Tibor Magura

Biodiversity Differences between Managed and Unmanaged Forests: Meta-Analysis of Species Richness in Europe

Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest-dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan-European scale. We used meta-analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management-intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta-analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.

Carabids and forest edge: spatial pattern and edge effect

The spatial distribution of carabid species and the effects of edge on their diversity were studied along two repeated forest-grass transects using pitfall traps in the Aggtelek National Park, in Hungary. The location of boundaries between the adjacent habitats (forest interior versus forest edge and forest edge versus grass) was identified using moving window-technique for carabid abundances. Five groups of carabid species were distinguished: habitat generalists, forest generalists, species preferring the grass, forest specialists, and edge-preferring species. Carabids of the forest interior, forest edge and grass also separated from each other by the ordination based on abundance, suggesting that all three habitats had a distinct species assemblage. Ground and surface temperatures, cover of the leaf litter, herbs, shrubs, the canopy closure and the abundance of the carabid preys were important factors for the spatial pattern of the dominant carabids. The study revealed a significant edge effect on carabids. Carabid species richness was significantly higher in the forest edge and the grass than in the forest interior. The herb cover was the most important factor determining the carabids species richness. The study suggested that the elevated species richness of the forest edges was due, in addition to the edge-preferring species, to the presence of species typical of adjacent habitats (forest interior and grass). This highlights the importance of forest edges in nature management, as they operate as a source habitats for dispersal processes contributing to the carabid recolonisation in the adjacent habitats, after habitat destruction. This result also emphasises the significance of forest edges during the succession and regeneration of adjacent disturbed habitats.

Forest edge and diversity: carabids along forest-grassland transects

Diversity relationships of carabids in forest edges and the neighbouring forest interior and the surrounding grassland are studied. Samples were taken along three replicated forest-grassland transects using pitfall traps in the Aggtelek National Park in Hungary during 2 years. The study revealed significant edge effect on the carabids. The Shannon diversity of carabids were significantly higher in the forest edge and the grassland than in the forest interior. Carabids of the forest interior, forest edge and grassland can be separated from each other by ordinations, both on the species composition and abundance, suggesting that all three habitats have a distinct species assemblage. Moreover, indicator species analysis detected significant edge associated species; based on the specificity and fidelity of the carabids we have distinguished five groups of species: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Multiple linear regression analyses showed that relative air moisture, temperature of the ground and cover of the herbs are the most important factors determining the diversity of carabids along the transects. Our results show that the high diversity of carabids in forest edges is due to the edge-associated species and the presence of species characteristic to the adjacent habitats. The significance of forest edges in nature conservation, serving as a source habitat for dispersal processes, contributing to the recolonization of carabids after habitat destruction or other disturbance is emphasized.

The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands

Aim The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge-preferring species modifies the species–area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species–habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge-preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge-preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge-preferring species were shown to influence the species–area relationship: the number of edge-preferring species increased with the edge:area ratio. Main conclusions Both generalist and edge-preferring species can considerably distort the species–area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.

Changes in carabid beetle assemblages along an urbanisation gradient in the city of Debrecen, Hungary

Responses of carabid beetles (Coleoptera: Carabidae) to urbanisation were studied along an urban-suburban-rural gradient representing decreasing intensities of humandisturbance. Carabids were collected by pitfall trapping during their activity period in lowland oak forest patches in the city of Debrecen, Eastern Hungary. The average number of carabid species was significantly higher in the rural and urban areas compared to the suburban one. The high overall species richness in the urban area was due to the presence of species preferring open habitats. The species richness of forest specialist carabids significantly increased along the urban-rural gradient. The overall carabid abundance was significantly higher in the rural than the other two areas. The results did not support the hypothesis that overall diversity should decrease in response to habitat disturbance. They also contradicted the intermediate disturbance hypothesis: species richness was not the highest in the moderately disturbed suburban area. In the urban area, opportunistic species dominated. The average carabid body size was significantly larger in the rural and suburban areas than in the more disturbed urban area. Multivariate methods detected changes in species composition and abundance structure along the urban-rural gradient. Significant proportion of the variation in abundance and species richness was explained by the heterogeneity of environmental variables (ground temperature, surface temperature, humidity, cover of decaying wood material, herbs, canopy layer, and by the amount of prey).

Diversity and composition of carabids during a forestry cycle

Carabids (Carabidae, Coleoptera) were studied for 2 yearsto explore changes in their diversity and composition during a forestry cycle inthe Hungarian Mountain Range. Native beech forest was clear-cut and replantedwith Norway spruce. Recently planted (5 years old), young (15years old), middle-aged (30 years old) and old spruceplantations (50 years old) were studied using pitfall traps. Ourresults showed that the species richness of carabids was significantly lower inthe plantations than in the native beech forest. Species composition washomogeneous in the beech forest. β-Diversity of the 15–30year old plantations was also low, and it increased slightlytowards ageing. Heterogeneity of the species composition of the 5year old plantation was higher than that of the others, whichreflected the patchiness of the young plantation. The main tendencies of thecompositional changes were as follows. Deciduous forest specialists decreasedsignificantly in abundance in the plantations; they appeared in highabundance only in the beech forest. Species characteristic of open habitatsincreased remarkably in abundance in the recently established plantation.Multiple linear regression between the diversity of carabids and of 12environmental variables shows that the pH and compaction of the soil, and thecover of leaf litter had a significant effect in determining diversity ofcarabids.

