Csaba Tölgyesi

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.

The effect of the invasive Asclepias syriaca on the ground-dwelling arthropod fauna.

Abstract The management of natural and seminatural systems often leads to disturbance associated with the appearance of non-native species. The spread of these species is increasing due to global environmental changes combined with local management interventions. These non-native species may establish self-sustaining populations influencing ecosystem functions, including the habitat use of native species. Here we explore the response of diplopods, spider and ant assemblages and the activity-density of individual species to the establishment of the non-native plant species, Asclepias syriaca in a disturbed poplar forest in Hungary. The relationship between the species richness of spiders and ants and the structural features of A. syriaca was weak. We found a significant relationship between the structural features of A. syriaca stands and the density and activity of the diplopod Megaphyllum unilineatum. We explain this relationship by the modified microclimate and litter quality of the habitats invaded by A. syriaca. The species composition of ant and spider assemblages was sensitive to A. syriaca. Asclepias syriaca had a negative local effect on the abundance of two spider species which were common in the studied forest. However, A. syriaca positively influenced the abundance of two ant species, most probably via indirect trophic relationships, as they feed on aphids living on A. syriaca. Our study shows that invasive plants can have mixed effects on local invertebrate assemblages. It is therefore crucial to understand how native assemblages respond to these changes in order to better manage these novel ecosystems and maximize their biodiversity benefits.

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.

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.

Can Naturalness Indicator Values Reveal Habitat Degradation? A Test of Four Methodological Approaches

ABSTRACT Assessing habitat naturalness belongs to the most current issues in conservation biology. It has been recognized that plants are able to indicate the naturalness of their habitat. Thus, species may be given relative naturalness indicator values (i.e. scores on an ordinal scale), reflecting their different tolerances against habitat degradation. In the present study, our first goal was to test whether relative naturalness indicator values are able to reveal known differences in naturalness levels. Our second purpose was to compare four different methodological approaches in order to identify which is the most reliable when analyzing habitat naturalness. We compared near-natural and degraded plots on the bases of (1) unweighted plot means, (2) plot medians, (3) unweighted naturalness indicator value populations, and (4) frequency-weighted naturalness indicator value populations. We found that relative naturalness indicator values performed well in differentiating among near-natural and degraded vegetation. Unweighted mean indicator values were the most reliable, but frequency-weighted indicator value populations were nearly as efficient as unweighted means. We conclude that relative naturalness indicator values provide a simple but reliable tool for estimating habitat deterioration.

The effects of micro-habitats and grazing intensity on the vegetation of burial mounds in the Kazakh steppes

Background: Burial mounds (kurgans) of Eurasian steppes are man-made habitat islands that have the potential to harbour rich plant diversity due to micro-habitats associated with their topography. Aims: We assessed whether kurgan micro-habitats harboured different species pools and functional groups from those found on the surrounding steppes. In addition, we asked if these mounds were affected by different grazing intensities from those on the surrounding vegetation. Methods: We surveyed kurgan micro-habitats (northern and southern slopes, surrounding ditch) and adjacent steppe plains in non-grazed, moderately grazed and heavily grazed sites in northern Kazakhstan. We analysed differences in species composition of four habitats under three grazing regimes using Generalised Linear Mixed Models, PCA ordination and indicator species analysis. Results: Kurgan micro-habitats had diverse vegetation and supported the co-existence of plant species with different environmental needs. We identified 16 steppe specialists confined to kurgan micro-habitats. Steppe vegetation was well-adapted to extensive grazing, although heavy grazing supported ruderals and a decline in steppe specialists. There was a significant interaction between grazing intensity and habitat type: heavy grazing supported ruderals and suppressed steppe specialists especially on the slopes. Conclusions: We highlighted that kurgans play an important role as maintaining high plant diversity locally in extensive steppe plains in Central-Asia by increasing environmental heterogeneity and supporting specialist species confined to these micro-habitats.

Hydrologic fluctuations trigger structural changes in wetland—dry grassland ecotones but have no effect on ecotone position

Ecotones between plant communities have received considerable attention among ecologists in the context of fragmentation, climate change and the management of heterogeneous landscapes. However, the predictability of ecotone dynamics is low and the processes taking place within ecotones are still poorly understood. In this study we aimed to characterize the positional and structural dynamics of thirteen ecotones in an ecotone-rich steppe–wetland landscape of Hungary in relation to the inter-annual fluctuations of water regime and the gradients of elevation and of soil composition. According to our results, the ecotones between steppe and wetland communities were sharp and their positions coincided with those places in the landscape where the rate of change in elevation was the highest, confirming that microtopography is a major determinant of ecotone position. Soil boundaries were also detected, mostly downhill to the ecotones. Interestingly, the fluctuations of the water supply had no effect on the position of the ecotones but significantly influenced a structural ecotone parameter, the compositional contrast bridged by the ecotones. High water supply caused high contrast, while low supply went along with low contrast. We explain these changes by asymmetric sensitivities to edge effects. When the water supply was low, the wetland edges became similar to the steppe edges due to the decrease of wetland specialists and to the increase of steppe specialists, but steppe edges did not exhibit an opposite change in wet years, suggesting that steppe communities dominated over wetland communities. The asymmetry in the interaction between the two communities may have pushed the soil boundaries downhill to the ecotones but the currently steppe-like soil of wetland edges could also make wetland edges more sensitive to edge effects; thus, the cause-effect relationship is difficult to disentangle.

Relationship of different feeding groups of true bugs (Hemiptera: Heteroptera) with habitat and landscape features in Pannonic salt grasslands

Eastern European grasslands are still inhabited by a rich arthropod fauna, but the drivers and mechanisms influencing their communities have to be understood to ensure their future survival. Heteroptera communities were studied in 20 plot-pairs in Pannonic salt steppe–salt marsh mosaics in Hungary. The effects of vegetation characteristics, landscape diversity and the proportion of surrounding grasslands on the composition, species richness and abundance of different feeding groups of true bugs (carnivores, specialist and generalist herbivores) were examined using ordinations and mixed-effect models. We found distinct herbivorous assemblages corresponding to microtopography-driven differences in water regime and vegetation between steppe and marsh plots, but this pattern was less pronounced in carnivorous assemblages. A higher species richness of true bugs was found in the more diverse steppe vegetation than in the salt marsh vegetation, while the abundance pattern of true bugs was opposite. Landscape diversity had a positive effect on the species richness and abundance of generalist herbivores and carnivores. Our results suggested that generalist herbivores and carnivores appear to drive diversity patterns in the local landscape due to their high dispersal abilities and the broader range of resources they can utilize. Specialist herbivores strongly influence the local insect biomass in relation to the distribution and density of their host plants. The present study highlights the importance of both habitat and landscape diversity for local insect diversity in Pannonic salt grasslands and suggests that the main threats for arthropod diversity are those processes and activities that homogenize these areas.