Secondary succession, soil formation and development of a diverse community of oribatids and saprophagous soil macro-invertebrates

Investigations into different stages of secondary succession (from a wheat field to a beechwood on Threstone; Northern Germany) demonstrated the formation of a carbon rich top soil in later successional stages. Parallel to changes in plant species and soil formation, there were also changes in species composition and diversity of saprophagous macro-invertebrates (Lumbricidae, Diplopoda, Isopoda) and oribatid mites (Acari: Oribatida). Diversity of diplopod and isopod species increased after cessation of cultivation, but in a late successional stage (ca 50 y-old fallow, ash-dominated wood) species number of diplopods and isopods declined strongly. In comparison with the other soil invertebrate groups, species composition of earthworms among the sites was more similar. Accumulation of soil C was assumed to be related to wood formation and occurrence of woody debris and recalcitrant leaf litter of beech trees. Incorporation of recalcitrant litter materials by earthworm species living in the upper mineral soil presumably contributed significantly to accumulation of soil C. Accumulation of soil C was accompanied by the development of an oribatid mite community rich in species. In early successional stages oribatids predominantly colonized the litter layer, while most oribatid mites of the beechwood inhabited the upper mineral soil. Maximum diversity of oribatid mites in the beechwood is assumed to be related to instability of the mineral soil caused by earthworm activity. Changes in species composition and diversity are discussed considering succession theory. Even soil invertebrates of similar trophic groups appear to respond very differently to successional changes. It is concluded that conservation strategies to maintain high diversity of soil invertebrates are most likely to be successful if a wide range of habitats of different successional stages is included.     

Assessing Bioindication with Earthworms in An Intensively Farmed Rural Landscape (Yuanqiao and Daqiao Villages in Qianjiang Municipality,Located in Hubei Province,Subtropical China)

Earthworm species diversity, population density, biomass, age-structure, average individual weight, and pattern of distribution were studied in the agricultural landscape of two adjacent villages in Qianjiang Municipality located on the Jiang-Han plain, Hubei Province, China. We identified 16 earthworm species belonging to three families (Moniligastridae, Megascolecidae, and Lumbricidae). Among these, Amynthas hupeiensis and Metaphire guillelmi were the dominant species, and were recorded in significant numbers in different seasons and in the various land-use systems studied. The highest earthworm population density was found in June, before the hot summer, and averaged 23?ind. M^?2, with a maximum of 239?ind. ^?2. Among the nine land-use systems distributed in the landscape, home gardens, intensively cropped commercial vegetable fields and their margins with weeds and bushes generally showed a higher earthworm population density, species diversity, and biomass compared with the other systems. These observations confirm the importance of an heterogeneous agricultural landscape in preserving earthworm abundance and diversity and the possibility to use this group as a possible tool for monitoring landscape changes and sustainability of agricultural practices.     

Bolete diversity in two relict forests of the Mexican beech (Fagus grandifolia var. mexicana; Fagaceae)

The current distribution of the endangered Mexican beech [Fagus grandifolia var. mexicana (Martinez) Little] is restricted to relict isolated populations in small remnants of montane cloud forest in northeastern Mexico, and little is known about its associated biota. We sampled bolete diversity in two of these monospecific forests in the state of Hidalgo, Mexico. We compared alpha diversity, including species richness and ensemble structure, and analyzed beta diversity (dissimilarity in species composition) between forests. We found 26 bolete species, five of which are probably new. Species diversity and evenness were similar between forests. Beta diversity was low, and the similarities of bolete samples from within and between forests were not significantly different. These results support the idea that the two forests share a single bolete ensemble with a common history. In contrast, cumulative species richness differed between the forests, implying that factors other than the mere presence of the host species have contributed to shaping the biodiversity of ectomycorrhizal fungi in relict Mexican beech forests.     

