Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.     

Evidence for human-mediated dispersal of exotic earthworms: support for exploring strategies to limit further spread

As potent ecosystem engineers, non-native earthworms are altering the fundamental structure and function of previously earthworm-free cold-temperate hardwood forests in North America. Discarded earthworms used for fishing bait has been presumed to be an important vector for the continued spread of non-native earthworms because epicentres of invasion often include boat landings, lakeshores and roads. However, controversy has remained about the overall importance of human-mediated spread vs. natural expansion of established earthworm populations. In this issue of Molecular Ecology , Cameron et al . explore the continued role of humans in dispersing non-native earthworms.     

Invasion Patterns of Lumbricidae Into the Previously Earthworm-free Areas of Northeastern Europe and the Western Great Lakes Region of North America

We examine the patterns of expansion of exotic European earthworms in northeastern Europe and the western Great Lakes region of North America. These areas share many ecological, climatic and historical characteristics and are devoid of indigenous earthworm fauna due to Quaternary glaciations. These regions are being colonized by a similar suite of exotic lumbricid species and it is unlikely that this is the result of chance, but rather indicates that these species have particular characteristics making them successful invaders. The present macro-scale distributions of earthworm species in northern Russia show little connection to the pattern of the last glaciation. Rather, the primary factors that determine the current distributions of earthworm species include climatic conditions, the life history traits of different earthworm species, the suitability of habitat and intensity and patterns of human activity. In the western Great Lakes region of North America, there are three primary factors affecting current distributions of exotic earthworm species including the patterns of human activity and land use practices, the composition of particular source populations of earthworms associated with different vectors of transport and the soil and litter properties of habitats across the region. Disturbance of a habitat does not appear to be a prerequisite to the invasion and establishment of exotic earthworms. Analysis of the macro-scale distributions of Lumbricidae species in northeastern Europe may provide important insights into the potential of invasive European earthworm species to spread in North America, and identify potentially invasive species.     

Earthworm invasions in the tropics

The effects and implications of invasive species in belowground terrestrial ecosystems are not well known in comparison with above-ground terrestrial and marine environments. The study of earthworm invasions in the tropics is limited by a lack of taxonomic knowledge and the potential for loss of species in native habitats due to anthropogenic land use change. Alteration of land use plays a major role in determining the abundance and community structure of earthworms and the establishment of exotic earthworms in areas previously inhabited by worms. Once an exotic species has become established into a new place, site and species characteristics seem to be key factors determining their spread. We reviewed the literature on the distribution and effects of exotic earthworms to understand the interactions of earthworm invasion and land use history in the tropics. Patterns in the abundance, effects and mechanisms of earthworm invasions on ecosystem processes in the tropics are elucidated using Pontoscolex corethrurus as a case study.     

Genetic structure of invasive earthworms Dendrobaena octaedra in the boreal forest of Alberta: insights into introduction mechanisms

Population genetic studies can help to determine whether invasive species are established via single vs. multiple introduction events and also to distinguish among various colonization scenarios. We used this approach to investigate the introduction of Dendrobaena octaedra , a non-native earthworm species, to the boreal forest of northern Alberta. The spread of non-native earthworms in forested systems is not well understood, although bait abandonment and vehicular transport are believed to be important. Mitochondrial DNA sequencing revealed that multiple introductions of this species have occurred in northern Alberta, although individual populations may have been established by either single or multiple invaders introduced on one or more occasions. There was no relationship between genetic distances and either geographical distances or distances along road networks, suggesting that human-mediated jump dispersal is more common than diffusive spread via road networks or via active dispersal. As well, genetic diversity was significantly greater at boat launches than roads, indicating that multiple introductions may be more likely to occur at those locations. Focusing management efforts on areas where multiple introductions are likely to occur may help to reduce invasive species' potential for adaptive evolution and subsequent rapid spread.     

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta‐analysis)

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta‐analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non‐native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non‐native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm‐invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long‐lasting effects on ecosystem functioning.     

