4

1,736

36.980

437.292

16.5

23.3

1,592

38.036

477.004

17.4

24.7

1,472

38.604

516.368

18.3

26.2

1,224

39.208

537.228

20.2

26.7

1,144

41.404

570.180

21.5

26.7

100

1,068

41.556

585.912

22.3

26.6

105

1,012

43.956

631.524

23.5

26.9

2,632

36.436

365.172

13.3

20.1

1,140

25.888

303.516

17.0

23.2

816

22.480

286.980

18.7

25.0

756

23.120

300.944

19.7

25.1

624

21.172

287.976

20.8

26.2

620

25.128

351.032

22.7

26.6

584

27.388

389.152

24.4

26.8

584

31.064

455.964

26.0

27.4

100

564

31.724

485.060

26.8

27.5

105

548

34.452

540.876

28.3

27.8

275

Table 2. Development of stand characteristics – continued Plot

Stand

Total Main

Age

V7b

Mean Diameter d1,3 [cm]

Height [m]

[year]

[pcs ha–1]

[m2 ha–1]

[m3 ha–1]

[dg]

[hg]

2,568

35.500

354.504

13.3

20.1

1,552

28.968

307.740

15.4

21.0

1,032

23.808

284.196

17.1

23.6

992

25.948

319.808

18.3

24.0

800

22.012

279.256

18.7

24.5

784

24.992

334.212

20.2

25.7

740

26.128

355.132

21.2

25.4

732

28.916

398.292

22.4

25.5

100

712

29.128

418.452

22.8

25.6

105

704

31.792

468.432

24.0

26.1

Total

2,308

40.060

444.152

14.9

21.6

Main

520

26.696

387.996

25.6

29.8

440

27.096

416.032

28.0

31.5

324

26.024

420.956

32.0

33.2

312

28.468

482.532

34.1

34.7

312

31.896

563.500

36.1

35.9

280

32.104

595.804

38.2

37.2

280

35.168

668.400

40.0

38.0

100

272

36.052

690.564

41.1

38.2

105

272

38.728

778.980

42.6

38.8

N, number of trees; G, basal area; V7b, volume of the timber to the top of 7 cm o.b. C → plot with thinning from below. H → plot with thinning from above, thinning interval 5 years. H2 → plot with thinning from above, thinning interval 10 years. 0 → control plot (with no treatment).

were performed in the whole vertical profile, which resulted in a better diameter differentiation. It was also confirmed by the values of indices of diameter differentiation (TMd) according to (Füldner, 1995), which were found the highest, just in the plots treated by the free crown thinning (for H = 0.578 and H2 = 0.516). The values above 0.500 represent the strong type of differentiation. For comparison, in the control plot it was 0.398 (medium type of differentiation) and the lowest values of indices were obtained in plot C (0.173 – little differentiation), where total suppressed level of the stand was removed by the treatment. Height structure The height (stand) structure of the investigated plots was expressed by the relative number in the growth (tree) classes (Fig. 3). The proportion of trees in the crown level of the stand (1st + 2nd growth class) and the suppressed level of the stand (3rd to 5th growth class) is very important from the silvicultural point of view. The 276

structure depends especially on site, tree species, stand age and tending measures (Šebík and Polák, 1990). In the initial stage of the research, the height structure was practically the same. The proportion of the suppressed level of the stand ranged from 28.4% on control plot to 29.9% on plot H2. The differences (shifts) in the height structure (proportion between the crown level of the stand and the suppressed one), after 45 years of investigation were found only at about 10% (plot 0 and H). Contrary to the mentioned plots, plot H2 remained unchanged. These results are in accordance with the outcomes published by Šebík and Polák (1990), who stated the shift of the trees to the higher growth (tree) classes, when heavy crown thinning was applied. The mentioned authors also concluded that decreased number of co-dominant trees in the stand with shade-bearing species is typical, together with increased amount of the suppressed ones. The most proportioned are being the fourth, or the 4th and the 5th growth class, which was also confirmed by our research on PRP Cigánka. The similar results were published by

Fig. 3. Relative number according to the growth classes on plots after 45 years in 2012.

Assmann (1968) for 102 years old beech stand tended by mild crown thinning, where proportion of the crown level of the stand and the suppressed level of the stand was found of 53.8% and 46.2%, respectively. The highest changes were registered on plot C (heavy thinning from below), where in a consequence of removed suppressed level of the stand remained only intermediate individuals (the 3rd growth class) with low proportion of 17.6%. Very interesting should be considered the fact, that control plot left to self-development showed practically the same height structure in comparison with the plots tended by the free crown thinning (Fig. 4). This was also confirmed by the statement published in the past (Štefančík, 2007), that according to its conception, the mentioned thinning method is very similar to principles of close to nature silviculture. It was also proved by the values of indices characterized the

vertical structure (APi) according to Pretzsch (1992). On the plots with the free crown thinning, the values were found of 0.791 and 0.758. For example, in the stand with a selection structure, the mentioned index is able to achieve the value of 0.900 (Pretzsch, 1992). For comparison, we suggest, that on the control plot (0) in PRP Cigánka, the index was found of 0.447. Consequently, the indices of the height differentiation (TMh) according to Füldner (1995) were found the highest for plots tended by the free crown thinning (H = 0,514 and H2 = 0,439), contrary to control plot (0.302) and plot C (0.037). As for the comparison of the values of the mean height (hg), after 45 years, the highest differences (statistical significant at the level α = 0.05) were found between plot C and each other plots. The rest three plots showed similar values, but differences among them were statistically insignificant.

Fig. 4. Stand structure on PRP Cigánka plot 0 (left) and plot H (right).

277

Development of quantitative production

nual increment on basal area and/or volume increment in 5 years periods (Figs 5 and 6). The total mean annual volume increment during the investigated period was found 14.3 m3 ha–1 on plot C, followed by plot H – 11.1 m3 ha–1, plot H2 – 9.0 m3 ha–1 and control plot – 8.7 m3 ha–1.

The development of stand characteristics during the investigated period is presented in Tables 2 and 3. At establishment of the plots, the highest initial number of trees (N) was found on control plot (0) and the lowest on plot C. After 45 years, the order was not changed, whereby on control plot remained 34.4% out of the initial number of trees, but on the plot C only 11.8%. As for the other stand characteristics (basal area – G, and volume of the timber to the top of 7 cm o.b. – V7b), the highest values were found on plot tended by heavy thinning from below and the lower on plots with the free crown thinning (H, H2). These results are in accordance with the experiences of numerous thinning experiments established in the past, concluded by Assmann (1968),Šebík andPolák (1990), Štefančík (1990). The analysis of the total decrease (thinning, selfthinning, abiotic injurious factors) according to G and V7b for the period of 45 years showed the highest percentage on plots tended by the free crown thinning (H, H2) and the lowest on control plot (Table 3). As for the total production (according to G and V7b), the highest values were found on plot with heavy thinning from below and the free crown thinning (thinning interval of 5 years). The same results were also obtained, by expression of growth index of the total production in investigated period. It suggests suitable effects of even though delayed tending measures in beech stands. Additionally, beech species is wellknown of its very good responses to liberation (releasing) up to the oldest period (Assmann, 1968; Šebík and Polák, 1990). It was fully confirmed by the results from PRP Cigánka. It should be concluded, that from quantitative point of view, the best results were obtained on plots tended by heavy thinning from below and the free crown thinning with thinning interval of 5 years, contrary to control plot (0), characterized by the worst outcomes. It was also confirmed by the values of the current an-

Development of target (crop) trees Information related to the target (crop) trees (TT) development, representing qualitative production in commercial forests is presented in Table 4. It can be seen, that from quantitative parameters point of view, in the initial stage of the research, the highest values were found on plot C and/or the lowest on plot H2. Number of TT ranged from 176 to 208 individuals per hectare. During the tending period of 45 years, the situation was changed unambiguously in favour of plots treated by the free crown thinning (H and H2). On the mentioned plots, double number of TT was cultivated in comparison to plot tended by heavy thinning from below (plot C). The same results were obtained, if we take into account the production parameters (basal area, volume of the timber to the top of 7 cm o.b.). The proportion of TT out of the main stand is considered to be a very important parameter. The plots managed by the free crown thinning showed also the best results according to the mentioned quantitative parameters in comparison with plots tended by heavy thinning from below, or control plot. The model of the future mature beech stand developed by Štefančík (1984) assumed at stand age of 110–130 years, in acid site, the number of TT presented 173 to 200 trees per hectare and 376 m3 ha–1 of volume of the timber to the top of 7 cm o.b. Its proportion had to be of 75% out of the main stand. Mean diameter d1,3 was assumed to achieve 40 cm. It can be seen, that the results from the PRP Cigánka obtained at stand age of 105 years are very close to the mentioned model, except for number of TT, which is much lower. It is a consequence of delayed tending, which started at stand age of 60 years. It is a generally

Table 3. Development of quantitative production of the stand for 45 years Plot

Age range

Total decrease of trees N [pcs ha–1]

Total production

G % of

[m2 ha–1]

V7b % of TP [m3 ha–1]

[years]

% of TP [pcs ha–1]

V7b

[m2 ha–1] Index of total stand

[m3 ha–1]

Index of total stand

60–105

1,928

65.6

12.856

22.6

98.616

13.5

2,940

56.812

1.633

730.140

2.164

60–105

2,084

79.2

29.688

46.3

321.528

37.2

2,632

64.140

1.760

862.404

2.362

60–105

1,864

42.6

27.192

46.1

291.424

38.4

2,568

58.984

1.662

759.856

2.143

60–105

2,036

88.2

27.884

41.9

307.660

28.3

2,308

66.612

1.663

1,086.640

2.447

N, number of trees; G, basal area; V7b, volume of the timber to the top of 7 cm o.b.; TP, total production. C → plot with thinning from below. H → plot with thinning from above, thinning interval 5 years. H2 → plot with thinning from above, thinning interval 10 years. 0 → control plot (with no treatment).

278

Current annual increment [m2 ha–1] Current annual increment [m3 ha–1]

Fig. 5. Current annual basal area increment in the 5 years period of investigation.

Fig. 6. Current annual volume increment in the 5 years period of investigation.

279

Table 4. Development of target (crop) trees Plot

Age

V7b % out of

[years]

[pcs ha ] –1

[m ha ] 2

–1

main

[m ha ] 3

–1

stand 0 H H2 C

Mean % out of

diameter

height

main

d1,3 [cm]

[m]

stand

[dg]

[hg]

200

6.688

19.2

80.992

24.0

20.6

25.4

105

108

11.420

26.0

191.048

30.3

36.7

32.4

188

6.428

25.2

79.308

29.1

20.9

25.6

105

124

18.332

53.2

320.988

59.3

43.4

33.5

176

6.512

24.0

81.312

29.4

21.7

26.0

105

132

16.724

52.6

282.404

60.3

40.2

32.7

208

10.372

38.3

138.636

40.1

25.2

27.5

105

13.303

34.3

277.620

35.6

49.9

40.0

known fact, that the best stand age in order to determine and cultivate the TT is considered at the period of 30–40 years (Štefančík, 1974, 1984). As it can be seen, the obtained results from PRP Cigánka showed that it is possible to achieve assumed quantitative production in case of delayed, but systematic tending. On the other hand, it is not possible to cultivate desired qualitative production represented by number of trees with the best quality (target trees), especially on plot managed by heavy thinning from below, or plot without tending. Conclusions Based on the 45 years of investigation of beech stand development managed by delayed tending, where different methods of tending were applied, it can be concluded: o The differences of diverse tending regime were increased between plots after 45 years of investigation in comparison with the initial stage of the experiment. The differences were found significant at the level α = 0.05 between plot C and each other plot. From diameter structure point of view, the order of plots remained unchanged. The highest mean diameter (dg) was found on plot managed by heavy thinning from below, from the initial stage up to now. The lowest one showed the control plot. o The differences (shifts) in the height structure (proportion of the crown level of the stand and the suppressed level of the stand) on plots during the investigated period of 45 years were found at about 10% on plot 0 and H. For plot H2 it remained unchanged. The highest changes were registered on plot C (heavy thinning from below), where due to remov-

280

ing of the suppressed level of the stand, only intermediate individuals (the 3rd growth class) remained in the stand with lower proportion of 17.6%. o The control plot, left to the self-development showed practically the same height structure like the plots tended by the free crown thinning (H and H2). o From quantitative point of view, the best results were found on plots tended by heavy thinning from below and the free crown thinning with thinning interval of 5 years. Consequently, the worst results were obtained from control plot. o As for the total production (expressed by basal area and volume of the timber to the top of 7 cm o.b.), the highest values were found on plot tended by heavy thinning from below and plot with the free crown thinning (thinning interval of 5 years). The same results were also obtained according to the index of the total production. It suggests suitable effect of tending, although delayed, in older beech stands. o The number of target (crop) trees in the initial stage of stand ranged from 176 to 208 individuals per hectare. At the stand age of 105 years, after tending for 45 years, the highest number of crop trees was showed by the plot tended by the free crown thinning (124 and 132 pieces per hectare), and the lower by the plots managed by heavy thinning from below and control plot (68 and 108 pieces per hectare, respectively). o The results, found by long-term investigation (period of 45 years) confirmed, that by systematic and intensive tending, although delayed, it is possible to achieve desired quantitative production, but not qualitative production, represented by the number of the best quality (target) trees, especially on control plot and plot tended by the free crown thinning.