Tree diversity drives abundance and spatiotemporal β‐diversity of true bugs (Heteroptera)

1. Spatiotemporal patterns of canopy true bug diversity in forests of different tree species diversity have not yet been disentangled, although plant diversity has been shown to strongly impact the diversity and distribution of many insect communities. 2. Here we compare species richness of canopy true bugs across a tree diversity gradient ranging from simple beech to mixed forest stands. We analyse changes in community composition by additive partitioning of species diversity, for communities on various tree species, as well as for communities dwelling on beech alone. 3. Total species richness (γ‐diversity) and α‐diversity, and abundance of true bugs increased across the tree diversity gradient, while diversity changes were mediated by increased true bug abundance in the highly diverse forest stands. The same pattern was found for γ‐diversity in most functional guilds (e.g. forest specialists, herbivores, predators). Temporal and even more, spatial turnover (β‐diversity) among trees was closely related to tree diversity and accounted for ～90% of total γ‐diversity. 4. Results for beech alone were similar, but species turnover could not be related to the tree diversity gradient, and monthly turnover was higher compared to turnover among trees. 5. Our findings support the hypothesis that with increasing tree diversity and thereby increasing habitat heterogeneity, enhanced resource availability supports a greater number of individuals and species of true bugs. Tree species identity and the dissimilarity of true bug communities from tree to tree determine community patterns. 6. In conclusion, understanding diversity and distribution of insect communities in deciduous forests needs a perspective on patterns of spatiotemporal turnover. Heterogeneity among sites, tree species, as well as tree individuals contributed greatly to overall bug diversity.     

Changes in Vegetation and Earthworm Populations under Free Grazing European Bison (<Emphasis Type="Italic">Bison bonasus</Emphasis>) in Broad-Leaved Forests of the Kaluzhskie Zaseki State Nature Reserve

This paper studies the impact of a free-ranging bison population on the vegetation and soil fauna in spatially adjacent biotopes (meadow-transition zone-surrounding forest) in the Kaluzhskie Zaseki State Nature Reserve. It is noted that bison maintains meadows in the absence of haymaking. Analysis of phytosociological relevés shows that the forest vegetation has been significantly transformed by bison over 10–15 years, as evident from (1) differences in the ecological-coenotic structure of the vegetation in the different biotopes, (2) a larger ecological-coenotic diversity of communities in the transition zones in comparison with surrounding forests, and (3) an increase in the participation of nitrophilous species in the vegetation. A fairly high earthworm species diversity (a total of seven species in the biotopes) is found; Aporrectodea caliginosa dominates in the earthworm density and biomass. The total earthworm density and biomass became lower in the transition zone than in neighboring forests and meadows, and the differences in earthworm density between the transition zone and the meadows were statistically significant.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning

Tree species composition and stand structural complexity are valuable indicators of sustainable forest management. This article aims to investigate the relative influence of forest overstorey composition and structural attributes on understorey composition and diversity, taking into account also site characteristics and broad-scale environmental variables. We sampled vascular plant species composition and forest structure in 132 plots in the Cilento and Vallo di Diano National Park (southern Italy). Spearman’s non-parametric correlation coefficients were calculated between overstorey and understorey diversity indices, beech percentage, and altitude and environmental indices. A complete partitioning of the variation in understorey composition was then performed through canonical correspondence analysis considering four sets of variables: (1) overstorey composition, (2) structural attributes, (3) topography, and (4) landscape abiotic variables. Finally, we constructed a regression tree analysis of understorey species richness using the same explanatory variables. Understorey diversity indices were positively correlated with overstorey diversity indices and with environmental indices (i.e., light and soil heterogeneity). Overstorey and understorey diversity indices were negatively correlated with both altitude and the dominance of beech in the overstorey. Compositional variation was due primarily to overstorey composition and secondarily to structural attributes. Regression tree analysis revealed that altitude, overstorey species richness, and structural attributes play an important role in determining understorey species richness. According to our results, understorey composition and diversity are strongly related to overstorey composition and structural attributes. Indeed, the latter proved to be effective indicators of understorey characteristics in the study area.     

