Species-rich Habitats Research Articles

Mass mortality hits gorgonian forests at Montecristo Island.

Mediterranean gorgonian forests are species-rich habitats, and like many other marine habitats they are threatened by anthropogenic disturbances and mass mortality events. These mortality events have often been linked to anomalies in the temperature profiles of the Mediterranean region. On 5 September 2017, colonies of the gorgonians Eunicella singularis and Eunicella cavolini exhibited rapid tissue loss, down to a depth of 30 m along the steep cliffs of Montecristo Island, Tuscan Archipelago National Park, Tyrrhenian Sea, Italy. Interestingly, Montecristo has previously been identified as a reference site for the ecological quality assessment of the western Mediterranean benthic assemblages on rocky bottoms. The observed mortality event occurred during a period of increased sea temperature. By utilising a combination of high-resolution oceanographic analysis, forecast models and citizen science initiatives, we propose that an early warning system for the concomitance of heat waves and mortality events can be put in place. A temperature-based coral disease surveillance tool could then be established for the entire Mediterranean Sea. Such a tool would allow for the timely study of mass mortality phenomena and the implementation of prompt mitigation and/or restoration initiatives. Finally, this specific mortality event, in a Marine Protected Area, offers a unique opportunity to monitor and assess the resilience of gorgonian populations and associated benthic assemblages in the absence of other, more directly, anthropogenic disturbances such as pollution and land runoff.

Effects of roads and adjacent areas on diversity of terrestrial isopods of Hungarian highway verges

AbstractTransportation infrastructure may be the most important driver of social and economic development, but it is a major cause of environmental change in landscapes. The main objective of this paper is to report road edge effects on isopods of Hungarian highway verges. We examined the isopod diversity along five highways (M0, M1, M3, M5, M7) while accounting for road edge proximity and the adjacent areas. Double-glass pitfall traps were set in a total of 15 sites and at three distances from the edge of the roads next to different types of adjacent areas. We found differences between ecological parameters of isopod communities in relation to adjacent areas as well as to road edge proximity. The highest diversity was observed near urban areas, while the lowest was near the arable fields. Isopod diversity increased with decreasing distance from a road. Species diversity of different types of verges based on adjacent areas varied strongly in relation to road edge proximity. A medium distance (40 m) from roads had a positive effect on species richness, while verges next to arable fields were the most species-rich habitats. The general conclusion of this study is that highway verges provided suitable environment conditions for generalist isopod species but may be a limiting factor for specialist isopods. Moreover, highway verges function as corridors for isopods. The proximity of roads and urban areas positively affected isopods, and verges close to roads and urban areas are considered as an attractive environment for isopods in heterogeneous roadside verges.

Especies vegetales colonizadoras de áreas perturbadas por la minería en bosques pluviales tropicales del Chocó, Colombia

The Choco (Colombia) is a region with high biodiversity but dramatically affected by mining. In this work, vascular plant species that colonize abandoned mines in the rain forests of Chocoan region were identified. In particular, plants were collected from different landforms in seven abandonment mines (3-15 years of abandoned after mining activity) of three municipalities in the Choco region. Sixty-six species, 47 genera and 22 families were identified. The most representative families were Cyperaceae (14.9 % genera and 25.8% species), Melastomataceae (14.9 and 15.2%) and Rubiaceae (10.6 and 12.1%), while the more species-rich genera were Cyperus (8.5% species), Rhynchospora (8.5%), Scleria (6.4%) and Spermacoce (6.4%). The dominant life form was herbaceous (80.3% species) and more species-rich habitats were non-flood plains (36.3% species), the mine-forest edge (34.8%) and flood depressions (31.8%). The flood depressions included more exclusive species (42.8%, n = 42). Early revegetation of mine depends on the life history of colonizing plants and factors associated with the substrate.

Integrative Profiling of Bee Communities from Habitats of Tropical Southern Yunnan (China)

Understanding and managing pollination service is hindered by taxonomic impediments and paucity of data, particularly in the tropics. Herein we apply integrative species delineation and taxonomy to test impacts of land use on the diversity of bee communities within Xishuangbanna (Yunnan, south China), a highly biodiverse tropical region which has undergone extensive land conversion to rubber plantation. 128 Operational Taxonomic Units (OTU) were inferred by an iterative and integrative approach. Bee activity differed significantly across land use samples, although community composition corresponded more to level of vegetation density, when accounting for spatial structure. Species diversity was high in young rubber plantations, although composition overlapped with other species-rich habitats (natural forest edge and river banks), and older plantations (>8 years) showed very low diversity under all measures. Community structures were similar between the natural forest interior and edge, although analysis indicated contrasting drivers of diversity, with clustering in the interior and overdispersion in the forest edge. Further, phylogenetic diversity and derived indices were underestimated when reference data were omitted from analysis. The description of bee communities herein permits more informed choices in land management with respect to ensuring continuation of essential services by bees.