Acknowledgements This work was supported by the Slovak Research and Development Agency under the contract No. APVV0262-11 and Technology Agency of the Czech Republic TA02021250 Silvicultural-ecological and economic optimum of forest stand tending.

References Assmann, E. 1968. Náuka o výnose lesa [Science on forest yield]. Bratislava: Príroda. 488 p. Fabrika, M. 2005. Návrh algoritmov pre prebierkový model rastového simulátora SIBYLA [Proposal of algorithms for thinning models of growth simulator Sibyla]. Lesn. Čas. – For. J., 51: 145–170. F üldner , K. 1995. Strukturbeschreibung in Misch beständen. Forstarchiv, 66: 235–240. Jurča, J., Chroust, L. 1973. Racionalizace výchovy mladých lesních porostů [Rationalization of thicket tending]. Praha: SZN. 239 p. Korpeľ, Š., Peňáz, J., Saniga, M., Tesař, V. 1991. Pestovanie lesa [Silviculture]. Bratislava: Príroda. 472 p. Pretzsch, H. 1992. Konzeption und Konstruktion von Wuchsmodellen für Rein- und Mischbestände. Forstliche Forschungsberichte, Nr. 115. München: Forstwissenschaftliche Fakultät der Universität München. 358 p. Réh, J. 1968. Štúdium štruktúry bukovej húštiny [A study in the structure of beech thicket]. Lesn. Čas., 14: 651–671. Réh, J. 1969. Príspevok kpoznaniu vývoja aniektorých morfologických znakov buka vhúštinách [Contribution to the knowledge of development and some morphological signs of beech in coppices]. In Zbor. ved. Prác Lesn. Fak. VŠLD Zvolen, XI (3): 67–82. Šebík, L. 1969. Vplyv miernej podúrovňovej a akostnej úrovňovej prebierky na vývoj výškového rastu vbukových žrďovinách [The influence of moderate low thinning and qualitz crown thinning upon the development of height growth in beech pole timber]. In Zbor. ved. Prác Lesn. Fak. VŠLD Zvolen, XI (1): 63–85. Šebík, L., Polák, L. 1990. Náuka o produkcii dreva [Science on wood yield]. Bratislava: Príroda. 322 p.

Štefančík, L. 1974. Prebierky bukových žrďovín [Thinnings in beech pole stands]. Lesnícke štúdie, 18. Bratislava: Príroda. 141 p. Štefančík, L. 1984. Úrovňová voľná prebierka – metóda biologickej intenzifikácie a racionalizácie selekčnej výchovy bukových porastov [Free crown thinning – a method of biological intensification and rationalization of the selection tending of beech stands]. In Ved. Práce Výsk. Úst. lesn. Hospod. Zvolen, 34: 69–112. Štefančík, L. 1985. Z histórie výchovy lesných porastov na Slovensku (s osobitným zreteľom na obdobie 1963–1982) [From the history of forest stands tending in Slovakia (with a special attention on the period of 1963–1982]. In Zbor. Lesn. drevár. poľov. Múz. Antol, 13: 3–40. Štefančík, L. 1990. Výskum pestovno-produkčných otázok prebierok vbučinách [Research of silvicultureproduction questions of thinning in beech stands]. Inaugural dissertation. Zvolen: Forest Research Institute. 10 p + appendices. Štefančík, I. 2007. Prebierky vbukových porastoch ako nástroj prírode blízkeho pestovania lesov [Thin nings in beech stands as a tool of close to nature silviculture]. In Prknová, H. (ed.). Význam přírodě blízkých způsobů pěstování lesů pro jejich stabilitu, produkční amimoprodukční funkce. Praha: ČZU, p. 126–133. Štefančík, I., Bolvanský, M. 2011. Pestovanie bukových porastov [Silviculture of beech stands]. In Barna, M., Kulfan, J., Bublinec, E. (eds). Buk abukové ekosystémy Slovenska. Bratislava: Veda, p. 435–456. Štefančík, L., Štefančík, I., Cicák, A. 1991. Zhodnotenie výskumu prebierok a zdravotného stavu nezmiešanej bučiny v imisnej oblasti [Evaluation of the research of thinnings and health state of unmixed beech stand in immission region]. In Ved. Práce Výsk. Úst. Lesn. Hospod. Zvolen, 40: 213–238. Štefančík, L., Utschig, H., Pretzsch, H. 1996. Paralelné sledovanie rastu aštruktúry nezmiešaného bukového porastu na dlhodobých prebierkových výskumných plochách vBavorsku ana Slovensku [Parallel observations of unmixed beech stand growth and structure on long range thinning research plots in Bavaria and Slovakia]. Lesnictví – Forestry, 42: 3–19.

Received February 6, 2013 Accepted April 8, 2013

281

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Results of an ecological-production research on forest ecosystems of woody plants introduced to Slovakia

Ferdinand Tokár Kalinčiakova 3, Zlaté Moravce, Slovak Republic, e-mail: [emailprotected]

Abstract Tokár, F. 2013. Results of an ecological-production research on forest ecosystems of woody plants introduced to Slovakia. Folia oecol., 40: 282–289. The work gives achronological list of the results obtained in an ecological and production research on 49 coniferous and 10 broadleaved exotic woody plants in 298 parks and woody subjects across Slovakia. The results can be used in orchard and forestry practice. Since 1971, the research has been oriented to assessment of forest ecosystems and phytotechnique for forest stands consisting of selected exotic woody plants Pinus nigra Arnold, Castanea sativa Mill., Quercus rubra L. and Juglans nigra L. In the area of the Little Carpathians the best results in growth and production were achieved in Pinus nigra Arnold at the age of 100 years under proportion rate up to 30% in the group of forest types (slt) Querceto-fa*getum (464 m3 ha–1), fa*getum pauper (463 m3 ha–1) andfa*geto Quercetum (432 m3 ha–1). In the pure stands the highest stock was observed in the group of forest types Querceto-fa*getum (310 m3 ha–1). In Castanetarium Horné Lefantovce the best results out of 86 Castanea sativa progenies were obtained in 15 progenies (Jelenec 2, Horné Lefantovce A, Tlstý Vrch 1, 2, 2', 3, 4, 9, Duchonka 2, 3, 5, 6, 10, 12, Bratislava 4) and the worst results were obtained in seed progenies Stredné Plachtince 5, Krná 3, Modrý Kameň 7. Following evaluation of phytotechnique impact on production of different stand types of Castanea sativa Mill. at age of 38 years, the highest stock was observed in mixed stands Tilia cordata Mill. (416 m3 ha–1, 190 t ha–1, total production 635 m3 ha–1, 333 t ha–1). In mixed stands of Juglans nigra L. (20%) and Quercus rubra L. (80%) in the locality Ivanka pri Nitre, the highest stock was observed at the age of 48 years (438 m3 ha–1, 263 t ha–1) and total production 662 m3 ha–1 and 410 t ha–1. In the locality Sikenica in pure stands of Juglans nigra L. the highest stock at the age of 64 years was found in the stand with the strong crown thinning (464 m3 ha–1, 195 t ha–1, total production 573 m3 ha–1 and 246 t ha–1). In addiction to these production characteristics also leaf area indices were assessed (LAI). Keywords ecology, exotic woody plant, forest ecosystems, production

Analysis of the issue and the research focus In the past in Slovakia attention was given predominantly to the growth and production of autochthonous woody plant species (Halaj, 1963; Halaj and Řehák, 1979; Šebík and Polák, 1990; Šmelko, 2000). In allochthonous woody plant species issues of growth, production and distribution were evaluated (Holubčík, 1968; Benčať, 1982). The aim of our work is to introduce chronological survey of obtained results from ecologicproduction research. The Department of system and ecology of woody plants of the former Institute of Dendrobiology SAS in 282

the Arboretum Mlyňany SAS focused their research on forest ecosystems of woody plants introduced to Slovakia on the following points: o Taxonomy of exotic species in selected dendrological subjects in Slovakia (assortment, mensurational data, fertility, natural regeneration) o Valorisation of structure and production (volume, mass) and quality of various stands of selected exotic woody plants in Slovakia (Castanea sativa Mill., Quercus rubra L., Juglans nigra L., Pinus nigra Arnold) o Assessment of effects of phytotechnique (thinning) on production, dendrochronology, quality, leaf area

index (LAI) and energy potential in a variety of stand types of woody plants introduced into Slovakia o Monitoring of physiological-biochemical aspects of biomass production in various stand types of exotic woody plants (fluorescence, contents of selected elements in soil and leaves) o Assessment of resistance of stands of selected exotic woody plants against biotic and abiotic harmful agents o Evaluation of herb vegetation in various stand types of exotic woody plants and changes to this component due to long term introduction (Castanetarium in Jelenec, Castanetarium Horné Lefantovce) o Evaluation of natural regeneration of stands of exotic species in Slovakia o Quantitative assessment of selected chemical elements accumulated in aboveground biomass and in soil in stands consisting of exotic woody plants in Slovakia.

Material and methods The ecological description of the exotic species distribution in parks and dendrological objects in Slovakia has been adapted from Benčať, 1982. With using our own measured data and the data from Forest management plans (FMP), we have accomplished ecological-production analysis for 613 stands of black pine (Pinus nigra Arnold) in the region Malé Karpaty Mts. We considered the group of forest types (gft), stand age and structure and black pine proportion (1–30%, 31–60%, 61–90% and100%). The results were processed with using the Korf growth function, on a computer TESLA 200 in the Computing centre of the Technical University in Zvolen. The phytotechnique of various stand types (pure and mixed stands with different rates of domestic and alien woody plants) of Castanea sativa Mill., Quercus rubra L. andJuglans nigra L. works with thinning from above applied in graded intensity (moderate, heavy), with positive selection, at different repetition intervals (5–10 year), focusing on tending promising trees on the permanent research plots series (PRP) Žirany (7 partial PRPs with hom*ogeneous and mixed stands of Castanea sativa Mill.), Ivanka pri Nitre (6 partial PRPs with hom*ogeneous and mixed stands of Quercus rubra L. andJuglans nigra L.), Sikenica (3 partial PRP with hom*ogeneous stands of Juglans nigra L.) andCastanetarium Lefantovce (86 seed progenies of Castanea sativa Mill. from 12 localities in Slovakia). The ecological description of the PRP series Žirany, Lefantovce, Ivanka pri Nitre andSikenica can be found in Tokár, 1987, 1998; theCastanetarium Lefantovce is characterised in Tokár, 2003; Tokár and Kukla, 2006. Biometrical measurements of the stand height, diameter d1,3 and standing volume (volume production)

were performed by methods commonly used in forestry practice (Halaj, 1963; Šmelko, 2000). For the calculation of the volume of large black pine timber, we used, due to the lack of our own tables, the mass tables for forest pine, red oak, black nut; for edible chestnut the mass tables for oak converted per one hectare. The aboveground wood biomass was obtained by the destructive method (method of sample trees). The total number of sample trees for each woody species in the stand was determined by stratified selection (Šmelko and Wolf, 1977). The mass of stem, branches, annual shoots and leaves was obtained by weighing on a scale Kamor in dry mass at 105 oC. Photosynthetically active leaf surface area was estimated with the aid of aphoto-planimeter EIJKELKAMP. The time dependence of the values of standing volume and mass as well as the values of overall volume and mass production (growing stock + thinning + mortality + other losses) andthe LAI values was fitted with a mathematical function – specific for each tree species and stand type (an exponential or a 2nd degree polynomial). On each PRP series, the production results expressed through growth index and through index increment per cent were compared with the control PRP (without intervention) and tested statistically with the t-test (Šmelko and Wolf, 1977). The principle of phytotechnique of stands of exotic woody plants is tending the promising trees (Tokár 1987, 1998). The contents of elements (Mg, Ca, K, Na, Zn, Pb, Fe, Cu, Mn) in the aboveground biomass and in the soil were assessed with the aid of an absorption spectre-photometer IL VIDEO 12 (Tokár and Konôpková, 1995).