赤子爱胜蚓对森林凋落物的分解效率

蚯蚓是帽儿山林区的主要土壤动物类群。饲养实验表明,赤子爱胜蚓(Eiseniafoetida)对不同种类、不同腐解程度的木本植物叶片的摄食量不同,对未分解叶的食性偏好依次为:白桦(Betulaplatyphylla)>杨(Populusdavidiana)>红松(Pinuskoraiensis)>蒙古栎(Quercusmongolica),对半分解叶的食性偏好依次为:杨>红松>蒙古栎>白桦,对同种叶片半分解叶的消耗能力大于未分解叶。这表明叶片种类和叶子腐化程度是影响蚯蚓摄食偏好的两个主要因素,混交林比纯林更有利于蚯蚓对难腐化叶片种类的分解。研究还表明,赤子爱胜蚓生物量在盛夏、春末以增重为主,夏初、秋初以失重为主,对落叶的消耗能力随种群生物量增大而增大,摄食量随温度的升高而增大,说明影响蚯蚓分解能力的另两个重要因素是种群生物量和温度。比较研究表明,从不适宜条件到适宜条件,蚯蚓消耗能力的理论范围约为6.378~27mg/(d·g)。能流研究表明,不同生境下,蚯蚓的各种生态效率变辐较大,即,毛增长效率:2.34%~9.92%,组织增长效率:28.95%~90.86%,同化效率:8.10%~34.27%,粪便率:65.73%~91.90%。赤子爱胜蚓的毛增长效率与同化效率约为该林区另一类主要土壤动物马陆的两倍,是帽儿山森林生态系统的重要分解者。因此,通过林木种类优化配置、温湿度条件的改变,可提高蚯蚓种群生物量及其分解能力,加快寒区厚积凋落物层的分解,加速林分更新。     

Diversity of lacewing assemblages (Neuropterida:Neuroptera) in different forest habitats and agricultural areas in the East Mediterranean area of Turkey

The objectives of this study were to understand lacewing diversity patterns and the role of environmental factors in determining lacewing diversity in four habitat types: agricultural area (crops and orchards), pine forest (Pinus nigra Arnold and, Pinus brutia Ten), natural forest (cedar and willow‐oak), and mixed forest (beech and oak). Several diversity patterns were evaluated, including evenness and dominance, in the East Mediterranean area of Turkey by analyzing the abundance and species composition of lacewing assemblages using the Simpson and Shannon Diversity Indices. It was questioned whether these communities are structured according to environmental conditions such as altitude, temperature, rainfall, wind and humidity. This question was explored using generalized linear models (GLMs) and principal component analysis (PCA) to detect the effect of ecological parameters on the species composition of the lacewing assemblages. The data suggest that the species abundance, composition, and diversity of lacewings are affected by different forest and agricultural habitat types. Based on the results of the present study, it is recommended that the area of farmland under cultivation is decreased and the area of sustainably managed forest and agricultural land is increased in order to better protect lacewing diversity in the East Mediterranean area of Turkey.     

Ecological description of ground beetle communities (Coleoptera, Carabidae) in northern taiga meadows of Arkhangelsk Region

Species composition and ecological structure of ground beetle population was studied in northern taiga meadows of the Arkhangelsk Region. Meadows in the northern forest zone proved to harbor 91 ground beetle species. Carabid complexes formed in the intrazonal biocenoses of the northern forest zone can be as rich as the family topical groups in the southern forest zone by the number of species and ecological diversity. Ecological properties of the fauna and ground beetle population proved similar in different parts of the forest zone. The proportion of stenobiotic meadow species proved to decrease while that of ecologically plastic ones increased from south to north. The proportion of the genus Harpalus decreased in the ground beetle population while the number of Amara species remained unaltered and their abundance increased. The changes in the species composition caused no transformation of the ecological structure of ground beetle population since they were limited to a single life form or guild.     

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. -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.     