Estimating the magnitude of morphoscapes: how to measure the morphological component of biodiversity in relation to habitats using geometric morphometrics.

Ecological indicators are currently developed to account for the different facets of loss of biological diversity due to direct or indirect effects of human activities. Most ecological indicators include species richness as a metric. Others, such as functional traits and phylogenetic diversity, account for differences in species, even when species richness is the same. Here, we describe and apply a different indicator, called morphoscape dimension, accounting for morphological variability across habitats in a geographical region. We use the case of ground beetles (Coleoptera: Carabidae) in four different habitats in the Po Plain in Northern Italy to exemplify how to quantify the magnitude of the morphological space (i.e. the dimension of the morphoscape) occupied by the species in each habitat using geometric morphometrics. To this aim, we employed a variety of metrics of morphological disparity related to univariate size, and more complex multivariate shape and form. Our 'proof of concept' suggests that metrics assessing size and form might largely tend to simply mirror the information provided by species richness, whereas shape morphoscape disparity may be able to account for non-trivial differences in species traits amongst habitats. This is indicated by the woodland morphoscape being on average bigger than that of crops, the most species-rich habitat, despite having almost 20% less species. We conclude suggesting that the analysis of morphoscape dimension has the potential to become a new additional and complimentary tool in the hands of conservation biologists and ecologists to explore and quantify habitat complexity and inform decisions on management and conservation based on a wide set of ecological indicators.

Phylogeography of a tough rock survivor in European dry grasslands.

Phylogeographic analyses of plants in Europe have revealed common glacial refugia and migration routes for several trees and herbs with arctic-alpine distributions. The postglacial histories of dry grassland species in central Europe have rarely been analyzed, even though the extremely species-rich habitat is threatened. Sedum album (Crassulaceae) is a common inhabitant of rocky sites in central European dry grasslands. We inferred the phylogeographic history of S. album over its distribution range in Europe. Genetic diversity within and differentiation between 34 S. album populations was examined using AFLP markers. Population isolation was indicated based on the rarity of the fragments and by isolation-by-distance effects. We sequenced the trnL-trnF region in 32 populations and used chloroplast microsatellites to analyze chloroplast haplotype distributions. Two distinct S. album lineages were detected. One lineage was comprised of populations from eastern and central parts of central Europe, and the Apennine Peninsula. A second lineage was comprised of populations from the Iberian Peninsula and western and northern parts of central Europe. Glacial refugia were identified based on the accumulation of ancient chloroplast haplotypes, high diversity of AFLP fragments within populations, and high levels of rare fragments in Liguria, Serbia, the Apennine and Iberian peninsulas. Cryptic refugia were detected in the Czech Republic and Slovakia. Isolation by distance was present all over the distribution range, and it was separately detected in southwestern and central Europe. In western Europe, where a contact zone between the two lineages can be expected, no isolation by distance was detected. Our results suggest migration routes of S. album northeastward from glacial refugia in southern Iberia, northward from the Apennine Peninsula, and northward and westward from the southeastern parts of central Europe. Therefore, central European grasslands were recently colonized by northern cryptic populations and source populations originating in the east and the Apennine Peninsula.

Island biogeography theory outweighs habitat amount hypothesis in predicting plant species richness in small grassland remnants