Human-mediated introduction of geoengineering earthworms in the Fennoscandian arctic

It is now well established that European earthworms are re-shaping formerly glaciated forests in North America with dramatic ecological consequences. However, few have considered the potential invasiveness of this species assemblage in the European arctic. Here we argue that some earthworm species (Lumbricus rubellus, Lumbricus terrestris and Aporrectodea sp.) with great geomorphological impact (geoengineering species) are non-native and invasive in the Fennoscandian arctic birch forests, where they have been introduced by agrarian settlers and most recently through recreational fishing and gardening. Our exploratory surveys indicate no obvious historical dispersal mechanism that can explain early arrival of these earthworms into the Fennoscandian arctic: that is, these species do not appear to establish naturally along coastlines mimicking conditions following deglaciation in Fennoscandia, nor were they spread by early native (Sami) cultures. The importance of anthropogenic sources and the invasive characteristics of L. rubellus and Aporrectodea sp. in the arctic is evident from their radiation outwards from abandoned farms and modern cabin lawns into adjacent arctic birch forests. They appear to outcompete previously established litter-dwelling earthworm species (i.e. Dendrobaena octaedra) that likely colonized the Fennoscandian landscape rapidly following deglaciation via hydrochory and/or dispersal by early Sami settlements. The high geoengineering earthworm biomasses, their recognized ecological impact in other formerly glaciated environments, and their persistence once established leads us to suggest that geoengineering earthworms may pose a potent threat to some of the most remote and protected arctic environments in northern Europe.     

Ecosystem effects of non-native earthworms in Mid-Atlantic deciduous forests

In many mid-Atlantic forests where both native and non-native earthworms exist, it is the non-native species that are the dominant component of the soil macrofauna. Few earthworm ecology studies, however, focus attention on these forest systems in order to determine the relative ecological roles and potential interactions of the native and non-native earthworms. In a series of field samplings and experimental manipulations we collected data on the effects of earthworms on below-and aboveground ecosystem processes. Earthworm abundance and the ecological processes measured were dynamic in space and time across the range of study sites. Leaf litter decay rates doubled at sites that had abundant non-native earthworms. Earthworms also altered the abundance of soil fungi, the activity of extracellular enzymes, soil respiration, and the growth of tree seedlings but the effects varied among sites depending on differences in land-use history and forest age. Red oak seedling growth was less at sites that had abundant earthworms but tulip poplar and red maple seedlings grew equally well with and without abundant earthworms. These preliminary results suggest that non-native earthworms have significant ecosystem effects, even in forests where native earthworms still occur. Land use history, however, plays an important role in determining what those effects will be, and these effects are likely to be dynamic, depending on the abundance of non-native earthworms.     

Refuting a controversial case of a human-mediated marine species introduction

Human activities have strongly impacted natural communities through the introduction of non-native species in historical times. A frequently cited marine example is Littorina littorea , a common intertidal gastropod that was first reported in North America in 1840. The seemingly sudden appearance and rapid geographical spread of this species southward from Nova Scotia has led many researchers to consider L. littorea a human-mediated species introduction. This is despite allozyme and subfossil evidence that the `European periwinkle' was in North America long before 1840. Our mitochondrial and nuclear DNA sequence data confirm that L. littorea has been in continuous residence in North America for at least 8000 years. It appears most likely that ecological interactions, rather than oceanographic or climatic forces, maintained the limited geographical distribution of L. littorea prior to the 19th century.     

Condition status and parasite infection of Neogobius kessleri and N. melanostomus (Gobiidae) in their native and non-native area of distribution of the Danube River

The success of introduced species is often facilitated by escape from the effects of natural predators and parasites. Introduced species can profit from this favourable situation, attaining higher population densities and greater individual sizes in novel areas. In this study, somatic condition and parasite infection were compared between native and non-native populations of Neogobius kessleri Günther; introduced only within the interconnected Danube and Rhine River system, and N. melanostomus (Pallas); widely introduced throughout several river systems in Europe and North America. Higher values of Fulton’s condition factor were observed in non-native populations of both goby species. Neogobius melanostomus attained higher gonadosomatic index values in non-native populations, indicating potential increased investment in reproduction in its new area. A lower splenosomatic index was observed in non-native populations, especially in N. melanostomus. Parasite infracommunity richness and mean abundance were higher in N. kessleri in both native and non-native populations, suggesting higher susceptibility of N. kessleri to these parasites. Non-native populations of both hosts showed higher infra-community richness as a result of acquiring parasites native to the new area, but lower parasite abundance. Differences in success of the introduction and establishment in new areas between the two fish species may be associated with a relatively low parasite infection rate and a higher gonadosomatic index in non-native populations of N. melanostomus in comparison to N. kessleri.     

