Zoltán Bátori

The conservation value of karst dolines for vascular plants in woodland habitats of Hungary: refugia and climate change

Limestone (karst) surfaces in Hungary are rich in dolines, in which many endangered vascular plant species occur. To date, the majority of studies dealing with doline vegetation have focused on the local rather than the landscape level, without using comparative data from other areas. However, in this study we aimed to compare the vegetation pattern and species composition of dolines under different climate regimes of Hungary with regard to regional species pools. The fieldwork was carried out between 2005 and 2012. Twenty dolines were selected in the Mecsek Mountains (southern Hungary) and nine dolines in the Aggtelek Karst area (northern Hungary). More than 900 vascular plants and more than 2000 plots were included in the study. The moving split window (MSW) technique, nestedness analysis and principal coordinates analysis (PCoA) were used to reveal the vegetation patterns in dolines. Although we found remarkable differences between the species composition of the two regions, dolines of both regions play a similar role in the preservation of different groups of species. Many plants, in particular mountain species, are restricted to the bottom of dolines where appropriate environmental conditions exist. In addition, depending on the doline geometry, many species of drier and warmer forests have colonized the upper slopes and rims. Thus, we can conclude that karst dolines of Hungary can be considered as reservoirs for many vascular plant species, therefore they are particularly important from a conservation point of view. Moreover, these dolines will likely become increasingly indispensable refugia for biodiversity under future global warming. Aggtelek Karst area; global warming; Mecsek Mountains; relict species; transects

Properties of shrubforest edges: a case study from South Hungary

Knowledge on edge properties is important from a conservation perspective. Our study was carried out in the ancient vegetation mosaic of the Villány Mts, South-Hungary. Sampling was conducted along eight transects, each running from a rock sward through a shrubforest patch into another rock sward. Unlike most studies, we identified edge position objectively, using a moving split-window analysis. Five habitat types along each transect were distinguished: north-facing rock sward interior, north-facing edge, shrubforest interior, south-facing edge, and south-facing rock sward interior. In the forty 2 m2 plots, a total of 157 species were found. Species richness and Shannon-diversity of the edges was higher than those of the shrubforest interiors, but not significantly different from the rock swards. Cover did not differ significantly among habitat types. We found only a few edge-related species. No differences between differently-oriented edges were revealed. Species composition of the edges was influenced mostly by the rock sward matrix. We hypothesize that ecological conditions of the edges resemble those of the rock sward interiors. Thus, sward species can penetrate into shrubforest edges, entailing a similar composition of edges and rock swards, resulting in similar diversities. Edges might be viewed as refugia for valuable plants of rock swards.

Ecological conditions, flora and vegetation of a large doline in the Mecsek Mountains (South Hungary)

Ecological conditions, flora and vegetation of a large doline in the Mecsek Mountains (South Hungary) Vegetation-environment relationships were investigated in a large doline of the Mecsek Mts (South Hungary). To reveal the vegetation pattern, we collected vegetation data and environmental variables along a 243 m long transect. Atotal of 144 vascular plant species and 4 vegetation types were identified in the doline. We found that both the species composition and the vegetation pattern are significantly influenced by air temperature, air humidity, soil moisture and altitude. Our results confirm the putative temperature and vegetation inversion in the doline.

Vegetation history, recent dynamics and future prospects of a Hungarian sandy forest-steppe reserve: forest-grassland relations, tree species composition and size-class distribution

Pannonian forest-steppes host a high number of endemic species and contribute to landscape-scale heterogeneity. Alterations in the proportion of forests and grasslands due to changes in land-use practice and climatic parameters can have serious nature conservation consequences. Hypotheses about forest-steppe dynamics have rarely been verified by detailed analyses, especially for the sandy forest-steppes. We integrated historical analysis, aerial photo interpretation and field investigation to determine how vegetation of a sandy forest-steppe has changed, how current dynamical processes operate and how native and exotic tree species regenerate under present conditions. The vegetation of the study area before the onset of major anthropogenic environmental transformations in the Carpathian Basin may have been a mosaic of forested and unforested patches. However, there is strong evidence that after heavy deforestation, the region was almost completely treeless between the 15th and the 19th centuries. Forest cover was able to recover by the 1800s but the lack of forested areas in the region for centuries explains why forest patches are still poor in species. Grasslands, which existed continuously, are more diverse, supporting several rare and endemic species. From 1953 till 2013, 72.45% of the area proved to be stable, but 27.55% showed clear dynamical character, changing either from forest to grassland, or vice versa. Thus, cyclic dynamics can occur in sandy forest-steppes. We found that forest patches of different size, differently exposed edges and grasslands provide different habitats for the tree species. Exotic species were present in large numbers, probably due to the small size of the reserve and the lack of a buffer zone.