ContextThe habitat amount hypothesis has rarely been tested on plant communities. It remains unclear how habitat amount affect species richness in habitat fragments compared to island effects such as isolation and patch size.ObjectivesHow do patch size and spatial distribution compared to habitat amount predict plant species richness and grassland specialist plant species in small grassland remnants? How does sampling area affect the prediction of spatial variables on species richness?MethodsWe recorded plant species density and richness on 131 midfield islets (small remnants of semi-natural grassland) situated in 27 landscapes in Sweden. Further, we tested how habitat amount, compared to focal patch size and distance to nearest neighbor predicted species density and richness of plants and of grassland specialists.ResultsA total of 381 plant species were recorded (including 85 grassland specialist species). A combination of patch size and isolation was better in predicting both density and richness of species compared to habitat amount. Almost 45% of species richness and 23% of specialist species were explained by island biogeography parameters compared to 19 and 11% by the amount of habitat. A scaled sampling method increased the explanation level of island biogeography parameters and habitat amount.ConclusionsHabitat amount as a concept is not as good as island biogeography to predict species richness in small habitats. Priority in landscape planning should be on larger patches rather than several small, even if they are close together. We recommend a sampling area scaled to patch size in small habitats.

Study of Plant Species Richness in Habitats with Different Grazing Intensities at Golestan National Park and Surrounding Area

Study of Plant Species Richness in Habitats with Different Grazing Intensities at Golestan National Park and Surrounding Area

The role of novel forest ecosystems in the conservation of wood-inhabiting fungi in boreal broadleaved forests.

The increasing human impact on the earth's biosphere is inflicting changes at all spatial scales. As well as deterioration and fragmentation of natural biological systems, these changes also led to other, unprecedented effects and emergence of novel habitats. In boreal zone, intensive forest management has negatively impacted a multitude of deadwood‐associated species. This is especially alarming given the important role wood‐inhabiting fungi have in the natural decay processes. In the boreal zone, natural broad‐leaved‐dominated, herb‐rich forests are threatened habitats which have high wood‐inhabiting fungal species richness. Fungal diversity in other broadleaved forest habitat types is poorly known. Traditional wood pastures and man‐made afforested fields are novel habitats that could potentially be important for wood‐inhabiting fungi. This study compares species richness and fungal community composition across the aforementioned habitat types, based on data collected for wood‐inhabiting fungi occupying all deadwood diameter fractions. Corticioid and polyporoid fungi were surveyed from 67 130 deadwood particles in four natural herb‐rich forests, four birch‐dominated wood pastures, and four birch‐dominated afforested field sites in central Finland. As predicted, natural herb‐rich forests were the most species‐rich habitat. However, afforested fields also had considerably higher overall species richness than wood pastures. Many rare or rarely collected species were detected in each forest type. Finally, fungal community composition showed some divergence not only among the different habitat types, but also among deadwood diameter fractions. Synthesis and applications: In order to maintain biodiversity at both local and regional scales, conserving threatened natural habitat types and managing traditional landscapes is essential. Man‐made secondary woody habitats could provide the necessary resources and serve as surrogate habitats for many broadleaved deadwood‐associated species, and thus complement the existing conservation network of natural forests.

Tree phylogenetic diversity promotes host-parasitoid interactions.

Evidence from grassland experiments suggests that a plant community's phylogenetic diversity (PD) is a strong predictor of ecosystem processes, even stronger than species richness per se This has, however, never been extended to species-rich forests and host-parasitoid interactions. We used cavity-nesting Hymenoptera and their parasitoids collected in a subtropical forest as a model system to test whether hosts, parasitoids, and their interactions are influenced by tree PD and a comprehensive set of environmental variables, including tree species richness. Parasitism rate and parasitoid abundance were positively correlated with tree PD. All variables describing parasitoids decreased with elevation, and were, except parasitism rate, dependent on host abundance. Quantitative descriptors of host-parasitoid networks were independent of the environment. Our study indicates that host-parasitoid interactions in species-rich forests are related to the PD of the tree community, which influences parasitism rates through parasitoid abundance. We show that effects of tree community PD are much stronger than effects of tree species richness, can cascade to high trophic levels, and promote trophic interactions. As during habitat modification phylogenetic information is usually lost non-randomly, even species-rich habitats may not be able to continuously provide the ecosystem process parasitism if the evolutionarily most distinct plant lineages vanish.

Assemblages of saproxylic beetles on large downed trunks of oak.

