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Vegetation of Oak-Hornbeam, Scree and Ravine Forests at Lower Altitudes

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Vegetation of Oak-Hornbeam, Scree and Ravine Forests at Lower Altitudes ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Tuexenia - Mitteilungen der Floristisch-soziologischen Arbeitsgemeinschaft Jahr/Year: 2017 Band/Volume: NS_37 Autor(en)/Author(s): Novak Pavel, Zukal Dominik, Vecera Martin, Pistkova Katerina Artikel/Article: Vegetation of oak-hornbeam, scree and ravine forests at lower altitudes in Transcarpathia, Western Ukraine 47-63 ©Floristisch-soziologische Arbeitsgemeinschaft; www.tuexenia.de; download unter www.zobodat.at Tuexenia 37: 47–63. Göttingen 2017. doi: 10.14471/2017.37.021, available online at www.tuexenia.de Vegetation of oak-hornbeam, scree and ravine forests at lower altitudes in Transcarpathia, Western Ukraine Vegetation der Eichen-Hainbuchen- sowie Hangschutt- und Schluchtwälder der unteren Lagen Transkarpatiens, West-Ukraine Pavel Novák*, Dominik Zukal, Martin Večeřa Ț Kateřina Píšťková Department of Botany and Zoology, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic *Corresponding author, e-mail: [email protected] Abstract Transitional between the Pannonian Basin and the Eastern Carpathians the Transcarpathian Ukraine (Transcarpathia) has a diverse geology including Quaternary sediments, volcanites, limestones and flysch, and its climate at low altitudes is relatively warm and humid. We conducted a field survey in June 2016 focusing on mesophilous forest vegetation along a transect covering almost the whole low- altitudinal part of Transcarpathia. We recorded 54 relevés in the field and further digitized 22 relevés from literature. Using classification analysis, we distinguished three clusters of oak-hornbeam forests (alliance Carpinion betuli; 1–3) and three clusters of scree and ravine forests (alliance Tilio platyphylli- Acerion; 4–6): (1) Slightly wet Pannonian oak-hornbeam forests occurring in the lowland part of the region (Circaeo-Carpinetum); (2) Mesophilous oak-hornbeam forests (Carici pilosae-Carpinetum); (3) Xero-mesophilous oak-hornbeam forests (a drier subtype of the Carici pilosae-Carpinetum and the association Primulo veris-Carpinetum); (4) Mesophilous scree forests (Phyllitido-Aceretum); (5) For- ests of steep slopes (Aceri-Tilietum) and transitions to mesophilous oak-hornbeam forests (Carici pi- losae-Carpinetum); (6) Cool and wet scree and ravine forests (Arunco dioici-Aceretum pseudoplatani). Using indirect ordination analysis, three environmental variables (altitude, heat load index and slope) were identified as factors of significant influence on the species composition. These factors well distin- guish oak-hornbeam forests from scree and ravine forests. Keywords: Carpinion betuli, Eastern Carpathians, forest vegetation, Pannonian Basin, phytosociology, syntaxonomy, Tilio platyphylli-Acerion Erweiterte deutsche Zusammenfassung am Ende des Artikels 1. Introduction Mesophilous forests are considered natural vegetation with a wide potential distribution across Central Europe (BOHN et al. 2000–2003). Due to deforestation of lowland regions and the occurrence of endangered biota, some types of these forests have been recorded on the European Red List of Habitats (JANSSEN et al. 2016). The Carpathians and their foothills are regarded as one of the centres of diversity of de- ciduous forest vegetation in Europe (OSZLÁNYI et al. 2004, WILLNER et al. 2009). Many regional and national phytosociological studies considering Carpathian forests have been conducted in all the Carpathian countries including Ukraine (e.g., ONYSHCHENKO 2007, Manuscript received 29 March 2017, accepted 22 July 2017 Co-ordinating Editor: Erwin Bergmeier 47 ©Floristisch-soziologische Arbeitsgemeinschaft; www.tuexenia.de; download unter www.zobodat.at ŠEBESTA et al. 2011). Yet relatively little attention has been paid to oak-hornbeam, scree and ravine forests, which typically occur below the beech forest belt (ONYSHCHENKO 2009). Therefore, our aim was to fill this gap and describe the diversity of forest vegetation at the lower altitudes of Transcarpathia, a region in Western Ukraine comprising the southwestern slopes of the Ukrainian Carpathians and the adjacent part of the Pannonian Basin. Local oak-hornbeam forests have been surveyed near the towns of Uzhhorod and Muka- cheve, though no attention has been paid to scree and ravine forests in the studies published so far (KRAMARETS et al. 1992, ONYSHCHENKO Ț LUKASH 2005, VOROBYOV et al. 2008, ONYSHCHENKO 2009). In contrast, the forest vegetation in adjacent areas of the Pannonian- Carpathian transitional zone in neighbouring countries has been investigated fairly intensive- ly in eastern Slovakia (e.g., HADAČ Ț TERRAY 1989) and north-western Romania (e.g., RAIU Ț GERGELY 1979, MARIAN 2008). The main goals of our study are: (1) to survey the studied vegetation based on numerical analyses and to compare it with syntaxa previously described in Ukraine and adjacent coun- tries; (2) to provide clearer understanding of the environmental factors that drive the floristic composition of the studied vegetation types in Transcarpathia. 