Results The ecology and production of exotic woody plants is an issue studied by the researchers in the „Arboretum Mlyňany“ – Institute of Dendrobiology SAS since 1966. Their activities began with an evaluation of growth and production performance of selected 59 exotic taxa (49 conifers, 10 broadleaves) in 298 parks and dendrological subjects in Slovakia. The results of this survey, useful for orchard management and, principally, for forest practices are in the works Tokár (1976, 1979). These results not only justify and confirm the success of introduction of these woody plants into our climatic conditions, mainly in terms of growth and production, but they also represent a new knowledge concerning fructification, natural regeneration and other important features (also concerning orchards – such as habitus). The results should be reputed as a valuable source of scientific knowledge about the gene pool of the cultural dendroflora in Slovakia waiting for use in dispersion, protection and saving of these taxa. 283

Beginning with 1976, the ecological-production research was oriented on valorisation of forest stands of exotic woody plants in the region Malé Karpaty Mts (in frame of the programme Man and Biosphere) and on phytotechnique of young forest stands of selected exotic woody plants (Castanea sativa Mill., Quercus rubra L. andJuglans nigra L.) on four PRP series (Žirany, Lefantovce, Ivanka pri Nitre andSikenica). In the region Malé Karpaty Mts (Tokár, 1985, 1991b), exotic woody plants are grown on 2,270 ha of the actual forest area and 1,579 ha of the reduced forest area. The major part concerns Pinus nigra Arnold (2,212 ha actual forest area and1,533 ha reduced forest area). Many minor proportions concern Pinus strobus L. (1.78 ha actual forest area), Pseudotsuga menziesii (Mirbel) Franco (26.07 ha), Aesculus hippocastanum L. (5.36 ha), Castanea sativa Mill. (17.53 ha), Quercus rubra L. (5.30 ha) andJuglans nigra L. (1.91 ha). As for the ecology, in the Malé Karpaty Mts, the exotic woody plants are most abundant in the groups of forest types (GFT): fa*geto-Quercetum (892 ha actual forest area), Corneto-Quercetum (490 ha), fa*getum pauper (302 ha) andQuerceto-fa*getum (169 ha). From the viewpoint of age, the first age class of 1–10 years (597 ha) and the sixth class encompassing 51–60 years (585 ha) are dominant. Black pine is mostly grown in the gft-s fa*getoQuercetum (867 ha), Corneto-Quercetum (490 ha), fa*getum pauper (295 ha), Corneto-fa*getum (156 ha), and Querceto-fa*getum dealpinum (100 ha). The results of the ecological-production analysis of the black pine in the Malé Karpaty Mts demonstrate that the most favourable conditions for growth and volume (mass) production in this region are in the gft-s Querceto-fa*getum, fa*geto-Quercetum andfa*getum pauper. The biggest overall standing volume is in the mixed stands – with the black pine proportion less than 30% (younger than 100 years, in gft FQ 432 m3 ha–1, in QF 464 m3 ha–1 andin Fp 443 m3 ha–1). In the pure stands, the highest standing volume was found in the gft-s Querceto-fa*getum (310 m3 ha–1) andfa*getoQuercetum (295 m3 ha–1). The mixed stands had by from 10% (Corneto-Quercetum) to 64% (fa*getum pauper) more volume stock than the pure stands. The results documenting the influence of phytotechnique on volume and mass production in the hom*ogeneous stands of Castanea sativa Mill. on the PRP series Žirany (Table 1, Figs 1–2) show that a stronger positive influence during the whole stand growth (years 1972–2001) was obtained in heavy thinning from above applied after each 10 years. The total mean increment in the 46-year-old trees was from 19.4 to 23.9 m3 ha–1 year–1 andfrom 10.5 to 13.3 t ha–1 year–1 (Tokár 1998, 2002). The phytotechnique (moderate thinning from above with positive selection and 5-year interval of repetition) in the pure (Fig. 3) and mixed stands of Castanea sativa Mill. on the PRP series Lefantovce (Table 284

2) resulted in better volume and mass production in the mixed (Castanea sativa Mill. + Tilia cordata Mill. (Fig. 4), Castanea sativa Mill. + Pinus sylvestris L.) stands than in the pure Castanea sativa Mill. stands (in volume by 5.31%–34.3%, in mass by 10.53%– 31.60%). The cause underlying the better production in the mixed stands compared to the pure ones should be assigned to the favourable allopathic relations and soil conditions developed in these stands. The total mean increments in the 38-year-old trees were from 14.42 to 19.17 m3 ha–1 year–1 and from 7.23 to 9.79 t ha–1 year–1 (Tokár, 2002; Tokár and Krekulová, 2003, 2004). Table 1. Volume and mass production in the pure stands of Castanea sativa Mill. on the PRP series Žirany in 2001 (stand age 46 years) PRP I II III IV V

Thinning degree Moderate

Growing stock Interval 5

Total production

m3 ha–1

t ha–1

m3 ha–1

t ha–1

536

271

939

493

Moderate

566

260

894

486

Heavy

749

372

1,102

612

Moderate

621

292

888

487

Heavy

755

384

1,094

597

Control

VII

Heavy

676 10

634

325

876

483

331

965

541

Fig. 1. Stem of a high-quality edible chestnut (Castanea sativa Mill.) tree on the PRP Žirany (photo F. Tokár).

V (Control)

214

105

358

204

220

124

316

173

Together

434

229

674

377

Castanea sativa

200

322

186

Pinus sylvestris L.

171

263

130

Together

371

168

585

316

Castanea sativa Mill. Pinus sylvestris L.

Mill.

Fig. 2. hom*ogeneous stand of edible chestnut (Castanea sativa Mill.) on the PRP Žirany (photo F. Tokár).

Table 2. Volume and mass production in various stand types of Castanea sativa Mill. on the PRP series Lefantov- ce in 2001 (stand age 38 years) Growing stock Partial PRP

Species

m3 ha–1

Total production m3

t –1

t ha–1

–1

174

548

I (Control)

Castanea sativa Mill.

356

Castanea sativa Mill.

336

152

556

282

Castanea sativa

195

411

211

221

325

149

Together

416

182

736

360

Castanea sativa

261

129

426

232

150

209

101

411

190

635

333

III (Control)

292

Mill. Tilia cordata Mill.

Mill. Tilia cordata Mill. Together

Fig. 3. hom*ogeneous stand of edible chestnut (Castanea sativa Mill) on the PRP Lefantovce (photo F. Tokár).

Valuable ecological-production results were attained in the Castanetarium Horné Lefantovce (14.38 ha) by improving growth and production of 86 seed progenies of edible chestnut from 12 localities in Slovakia (Benčať and Tokár, 1978). In a tree age of 35 years, very good results were obtained in 15 seed progenies (Tokár, 2003). The production and resistance potential has been evaluated in Tokár et al., 2004. The results of assessment of soils and phytocoenoses in the Castanetarium Horné Lefantovce and in the Castanetarium Jelenec showed that the edible chestnut was an important factor causing changes in the phyto-

285

Fig. 4. Thinning PRP of edible chestnut and small-leaf linden on the PRP Lefantovce (photo F. Tokár).

coenoses. The phytocoenoses in these localities belong into the 3rd forest vegetation tier, the group of forest types fa*getum pauper inferiora (Tokár and Kukla 2005, 2006). In the pure (Fig. 5) and mixed stands of Quercus rubra L. andJuglans nigra L. on the PRP series Ivanka pri Nitre (Table 3), the overall production was most effectively controlled by moderate thinning from above

with positive selection and repetition interval of 5 years in the mixed stands of Juglans nigra L. andQuercus rubra L. or Tilia cordata Mill. (Fig. 7). The overall mean increments in the trees aged 48 years were from 12.76 to 16.29 m3 ha-1 year-1 and from8.16 to 11.54 t ha-1 year-1 (Tokár 1991a, 1998, 2005). In the pure stands of Juglans nigra L. on the PRP series Sikenica (Table 4, Fig. 6), stronger posi-

Table 3. Volume and mass production of various stand types of Quercus rubra L. and Juglans nigra L on the PRP series Sikenica in 2003 (stand age 48 years) Partial PRP I

II III

Species

Proportion

Age

[%]

[years]

m ha

t ha

m ha–1

t ha–1

Quercus rubra L.

Juglans nigra L

402

343

552

421

434

367

613

469

438

263

662

410

Together

100

Quercus rubra L.

100

Growing stock 3

–1

Total production –1

Quercus rubra L.

304

216

460

331

Juglans nigra L.

175

125

242

162

Together

100

479

341

702

493

Juglans nigra L.

100

430

320

630

416

Juglans nigra L.

369

258

407

281

Tilia cordata Mill. Together VI (Control)

132

180

501

314

581

367

Quercus rubra L.

426

293

505

355

Juglans nigra L.

261

194

287

206

687

487

792

561

Together

286

100

Fig. 5. hom*ogeneous stand of red oak (Quercus rubra L.) on the PRP Ivanka pri Nitre (photo F. Tokár).

Fig. 7. Mixed stand of blacknut with small-leaf linden on the PRP Ivanka pri Nitre (photo F. Tokár).

tive impacts on the overall volume and mass production in years 1979–2003 were found for heavy thinning from above with positive selection and 5-year interval of repetition. The overall mean increments in the 64-year-old trees were from 7.22 to 8.95 m3 ha–1 year–1 and from3.31 to 3.84 t ha–1 year–1 (Tokár 1992, 1998; Tokár and Krekulová 2005).

Table 4. Volume and mass production in the pure stands of Juglans nigra L. on the PRP series Sikenica in 2003 (stand age 64 years) PRP

Thinning degree

Growing stock 3

–1

m ha

–1

t ha

Total production m3 ha–1

t ha–1

III

Moderate

381

173

468

215

Heavy

464

195

573

246

Control

454

208

462

212

287

References

Fig. 6. hom*ogeneous stand of blacknut (Juglans nigra L.) with natural regeneration on the PRP Sikenica (photo F. Tokár).

On all PRP series, we used thinning methods focused on tending promising trees, selected from the trees with suitable quantitative and qualitative parameters (Tokár, 1987, 1998). The content of elements in aboveground biomass and in soil in the forest stands composed of exotic woody plants varied with the plant species and the biomass compartment (e.g. Ca bark, stem; Kleaves, Na stem xylem) (Tokár and Konôpková, 1995). In the forests of Slovakia (primarily in southern areas ofgft Carpineto-Quercetum), black locust (Robinia pseudoacacia L.) – one of the first woody plants introduced to Europe, has aspecific status. Today the black locust forest stands represent about 34,000 ha, which is 1.87% of the total forest land area in Slovakia. The black locust production in forest stands in SW Slovakia was evaluated by Benčať (1988). The destructive method used (sample trees) for assessment of aboveground biomass production in model stands of exotic woody plants was also suitable for deriving eco-physiological characteristics of these stands and woody plants (leaf area index – LAI, biomass production per leaf area unit, and similar) (Tokár 1987, 1998; Konôpková, 2003; Kmeť and ŠalgovičovÁ, 2003; Šalgovičová and Kmeť, 2004).

288

Benčať, F. 1982. Atlas rozšírenia cudzokrajných drevín na Slovensku arajonizácia ich pestovania [Atlas of distribution of exotic woody species in Slovakia and zoning of their cultivation]. Bratislava: Veda. 368 p. Benčať, T. 1988. Black locust biomass production in Southern Slovakia. Bratislava: Veda. 192 s. Benčať, F., Tokár, F. 1978. Výsledky fenologického pozorovania gaštana jedlého (Castanea sativa Mill.) na experimentálnej ploche vHorných Lefantovciach [Results on phenological observations of the chestnut tree (Castanea sativa Mill.) on experimental plot in Horné Lefantovce. Folia dendrol., 4: 49–89. Halaj, J. 1963. Tabuľky na určovanie hmoty a prírastku porastov [Tables to the determination of mass and increment of forest stands. Bratislava: SVPL. 328 p. Halaj, J., Řehák, J. 1979. Rastové tabuľky hlavných drevín ČSSR [Yield tables of main woody plants in the ČSSR]. Bratislava: Príroda. 325 p. Holubčík, M. 1968. Cudzokrajné dreviny vlesnom hospodárstve [Exotic woody plants in forest management]. Bratislava: SVPL. 371 p. Kmeť, J. Šalgovičová, A. 2003. Ecophysiological aspect of growth of the European chestnut (Castanea sativa Mill.) in Slovakia. Folia oecol., 30: 141–147. Konôpková, J. 2003. Produkcia, energetický ekvivalent a obsah živín vybraných drevín. Autoreferát dizertačnej práce [Production, energy equivalent and nutrient content of selected woody plants. Shortened version of PhD thesis]. Nitra: Slovenská poľnohospodárska univerzita vNitre. 32 p. Šalgovičová, A., Kmeť, J. 2004. Influence of site conditions on physiological status of black walnut (Juglans nigra L.) stands. Folia oecol., 31: 100–110. Šebík, L., Polák, L. 1990. Náuka oprodukcii dreva [Wood production science]. Bratislava: Príroda. 322 p. Šmelko, Š. 2000. Dendrometria [Dendrometry]. Zvolen: Technická univerzita vo Zvolene. 399 p. Šmelko, Š., Wolf, J. 1977. Štatistické metódy vlesníctve [Statistical methods in forestry]. Bratislava: Príroda. 330 p. Tokár, F. 1976. Rastové a produkčné schopnosti vybraných lesnícky upotrebiteľných cudzokrajných ihličnatých drevín v parkoch na Slovensku [Yield and production abilities of selected forestry usable exotic coniferous trees in parks in Slovakia]. In Benčať, F. Štúdie o ihličnatých drevinách. Zborník Dendrologickej sekcie Československej botanickej spoločnosti. Bratislava: Veda, p. 201–211. Tokár, F. 1979. Zhodnotenie vybraných cudzokrajných listnatých drevín na Slovensku z hľadiska ich rastu a možnosti pestovania [Evaluation of selected exotic broadleaved woody plants from viewpoint of their growth and cultivation possibilities]. Acta dendrobiologica, 1–2. Bratislava: Veda, p. 119–146.