Old living oaks (Quercus robur) are known as a very species‐rich habitat for saproxylic beetles, but it is less clear to what extent such veteran trees differ from an even rarer feature: downed trunks of large oaks. In this study, we set out to sample this habitat, using window traps, with two aims: (1) to describe the variation of assemblages among downed trunks of different type and (2) to compare beetles on downed oaks with data from veteran standing trees. The results showed that trunk volume and sun exposure better explained assemblages as well as species numbers on downed trunks than did decay stage. Furthermore, species classified as facultative saproxylic species showed weak or no differentiation among downed trunks. Species with different feeding habits showed no apparent differentiation among downed trunks. Furthermore, species composition on dead, downed oak trunks differed sharply from that of living, veteran oaks. Wood or bark feeders were more common on veterans than downed trunks, but there was no difference for those species feeding on fungi or those feeding on insects and their remains. In conclusion, for a successful conservation of the saproxylic beetle fauna it is important to keep downed oak trunks, and particularly large ones, in forest and pastures as they constitute a saproxylic habitat that differs from that of living trees.

Advances in ecological studies of epiphytes using canopy cranes

Forest canopies are one of the most species-rich habitats, but among the least explored in the biosphere. They play a crucial role in the process of material and energy exchange between the forest and atmosphere. Individual ecosystem members (e.g., epiphytes) and the ecological function of canopies have been given insufficient attention because of inaccessibility. Canopy cranes have been successfully used to guarantee non-destructive and reiterated sampling of epiphytes, thus offering a top-down perspective of the entire canopy. These cranes have become the symbol of canopy research and enable epiphyte research. Globally, western developed countries have conducted many studies of diversity and spatial distributions of epiphytes using canopy cranes, thus accumulating an abundance of valuable results. This review summarizes the structure, development history, and distribution of canopy cranes as well as general information about international canopy research organizations. Ecological studies of epiphytes performed around the world using these canopy cranes are also reviewed. Additionally, the development of canopy ecology and the construction of canopy cranes in China are introduced briefly. In analyzing current research trends in ecological studies of epiphytes in China and globally, the following aspects were considered: biodiversity, spatial patterns and maintenance mechanisms, ecological adaptations of epiphytes, their relationship with canopy animals, and their responses to climate change.

Rainfall fluctuations and vegetation patterns in alkali grasslands - using self-organizing maps to visualise vegetation dynamics

Knowledge about the drivers of vegetation dynamics in grasslands is fundamental to select appro-priate management for conservation purposes. In this study, we provide a detailed analysis of vegeta-tion dynamics in alkali grasslands, a priority habitat of the Natura 2000 network. We studied vegetation dynamics in five stands of four alkali grassland types in the Hortobagy National Park (eastern Hunga-ry), between 2009 and 2011. We analysed the effect of fluctuations in precipitation on both the overall vegetation composition and on the cover of each species using Self Organizing Map neural networks (SOM). We found that SOM is a promising tool to reveal plant community dynamics. As we analysed species cover and overall vegetation composition separately, we were able to identify the species re-sponsible for particular vegetation changes. Fluctuations in precipitation (a dry season, followed by a wet and an average season) caused quick shifts in plant species composition because of an increasing cover of halophyte forbs, probably because of salinisation. We observed a similar effect of stress from waterlogging in all studied grassland types. The species composition of Puccinellia grasslands was the most stable over the three years with varying precipitation. This was important as this grassland type contained many threatened halophyte species. Self-organising maps revealed small-scale vegetation changes and provided a detailed visualisation of short-term vegetation dynamics, thus we suggest that the application of this method is also promising to reveal community dynamics in more species-rich habitat types or landscapes.

Rastlinski Svet Dolomitov

In Europe, the management of semi-natural grasslands is changing steadily in extent and intensity. These changes represent a serious threat to one of the most species-rich habitats in Europe; therefore, the related shifts in species composition should be assessed by means of sound scientific. We describe a new methodology aimed at: (i) quantifying the temporal changes in vascular plant species composition of semi-natural grasslands; (ii) identifying the environmental and management variables that drive patterns of change. Our approach consists of a diachronic analysis based on historical phytosociological data associated with detailed vegetation maps. To describe and test our methodology, in 2013 we re-visited 24 historical plots of Bromus erectus grasslands that were first sampled in 1982. To designate the new sampling sites, we combined the topographic and typological information available for the historical plots with the spatial information of the associated map. The degree of change in species composition was calculated through ordination techniques; multivariate distances were used in a regression analysis to identify the environmental variables responsible for patterns of compositional change. Our worked example demonstrated that species composition has changed significantly in the last 30 years, with patterns that have been substantially influenced by topography and landscape structure.