The influence of habitat provisioning: use of earthworm burrows by the terrestrial salamander, Plethodon cinereus

Studies examining the influence of habitat provisioning by one species on the behavior of other species can provide key insights regarding impacts of ecosystem engineers on the availability of resources to other species. More specifically, an organism’s use of additional habitat provided by ecosystem engineers may affect the interpretation of observational or demographic data. We chose to examine the possible influence of earthworms, as ecosystem engineers, on the behavior of terrestrial salamanders, common forest vertebrates in North America. We conducted two experiments in microcosms to examine whether the earthworm Lumbricus terrestris acted as an ecosystem engineer by providing additional habitat (burrows) for Plethodon cinereus, a common woodland salamander. We also examined whether the behavioral changes of the non-burrowing, fossorial P. cinereus differed across age classes. Adults and juveniles responded similarly, with both adults and juveniles found under cover objects more often when earthworms were absent and using earthworm burrows when available. Our field data supported our experimental data in that greater numbers of earthworms on field plots were associated with a lower proportion of cover objects being occupied by salamanders (i.e., salamanders were likely underground). We found no differences in the response of adults and juveniles to the presence of earthworms. By providing underground habitat for a common terrestrial amphibian, the presence of earthworms may impact demographic studies on salamanders and conclusions drawn from those studies regarding aboveground faunal communities in eastern North America.     

Do non-native earthworms in Southeast Alaska use streams as invasional corridors in watersheds harvested for timber?

Exotic earthworms from Europe and Asia have invaded previously earthworm-free areas of North America where they consume leaf litter, mix soil horizons, and alter nutrient cycling. Primarily, earthworm introductions occur through human activities; we hypothesized that the combination of logging (i.e., road construction and soil disturbance) and stream transport (i.e., hydrochory) allows earthworms to invade new ecosystems and spread within watersheds. On Prince of Wales Island, AK, we surveyed riparian zones in 11 watersheds with varying timber harvest intensity for terrestrial oligochaetes. Additionally, common invasive earthworms were experimentally submerged in a local stream to test for tolerance to prolonged immersion: all taxa survived immersion for at least 6 days. Using principal components analysis, watershed and harvest variables describing the watersheds upstream of our sampled riparian areas were reduced to two principal components describing harvest intensity (PC1) and harvest style (PC2). Logistic models successfully predicted earthworm abundance (r ²  = 0.70) from PC1, which indicated that watersheds with older, intense upstream timber harvest contained larger earthworm populations. Earthworm species richness was best predicted by PC2 (r ²  = 0.39), which suggested that earthworm communities in watersheds containing large clear-cut stands were more species-rich. Collectively, these results suggest that (1) invasive earthworms may use streams for dispersal and (2) upstream introductions via timber harvest can initiate downstream earthworm invasions. Hydrochory would allow invasive earthworms to spread at rates (tens of km d⁻¹) that are much greater than previously reported rates of terrestrial spread (5–10 m y⁻¹). Effective control of exotic earthworms in riparian zones will require watershed-level management and surveillance.     

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.     

Northward range extension of an endemic soil decomposer with a distinct trophic position

Ecological niche theory asserts that invading species become established only if introduced propagules survive stochastic mortality and can exploit resources unconsumed by resident species. Because their transportation is not controlled by plant health or biosecurity regulations, soil macrofauna decomposers, including earthworms are probably introduced frequently into non-native soils. Yet even with climatic change, exotic earthworm species from southern Europe have not been reported to become established in previously glaciated areas of northern Europe that already have trophically differentiated earthworm communities of ‘peregrine’ species. We discovered established populations of the earthworm Prosellodrilus amplisetosus (Lumbricidae), a member of a genus endemic to southern France, in six habitats of an urban farm in Dublin, Ireland, about 1000 km north of the genus''s endemic range. Not only was P. amplisetosus the dominant endogeic (geophagous) earthworm species in two habitats, it also occupied a significantly different trophic position from the resident species, as evinced by stable isotope ratio analysis. The suggested ability of this non-native species to feed on and assimilate isotopically more enriched soil carbon (C) and nitrogen fractions that are inaccessible to resident species portends potential implications of decomposer range expansions for soil functioning including C sequestration.     