Tokár, F. 1985. Rozšírenie cudzokrajných drevín v lesných porastoch Malých Karpát a ekologickoprodukčná analýza ich hlavných druhov [Distribution of exotic woody plants in the Little Carpathian stands and an ecological-production analysis of the main species]. Lesnictví, 31, 6, s. 501–518. Tokár, F. 1987. Biomasa vybraných cudzokrajných drevín v lesných porastoch juhozápadného Slovenska [Biomass of selected exotic woody plants in forest stands of south-western Slovakia]. Acta dendrobiologica. Bratislava: Veda. 116 p. Tokár, F. 1991a. Vplyv úrovňových prebierok na objemovú a hmotnostnú produkciu nadzemnej biomasy rôznych typov porastov Quercus rubra L. a Juglans nigra L. [Influence of crown thining on volume and weight production of above ground biomass of various stand types consisting of red oak (Quercus rubra L.) and black walnut (Juglans nigra L.)]. Lesn. Čas., 37: 349–362. Tokár, F. 1991b. Výskyt a produkcia vybraných cudzo krajných drevín v lesných ekosystémoch Malých Karpát [Occurence and production of selected exotic woody plants in forest ecosystems in the Litte Carpathians]. Acta dendrobiologica. Bratiskava: Veda. 128 p. Tokár, F. 1992. Vplyv prebierok na vývoj objemovej a hmotnostnej produkcie u nezmiešaných porastov orecha čierneho (Juglans nigra L.) [Influence of thinning on development of volume and weight production of pure black walnut (Juglans nigra L.)]. Lesn. Čas. – For. J., 38: 189–203. Tokár, F. 1998. Fytotechnika a produkcia dendromasy porastov vybraných cudzokrajných drevín na Slovensku [Phytotechnics and dendromass production of selected exotic woody plants in Slovakia]. Acta dendrobiologica. Bratislava: Veda. 157 s. Tokár, F. 2002. Growth and production of dendromass in European chestnut stands (Castanea sativa Mill.) in Slovakia and their phytotechnics. Ekológia (Bratislava), 21, Suppl. 2: 124–142. Tokár, F. 2003. Growth, production and quality of 35-year old seed progenies of European chestnut (Castanea sativa Mill.). Folia oecol., 30: 99–106.

Tokár, F. 2005. Dub červený (Quercus rubra L.) a orech čierny (Juglans nigra L.) – významné cudzokrajné dreviny vsadovníctve, lesnej a poľnohospodárskej krajine [Red oak (Quercus rubra L.) and black wall nut (Juglans nigra L.) – important exotic woody plants in landscaping, forest and agricultural landscape]. In Bernadovičová, S., Juhásová, G. (eds). Dreviny vo verejnej zeleni. Zborník z konferencie s medzinárodnou účasťou. Bratislava, 10. – 11. 5. 2005. Zvolen: Ústav ekológie lesa SAV, p. 215–222. Tokár, F., Juhásová, G., Bernadovičová, S., Adamčíková, K., Kobza, M., Pavlíková, A. 2004. Production and resistance potential of European chestnut (Castanea sativa Mill.) in the Castanetarium Horné Lefantovce. Folia oecol., 31: 40–52. Tokár, F., Konôpková, J. 1995. Fytotechnika a dynamické zmeny obsahu vybraných chemických prvkov v nadzemnej dendromase rovnorodých porastov gaštana jedlého (Castanea sativa Mill.) [Silvicultural practices and dynamic changes in the content of some chemical elements in aboveground dendromass of pure stands of the Spanish chestnut (Castanea sativa Mill]. Lesnictví – Forestry, 41: 125–131. Tokár, F., Krekulová, E. 2004. Aboveground biomass production and leaf area index in various types of chestnut (Castanea sativa Mill.) stands in Slovakia. Ekológia (Bratislava), 23: 342–352. Tokár, F., Krekulová, E. 2005. Influence of phytotechnology on growth, production and leaf area index of black walnut (Juglans nigra L.) monocultures in Slovakia. J. Forest Sci., 51: 213–224. Tokár, F., Kukla, J. 2005. European chestnut (Castanea sativa Mill.) aboveground dendromass and its impact on composition of phytocoenoses in Jelenec Castanetarium PA. Ekológia (Bratislava), 24: 217–230. Tokár, F., Kukla, J. 2006. Ecological conditions in the Castanetarium Horné Lefantovce and growth of European chestnut (Castanea sativa Mill.). Ekológia (Bratislava), 25: 188–207.

Received December 12, 2012 Accepted May 27, 2013

289

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Short communication

Preservation and restoration of living plant collections on the example of the Buda Arboretum of Corvinus University, Budapest Gábor Schmidt1, Magdolna Sütöri-Diószegi2 Department of Floriculture and Dendrology, Corvinus University of Budapest Faculty of Horticultural Sciences, 1118 Budapest, Villányi út 29-43, tel: +36 1/482 6461, Hungary, 1 e-mail: [emailprotected], 2e-mail: [emailprotected]

Abstract Schmidt, G., Sütöri-Diószegi, M. 2013. Preservation and restoration of living plant collections on the example of the Buda Arboretum of Corvinus University, Budapest. Folia oecol., 40: 290–294. The Buda Arboretum was initiated in the winter of 1893/94. Now it covers 7.5 hectares and is surrounded by the constantly growing city of Budapest. At present, the Arboretum is under very strong urban effect. Within the framework of a EU-project “Preservation and Restoration of Living Plant Collections and Historical Gardens” the Buda Arboretum was profoundly reconstructed and developed between 2010– 2012. There were reconstructed selected objects serving to special purposes, e.g.: 1. Special biotypes (garden pond and the surrounding wetland, rock-gardens, pergolas for the climbing plants; a retaining wall giving shelter for the Mediterranean collections; greenhouse as a biotope for tropical and subtropical plants), 2. The historic geometrical garden part (called Parade Square), 3. Ecological solutions for water supply, 4. Suppression of invasive species and development of Laurocerasus, Malus, Potentilla, Prunus, Syringa collections, wetland-perennials, collection of Hungarian bred woody ornamentals introduction and trial of new Mediterranean species, etc. After reconstruction, the plant material includes over 1,900 woody species and cultivars, more than 240 kinds of bulb-flowers, 500 different perennials, 250 annuals and round about 300 tropical and subtropical (greenhouse) taxa. Keywords arboretum, Buda Arboretum of Corvinus University of Budapest, draught- and pollution-tolerance, global warming-up, heat-tolerance, woody ornamentals

Introduction and review of literature The Buda Arboretum is one of the richest plant collections in Hungary. It was initiated in the winter of 1893/94 on 3 hectares, on the premises of the Horticultural School (the predecessor of the present Faculty of Horticultural Sciences) (Räde, 1943). The other parts of the territory were utilized by orchards, vineyards, and glasshouses for ornamental plants and vegetables, according to the profile of the School. Later, the fruit- and vine-plantations and the glasshouses were moved to the outskirts of the city and the whole site was reverted to 290

an arboretum. The different steps of the process were described in works of Schmidt (1994), Zalainé (2003), Probocskai (1994), Hámori and Schmidt (2003). Now it covers 7.5 hectares and is surrounded by the constantly growing city of Budapest. The site is situated on the southern foothill of the 235 m high Hill of Gellért. The original vegetation was probably a mixed carstwood forest (Ceraso mahaleb-Quercetum and Orno-Quercetum), with some elements of mixed floodplain hardwood forest (Fraxino pannonicae-Ulmetum) (Facsar, 2008). At present, the Arboretum is under very strong urban effect: the summer is hot, the

winter is mild, the air is polluted. The Buda Arboretum has been protected by law as a natural reserve (living gene collection of woody plants) since 1974 and also as a historical garden since 2005 (Csepely-Knorr and Sárospataki, 2009). The collections serve three main purposes: 1) education of students and public (a “living textbook”); 2) display of Hungarian-bred woody ornamental cultivars, and 3) testing, examination and trying out of plants of subtropical and Mediterranean origin in order to show the possible benefits of urban microclimate and also as potential plant materials for the case of global warming (Schmidt, 2008). In 2010, a considerable EU-fund was earned (Project No KMOP-3.2.1/B-09-2009-0003) for the reconstruction and the development of the Arboretum. The first publications reporting on the funding and the preliminary results were published in Hungarian language by Schmidt and Sütöri-Diószegi, M., 2011; Honfi et al., 2012a; Honfi et al., 2012b; Schmidt and Sütöri-Diószegi, 2010. The head of project management was prof. Károly Hrotkó, the head of the reconstruction and planting was prof. Gábor Schmidt, the coordinators were dr. Peter Honfi and dr. Magdolna Sütöri-Diószegi.

and humid alpine biotypes (rock-gardens with collections from plants of dry native hills and also true alpine plants) were reconstructed on 1,400 m2; 1.3. Pergolas for the climbing plants were reconstructed on 240 m2 (Fig. 2.); and 1.4. South-facing retaining wall as a biotope for the open-ground Mediterranean woody plant collections. A retaining wall giving shelter from the north, is extremely dry and warm and offers excellent conditions for true Mediterranean plants like cypresses (Cupressuses), Yuccas, pomegranates (Punica granatum L.), hardy cactuses (Cactaceae Ivss) and others. The Albizia julibrissin (Willd.) Durazz. tree brings a profusion of soft pink mimosa-like flowers from July through September. Also here grow specimens of the bead-tree (Melia azedarach L.) and the holly oak (Quercus ilex L.). Before reconstruction the wall was partially ruined and dangerous for life. After reconstruction it became safe and the area for Mediterranean collection increased by 600 m2 (list of plants see later).

Materials and methods The reconstruction-project started on 1 June 2010 and ended on 31 March 2012. The main parts (sub-projects) of the project were as follows: 1. Reconstruction of special biotypes 2. Reconstruction of a historical geometric part of the garden 3. Ecological solutions for heating and for water supply 4. Suppression of invasive species in the hardy plant collections. Each of the mentioned elements needed different approach and methods. For the sake of simplicity, these methods will be described in the next chapter only.

Fig. 1. Garden pond.

Results and discussion The results (and also the lessons) of the reconstruction project are as follows (see also the Figs 1–6). 1. Reconstruction of special biotypes Because of the limited space, the present paper will concentrate mainly on the most important woody plant collections. The biotypes of herbaceous collection and those for minor woody collections will be shortly mentioned only, and illustrated by some photos. Such are: 1.1. Wetland biotypes (the garden pond and the surrounding artificial wetland, Fig. 1.); 1.2. Dry carstland-

Fig. 2. Pergola for the climbing plants.

291

2. Reconstruction of the historical geometric garden – part called Parade Square. The Buda Arboretum is maintained as a natural plant protection and also is registered and protected as a historical garden. The most characteristic part of it is the 3,000 m2 large geometrical garden section called Parade Square. In the past, the square was fully planted with herbaceous flower-beds as well as with roses – hence the name. Now that park-maintenance became too costly, the former baroque style is just symbolised by two symmetrically arranged groups of arborvitae (Thuja ssp.), the regular outlines of the lawn and by some adjacent bedding plants. The statue in the upper centre (in front of Building F) commemorates the famous fruit-breeder Máté Bereczki (Fig. 3).

Fig. 4. Greenhouse for tropical and subtropical plant collection.

Fig. 3. Parade Square

3. Environmental-friendly solutions for heating and for water supply 3.1 Energy-saving solutions for heating and cooling The Arboretum contains a relatively small glasshouse (110 m2) for the tropical and subtropical ornamental plants. The glasshouse is 20 years old and, before reconstruction, it was far outdated and in a very bad condition. The heating during the winter (with gas) needed a lot of energy and money, and the cooling in the summer was carried out with outdated heaters (pipes) and methods. Simply said, the air-conditioning was insufficient for the plants and yet, very expensive (Fig. 4). 3.2 Reutilization of run-off water from the roofs of the buildings Several solutions were used for reutilization of run-off rainwater, the best of which are shown on Fig. 5.

292

laurocerasus L. collection: 34, other Prunus collections include the following sub-genera: Amygdalus, Cerasus, Padus, Prunus: 74 taxa; Syringa collection: 46 taxa.

Fig. 5. Collection of run-off rainwater from the roofs of the buildings.

4. Suppression of invasive species and development of hardy perennial and woody plant collections 4.1 Suppression of invasive species This work included the regional removing of herbaceous weed and also the moving woody species, first of all: Ailanthus altissima (Mill.) Swingle, Acer negundo L., Clamatis vitalba L., Fraxinus pennsylvanica Marsh., Parthenocissus quinquefolia (L.) Planch., Cotoneaster multiflorus Bunge, Diospyros lotus L. 4.2 Development of hardy plant collections 4.2.1 Woody plant collection Hibiscus collection: 26 taxa (Fig. 6 a); Malus collection: 42 taxa (Fig. 6 b); Potentilla collection: 38 taxa; Prunus

Fig. 6a. Hibiscus syriacus ’Minerva’.

Fig 6b. Malus ’Professor Sprenger’.