Local topography affects seed bank successional patterns in alluvial meadows

Seed banks may play a crucial role in the maintenance of community diversity, but their role on semi-natural grasslands, one of the most species rich habitats in Europe, is usually unexplored in population and community studies. We aim to clarify how local factors and topography influence seed bank successional patterns on semi-natural alluvial meadows. All 35 meadows were situated in the Eagle (Orlické) Mountains in the Czech Republic and were divided according to fallow time. Paired plots, represented low and high topography, were established on each meadow. We recorded plant species occurrence in vegetation and in the seed bank to ascertain how seed abundance, diversity and the similarity between seed bank and vegetation were influenced by topography and management. Seed abundance significantly changed, whereas seed diversity remained stable after meadow abandonment. Seed abundance was high in the low topography plots probably due to higher aboveground biomass which promoted seed accumulation. In the high topography plots, seed abundance was low in managed meadows, but seeds were accumulated after abandonment. Similarity between the seed bank and vegetation was low, but over half of the typical meadow species survived in the seed bank for long periods of time, even when they had disappeared from the aboveground vegetation. In conclusion, the local topography of alluvial meadows influenced seed bank successional patterns, mainly seed abundance, similarity between the seed bank and vegetation and responses of individual species in the seed bank. Seed bank successional patterns are dependent on specific habitat conditions and communities with high seed bank accumulation are more stable and better restored than communities where the seed bank has become severely depleted.

Identifying Shifts in Leaf-Litter Ant Assemblages (Hymenoptera: Formicidae) across Ecosystem Boundaries Using Multiple Sampling Methods.

Global or regional environmental changes in climate or land use have been increasingly implied in shifts in boundaries (ecotones) between adjacent ecosystems such as beech or oak-dominated forests and forest-steppe ecotones that frequently co-occur near the southern range limits of deciduous forest biome in Europe. Yet, our ability to detect changes in biological communities across these ecosystems, or to understand their environmental drivers, can be hampered when different sampling methods are required to characterize biological communities of the adjacent but ecologically different ecosystems. Ants (Hymenoptera: Formicidae) have been shown to be particularly sensitive to changes in temperature and vegetation and they require different sampling methods in closed vs. open habitats. We compared ant assemblages of closed-forests (beech- or oak-dominated) and open forest-steppe habitats in southwestern Carpathians using methods for closed-forest (litter sifting) and open habitats (pitfall trapping), and developed an integrated sampling approach to characterize changes in ant assemblages across these adjacent ecosystems. Using both methods, we collected 5,328 individual ant workers from 28 species. Neither method represented ant communities completely, but pitfall trapping accounted for more species (24) than litter sifting (16). Although pitfall trapping characterized differences in species richness and composition among the ecosystems better, with beech forest being most species poor and ecotone most species rich, litter sifting was more successful in identifying characteristic litter-dwelling species in oak-dominated forest. The integrated sampling approach using both methods yielded more accurate characterization of species richness and composition, and particularly so in species-rich forest-steppe habitat where the combined sample identified significantly higher number of species compared to either of the two methods on their own. Thus, an integrated sampling approach should be used to fully characterize changes in ant assemblages across ecosystem boundaries, or with vegetation change over time, and particularly so in species-rich habitats such as forest-steppe ecotones.

Combining Approaches to Support High Nature Value Farmland in Southern Transylvania , Romania

Abstract The region of Tarnava Mare in Southern Transylvania contains extensive semi-natural open landscapes maintained by predominantly low-intensity farming, which is widespread in Romania and indeed many areas of Eastern Europe. Threats to these species-rich habitats from agricultural intensification and land abandonment have been increasing in recent years, to a large extent linked with Romania’s accession to the EU in 2007. At the same time, however, the opportunities for biodiversity conservation in the area have expanded. In 2008, the region became a Site of Community Importance (SCI) as part of the Natura 2000 network, and farmers have applied agri-environment schemes as part of the EU Common Agricultural Policy since 2006. Furthermore, the Tarnava Mare region has been the location of several EU and nationally funded projects combining research, practical and information measures. In this article, we review these various instruments from the practical perspective of an NGO that has been working since 2004 to support High Nature Value farmland and rural communities in this region. We focus on three major support measures - agri-environment schemes, Natura 2000, and publicly funded conservation projects - and consider their effects individually and collectively. We conclude that the presence of multiple instruments can have synergistic effects on the conservation of semi-natural open habitats such as HNV farmland, and that this overlap provides a certain amount of resilience: if one instrument fails, another may fill the gap. Cross-cutting projects combining research with activities to tackle the “problem” of the socio-economic undesirability of low-intensity farming as well as the “symptom” of the loss of HNV farmland are also particularly important in this context.