Threat of non-native crayfish introductions into Turkey: global lessons

Introductions of crayfish species from their home range to new environments have been carried out in many parts of the world. The most important introduced crayfish species are Procambarus clarkii, Pacifastacus leniusculus, Cherax destructor, C. quadricarinatus, Orconectes limosus, O. rusticus and Astacus leptodactylus. The environmental impact of crayfish introductions can be positive, negative or neutral. However, native crayfish populations in Europe have been negatively affected by introductions of non-indigenous crayfish species from America. Negative effects of non-native crayfish introductions included displacement of native crayfish species, transfer of disease (crayfish plague), consumption of fish eggs, reduction of fish stocks, consumption of large amounts of macrophytes, indirect and direct effects on other invertebrates and upsetting production in rice fields. As a result of non-native crayfish introductions, the natural harvest and crayfish industry in Europe have been severely affected. Large quantities of Turkish A. leptodactylus were harvested (approximately 7,000 tonnes annually) and exported to Europe before the crayfish plague was observed in these populations. The total harvest of A. leptodactylus in Turkey reduced dramatically to 320 in 1991 after the plague. Therefore, although Turkey currently has no known non-native crayfish species, there is a threat of non-native crayfish introduction in order to increase crayfish productions and subsequent harvest. The North American spiny-cheek crayfish, O. limosus, has been spreading quickly down the River Danube and could soon reach neighboring countries including Turkey. The North American signal crayfish, P. leniusculus is known from Greece and could be a threat to native stocks if it is introduced into Turkey for aquaculture. Additional threats may come from the release of other North American species, which are widely available through the aquarium trade. We conclude that the spread of non-native crayfish introductions throughout Turkey will increase local problems, because introductions of non-native crayfish in many parts of the world have been known to have caused important reductions in population density and numbers of native crayfish species. Furthermore, freshwater ecosystems may be altered by such introductions and the economic viability of native crayfish species fisheries could be severely reduced in Turkey.     

Non-native earthworms in riparian soils increase nitrogen flux into adjacent aquatic ecosystems

Riparian zones are an important transition between terrestrial and aquatic ecosystems, and they function in nutrient cycling and removal. Non-native earthworms invading earthworm-free areas of North America can affect nutrient cycling in upland soils and have the potential to affect it in riparian soils. We examined how the presence of earthworms can affect riparian nutrient cycling and nutrient delivery to streams. Two mesocosm experiments were conducted to determine how (1) the biomass of earthworms and (2) earthworm species can affect nutrient flux from riparian zones to nearby streams and how this flux can affect streamwater nutrients and periphyton growth. In separate experiments, riparian soil cores were amended with one of four mixed earthworm biomasses (0, 4, 10, or 23 g m(-2) ash-free dry mass) or with one of three earthworm species (Aporrectodea caliginosa, Lumbricus terrestris, L. rubellus) or no earthworm species. Riparian soil cores were coupled to artificial streams, and over a 36-day period, we measured nutrient leaching rates, in-stream nutrient concentrations, and periphyton growth. Ammonium leaching increased with increasing biomass and was greatest from the A. caliginosa treatments. Nitrate leaching increased through time and increased at a greater rate with higher biomass and from cores containing A. caliginosa. We suggest that the overall response of increased nitrate leaching [90% of total nitrogen (N)] was due to a combination of ammonium excretion and burrowing by earthworms, which increased nitrification rates. During both experiments, periphyton biomass increased through time but did not differ across treatments despite high in-stream inorganic N. Through time, in-stream phosphorus (P) concentration declined to <5 microg l(-1), and periphyton growth was likely P-limited. We conclude that activities of non-native earthworms (particularly A. caliginosa) can alter biogeochemical cycling in riparian zones, potentially reducing the N-buffering capacity of riparian zones and altering stoichiometric relationships in adjacent aquatic ecosystems.     