– Collection of Hungarian bred woody ornamentals: 74 taxa: Betula pendula Roth cv. Karaca, Buxus microphylla Sieb. et Zucc. cv. Betlér, Campsis × tagliabuana (Vis.) Rehd. cv. Galen Select, Campsis radicans (L.) Seem. ex Bureau cv. Barack, Chamaecyparis lawsoniana (A. Murray) Parl. cv. Tekeres, Cotoneaster salicifolius Franch. cv. Rózsaszín Füzér × Cupressocyparis notabilis (A. F. Mitchell.) Farjon cv. Márta, Hedera helix L. cvs. Arács, Balkon, Blue Star, Börzsöny, Csocsoszan, Duna, Krokó, Marble, Negro, Perint and Zebegény, Juniperus × media Van Melle cv. Mint Julep Tarka, Juniperus chinensis L. cvs. Eldorado, Favorit, Gold Rush, Juniperus communis L. cv. Fancsika, Juniperus conferta Parl. cv. Sláger, Juniperus sabina L. cvs. Báránd, Szőke Tisza and Tarka, Juniperus virginiana L. cvs. Golden Rain and Little Mityu, Picea pungens Engelm. cv. Edith, Pinus sylvestris L. cv. Sé, Prunus cerasifera Ehrh. cv. Colos, Prunus laurocerasus L. cvs. Ani, Antonius, Cipora, Cleopátra, Gabi, Hagar, Leander, Parviflora and Zita, Prunus padus L. cvs. 1/a sz. klón, 6 sz. klón, Aurora, Piros Oszlop (13. sz. klón) and Rózsaszín Május, Prunus persica cv. Orlóci Kiméra, Pyrus pyraster cv. Bihar, Rosa hybrids: cvs. Arany János, Bethlen Gábor, Házsongárd, Hild József, Máramaros, Nagyhagymás, Nyitra, Regéc, Szent Imre and Szent Margit, Salix matsudana Koidz. cv. Tarkabarka, Sorbus bakonyensis Jáv., S. borbasii Jáv. cv. Herkulesfürdő, S. borosiana Kárp. cv. Alba Regia, S. 293

cv. Hainburg, Taxus baccata cv. Zöld, Thuja occidentalis L. cvs. Miki, Romantika, Szőllősi Klón (Malonyana Aurea), Thuja orientalis cvs. Dundi, Hunor, Jászkiséri, Lakatos and Thuja orientalis L. cv. Telihold, Tilia platyphyllos Scop. cv. Favorit, Tilia platyphyllos Scop. cv. Pannonia. – Introduction and trial of new Mediterranean species: 61 taxa. In 2011–12, the following new tender species and cultivars were planted and tired: Acca sellowiana (O. Berg) Burret, Albizia julibrissin Durazz. cv. Summer Chocolate, Berberis darwini Hook.; Caesalpinia gilliesii (Wallich ex Hook.) Wallich ex D. Dietr.; Callistemon citrinus (Curtis) Skeels; Ceanothus delilianus Spach. cv. Gloire de Versailles; cv. Henri Defossé; Ceanothus pallidus Lindl. cvs. Marie Simon; Perle Rose, Cistus corbariensis Pourr.; C. pulverulentus Pourr. cv. Sunset; C. purpureus Lamn.; Cistus purpureus Lamn. cv. Alan Frad; Cordyline australis (Forst. f.) Hook. f.; Cotoneaster lacteus W.W.Sm.; Elaeagnus × ebbingei Boom ex Doorenb. cvs. Clône Erigé, Compacta, Eleador, Gilt Edge, Limelight; Escallonia cvs. Apple Blossom, Crimson Spire, Donard Seedling; Eucalyptus gunnii Hook f., Hebe arts; Itea virginica L. cv. Little Henry, Jasminum officinale L., Lagestroemia hybrids: cvs. Appalache, Hopi, Nivea, Pecos, Petite Pink, Rosea Nova, Rouge, Togo, Tonto, Lavandula angustifolia cvs. Hidcote, Munstead, Rosea; Lavandula × intermedia Loisel. cvs. Abrial, Chamallow, Edelweiss, Grosso, Imperial Gem; Leycesteria formosa Wall., Mahonia japonica Thunb.; Nandina domestica Thunb. cvs. Firepower, Richmond and Wood’s Dwarf; Osmanthus heterophyllus (G. Don) P. S. Green cvs. Goshiki, Purpureus, Phormium tenax J. R. Forst. & G. Forst.; Photinia × fraseri Dress cvs. Camilvy, Nana, Pink Marble; Prunus lusitanica L.; Punica granatum L. cvs. Chico; Maxima Rubra, Teucrium fruticans L.; Viburnum tinus L. cvs. Eve Price and Gwenlian. 4.2.2 Perennial plant collections Dryland perennials: from 60 to 75 taxa, wetland-perennials: from 20 to 70 taxa, alpine perennials: from 40 to 110 taxa.

Acknowledgement The project was supported by KMOP3.2.1/B-09-2009-0003 and TÁMOP-4-2.1.B-09/1/ KMR- 2010-0005 EU-project. The Authors would like to express their special thanks to the organizers of the excellent Conference in Mlyňany Arboretum (Sept. 18– 19., 2012) and also to Stephan Bakay for completing the Slovakian abstract of this paper.

References Csepely-Knorr, L., Sárospataki, M. 2009. “Gellérthegyi Paradicsom” – A Budai Arborétum Felső kertjének építéstörténete a II. világháborúig [The building history of the Upper Garden of the ’Budai Arboretum’ until world war II]. 4D Tájépítészeti és Kertművészeti, 14: 2–25 Facsar, G. 2008. Az eredeti növényzet rekonstrukcciója [Reconstruction of the original vegetation]. In Schmidt, G. A Budapesti Corvinus Egyetem Budai Arborétuma. Budapest: Mezőgazda Kiadó, p. 5. Hámori, Z., Schmidt, G. 2003. A Budai Arborétum története [History of the Buda Arboretum]. Lecture. Lipai, J., Ormos, I., Vas, K. (persons responsible for session). Tudományos Ülesszak. Budapest, 6–7. 11. 2003. Honfi, P., Sütori-Diószegi, M., Schmidt, G. 2012a. Megújult zöld oázis. A Budai Arborétum [A reconstructed oasis: The Buda Arboretum]. Természetbúvár, 67 (3): 36–38. Honfi P., Czigány K., Kohut I., Schmidt G., SütörinéDiószegi, M. 2012b. A megújult Budai Arborétum [The Buda Arboretum after reconstruction]. Budapest: Budapesti Corvinus Egyetem, Kertészettudományi Kar, p. 4–32. Probocskai, E. 1994. Adatok a kertészeti és élelmiszeripari egyetem 100 éves Arborétumának történetéhez [Additional information to the history of 100 years old Buda Arboretum]. Publ. Univ. Hort. Indusriaeque Alimentariae. Budapest, 44: 2–6. Räde, K. 1943. Ötven évvel ezelőtt [Fifty years ago]. Kert. Sz., 67–70. Schmidt, G. 1994. The Buda Arboretum of the University of Horticulture and Food Industry. Budapest: Marton Press, p. 1–68. Schmidt, G. 2008. A BCE Budai Arborétuma [The Buda Arboretum of Corvinus University of Budapest]. Budapest: Corvinus University KeTK, p. 3–24. Schmidt G., Sütöri-Diószegi M. 2010. Testing urban climate with heat-tolerant woody plants in the Buda Arboretum. Acta Horticulturae et Regiotecturae, 13, Spec. 2: 37–41. Schmidt, G., Sütöri-Diószegi, M. 2011. Magyar nemesítésű díszfák-díszcserjék gyűjteményének fejlesztése a BCE Budai Arborétumában [Development of Hungarian woody cultivar collections in the Buda Arboretum]. Kertgazdaság, 43 (4): 60–68. Zalainé Kovács, É. (ed.) 2003. 150 év a kertészettudományi, élelmiszertudományi és tájépítészeti oktatás szolgálatában [150 years in the service of horticultural, food industrial and landscape sciences]. Budapest: BKÁE Kertészettudományi Kar, Élelmiszer-tudományi Kar, Tájépítészeti, -védelmi és -fejlesztési Kar, p. 13–18., p. 141–164. Received December 6, 2012 Accepted March 22, 2013

294

FOLIA OECOLOGICA – vol. 40, no. 2 (2013). ISSN 1336-5266

Survey paper

The Primeval Beech Forests of the Carpathians and Ancient Beech Forests of Germany: joint natural heritage of Europe Ivan Vološčuk1, Viliam Pichler2, Magdaléna Pichlerová3 1

Institute for Research of Landscape and Regions, Faculty of Natural Sciences, Matej Bel University, Cesta na amfiteáter 1, 974 00 Banská Bystrica, Slovak Republic, e-mail: [emailprotected] 2 Faculty of Forestry, Technical University in Zvolen, T. G. Masaryka 24, 960 53 Zvolen, Slovak Republic, e-mail: [emailprotected] 3 Faculty of Ecology and Environmental Sciences, Technical University in Zvolen, T. G. Masaryka 24, 960 53 Zvolen, Slovak Republic, e-mail: [emailprotected]

Abstract Vološčuk, I., Pichler, V., Pichlerová, M. 2013. The Primeval Beech Forests of the Carpathians and Ancient Beech Forests of Germany: joint natural heritage of Europe. Folia oecol., 40: 295–303. The European beech fa*gus sylvatica L. ssp. sylvatica L. is exclusively found in Europe. The beech survived the last ice age in small refuges in the south and south-east Europe and went on the colonisation of large parts of the continent. The post ice colonization of the landscape by the beech took place parallel to the settlement of land by humans and the formation of amore complex society. For centuries much of the Carpathian mountain forests remained untouched. Virgin forests constitute a natural heritage of global significance. In 2007 the primeval beech forests of the Carpathians (Slovakia, Ukraine) were added to UNESCO´s World Heritage List. On 25 June 2011, the UNESCO World Heritage Committee added five of Germany´s beech forest regions to the World Heritage List. This extended the transboundary world natural heritage site „Primeval Beech Forest of the Carpathians“, located in the Slovak Republic and Ukraine, to include anumber of German forest regions, and renamed it „Primeval Beech Forests of the Carpathians and Ancient Beech Forests of Germany“. The paper is aimed at the presentation of the outstanding universal value of the ecological processes in the Joint World Heritage Sites, and present principles of their Integrated Management Plan. Ultimate goal is to achieve that management and socio-economic sustainable development practices are in harmony with primary objectives of WHS protection, biodiversity conservation, ecosystem and landscape stability, rational use of natural resources, ecotourism development and with potential of the landscape in largest possible extend. Keywords ancient beech forest, Carpathy, Germany, primeval beech forest, World Heritage

Introduction Europe´s beech forests are deciduous forests which are dominated by the European Beech (fa*gus sylvatica L.). The beech is endemic to Europe and beech forests are limited to Europe (Gömöry et al., 2011). Such forests therefore share the fate of all deciduous forests of the

northern hemisphere´s nemoral zone. They have been exposed to an enormous development pressure (settlement, utilisation) for centuries so that natural forests have become scarce (Britz et al., 2009). Beech is one of the most important elements of forests in the Temperate Broad-leaf Forest Biome and represents an outstanding example of the re-colonisation and develop295

ment of terrestrial ecosystems and communities after the last ice age, aprocess which is still ongoing (Knapp, 2011). Forest communities built up and dominated by the beech are widespread across major parts of Central Europe. Potentially forming the predominant zonal vegetation in Western and Central Europe in terms of area, they are found at the montane level of the South European mountain ranges. They show the widest amplitude of soil trophic levels and altitude distribution, of all deciduous forests in Europe potentially occupying the largest area (Bohn and Neuhäusl, 2003). The European beech forests stand out due to an exceptional variety of types. According Bohn and Neuhäusl (2003), atotal of 86 different biocoenotic units of the beech and mixed beech forests are found in the beech forest area, subdivided according to trophic and altitude levels as well as geographical and local forms. Of these units, 14 cover more than 50% of the potential natural range, with as many as eight units being also widespread in Germany with significant proportions of the overall area. Atotal of 28 biocoenotic units, which roughly equals one-third of all European units, are widespread in Germany, which emphasises Germany´s particular responsibility for the preservation of the beech forests worldwide (Britz et al., 2009). The European beech forests show adecline in vascular plant species numbers from glacial refuges in Southern Europe to the north and northwest, in which directions they were advancing. Their centres of diversity lie in the Eastern Carpathians, the Dinaric Alps, and the Pyreneans (Dierschke and Bohn, 2004). The particular evolutionary connection clearly reflects in the entire Central European Flora. The different beech forest types are home to 20% of the terrestrial fauna in Central Europe – 7,000 to 10,000 animal species (Otto, 1994) that have mostly adapted their rhythm of life to the seasonal cycle. Alongside with the plants, fungi, and microorganisms, they are the determining factors in the beech forest system. The history of the beech forests is closely linked with the history of European civilisation (Bennett, 1994; Britz et al., 2009). The post-glacial colonisation of the landscape by the beech tree ran in paralel with the establishment of communities by mankind and the formation of more highly organised forms of society. That is why the beech is deeply rooted in European culture (Pichler et al., 2007a).

Material and methods The beech ecosystem research which has been the basis for elaborating on the World Heritage Nomination Project was carried out in two regions: in the Carpathian Mts and in the German Lowlands.