Influence of forest development phase on centipede diversity in managed beech forests in Slovenia

Biodiversity in managed forests depends on management practices that determine the forest structure. The influence of stand structure on centipede diversity was investigated by comparing centipede communities from different forest development phases. Using the quadrat counts method, soil samples were taken thrice during 2000 (in spring, summer and autumn) from a deforested area and three evenly aged beech stands (juvenile, pole and timber phases) in managed beech forests in Slovenia. The characteristics of centipede communities and the similarities among them were estimated. The collection comprised 1150 centipedes representing 32 species. At a single site, 10–24 species were found. The average density was between 22 and 808 individuals per m². Differences among communities from different sites were found. The highest species number and abundance were found in the juvenile phase and the lowest in the deforested area. The occurrence of species differed among phases, but none of the species found with a high incidence in one development phase was restricted to that phase. The composition of the centipede community was more sensitive to the successional status of the forest than to the season. A comparison with differently structured forest was made, which suggests that homogenisation leads to reduced centipede diversity. To preserve biodiversity and healthy forest functioning, the maintenance of heterogeneity within a forest stand should be an important focus of management operations.     

Earthworm community structure and diversity in experimental agricultural watersheds in Northeastern Ohio

Earthworms are known to have an important impact on soil fertility but much remains to be known about the factors that influence earthworm abundance and species diversity in agricultural soils and the impact of earthworm diversity on soil processes in those soils. We have studied factors that influence earthworm community structure and biodiversity in experimental agricultural watersheds at the North Appalachian Experimental Watershed near Coshocton, Ohio. We sampled earthworm communities in seven such watersheds from 1990 to 1992. Six earthworm species were present:Aporrectodea caliginosa, A. trapezoides, A. tuberculata, Lumbricus rubellus, Lumbricus terrestris and Octolasion tyrtaeum. The total earthworm biomass ranged from 2 to 32 g m^-2 and population levels ranged from 10 to 350 worms m^-2. Earthworm community structure and diversity differed among watersheds and was influenced by cropping patterns, geographic location and tillage. The greatest earthworm diversity and highest earthworm population levels occurred in a no-tillage watershed and a watershed that had previously been in ryegrass and long-term no-till. High earthworm populations were also observed in a chisel-plowed watershed. Watersheds that had been managed identically for 50 years had much different earthworm communities, indicating that factors other than management were important in determining species composition. The three watersheds with the lowest populations and diversity were adjacent to one another at one end of the station, indicating that at this site geographic location had the predominant influence on earthworm communities. A severe drought in 1991 greatly reduced earthworm populations and biomass. However, earthworm species differed in their response to drought withLumbricus rubellus appearing to be the most drought-sensitive andAporrectodea spp. the most drought-tolerant.     

Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Exotic earthworms dispersion through protected forest areas and their potential impacts on nitrous oxide production

It is generally accepted that human activities are responsible for the dispersal of exotic earthworms in northeastern North America. We know little, however, about the relative effects of concurrent human activities on the structure of these earthworm communities in protected forest areas, nor on their impacts on soil biological activities. Our first objective was to infer the relative importance of recreational fishing and road traffic on the structure of Lumbricidae communities in Mont-Tremblant National Park, the oldest conservation area in the province of Quebec, Canada. Our second objective was to test the relationship between earthworm species abundances and soil properties related to microbial and nitrogen dynamics. We sampled earthworm communities around 61 lakes, which included 23 heavily-fished lakes and 20 non-fished lakes located near roads, as well as 18 non-fished lakes located in remote areas of the park. Our results revealed that fishing and proximity to roads both have a positive effect on the abundance of earthworms, as does the soil pH. Fishing activities had a greater effect than road proximity on the abundance and diversity of earthworm communities, notably on the abundance of the anecic species Lumbricus terrestris. To assess at a finer scale the effects of earthworm community structure on soil microbial and nitrogen dynamics, we collected and analyzed soils from 47 sampling points around two lakes with high earthworm densities. Exploratory redundancy analysis found a negative correlation between epigeic and anecic earthworm species, with the former correlating positively to microbial biomass and the latter correlating positively to nitrification and denitrification. Confirmatory path analysis established a positive indirect effect of Lumbricus terrestris, the preferred fishing bait, on potential soil nitrous oxide emissions. We conclude that the human-mediated dispersion of earthworms in the most pristine ecosystems of Quebec affects ecosystem functioning and thus requires a review of current policies regarding the use of live-bait by fishermen.     