Large- and small-scale environmental factors drive distributions of cool-adapted plants in karstic microrefugia.

Background and aims Dolines are small- to large-sized bowl-shaped depressions of karst surfaces. They may constitute important microrefugia, as thermal inversion often maintains cooler conditions within them. This study aimed to identify the effects of large- (macroclimate) and small-scale (slope aspect and vegetation type) environmental factors on cool-adapted plants in karst dolines of East-Central Europe. We also evaluated the potential of these dolines to be microrefugia that mitigate the effects of climate change on cool-adapted plants in both forest and grassland ecosystems. Methods We compared surveys of plant species composition that were made between 2007 and 2015 in 21 dolines distributed across four mountain ranges (sites) in Hungary and Romania. We examined the effects of environmental factors on the distribution and number of cool-adapted plants on three scales: (1) regional (all sites); (2) within sites and; (3) within dolines. Generalized linear models and non-parametric tests were used for the analyses. Key Results Macroclimate, vegetation type and aspect were all significant predictors of the diversity of cool-adapted plants. More cool-adapted plants were recorded in the coolest site, with only few found in the warmest site. At the warmest site, the distribution of cool-adapted plants was restricted to the deepest parts of dolines. Within sites of intermediate temperature and humidity, the effect of vegetation type and aspect on the diversity of cool-adapted plants was often significant, with more taxa being found in grasslands (versus forests) and on north-facing slopes (versus south-facing slopes). Conclusions There is large variation in the number and spatial distribution of cool-adapted plants in karst dolines, which is related to large- and small-scale environmental factors. Both macro- and microrefugia are therefore likely to play important roles in facilitating the persistence of cool-adapted plants under global warming.

IMPORTANCE OF KARST SINKHOLES IN PRESERVING RELICT, MOUNTAIN, AND WET-WOODLAND PLANT SPECIES UNDER SUB-MEDITERRANEAN CLIMATE: A CASE STUDY FROM SOUTHERN HUNGARY

Species composition and the vegetation pattern of the understory were investigated in different sized solution sinkholes in a woodland area of the Mecsek Mountains (southern Hungary). Vegetation data together with topographic variables were collected along transects to reveal the vegetation patterns on the slopes, and a species list was compiled for each sinkhole. The results indicate that the vegetation pattern significantly correlates with sinkhole size. In smaller sinkholes, vegetation does not change substantially along the transects; in larger sinkholes, however, vegetation inversion is pronounced. We also found that sinkhole size clearly influences the number of vascular plant species, in accordance with the well-known relationship between species number and area. In the forest landscape, many medium-sized and large sinkholes have developed into excellent refuge areas for glacial relicts, mountain, and wet-woodland plant species.

The Importance of Forest Patches in Supporting Steppe-Species: A Case Study from the Carpathian Basin

ABSTRACT Moderating effects of trees on the environment in their immediate proximity are considered an important force in structuring plant communities, especially in harsh environments. In the semi-arid regions of the middle Carpathian Basin, such facilitative influences are expected to become crucial for the survival of several plant species, given the current warming and drying tendencies. We used 20 × 20 m plots to analyze whether grassland species adapted to mesic conditions penetrate forest patches, where they are able to survive. Using transects and the moving split window analysis, we also investigated how far the positive effects of the forest patches extend into grasslands, and whether this enables the existence of a steppe community that cannot tolerate extreme dry conditions and unfavorable soils. We found that beside forest-related species, forest patches hosted large numbers of grassland-related species. Among them, plants of closed steppe grasslands were the most numerous, which usually cannot tolerate the harsh conditions of open sandy grasslands, and are often confined to areas with better water and soil conditions. Our results showed that there is a 5–8 m wide closed steppe zone around the forest patches. Some species that are not able to survive in open xeric sandy grasslands are restricted to this zone. Unfortunately, while considerable attention is paid to the research, protection and restoration of sandy grasslands, forest patches are usually neglected. Our results emphasize that the establishment of individual trees and groups of trees should be actively promoted, because they have considerable nature conservation benefits by supporing closed steppe species.