296

The complete ecological research of the mountain Primeval Beech Forests of the Carpathians started in the first half of the 20th century due to the famous Czech botanist and forest ecologist Professor Alois Zlatník (Zlatník, 1934, 1935, 1936; Zlatník and Hilitzer, 1932; Zlatník et al., 1938). Valuable knowledge concerning ongoing ecological processes in the Carpathian primeval beech forest ecosystems has been obtained after Second World War during the past years (Leibundgut, 1978; Jaworski et al., 1994a, 1994b; Korpeľ, 1989, 1995; Kricsfalusy et al., 2001; Commarmot et al., 2000; Bublinec and Pichler, 2001; Saniga, 2011; Saniga and Schütz, 2002; Stoyko et al., 1982; Stoyko and Tasenkevitch, 1993; Stoyko, 2002; Brändli and Dowhanytsch, 2003; Vološčuk, 1992, 1994, 1995, 1999, 2003; Hamor and Commarmot, 2005; Commarmot et al., 2000; Pichler et al., 2007b) and utilized for practical forest and conservation management (Vološčuk 1994, 1995; Pichler et al., 2007a). The phytocoenological releves (stationary plots) were decribed according to Zlatník (1976) and geobiocoenoses were classified according to Zlatník (1959). In Primeval Beech Forests of the Carpathians prevail the group of forest types fa*getum pauper, fa*getum typicum, fa*getum tiliosum, Abieto-fa*getum and fa*geto-Aceretum. The ecological research in Ancient Beech Forests of Germany (lowlands) was carried out during the past 40–50 years (Assmann et al., 2008; Dörfelt, 2008; Plachter et al., 2008; Knapp, 2011; Britz et al., 2009). Characteristics of the localities The World Natural Heritage „Primeval Beech Forests of the Carpathians“ is situated in the biogeographic region „Carpathian beech forests“ (Brändli and Dowhanytsch, 2003) with acentre of diversity in the Eastern Carpathians. It is a part of the Inner Carpathians, which form acontinuous mountain range over 1,300 km in length, 100 to 350 km in width, and up to 2,600 m in height. In the periphery and the montane-altomontane zone, large portions of this richly wooded mountain range are characterised by specious beech and mixed beech forests. The potential natural range of the beech forests therefore comprises an area of approx. 92,000 km2 throughout the Carpathian centre zone, which corresponds to roughly one-tenth of the pan-European beech forest area. These areas, located in mountainous and sub-alpine altitudes (400–1,940 meters a.s.l.), are primarily representative of mountain beech forest. The geographic coordinates of Primeval Beech Forests of the Carpathians are: N 47o–49o, E 22 o–24 o (Table 1). The last extensive primeval beech forests can now only be found in the Carpathians. This is the only place where there can still be experienced the uninterrupted dynamics of the coming and decline of beech forests since the last Ice Age. The great biodiversity of the

Table 1. Location and area of the component parts of the Primeval Beech forests of the Carpathians Component parts

Country

Core area ha

Buffer zone ha

Elevation a.s.l. m

Chornohora

Ukraine

2,476.8

12,925.0

640–1,550

Kuzyi-Trybushany

Ukraine

1,369,6

3,163.4

420–1,087

Maramorosh

Ukraine

2,243.6

6,230.4

600–1,470

Svydovets

Ukraine

3,030.5

5,639.5

720–1,500

Uholka – Shirokiy Luh

Ukraine

11,860.0

3,301.0

400–1,350

Stuzhitsia – Uzhok

Ukraine

2,532.0

3,615.0

600–1,221

Stužica – Buk. vrchy

Slovakia

2,960.0

11,300.0

512–1,210

Havešová

Slovakia

171.3

63.9

442– 741

Rožok

Slovakia

67.1

41.4

440– 789

Vihorlat

Slovakia

2,576.0

2,413.0

517–1,076

29,278.9

48,692.7

Total area ha

beech forests has managed to endure here. The World Heritage Site „Primeval Beech Forests of the Carpathians“ represents the beech forest of the mountain range in ten component parts. Four areas are located in the Slovak Republic, six are located in the Ukraine. The smallest area is 67 hectares in size, the largest approx. 12,000 hectares. They are located in the Eastern Carpathians, one of the most unspoilt habitats in Europe. All the component parts are remnants of primeval forests which are embadded in beech forests that are extensively managed. Germany is at the centre of distribution of the beech forests. If nature had its way they would cover approx. two thirds of the land area of Germany extending from the Alps over high and low mountains ranges and down to the lowlands at the sea coastlines. Now only approx. seven per cent of this surface is covered with beech forests due to deforestation and forest conversion. Larger contiguous forest areas are rare. The remaining forests are used in the forestry industry and beeches of approx. 120 years of age are harvested. The senescent and decay phases of alifecycle that is naturally of more than 300 years duration are absent and thus also the living spaces that emerge in these phases as tree hollows and dead wood with their typical biocoenosis. Primeval beech forests have long since disappeared barring afew miniscule remnants and with them also species that are dependent upon them. The Decision of the 35th Session of the World Heritage Committee, Paris 25 June 2011, approved the extension of the Primeval Beech Forests of the Carpathians (Slovakia and Ukraine), to include the Ancient Beech Forests of Germany, and becomes the Primeval Beech Forests of the Carpathians and the Ancient Beech Forests of Germany (Slovakia, Ukraine and Germany), on the basis of criterion (ix): outstanding examples rep-

resenting significant on-going ecological and biological processes in the evolution and development of ecosystems and communities of plants and animals. The German extension in 2011 is another major step towards protecting this unique ecosystem for the long term. The German part includes selected forest regions of the National Parks Hainich in Thuringia, KellerwaldEdersee in Hesse, Jasmund and Müritz in MecklenburgWestern Pomerania, and the forest of Grumsin in the Schorfheide-Chorin Biosphere Reserve in Brandenburg. These are the most valuable remaining examples of large, undisturbed beech forests in Germany. These German sites with their beech forests in the lowlands and central uplands are aperfect component to the mountain beech forests located in the Carpathians. This component part of the World Natural Heritage represents the characteristics and the natural processes of European beech forests under various ecological conditions. The development history of beech forests since the Ice Age, the enormous competitiveness of beech fa*gus sylvatica and the diversity of geographical, geological and ecological beech forest variations are aunique global phenomenon. The Ancient Beech Forests of Germany are indispensable to documenting the postglacial colonisation by fa*gus sylvatica from south to north, from east to west, and spanning the entire spectrum of altitudinal zones from the sea-shore, to the lowlands and the submontane belt, to the upper timber line in the mountains (Knapp, 2011). German´s component parts are the most outstanding examples worldwide of the respective beech forest types. Each component part has its own specific characteristics and local peculiarities that make it unique and irreplaceable. Jasmund: size 492.5 ha, buffer zone 2,510.5 ha, N 54o32´53´´ E 13o38´43´´ (0–131 a.s.l.). Jasmund is

297

a representative of the beech forest of the lowlands type. Half of Jasmund´s property border follows to coastline. Although this border is subject to very slow natural dynamic changes based on the denudation of the steep coast, it is clearly identifiable by distinctive habitat limits at any given point. Jasmund represents the beech forests of the lowlands on lime and boulder clay. Beech forests, chalk cliffs and sea form afascinating backdrop. The harsh coastal climate and the interaction of topography and climate lead to abroad range of different beech forest communities which are interspersed with streams and moors. Rare orchids, the great horsetail and the coral root are typical here. Serrahn: size 268.1 ha, buffer zone 2,568.0 ha, N 53o20´24´´, E 13o11´52´´ (67–124 m a.s.l.). The best structured lowland beech forests in Europe. Demarcation in Serrahn has produced acompact core area of beech-dominated forests. In the Serrahn part the forest of the Müritz National Park lowland beech forests grow on sands from the Ice Age. In the midst of an extended forest and lake landscape this old beech forests help us to imagine what the German beech forests once looked like. Lakes and mires enrich the forest landscape, create arich diversity of habitats and form the basis for agreat amount of biodiversity. The beech forest of Serrahn is consequently documenting moisture-related distribution limits in an outstanding manner. Grumsin: size 590.1 ha, buffer zone 274.3 ha, N 52o59´11´´, E 13o53´44´´ (76–139 m a.s.l.). Grumsin represents the beech forests of the lowlands on glacial sands and clay. The demarcation of the Grumsin component part largely follows the core area border of the Schorfheide-Chorin Biosphere Reserve, which was designated in 1990. Minor marginal zones which predominantly consist of pine woods rather than nearnatural deciduous forests and were likewise abandoned to natural development in 1990 have been assigned to the buffer zone. Water and forests are closely linked in Grumsin. Lakes, forest marches and moores in deep valleys interchange with marked ridges and conjure up atmospheric forest images in the ancient beech forests. These different structures in the most confined spaces form the basis for an exceptionally rich range of animal and plant species. The area represents an exceedingly textured young moraine landscapes with altitudes of between 60 and 140 m above sea level and all the typical elements in aunique fashion. Hainich: size 1,573.4 ha, buffer zone 4,085.4 ha, N 51o04´43´´, E 10o26´08´´ (290–490 m a.s.l.). Hainich National Park encompasses what is, at present, the largest unmanaged deciduous forest area in Germany. Hainich represents the best reference area for the specious eutraphent beech forests of the European collinesubmontane zones with their ground vegetation rich in geophytes and the exceedingly attractive floral display in early spring, representing the seasonality of Central European deciduous forests in aunique manner. The 298

most valuable beech forests that offer avery rich range of species grow on the central mountain ranges on limestone. It impresses through its extensive range of tree species and reveals lime beech forests of amagnitude, unspoilt nature and form that you will be unable to find in any other area. The demarcation in Hainich follows the distribution of the best-preserved beech forests with old growth stands. The buffer zone comprises the core area of the national park. The Hainich beech forest is unique proof of the currently ongoing ecological processes associated with the present climate change. Kellerwald: size 1,467.1 ha, buffer zone 4,271.4 ha, N 51o08´43´´, E 8o58´25´´ (245–626 m a.s.l.). The acidophilous beech forests of the lower mountain ranges grow on slate and geywacke in the Kellerwald. No roads and no settlement cut through the exceptionally old, extensive forests of the Kellerwald in which unique primeval forest relics have survived. The beech reaches its natural forest boundary at the rocky and scree slopes and forms abizarrely formed forest landscape. More of than 500 of the purest springs and streams form additional valuable habitats. In Kellerwald, the border was established taking into account the specific qualities of the component part, such asthe high relief energy, the disjointed occurrence of small primeval-forest like steep slopes, and the spatial distribution of valuable beech forests. Acoherent complex of valuable oldgrowth beech forests has been included. The demarcation of buffer zone follows the national park border. No buffer has been designated in avery small plot located on the northern border in order to integrate one of the primeval beech forest slopes into the property. Kellerwald contains the largest protected area of oligotraphent and mesotraphent beech forests, where undisturbed ecological and biological processes occur and is aperfect illustration of acidophilous beech forests.

Results and discussion Specific peculiarities of the Carpathian forests include the richness in endemic species, the occurrence of Europe´s largest population of predatory mammals with some 8,000 brown bears, 4,000 wolves and 3,000 lynxes as well as the most significant large-scale primary forest on the periphery of the European beech forests´distribution range. Representing its remaining primeval forests, the World Natural Heritage „Primeval Beech Forests of the Carpathians“ is an essential part of these unique beech forests landscapes. These undisturbed, complex temperate forests exhibit the most complete and comprehensive ecological patterns and processes of pure stands of fa*gus sylvatica across avariety of environmental conditions. The Carpathians Primeval Forests show abroad range of possible forest development stages from rejuvenation to decay (Pichler et al., 2007a).

Ukraine and the Slovak Republic have taken on a pioneering role with the inscription of the Primeval Beech Forests of the Carpathians in the World Heritage List in 2007. The Carpathian Mountains are home to the last remaining large-scale primeval beech forests in Europe. Since the end of the last Ice Age, the forests here have been able to develop undisturbed. Mightly beech trees up to 50 meters high dominate the structurally rich forests (Brändli and Dowhanytsch, 2003; Vološčuk, 2003). The dynamics of the primeval beech forests, the natural comings and goings, are able to play out entirely free from anthropogenic influences here. Globally endangered species of fauna, fungi and flora have been able to preserve their natural gene pool. The model of the main natural successional phases occuring in Central Europe (Korpeľ 1995, Pichler et al., 2007a): growing-up stage, optimal stage, decaying stage. In the growing-up stage, trees are found in all three layers – upper, middle and lower, and the crown closure is dense. As there is low mortality in trees of this age, there is little dead wood. At the end of phases, however, the competition between individuals is so great that strong dying off of juveniles occurs. In the following optimal stage, the maximum timber stock is reached, but the number of trees per area unit is low. With the lack of an understorey, the attainment of maximum height and aclosed canopy, the forest in this phase is known as „hall-forest“, being reminiscent of the interior of acathedral or great hall, and also bears some resemblance to acommercial forest. During the transition to the decaying stage tree vitality decreases and the proportion of dead wood increases considerably. In this phase, the number and size of gaps between tree clusters increases and regeneration of climax tree species starts again. An alternative view (Holling, 2001) suggests that the complexity of living systems of people and nature emerges not from arandom association of alarge number of interacting factors rather from asmaller number of controlling processes. These systems are self-organized, and asmall set of critical processes create and maintain this self-organization. „Self-organization“ is aterm that characterizes the development of complex adaptive systems, in which multiple outcomes typically are possible depending on accidents of history. According to Holling (2001) there are three properties that shape the adaptive cycle and the future state of asystem: wealth, controllability, and adaptive capacity. The adaptive cycle includes 4 phases: (1) long period of slow accumulation and transformation of resources, and (2) conservation (growing-up stage and optimum stage according to Korpeľ, 1995), (3) shorter period of collapse that creates opportunities for (4) innovation (from release to reorganization), or decaying stage with regeneration phase according to Korpeľ, 1995). The European natural beech forests stand out due to ahighly peculiar natural dynamism which is deter-