Non-native earthworms alter the assembly of a meadow plant community

Non-native earthworms can alter ecosystems by modifying soil structure, depredating seeds and seedlings, and consuming soil organic matter, yet the initial responses of plant communities to earthworm invasions remain poorly understood. We assessed the effect of non-native earthworms on seedling survival during germination and after establishment using six native and six non-native plant species grown from seed in single- and multi-species experimental mesocosms. We examined the extent to which earthworms (1) influenced seedling survival, (2) selectively depredated native versus non-native plants, (3) impacted establishment based on seed size and/or root morphology, and (4) shaped community assembly. The effect of earthworms on seedling survival varied temporally and among species but inconsistently with respect to species origin. Differences in seed/seedling survival translated to changes in community assembly. Earthworms tended to reduce species abundance, richness, evenness, and diversity in multi-species mesocosms and led to the divergence of communities by treatment. In general, species with large seeds and fibrous roots dominated communities with earthworms present, whereas species with small seeds and taproots only persisted in multi-species mesocosms without earthworms. Our findings suggest that earthworms act as ecological filters in the early stages of invasion to shape community composition based on plant morphological traits.

     

Spatial variability of an invasive earthworm (<Emphasis Type="Italic">Amynthas agrestis</Emphasis>) population and potential impacts on soil characteristics and millipedes in the Great Smoky Mountains National Park,USA

European and Asian earthworm invasions are widespread in North America. European earthworms especially are well-known to cause dramatic changes in ecosystems in northern, formerly glaciated portions of the continent, but less is known about the impacts of earthworm invasions in unglaciated areas inhabited by indigenous earthworms. We monitored fluctuations in the spatial extent of an Amynthas agrestis (Megascolecidae) population in the Great Smoky Mountains National Park in eastern Tennessee, USA. Two years of monthly growing-season observations revealed that the distribution of the earthworm population was dynamic, but overall distribution was closely linked to temperature and moisture with dramatic reductions of earthworm numbers associated with very dry conditions. In plots where A. agrestis were more often detected, we measured increased A-horizon soil aggregation and decreased thickness of the Oe/Oa-horizon. However, A. agrestis was not related to A-horizon microbial biomass, A-horizon C:N, Oi-horizon (litter) thickness, or mass of forest floor (O-horizon). Reductions in millipede species richness and density were associated with frequency of A. agrestis presence, possibly due to direct competition for food resources (Oe/Oa material). This evidence for potentially negative interactions between millipedes and A. agrestis suggests that expansion of the non-native earthworm into new habitats in the Park may alter soil physical properties and could pose a threat to native millipede diversity.     

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.     

Disturbances in deciduous temperate forest ecosystems of the northern hemisphere: their effects on both recent and future forest development

Disturbances in forests can kill mature trees, but also create the conditions necessary for the establishment of new tree cohorts and create micro-habitats for new plant and animal species, thereby increasing the species diversity compared to undisturbed stands. We review the types and intensities of disturbances on forests in three regions of the temperate zone of the northern hemisphere: northeastern North America, Central Europe, and East Asia. We focus on (1) the ways in which disturbances affect forest stand development; (2) the differences among the three areas in this regard; (3) the consequences for future forest management. In both northeastern North America and East Asia, hurricanes and typhoons represent the major mode of natural disturbance, while in Central Europe winter windstorms occur after deciduous trees have lost their leaves. Tornadoes can have even greater destructive power (but affect relatively narrow strips of land), and the more severe of these mainly occur in North America. The general disturbance patch system therefore is relatively large in northeastern North America, small in Central Europe, and of intermediate size in temperate East Asia. In addition to wholly natural disturbance factors, human commerce and globalization have enabled new disturbance types by introducing pests and diseases from one region to another. In North America especially, several of the most important foundation species in temperate forests are strongly affected, so that not just the species composition but also the whole forest structure is changing fundamentally. In all three areas in the past the change in land use by growing human populations strongly affected the structure as well as the species composition of forests. Nearly all the recent forest stands of the temperate zone had been used in the past in a particular way, and many of today’s forests had previously been converted into agricultural land. Finally climate change is superimposing itself on forest development worldwide. Nevertheless, climate change is not a new phenomenon, so forest ecosystems in all time periods have been exposed to changing climatic conditions and have had to adapt. Each forest stand therefore represents a unique recent expression of the interaction of environmental conditions and plant species, a “snapshot” of the relevant abiotic and biotic factors, including human impact.