Ground beetles (Carabidae) and edge effect in oak-hornbeam forest and grassland transects

We evaluated ground beetle diversity in relation to forest edge between an oak-hornbeam forest and adjacent herbaceous grassland. To test our hypothesis that the diversity of ground beetles was higher in the forest edge than the interior, pitfall trap samples were taken along two forest-grassland transects in northern Hungary. The diversity of ground beetles was significantly higher at the forest edge and in the grassland than in the forest interior. Ground beetle assemblages in the forest interior, forest edge and grassland could be separated from each other by ordination. Indicator species analysis detected five groups of species: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Rank correlation indicated leaf litter, herb, canopy cover, and prey abundance as the most important factors influencing carabid diversity. The high diversity of the forest edge resulted from the presence of edge-associated species and of species characteristic of adjacent habitats. Forest edges seem to play an important role in maintaining diversity. Serving as source habitats, edges also contribute to the recolonisation by ground beetles after habitat destruction or other disturbance in the adjacent habitats.

Impacts of Leaf-litter Addition on Carabids in a Conifer Plantation

As a fine-scale, manipulative model experiment leaf litter was added in plots to increase habitat heterogeneity in a 50-year-old Norway spruce plantation, established after the clear-cutting of a native beech forest, during a 2-year period in the Hungarian Mountain Range. Pitfall trap catches of carabids from leaf-litter plots were compared with those from control plots to explore the effect of leaf-litter addition. Difference in the species composition was revealed by ordination; scores of the samples of the two plot types were significantly separated along the first MDS axis. The most numerous species (Pterostichus oblongopunctatus) was significantly more abundant in the leaf-litter plots. However, there were no significant differences for the other most frequently obtained species. Habitat generalist species were the most abundant, followed by forest generalists, then forest specialists, and there were some open habitat species. Enhanced habitat heterogeneity (leaf-litter addition) in homogeneous plantations influenced the spatial distribution and composition of carabids, through altered abiotic (lower ground temperature in the leaf-litter plots) and biotic (more prey items) factors. Differences in abundance, species richness and Shannon diversity were not significant between the control and the manipulated plots, although carabid catch was higher in the leaf-litter plots during both years. Forestry practices to increase habitat heterogeneity should be considered to enhance biodiversity in managed forests.

Effects of urbanization on ground-dwelling spiders in forest patches, in Hungary

Effects of urbanization on ground-dwelling spiders (Araneae) were studied using pitfall traps along an urban-suburban–rural forest gradient in Debrecen (Hungary). We found that overall spider species richness was significantly higher in the urban sites compared to the suburban and rural ones. The increased diversity was due to the significantly more open-habitat species in the assemblages at the urban sites. This suggests that species from the surrounding matrix (grasslands and arable lands) penetrated the disturbed urban sites. The ratio of forest species was significantly higher in the rural sites than in the suburban and urban ones, suggesting that forest species are indeed sensitive to the disturbance caused by urbanization. Canonical correspondence analysis revealed that the species composition changed remarkably along the urbanization gradient. Open-habitat spiders were associated with the urban sites of higher ground and air temperature. Forest spiders were characteristic of the rural sites with higher amount of decaying woods. Our findings suggest that the overall diversity was not the most appropriate indicator of disturbance; species with different habitat affinity should be analyzed separately to get an ecologically relevant picture of the effect of urbanization.

Carabids (Coleoptera: Carabidae) in a forest patchwork: a connectivity analysis of the Bereg Plain landscape graph

For many species, one important key to persistence is maintaining connectivity among local populations that allow for dispersal and gene flow. This is probably true for carabid species (Coleoptera:Carabidae) living in the fragmented forests of the Bereg Plain (NE Hungary and W Ukraine). Based on field data, we have drafted a landscape graph of the area representing the habitat network of these species. Graph nodes and links represented two kinds of landscape elements: habitat (forest) patches and corridors, respectively. The quality of habitat patches and corridors were ranked (from low (1) to high (4)), reflecting local population sizes in the case of patches and estimated permeability in the case of corridors. We analysed (1) the positional importance of landscape elements in maintaining the connectivity of the intact network, (2) the effect of inserting hypothetical corridors into the network, (3) the effects of improving the quality of the existing corridors, and (4) how to connect every patch in a cost-effective way. Our results set quantitative priorities for conservation practice by identifying important corridors: what to protect, what to build and what to improve. Several network analytical techniques were used to account for the directed (source-sink) and highly fragmented nature of the landscape graph. We provide conservation priority ranks for the landscape elements and discuss the conditions for the use of particular network indices. Our study could be of extreme relevance, since a new highway is being planned through the area.