mined by the cycle of growth and decay of one single tree species, which is the beech. Old beech stands will regenerate with the crowns of individual trees gradually dying back to allow more light to the ground. Either there already is young beech wood that will now emerge, or the next generation of saplings will close the void within aperiod of afew years. The beech once again forms the upper crown canopy later on, thus resetting the cycle, which has been described as the small development cycle (Zukrigl et al. 1963, Leibundgut 1978, Korpeľ 1989, 1995). In the wake of major disruptions, however, the cycle may also involve the formation of an early successional forest made up of pioneer species such aspines, birches, goat willows or rowans, which is later on infiltrated by medium-shade and shade tree species. This big successional cycle may take several decades longer than the small one. Variation incorporating elements of both big and small cycle are possible. This endogenous cycle of development meets the diversity of sites resulting from the glacial and postglacial periods, producing the considerable structural variety as basis for the species-rich, complex system. Rooted in the beech´s enormous ecological plasticity, the high ecological stability results in abiodiversity-promoting continuity of the forest´s character, which makes the dynamics of the beech forest persistently „predictable“ for the forest dwellers. Old beech forests are, for example, home to amultitude of flightless ground beetles that would drop the ability to fly due to the habitat being continuously available or changing only at asmall scale (Britz et al., 2009, Plachter et al., 2008). Asignificant feature of the beech forests is decline in floristic diversity, which is aresult of the history of flora and vegetation, from the former glacial refuges in Southern and Southeastern Europe up the northern and northwestern subterritories. Old beech trees can form ahighly diverse habitat for fauna. The beech is akey species which creates its own internal forest climate and crucially influences soil formation, regeneration cycle, food chains and structures and reveals stonishingly specific diversity of plants, vertebrates, insects, molluscs and fungi. This diversity is described in terms of its ecological role in the ecological processes of beech forest ecosystems – trees and shrubs, mycorrhizae, geophytes, other herbaceous plants, lianas, herbivores, carnivores, dead wood inhabitants, destruens, etc. (Assmann et al., 2008). As opposed to the climatic patterns of tropical rainforests, the climate of the temperate zone is distinguished by its seasonal changes together with the phenological floral cycle involved. From a physiognomic perspective, the most striking feature of deciduous trees is the fall of leaves, which will further accentuate the seasonal differences and conditions of the biotopes respectively. However, the foliage changing with the seasons does not take place abruptely. In pure beech forests this process is accompanied by unique changes in 299

colour (Knapp, 2011). The most dramatic consequence of leaf fall is the light climate´s periodicity. This sets deciduous forests apart from all non-deciduous forest types, permitting the intermittent occurence of aherb layer that shows different specific adaptations. Spring geophytes exploiting the brief warm spring period prior to leafing for development are particularly well adapted and transform the soils of richer beech forests into acarpet of flowers. The association that has given rise to geophyte-rich beech forests is aresult of ecosystemary continuity as well as the inner functional and structural differentiation of the development cycle of deciduous forests. In this particular shape, it is without paralel in the world (Knapp, 2011). Amultitude of fungi are involved in dead wood decomposition, with anumber of species being specialised in the metabolisation of specific wood types. The species of the genus fa*gus are highly mycotrophic; in other words, much of their nutrient supply comes from fungi. Their survival is directly dependent on the mycobionts of ectotrophic mycorrhizae. The dominant mycorrhizal fungi associated with fa*gus sylvatica are Agaricomycetidae, asubclass of Basidiomycetes (hom*obasidiomycetes) from the genera Amanita, Boletus, Cortinarius, Inocybe, Laccaria, Lactarius, Tricholoma, Russula and Xerocomus. Soil acidity plays an important role in relation to the species spectrum of the mycorrhizal partners of fa*gus sylvatica (Dörfelt, 2008). Species typical of the beech include Fomes fomentarius (wood-inhabiting fungi), Ganoderma applanatum (wood decaying), Neobulgaria pura, Oudemansiella mucida, which is indicative of extensive matured wood pools, and Hericium coralloides, which, although widespread throughout the northern hemisphere and also growing on other trees, is only found in very old, mature beech forests. Dead beechwood is colonised very swiftly by very many lignocolous fungi. Three phases characterise decomposition of beech stups: initial, optimal and final phase. There are more than 10 parasitic biotrophic fungi which infect Europe´s beeches. Avery large number of fungi are involved in the decomposition of fallen beech leaves, fruits, mast (cupulae) and twigs (Dörfelt, 2008). An especially important symbiosis has been evolved between fungi and plants in the rhizosphere, which is called mycorrhiza. Forests of the temperate zone are home to fungi that will enter into specific symbioses with one or few tree species (Britz et al., 2009). Despite the beech´s absolute dominance, the beech forests show outstanding diversification and are unique in function and structure (Pichler et al., 2007a). Nortwithstanding the geologically short time of afew thousand years, ahighly characteristic faunistic biocoenosis has been evolved postglacially which is just aglobally unique as isthe plant community. The fauna can exist in all its diversity, and the postglacial evolutionary processes can take place only if each forest develop300

ment stage of the natural regeneration cycle is available – which is the case in the Primeval Beech Forests of the Carpathians (Pichler, 2007; Plachter et al., 2008). The Principles of Joint Management Plan Long-term protection and management of the World Heritage Sites is ensured through national legal protection as national parks or core areas of a biosphere reserve. Effective implementation of the integrated management plan and the trilateral integrated management system is required to guide the planning and management of this World Heritage Sites. The general objectives of the Integrated Management Plan are (Pichler et al., 2007a): o To ensure the most effective conservation of the WHS properties with all their abiotic and biotic components, geo- and biodiversity and ecological processes. To secure a lasting homeostasis and selfreproduction of the respective ecosystems and their protection both against anthropogenic factors. o To maintain and expand the existing, ecologically connected complex of primeval and natural beech forests that encompass and connect (link) the WHS on both the Slovak and the Ukrainian sides – within the corridors connecting the WHS. Supporting the succession of managed beech semi-natural forests. o To use WHS for scientific research in order acquire knowledge transferable and applicable on the level of sustainable. To use WHS for enhancement of landscape ecological stability. o To use WHS for enhancement of ecological and environmental education, awareness of primeval forests – chosen to maintain integrity and conservation of the existing sites, to preserve their naturalness and uniqueness. o To support of traditional crafts, products and ecotourism. Common elements of an effective management system could include: a) a thoroughly shared understanding of the property by all stakeholders; b) a cycle of planning, implementation, monitoring, evaluation and feedback; c) the involvement of partners and stakeholders; d) the allocation of necessary resources; e) capacity-building; and f) an accountable, transparent description of how the management system functions.

Conclusions Joint World Natural Heritage “The Primeval Beech Forests of the Carpathians and the Ancient Beech Forests of Germany” is indispensable to understanding the history and evolution of the genus fa*gus, which, given its wide distribution in the Northern Hemisphere and its ecological importance, is globally significant. These undisturbed, complex temperate forests exhibit the

most complete and comprehensive ecological patterns and processes of pure stands of European beech across a variety of environmental conditions and represent all altitudinal zones from seashore up to the forest line in the mountains. Beech is one of the most important elements of forests in the Temperate Broad-leaf Forest Biome and represents an outstanding example of the re-colonisation and development of terrestrial ecosystems and communities after the last ice age, a process which is still ongoing. They represent key aspects of processes essential for the long term conservation of natural beech forests and illustrate how one single tree species came to absolute dominance across a variety of environmental parameters. Furthermore, it is not enough for a site to meet the World Heritage criteria, but it must also meet the conditions of integrity and/or authenticity and must have an adequate protection and management system to ensure its safeguarding.

Acknowledgement This study was supported by the grant from the Slovak Grant Agency for Science VEGA no. 1/0364/10 and no. 1/0252/11.

References Assmann, T., Drees, C., Schroder, E., Ssymank, A. 2008. Low species diversity of beech forests – amyth. In Knapp, H.D. (ed.). Beech Forests – aGerman contribution to the global forest biodiversity. Skripten, 233. Bonn: Bundesamt für Naturschutz (BfN), Federal Agency for Nature Conservation, p. 25–31. Bennett, G. (ed). 1994. Conserving Europe´s natural heritage: towards a European Ecological Network. London: Graham and Trotman. 334 p. Bohn, U., Neuhäusl, R. 2003. Karte der natürlichen Vegetation Europas – Maßstab 1 : 2 500000. Bonn – Bad Godesberg: Bundesamt für Naturschutz. Brändli, U.B., Dowhanytsch, Y. (eds). 2003. Urwälder im Zentrum Europas. Birmensdorf, Switzerland: Eidgenössische Forschungsanstalt WSL; Rakhiv, Ukraine: Karpaten-Biosphärenreservat; Bern, Stuttgart, Wien: Haupt. 192 p. Britz, H., Dieckmann, O., Engels, B., Frede, A., Geisel, T., Grossmann, M., Kaiser, K., Knapp, H.D., Luthardt, M.E., Seuring, J. 2009. Nomination of the “Ancient Beech Forests of Germany” as extension to the World Natural Heritage “Primeval Beech Forests of the Carpathians”. Nationale Naturlandschaften, Federal Republic of Germany. Specialised editing Cognitio Kommunikation & Planung, Niedenstein. 180 p.

Bublinec, E., Pichler, V. 2001. Slovenské pralesy: diverzita a ochrana. Slovak primeval forests: diversity and conservation. Zvolen: Ústav ekológie lesa SAV. 196 p. Commarmot, B., Duelli, P., Chumak, V. 2000. Urwaldforschung – Beispiel Biosphärenreservat Transcarpatien. Birmensdorf, Switzerland: Naturwerte in Ost und West. Publ. zur Tagung “Forum für Wissen”, WSL, p. 61–68. Dierschke, H., Bohn, U. 2004. Eutraphente Rotbuchenwälder in Europa. Tuexenia, 24: 19–58. Dörfelt, H. 2008. Fungi of beech forests. In Knapp, H.D. (ed.). Beech forests – aGerman contribution to the global forest biodiversity. BfN Skripten, 233. Bonn: Bundesamt für Naturschutz (BfN), Federal Agency for Nature Conservation, p. 33–36. Gömöry, D., Kukla, J., Schieber, B. 2011. Taxonómia, fylogenéza arozšírenie buka vEurópe ana Slovensku [Taxonomy, phylogeny and distribution of beech in Europe and in Slovakia]. In Barna, M., Kulfan, J., Bublinec, E. (eds). Buk abukové ekosystémy Slovenska. Beech and beech ecosystems of Slovakia. Bratislava: Veda, p. 19–36. Hamor, F., Commarmot, B. (eds). 2005. Natural forests in the temperate zone of Europe – values and utilisation. International Conference in Mukachevo, Transcarpathia, Ukraine, October 13–17, 2003. Rakhiv, Ukraine: Carpathian Biosphere Reserve; Birmensdorf, Switzerland: Swiss Federal Research Institute WSL. 112 p. Holling, C.S. 2001. Understanding the complexity of economic, ecological and social systems. Ecosystems, 4: 390–405. Jaworski, A., Kaczmarski, J., Skrzysewski, J., Swiatkowski, W. 1994b. Structure and dynamics of lower subalpine timber stands of Carpathian Mts of primeval character. In Vološčuk, I. (ed). Research and management of the Carpathian natural and primeval forests. Ustrzyki Gorne, Poland: Bieszczady National Park, Association of the Carpathian National Parks and Protected Areas, p. 23–39. Jaworski, A., Skrzyszewski, J., Pach, M. 1994a. Characteristic of Acer paeudoplatanus L. and fa*gus sylvatica L. virgin type forests in Bieszczady National Park. In Vološčuk, I. (ed). Research and management of the Carpathian natural and primeval forests. Ustrzyki Gorne, Poland: Bieszczady National Park, Association of the Carpathian National Parks and Protected Areas, p. 40–49. Knapp, H. D. 2011. European beech forests and their biogeographical position. In Knapp, H. D., Fichtner, A. (eds). Beech forests. Joint Natural Heritage of Europe. BfN Skripten, 297. Bonn: Bundesamt für Naturschutz (BfN), Federal Agency for Nature Conservation, p. 9–14. Korpeľ, Š. 1989. Pralesy Slovenska [The Primeval forests of Slovakia]. Bratislava: Veda. 332 p. 301

