Kai Vellak

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.

Diversity of bryophyte vegetation in some forest types in Estonia: a comparison of old unmanaged and managed forests

The bryophyte vegetation of 3 pairs of unmanaged and managed forest stands, representing Oxalis drained peatland, Aegopodium and Oxalis forest site type, were compared. The total number of bryophyte species in unmanaged stands was 74 and that in managed stands 54. Out of the 20 species occurring only in unmanaged forests, 9 grow on decaying wood, and 3 on trunks or bases of big trees; 13 of them were hepatics. In unmanaged forests 11 hemerophobic species were recorded altogether. Although the difference in substrate and species diversity between unmanaged and managed stands is not statistically significant, in unmanaged forests more substrates characteristic for an old-growth stand are available, and the percentage of species preferring dryer habitats or prolonged humidity is a bit higher than in managed forests; the percentage of species associated with better illuminated habitats is higher in managed forests. Analysis of classification structure of the bryophyte layer synusia shows that the number of societies is also higher in unmanaged forests. This is associated with more numerous microhabitats; the average light and humidity indices calculated for every society, confirm this conclusion. The large discrepancy in bryophyte layer classification structure in old-growth and managed forests of the same forest site type is manifested not so much by species content in synusia as by their abundance proportions. The larger diversity of classification units in unmanaged forests is also seen at the synusia facies level; four of nine facies are confined exclusively to unmanaged stands. This is a strong argument for the informativeness of bryophyte layer classification structure for purposes of indication and monitoring as well.

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).

Relationships between species richness patterns in deciduous forests at the north Estonian limestone escarpment

Abstract The flora of the deciduous forests at the base of the north Estonian limestone escarpment is species rich, with an exceptionally high number of rare bryophyte species. Relationships between species richness of bryophyte and herb layers and biotic and environmental conditions were studied, using General Linear Mixed Models. Human disturbance (waste deposit, tree damage etc) was significantly negatively correlated with species richness of both plant layers. Soil nitrogen content was negatively and soil water retention positively correlated with bryophyte species richness, while herb richness was unrelated to soil factors. After eliminating the effects of environment, negative correlations in species richness and cover between the bryophyte and herb layers were discovered on finer scales (1 m2), referring to biotic interactions. This relationship was obscured with the simple correlation analysis. On the other hand, the positive correlation in species pools between the bryophyte and herb layers (0.1 ha) was insignificant. The species pools of both bryophyte and herb layers were significantly positively correlated with the species richness of the tree layer. In summary, bryophyte and herb layer richness responded differently to environmental conditions, but human disturbance significantly decreased the richness of both layers. Due to the uniqueness and small area of these forests we recommend protection and restoration of disturbed sites. Nomenclature: Ingerpuu et al. (1994) for bryophytes, Leht (1999) for phanerogams. Abbreviation: GLMM = General Linear Mixed Model.

Bryophyte vegetation in a wooded meadow: relationships with phanerogam diversity and responses to fertilisation

In the Laelatu wooded meadow in Estonia, famous for its phanerogam diversity, the bryophyte community has been investigated in order to compare its flora and diversity relationships with those of the vascular plant community. Ninety-six bryophyte species were found, 13 of them are hepatics; the majority of the bryophytes are epigeic species common to meadows and forests, including many calciphilous species. Vascular plants and bryophytes display opposite responses to fertilisation. For vascular plants, fertilisation increases the coverage and diminishes the number of species, while for bryophytes it diminishes coverage and increases the number of species. The relationship between the number of species in small plots and the total number of species in the area is similar for vascular plants and bryophytes. No significant changes in the bryophyte community in Laelatu wooded meadow has been detected during the last 30 years.

Management Effects on Bryophytes in Estonian Forests

Abstract.Bryophyte species richness and composition of six different forest types and three different management rates was analysed in 36 Estonian forests. Analyses were done separately for six different bryophyte species groups: total species number, hepatic, hemerophobic, common, non-epigeic and epigeic species. The differences in species richness between unmanaged and moderately managed forests were significant for all species groups, while the differences between moderately and intensively managed forests were significant only for total species number, and numbers of common and non-epigeic species. Our results showed also that in comparison of unmanaged and moderately managed forests, management effect was significant for species richness of all species groups except non-epigeic species, while in comparison of moderately and intensely managed forests, management rate remained significant only for total species richness and richness of common species. Also the species composition was affected by forest management. Only the compositions of unmanaged forest types were found to be significantly different between each other. In conclusion, even moderate management eliminates a significant number of bryophyte species, and the species composition of different forest types become more similar.

Vascular plant and bryophytes species representation in the protected areas network on the national scale

The complexity of nature conservation raises questions about biodiversity protection at the level of species as well as their spatial distribution between differently designated nature conservation areas. We have concentrated on comparison of the existing protected areas and recently established conservation initiative areas—Important Plant Areas. We have estimated how well these areas support the protection of two plant groups—bryophytes and vascular plants. We sought answers to the following questions: (a) are there any trends in the distribution of protected bryophyte and vascular plant species in the protected areas network, and (b) does the Important Plant Areas network promote better protection of bryophyte species compared with the existing protected areas network. Our results demonstrated that bryophytes need special care in nature conservation decisions to reach the reasonable conservation target. Important Plant Areas that were targeted to vascular plants have less importance in preserving bryophyte diversity than already existing conservation areas system. Conservation programs like IBA, IPA etc. have their specific tool and outcome to add conservation values to the existing protected areas system.

Growth depends on neighbours: experiments with three Sphagnum L. species

Abstract Our aim was to search for reasons why some peat mosses (Sphagnum), despite having wide distribution areas, consistently occur in small and distantly scattered populations. The effect of interspecific interactions was proposed as the main hypothesis. Three Sphagnum species exhibiting different distribution frequencies (S. wulfianum Girg., S. teres (Schimp.) Ångstr., and S. magellanicum Brid.) were selected, and two experiments in controlled conditions were established. In the first experiment, the peat mosses were grown in mono-species, two-species, and three-species mixtures. Only the growth of the species with the most restricted distribution (S. wulfianum) responded significantly to the presence of the other Sphagnum species. In the second experiment, shoots of S. wulfianum were watered with the exudates and extracts of the other two species. Significant effects were observed on the growth of S. wulfianum. We conclude that neighbour species can suppress the growth of some bryophyte species and possibly limit their natural distribution.

Compromises in Data Selection in a Meta-Analysis of Biodiversity in Managed and Unmanaged Forests: Response to Halme et al.

Meta-analysis(MA)isapowerfultoolforrevealinggeneraltrends and quantitatively synthesizing the results of inde-pendent studies. Nevertheless, the procedure has beencriticized, particularly when it has been applied to eco-logical and conservation biology studies.To provide an updated picture of the effect of forestmanagement on biodiversity, we performed an MA with

Drastic Turnover of Bryophyte Vegetation on Bog Microforms Initiated by Air Pollution in Northeastern Estonia and Bordering Russia

Human influence on bogs, including air pollution, causes changes in vegetation leading to the degradation of an ombrotrophic bog ecosystem into a more uniform transitional mire-like system. We have hypothesized that intensive atmospheric alkaline pollution will cause an increase in water pH and convergence of bryophyte species composition among microforms. We also expected that bog-specific acidophilic species will be replaced by species indigenous to neutral pH habitats. Through GLM and DCA analyses, we found that although natural acidic bogs are more species poor than polluted bogs, the increase in pH can lead to a decrease in bog-specific vegetation. In polluted bogs, the species composition in different bog microforms will become similar; in particular bog-specific Sphagnum mosses will be increasingly replaced by more tolerant brown mosses, particularly in lawns.