Size-assortative mating in the earthworm Eisenia fetida (Oligochaeta, Lumbricidae)

In many species of simultaneous hermaphrodites, body size correlates with fecundity, and larger partners are preferred to small ones. Since sperm exchange is usually reciprocal, small individuals may be rejected by larger partners resulting in size-assortative mating. We studied the mating patterns in a natural population of the simultaneous hermaphroditic earthworm Eisenia fetida (Oligochaeta, Lumbricidae). We found that size-assortative mating processes existed, with variance in body weight within pairs lower than between pairs in mating earthworms. This non-random mating pattern probably reveals the existence of mate selection in this species, which lives at elevated densities with high availability of potential mates.     

The impact of beech thickets on biodiversity

Beech bark disease has dramatically altered hardwood forest structure and composition across northeastern North America. Extensive overstory mortality has resulted in prolific root-sprouting in some stands leading to the development of understory thickets of clonal small-stemmed beech. Beech thickets may impact local forest biodiversity, but this has not been adequately evaluated. We hypothesized significant differences in diversity of groundcover flora, craneflies, amphibians, and small mammals between plots with and without beech thickets. Paired plots were established in uneven-aged northern hardwood forest stands with no recent management history at two sites in the Adirondack Mountains of New York State. Groundcover plants, terrestrial craneflies, amphibians and small mammals were sampled on twenty paired plots. Discriminant analysis showed a significant difference between thicket and non-thicket (control) areas; significant variables in plot type separation were beech sapling abundance, leaf litter depth, and coarse woody debris volume. Groundcover plant cover, richness, and diversity were significantly lower in thicket compared to non-thicket plots, while beech sapling density explained 17–38 % in groundcover plant species diversity. There were no significant differences between the diversity of cranefly, amphibian and small mammal communities of each plot type. Beech thickets are important determinants of local biodiversity.     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Evidence for ongoing introduction of non-native earthworms in the Washington,DC metropolitan area

Earthworm introductions and invasions are ongoing, with significant consequences for ecological characteristics and function where populations of invasive species reach high densities. In North America the influx of people, goods and materials to coastal cities has long been recognized to be related to introduction and establishment of many different invasive organisms. We conducted surveys for soil invertebrates in the Washington, DC area along the Potomac River corridor to examine the influence of historic soil profile disrupting disturbances on the composition of soil invertebrate communities. Here we report three earthworm taxa that either (1) had never been previously reported in North America (Lumbricidae: Helodrilus oculatus), (2) had never been reported from “wild” caught samples in forested soils (Lumbricidae: Eisenia fetida), or (3) represented a notable range expansion for an introduced species (Lumbricidae: Murchieona muldali). All three species reported here have attributes that give reason for concern over their expansion into North American soils, not least of which is their potential for competitive interactions with the remaining native earthworm species.     

Phytogeographical evidence for post‐glacial dispersal limitation of European beech forest species

The post‐glacial migration of European beech Fagus sylvatica has been addressed by many studies using either genetic or fossil data or a combination of both. In contrast to this, only little is known about the migration history of beech forest understorey species. In a review of phytosociological literature, we identified 110 plant species which are closely associated with beech forest. We divided the distribution range of European beech forests into 40 geographical regions, and the presence or absence of each species was recorded for each region. We compared overall species numbers per region and numbers of narrow‐range species (species present in <10 regions). A multiple regression model was used to test for the explanatory value of three potential diversity controls: range in elevation, soil type diversity, and distance to the nearest potential refuge area. A hierarchical cluster analysis of the narrow‐range species was performed. The frequency of range sizes shows a U‐shaped distribution, with 42 species occurring in <10 regions. The highest number of beech forest species is found in the southern Alps and adjacent regions, and species numbers decrease with increasing distance from these regions. With only narrow‐range species taken into consideration, secondary maxima are found in Spain, the southern Apennines, the Carpathians, and Greece. Distance to the nearest potential refuge area is the strongest predictor of beech forest species richness, while altitudinal range and soil type diversity had little or no predictive value. The clusters of narrow‐range species are in good concordance with the glacial refuge areas of beech and other temperate tree species as estimated in recent studies. These findings support the hypothesis that the distribution of many beech forest species is limited by post‐glacial dispersal rather than by their environmental requirements.