Korpeľ, Š. 1995. Die Urwälder der Westkarpaten. Stuttgart: Fischer. 310 p. Kricsfalusy, V., Ivaneha, I., Lugovoj, A., Budnikov, G., Mező-Kricsfalusy, G., Mateleshko, A., Popov, S., Syvokhop, J., Pavlej, J., Lesjo, I. 2001. Uzhanskyi National Nature Park. Uzhgorod: Karpaty. 117 p. Leibundgut, H. 1978. Über die Dynamik europäischer Urwälder. Allg. Forstz., 24: 686–690. Otto, H. 1994. Waldökologie. Stuttgart: Eugen Ulmer. 391 p. Pichler, V., Hamor, F., Vološčuk, I., Sukharyuk, D. 2007a. Outstanding universal value of the ecological processes in the primeval beech forests of the Carpathians and their management as World Heritage Sites. Acta ecol. Bratislava: Veda. 62 p. Pichler, V., Soroková, M. 2005. Utilisation of natural forests for ecotourism: matching the goals and reality. Forest Snow Landsc. Res., 79: (1/2): 185–194. Pichler, V., Vološčuk, I., Bublinec, E. 2007b. Designation of corridors connecting primeval forests properties within the East Carpathian Biosphere Reserve (Slovak Part). In Guziová, Z. (ed). Priorities for conservation of biodiversity in biosphere reserves in changing conditions. Proceedings from the international conference. Stará Lesná, Tatry Biosphere Reserve, Slovakia, 2–6 June. Bratislava: Slovak National Committee for UNESCO MAB Programme, p. 121–126. Plachter, H., Hoffmann, A. Panek, N., Schmidt, P.A. 2008. European beech forests as anatural site on the World Heritage List of UNESCO. In Knapp, H. D. Beech Forests – aGerman contribution to the global forest biodiversity. BfN Skripten, 233. Bonn: Bundesampt für Naturschutz (BfN), Federal Agency for Nature Conservation, p. 53–60. Saniga, M. 2011. Primeval beech forests. In Barna, M., Kulfan, J., Bublinec, E. (eds). Buk a bukové ekosystémy Slovenska. Beech and beech ecosystems of Slovakia. Bratislava: Veda, p. 209–226. Saniga, M., Schütz, J.P. 2002. Relation of dead wood course within the development cycle of selected virgin forests in Slovakia. J. Forest. Sci., 48: 513–528. Stoyko, S. 2002. Pralisovi ekosystemy Ukrajiny, jich polifunkcionalne znacenja dlja ochorony pryrody [The virgin ecosystems of Ukraine, their polyfunctional significance for nature protection]. Naukovi praci Lisivničoji akademii nauk Ukrajiny, Ľviv, Ukraine, 2002: 27–31. Stoyko, S., Sayik, D.S., Tatarinov, K.A., Tretyak, P.R., Tasenkevitsch, L.O., Malinovskij, K.A., Schewtschenko, S.V., Sukhariuk, D.D., Milkina, L.I., Komendar, V.I., Miller, M.P., Manyiko, M.P. 1982. The Carpathian reserve. Uzhgorod: Karpaty. 128 p. Stoyko, S., Tasenkevitch, L. 1993. Some aspects of endemism in the Ukrainian Carpathians. Fragm. flor. geobot., Suppl. 2 (1): 343–353. 302

Vološčuk, I. 1992. Biological diversity in the Carpathians. Oecologia mont., 1 (2): 43–47. Vološčuk, I. 1994. Conservation and rational use of forest ecosystems in Carpathian mountains. In Vološčuk, I. (ed.). Research and management of the Carpathian natural and primeval forests. Ustrzyki Gorne, Poland: Bieszczady National Park, Association of the Carpathian National Parks and Protected Areas, p. 5–10. Vološčuk, I. 1995. Long-term ecological research and monitoring on Carpathian national parks and biosphere reserves. In Hamor, F., Vološčuk, I. (eds). Methods of monitoring of the nature in the Carpathian national parks and protected areas. Rakhiv, Ukraine: Carpathian Biosphere Reserve, Association of the Carpathian National Parks and Protected Areas, p. 10–22. Vološčuk, I. (ed). 1999. The National parks and biosphere reserves in Carpathians. The last nature paradises. Tatranská Lomnica, Slovak Republic: Association of the Carpathian National Parks and Biosphere Reserves. 244 p. Vološčuk, I. 2003. Geobiocenologický výskum prírodných lesných ekosystémov v chránených územiach Karpát = The geobiocenological research in the natural forest ecosystems of the Carpathian protected areas. Monografické štúdie o národných parkoch; 3. Banská Bystrica: Štátna ochrana prírody Slovenskej republiky, 2003. 122 p. Zlatník, A. 1934. Studie o státních lesích na Podkarpatské Rusi. Díl první. Příspěvek k dějinám státních lesů a lesnictví na Podkarpatské Rusi [The study of the state forests in Transcarpathian Ruthenia. First volume. The contribution to the history of state forests and forestry in Transcarpathian Ruthenia]. Sbor. Výzk. Úst. zeměd. ČSR, sv. 126. Praha: Ministerstvo zemědělství republiky Československé. 109 p. Zlatník, A. 1935. Studie o státních lesích na Podkarpatské Rusi. Díl druhý. Přírodní podmínky státních lesů a polonin na Podkarpatské Rusi. Díl třetí. Vývoj a složení přirozených lesů na Podkarpatské Rusi a jejich vztah ke stanovišti. [The study of the State Forests in Transcarpathian Ruthenia. Second volume and third volume]. Sbor. Výzk. Úst. zeměd. ČSR, sv. 127. Praha: Ministerstvo zemědělství republiky Československé. 206 p. Zlatník, A. 1936. “Lužanský prales” na Podkarpatské Rusi, největší československá pralesová reservace [Luzhanskyi virgin forest in Subcarpathian Ruthenia, the largest Czechoslovakian virgin forest]. Krása našeho domova, 28: 110–118. Zlatník, A. 1959. Přehled slovenských lesů podle skupin lesních typů [Asurvey of Slovak forests by groups of forest types]. Brno: Lesnická fakulta Vysoké školy zemědělské vBrně. 92 p. Zlatník, A. 1976. Lesnícka fytocenologie [Forest phytocoenology]. Praha: Státni zemědělské nakladatelství. 495 p.

Zlatník, A., Hilitzer, A. 1932. Přehled přírodních reservací a jejich návrhů vPodkarpatské Rusi [The review of nature reserves and their proposals in Transcarpathian Ruthenia]. Sborník map, ročník VI., č. 2. Praha: nákladem Masarykovy akademie práce. 84 p. Zlatník, A., Korsuň, F., Kočetov, F., Kseneman. M. 1938. Průzkum přirozených lesů na Podkarpatské Rusi. Díl první [The investigation of natural for-

ests in Transcarpathian Ruthenia. First volume]. Sbor. Výzk. Úst. zeměd. RČS. Brno, sv. 152. Brno: nákladem Ministerstva zemědělství republiky Československé. 525 p. Zukrigl, K., Eckhardt, G., Nather, J. 1963. Standortskundliche und waldbauliche Untersuchungen in Urwaldresten der niederösterreichischen Kalkalpen. Mitt. Forstl. Bundesvers.-Anst. Wien, Heft 62. Wien: Österreichischer Agrarverlag in Komm. 244 p.

Received December 17, 2012 Accepted February 14, 2013

303

Instructions for authors Folia oecologica is ajournal devoted to publishing original scientific papers from the fields of ecology of forest ecosystems, communities and populations of plants, fungi and animals associated with forest environment and also the ecology of woody plants growing in both forest and non-forest environment, human settlements included. The journal also publishes short communications, methodological and survey papers in the area, book reviews, personalia and information about scientific events. The manuscripts are submitted to reviewers for evaluation of their significance. Manuscript layout. The manuscripts should be written in English, well-arranged, not exceeding amaximum extent of 20 pages, including tables and figures. The authors are responsible for the quality of the text, manuscripts written in poor English will be returned. Please, send two copies of the manuscript (A4 format, type size 12 font Times New Roman, double-space lines, 3 cm margins on each edge of the page) together with all figures and tables (each on a separate sheet) to the editorial office. Avoid dividing the words, smoothing right text margins; do not define the styles and paragraphs. Do not use either spacing or tabulator for beginning of a paragraph. If the paper has been submitted for publication, send two printed copies and one copy in electronic form (CD or DVD) as a Microsoft Word file (DOC, or RTF format). An original scientific paper should comprise: 1. The title. 2. The author’s name: full first name and second name. 3. Address: complete address and e-mail address (if available) of all the authors. 4. Abstract: in one paragraph, without references to tables, figures and literature, not exceeding 15 lines (900 characters). 5. Key words (maximum 6). 6.Introduction. 7. Material and methods. 8. Results. 9. Discussion (or Results and discussion). 10. Acknowledgement. 11. References. 12. Summary in Slovak (or in Czech): not exceeding an extent of one page, including the title of the paper in Slovak. In the papers, it is necessary to use SI symbols. Non-integer numbers should be provided with a decimal point, (e.g. 1.7), not a comma (1,7), the thousands (with exception of years) are separated with a comma: 5,600. The variables in mathematical formulae and expressions should be written in italics, the symbols for functions and constants in the normal font, the matrices in bold capitals, the vectors in bold small letters. Latin names of genera, species, subspecies and varieties are written in italics, the name of the author of the description (or his abbreviation) normally: e.g. Lymantria dispar (Linnaeus, 1758), Lymantria dispar (L.), Abies cephalonica Loud. var. graeca (Fraas) Liu. The names of cultivars are written normally, e.g. Olea europea L. cv. Chalkidikis. All the tables and figures must be referred to in the text: Table 1, Tables 2–4, Figs 2–4. The authors are asked to indicate placing of the tables and figures on the text margins. Literature citations. The literature cited in the text should conform to the following patterns: one author – Funke (1971) or (Funke, 1971), two authors – Sokal and Rohlf (1995) or (Sokal and Rohlf, 1995), three and more authors – Alstad et al. (1982) or (Alstad et al., 1982). More than one work written by the same author is to be distinguished with small letters appended after the year: Novák (1950a, 1950b). If the document does not contain either the name of the responsible person or the corporation and if it is not possible to conclude about the author with certainty from other authorities, the work should be cited as written by an Anonymus. References in the final list are to be provided with the full title and names of all authors; ordered alphabetically and according to the publication year. Latin names of genera, species and sub-species cited in the list of references are to be written in standard type. The titles are to be cited in the original language appended by an English translation (in brackets). The issue number (except the volume number) should be given (in parentheses) only in the case when the volumes are not paginated continually. The titles of periodicals should be cited in shortened form, according to the international rules, conform to the World list of scientific periodicals. The basic instructions can be found in Bojňanský et al. (1982) Periodiká z oblasti biologicko-poľnohospodárskych vied, ich citácia askratky [Periodicals in area of biological and agricultural sciences, their citations and abbreviations]. Bratislava: Slovenská spoločnosť pre poľnohospodárske, lesnícke apotravinárske vedy pri SAV. 704 p. In the case of a possible ambiguity, cite the periodical under the full name. Titles in languages not using the Latin alphabet should be transliterated keeping with the British Standard 2979 (in the case of the Cyrilic e.g. ж = zh, х = kh, ц = ts, ч = ch, ш = sh, щ = shch, ю = yu, я = ya). (The basic rules can be found e.g. in Bojňanský et al. 1982).

304

The following form of citation should be used: Work in a periodical Sharov, A.A., Liebhold, A.M., Ravlin, F.W. 1995. Prediction of gypsy moth (Lepidoptera: Lymantriidae) mating success from pheromone trap counts. Envir. Ent., 24: 1239–1244. Eiberle, K., Nigg, H. 1984. Zur Ermittlung und Beurteilung der Verbissbelastung. Forstwiss. Cbl., 103: 97–110. Book Szujecki, A. 1983. Ekologia owadów leśnych [Ecology of forest insects]. Warszawa: Państwowe Wydawnictwo Naukowe. 604 p. Miller, J.R., Miller, T.A. (eds) 1986. Insect-plant interactions. New-York: Springer-Verlag. 342 p. Work published in a book or in a proceedings Basset, Y., Springate, N.D., Aberlenc, H.P., Delvare, G. 1997. A rewiew of methods for sampling arthropods in tree canopies. In Stork, N.E., Adis, J., Didham, R.K. (eds). Canopy arthropods. London: Chapman & Hall, p. 27–52. Ciberej, J., Kováč, G., Bilá, A. 1999. Faktory ovplyvňujúce početný stav kamzíka vrchovského v TANAP-e [Factors influencing game populations in chamois (Rupicapra rupicapra L.) in the High Tatra National Park]. In Koreň, M. (ed.). Päťdesiat rokov starostlivosti o lesy TANAP-u. Zborník referátov z konferencie. Poprad: Marmota Press, p. 111–116. Dissertation Chromová, L. 2002. Pôdne a vegetačné zmeny lesných spoločenstiev okolia obce Brusno (Veporské vrchy) [Changes in soils and vegetation of forest communities of the Brusno village (the Veporské Mts.)]. PhD thesis. Bratislava: Comenius University, Faculty of Natural Sciences. 122 p. Tables. The tables should be submitted on separate sheets, not included into the text. The sheets must not be folded. The tables are to be numbered, each after other, with Arabic numerals (Table 1, Table 2…). The text in the caption should always begin with a capital letter. The tables can be self-explicable, not requiring references in the text. The numbering and captioning should be placed over the table, commentaries, if any, under the table. Submitted are only tables prepared in Word and Excel, without vertical grid lines. Use the font size 9. Table width should be of one or two text columns (77 and 160 mm) or 235 mm. Avoid doubling the information in tables and plots. Figures. Submitted are only high-quality drawings, plots and photographs in black, each on a separate A4 sheet. They can be prepared manually or printed with a laser or an ink printer. Please use only hatching, not shading. Avoid three-dimensional graphs, if possible. In captions use the Arial font. The font size should not exceed 11, the recommended size is 9. If possible, use the unified size. Figure width should be 77, 160 or if necessary, 235 mm. The lines must be well clean-cut and the written text must be distinctly readable also after the diminution. For the electronic version, only MS Excel is acceptable. The backside of sheet should be provided with the number of the figure and the author’s name. The graphs and ink drawings must be self-explicable and readable with captions and appended keys of symbols only, without necessity to seek explanations in the text. Off-prints. Each author and co-authors will obtain one electronic copy of the published paper. Editorial office. Institute of Forest Ecology SAS, Centre of Scientific Information – Library, Štúrova 2, 960 53 Zvolen, Slovak Republic, e-mail: [emailprotected] Manuscripts should be sent to the editorial office.

305

The shrub and Black Locust communities of chosen parts of the Hron downs, the Slovak Republic - KIPDF.COM (2024)