A comparison of the vegetation of forested and non-forested solution dolines in Hungary: a preliminary study

The present study compares the vegetation characteristics of two large forested and one large non-forested solution dolines in Hungary. We investigated the species composition and vegetation pattern along north to south transects (across the doline bottoms) and compared the richness of different species groups (dry and wet groups) on the doline slopes. We applied linear regression models for each slope to explore the effects of topography on species richness, and Detrended Correspondence Analysis (DCA) to detect the major gradients of floristic variation within each site. We found that the vegetation changed significantly along all transects; and, regardless of the vegetation cover, the doline bottoms contained several cool-adapted species. Variations within the two species groups were more pronounced on the south-facing slopes. The changes were similar in the forested dolines, indicating the role of forest cover in maintaining many cool-adapted species on the north-facing slopes as well. However, the number of cool-adapted species increased significantly along both slopes of the non-forested doline from the upper edge to the bottom. Contrary to our expectations, the species turnover along the slopes of the non-forested doline was lower than that along the slopes of the forested ones. We conclude that both the forested and non-forested dolines serve as refuges for many plant species adapted to different environmental conditions. Apart from providing an understanding of population patterns along environmental gradients, our results may also contribute to our understanding of an even more fundamental question for a future research agenda: the probable effects of climate change on vegetation characteristics in climatic islands with environmental conditions substantially different from the surrounding areas.

Transitions between community complexes: a case study analysing gradients through mountain ridges in south Hungary

Abstract The study of boundaries is a recurring theme in ecology. However, boundaries have been examined mainly on fine scales (between communities) and on coarse scales (between biomes), while boundaries of intermediate scales (e.g. between community complexes) are quite neglected. In this study, we analysed boundaries between mesic and xeric community complexes in a sub-Mediterranean karst area of South Hungary. We applied the moving split window (MSW) technique for boundary analysis. First, since the behaviour ofMSWconcerning complex vegetation patterns is not fully understood, we prepared artificial datasets (simulated communities) to test its capacities. Second, we established north-south oriented belt transects across mountain ridges of the Villány Mts, and investigated the transition between the community complexes of differently exposed slopes. UsingMSW, we were able clearly to distinguish between transitional zones and zones that do not represent real transitions: peaks in the Z-score profile of MSW merge only in the case of transitional zones. Moreover, we found that peaks merge depending on the independence (distinctness) of the transitional zone: when it is distinct, peaks merge only at the largest window widths. In the Villány Mts, transitions seem to occur mostly in the grasslands north of the ridges. We demonstrated that these grasslands can be regarded as boundaries between mesic and xeric complexes or as zones in their own right, with their own two boundaries. Interpretation depends upon the scale of observation.

Shrub Encroachment Under the Trees Diversifies the Herb Layer in a Romanian Silvopastoral System

ABSTRACT Rangelands with scattered trees are complex and dynamic systems with a long history in Europe. Generally referred to as “wood-pastures,” they are considered to have outstanding conservation value. Thorny shrubs are important for supporting the biodiversity of these wooded rangelands, as well as facilitating the regeneration of trees by acting as nurse species. We assess the direct effects of temporary shrub encroachment under the cover of mature sparse trees on overall plant and habitat diversity. We surveyed the herb layer of the main landscape features of a wood-pasture: open pasture, trees with a grass understory, trees with shrubs, and adjacent forest edges. The herb layer under trees with shrubs resembled that of forest edges more than open pastures and trees with grass. Trees with grass had a higher cover of ruderal species than trees with shrubs, while forest edges and open pastures had a lower cover of them. Forest species were absent from open pastures but were well represented in the other sites. The herb layer of trees with shrubs and forest edges had similar cover values, while trees with grass had a significantly lower cover of herbs than the other types. Trees with shrubs had higher species richness than any of the other three landscape features and had a much higher proportion of diagnostic species. We conclude that the scattered trees and shrubs of the studied silvopastoral system have additive facilitative effects on their understory, probably through modifying the microenvironment and grazing pressure, leading to the formation of temporary diversity hot spots with distinct vegetation. Thus maintaining a moderate level of shrub-encroachment under sparse trees is recommended for not only creating safe havens for tree recruitment but also increasing the overall species and habitat diversity of wood-pastures.