Nitrogen, phosphorus, and cation use efficiency in stands of regenerating tropical dry forest

Plants on infertile soils exhibit physiological and morphological traits that support conservative internal nutrient cycling. However, potential trade-offs among use efficiencies for N, P, and cations are not well explored in species-rich habitats where multiple elements may limit plant production. We examined uptake efficiency and use efficiency of N, P, K, Ca, Mg, Al, and Na in plots of regenerating tropical dry forests spanning a gradient of soil fertility. Our aim was to determine whether plant responses to multiple elements are correlated, or whether there are trade-offs among exploitation strategies across stands varying in community composition, soil quality, and successional stage. For all elements, both uptake efficiency and use efficiency decreased as availability of the corresponding element increased. Plant responses to N, Na, and Al were uncoupled from uptake and use efficiencies for P and essential base cations, which were tightly correlated. N and P use efficiencies were associated with shifts in plant species composition along the soil fertility gradient, and there was also a trend towards increasing N use efficiency with stand age. N uptake efficiency was positively correlated with the abundance of tree species that associate with ectomycorrhizal fungi. Taken together, our results suggest that successional processes and local species composition interact to regulate plant responses to availability of multiple resources. Successional tropical dry forests appear to employ different strategies to maximize response to N vs. P and K.

Historic hay cutting dates from Sweden 1873–1951 and their implications for conservation management of species-rich meadows

Semi-natural hay meadows are species rich habitats, formed by a long history of management and they have experienced a drastic decline all over Europe. There is a vast literature on conservation and species diversity of semi-natural hay-meadows, but very limited information on historic timing of hay cutting. We analyzed data collected between 1873 and 1951 on hay cutting dates and phenology of six plant species from farms distributed across Sweden. The data set comprised 16,015 observations from 175 sites. Results show that date of start and end of hay cutting varied across Sweden. The start of hay cutting was generally delayed by 2.2days per latitudinal degree and 1.5days per 100m altitude, while the end of hay cutting was generally delayed by 2.9days per latitudinal degree and 2.5days per 100m altitude. The average hay cutting period was 18.5±6.6days, and became slightly shorter northwards. Site-specific factors had a great impact on when hay cutting was performed, as indicated by a significant correlation between flowering (and leafing) phenology in other species and start date of hay cutting. Today, management for conservation is usually related to a calendar date (e.g. regulated in eligibility criteria and requirements for payment in agri-environment programs in EU). In order to mimic historic management that formed this habitat, management should instead account for latitude and altitude, between-year variation in timing of hay cutting, variation in both start and end dates of hay cutting and if possible local phenological conditions.

Modeling Wood Encroachment in Abandoned Grasslands in the Eifel National Park - Model Description and Testing.

The degradation of natural and semi-natural landscapes has become a matter of global concern. In Germany, semi-natural grasslands belong to the most species-rich habitat types but have suffered heavily from changes in land use. After abandonment, the course of succession at a specific site is often difficult to predict because many processes interact. In order to support decision making when managing semi-natural grasslands in the Eifel National Park, we built the WoodS-Model (Woodland Succession Model). A multimodeling approach was used to integrate vegetation dynamics in both the herbaceous and shrub/tree layer. The cover of grasses and herbs was simulated in a compartment model, whereas bushes and trees were modelled in an individual-based manner. Both models worked and interacted in a spatially explicit, raster-based landscape. We present here the model description, parameterization and testing. We show highly detailed projections of the succession of a semi-natural grassland including the influence of initial vegetation composition, neighborhood interactions and ungulate browsing. We carefully weighted the single processes against each other and their relevance for landscape development under different scenarios, while explicitly considering specific site conditions. Model evaluation revealed that the model is able to emulate successional patterns as observed in the field as well as plausible results for different population densities of red deer. Important neighborhood interactions such as seed dispersal, the protection of seedlings from browsing ungulates by thorny bushes, and the inhibition of wood encroachment by the herbaceous layer, have been successfully reproduced. Therefore, not only a detailed model but also detailed initialization turned out to be important for spatially explicit projections of a given site. The advantage of the WoodS-Model is that it integrates these many mutually interacting processes of succession.