2. Study area The study area comprises the flat margins of the Pannonian Basin filled by Quaternary sediments and the Carpathian foothills. The Carpathian foothills are formed of three geologi- cally distinct zones (from lowland to upland parts) starting with the Vihorlat–Gutâi Belt composed of volcanic hills and ridges with prevailing andesite, basalt and dacite. Towards the north-east, this belt is followed by a large region of Carpathian flysch comprising the highest peaks of the Ukrainian Carpathians. Between these two zones, the Pieniny Klippen Belt composed of Mesozoic limestone appears locally at the surface (ANDÓ 1999). Relevés were recorded at sites of various types of bedrock with the exception of flysch. The region is situated in a climatically transitional zone between the continental Panno- nian Basin with dry and hot summers and the mountain climate of the Carpathians with lower mean temperatures and higher precipitation (800–1,000 mm). The mean annual tem- perature ranges between 7 and 9.5 °C, the mean July temperature between 17 and 20 °C and the mean January temperature between -4 and -2 °C (ANDÓ 1999). Transcarpathia is one of the most forested regions of Ukraine, though the Pannonian part has been strongly deforested in the past and current forest cover is predominantly restricted to patches in the agricultural landscape. Alluvial hardwood and willow-poplar forests in the flat lowlands and oak-hornbeam forests in the lower parts of the Carpathian foothills are regarded as natural vegetation. Beech forests prevail at higher altitudes (BOHN et al. 2000–2003, DIDUKH Ț SHELYAG-SOSONKO 2008). Thermophilous oak forests are locally developed on the steep and sunny slopes of volcanic hills (FODOR 1958, STOYKO 2009). Relevés were recorded along the whole altitudinal gradient of oak-hornbeam forests and in the lower part (120–625 m a.s.l.) of the altitudinal range of scree and ravine forests. Flora and vegetation of the Transcarpathian lowlands differ significantly from the rest of Ukraine due to strong influences from neighbouring Pannonian and Carpathian regions (e.g., FODOR 1958, 1974, KRICSFALUSY 1999, ONYSHCHENKO 2009). This is reflected in the geo- botanical zonation of Ukraine, where the lower altitudes of Transcarpathia belong to the distinctive Pannonian Province, which is part of the Eurasian Steppe Region (DIDUKH Ț SHELYAG-SOSONKO 2008). 48 ©Floristisch-soziologische Arbeitsgemeinschaft; www.tuexenia.de; download unter www.zobodat.at 3. Methods 3.1 Vegetation sampling In June 2016, we conducted an extensive field survey in a transect 80 km long and 30 km wide be- tween the cities of Mukacheve and Bushtyno (48°03'–48°35' N, 22°36'–23°37' E) and the study area (approximately 2,400 km2) thus comprises a major part of the low-altitudinal areas of Transcarpathia. The survey focused on the vegetation of oak-hornbeam (alliance Carpinion betuli) and scree and ravine forests (Tilio platyphylli-Acerion). These forests have a tree layer dominated by Carpinus betulus, Quercus petraea, Q. robur or noble hardwood trees such as Acer spp., Fraxinus excelsior, Tilia spp. and Ulmus glabra. Vegetation was sampled following the Braun-Blanquet approach (DENGLER et al. 2008). Each relevé (n = 54) had a uniform area size of 100 m2. First, the percentage cover of each vegetation layer was estimated. Subsequently, the cover of each species was estimated using the extended nine-degree Braun-Blanquet cover-abundance scale (DENGLER et al. 2008). Cryptogams were not determined. Basic environmental variables (aspect and inclination of slope, percentage of rocks) were recorded for all relevés, as well as coordinates (WGS–84) taken using a GPS receiver. Furthermore, we took a mixed soil sample within each relevé (from the uppermost 15 cm at four places within the site). Subsequently, the pH of dried soil samples was measured in a water suspension (2:5) by portable instruments (GMH Greisinger). The heat load index (HLI) indicating the potential heat load of a given site based on its slope and aspect was calculated for each relevé using Equation 3 in MCCUNE Ț KEON (2002). Relevés from our field work were saved in TURBOVEG 2.0 (HENNEKENS Ț SCHAMINÉE 2001) and further analysed, primarily with the use of JUICE 7.0 (TICHÝ 2002). To obtain a more representative dataset, we included and digitized relevés of oak-hornbeam or scree and ravine forests (n = 22) from the study area published by ONYSHCHENKO Ț LUKASH (2005). Although
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