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We propose to consider semi-natural habitats—hotspots for biodiversity—being caught in a socio-ecological extinction vortex, similar to the phenomenon described for species threatened with extinction. These habitats are essentially socioecological systems, in which socioeconomic drivers are interlinked with ecological processes. We identify four highly interlinked and mutually reinforcing socio-economic processes, pertaining to the importance of semi-natural habitats for (i) agricultural production, (ii) policy, research and development; (iii) vocational education in the fields of agricultural sciences and (iv) public’s experiences with semi-natural habitats. Evidence from six countries in the boreal region demonstrates that recent slowing down or even reversal of two processes are insufficient to stop the extinction vortex phenomenon. We suggest research directions to ascertain the phenomenon, monitor its development and develop proactive actions to weaken the vortex. It is highly plausible that interventions directed at most, if not all, of the key vortex processes are needed to reverse the overall deteriorating trends of a socio-ecological system.

Sustainability of livestock production is a highly contested issue in agricultural sustainability discourse. This study aimed to assess the environmental impact of farms using semi-natural grasslands in Finland, or so-called High Nature Value (HNV) farms. We estimated the environmental impact of 11 such farms, including greenhouse gas emissions (GHG), nitrogen (N) balance, land occupation, and carbon storage. We also accounted for unique biodiversity, defined in this study as communities that are dependent on semi-natural grasslands. We compared these to the alternative states of the farms, specifically a hypothetical farm with the same production output but without access to semi natural grasslands. GHG emissions at the farm level (tCO(2eq)/ha) in HNV farms were 64% lower than on the alternative farms; GHG emissions at the product level (tCO(2eq)/t LW) and N balance (N kg/ha) were 31% and 235% lower, respectively. The carbon stocks were 163% higher at farm level. Biodiversity values, indicated by the share of semi-natural grassland in management, ranged from 23% to 83% on HNV farms. Six out of eleven farms would need to increase their arable land occupation by an average of 39% of arable land to fulfil their needs for animal feed if they did not utilize semi-natural grassland. This study contributes to growing evidence that HNV farming systems can support sustainable production by minimising arable land occupation, reducing nutrient loses, and increasing carbon storage while maintaining unique biodiversity.

The European continent contains substantial areas of semi-natural habitats, mostly grasslands, which are among the most endangered habitats in Europe. Their continued existence depends on some form of human activity, for either production or conservation purposes, or both. We examined the share of semi-natural grasslands within the general grassland areas in boreal Europe. We reviewed research literature across the region to compile evidence on semi-natural grasslands and other semi-natural habitats, such as wooded pastures, in respect to a range of topics such as ecology, land-use change, socioeconomics, and production. We also explored drivers of the research agenda and outlined future research needs. Challenges are faced when defining and quantifying semi-natural habitats even across a restricted region. Agricultural development and other policies clearly impact the research agenda in various countries. There are recent signs of a shift from classical ecological studies toward more multidisciplinary and integrated research. To sufficiently address the threats faced by semi-natural habitats, political and research frameworks in the European Union should pay more attention to the social-ecological complexity inherent in their management and should support the engagement of various actors into participatory governance processes. This is in line with a full-farm approach implicit in high nature value farming systems.

Questions: Small, remnant habitats embedded in degraded, human-dominated landscapes are generally not a priority in conservation, despite their potential role in supporting landscape-scale biodiversity. To warrant their inclusion in conservation management and policy, we question under which conditions they may exhibit the largest conservation value. Location: Nine landscapes spread across the counties of Stockholm and Sodermanland, Sweden. Methods: Per landscape, plant communities were surveyed in 6 and 12 1 × 1 m2 plots across large, intact semi-natural grasslands and small remnant grasslands, respectively. These two contrasting grassland types served as a model system. A topsoil sample was taken in each plot to determine habitat quality in terms of soil pH, plant-available P, and C:N ratio. We used a joint species distribution model to analyse the extent to which grassland type and habitat quality define and predict resident community diversity and composition, including whether they support grassland specialists. Results: At the landscape scale, the combined remnant grasslands sustained diverse plant communities which did include a significant subset of habitat specialists. Yet, the contribution of individual remnants clearly varied with local-scale habitat quality; soil phosphorus availability lowered plot-level species richness, mostly by constraining the occurrence of grassland specialists. Semi-natural grassland communities were comparatively insensitive to variation in soil phosphorus availability. Conclusions: The combined habitat amount and the significant number of habitat specialists sustained by remnant grasslands with high habitat quality, shows they can represent a valuable resource to support landscape-scale biodiversity conservation. This offers no wildcard to neglect the continued biotic and abiotic threats on semi-natural grassland plant diversity such as chronic and accumulating P eutrophication, discontinuation of management or poor matrix permeability, as semi-natural grasslands harbour the majority of habitat specialists, while sourcing surrounding remnant grassland communities.

Context One approach to maintain the resilience of biotic communities is to protect the variability of abiotic characteristics of Earth’s surface, i.e. geodiversity. In terrestrial environments, the relationship between geodiversity and biodiversity is well recognized. In streams, the abiotic properties of upstream catchments influence stream communities, but the relationships between catchment geodiversity and aquatic biodiversity have not been previously tested. Objectives The aim was to compare the effects of local environmental and catchment variables on stream biodiversity. We specifically explored the usefulness of catchment geodiversity in explaining the species richness on stream macroinvertebrate, diatom and bacterial communities. Methods We used 3 geodiversity variables, 2 land use variables and 4 local habitat variables to examine species richness variation across 88 stream sites in western Finland. We used boosted regression trees to explore the effects of geodiversity and other variables on biodiversity. Results We detected a clear effect of catchment geodiversity on species richness, although the traditional local habitat and land use variables were the strongest predictors. Especially soil-type richness appeared as an important factor for species richness. While variables related to stream size were the most important for macroinvertebrate richness and partly for bacterial richness, the importance of water chemistry and land use for diatom richness was notable. Conclusions In addition to traditional environmental variables, geodiversity may affect species richness variation in streams, for example through changes in water chemistry. Geodiversity information could be used as a proxy for predicting stream species richness and offers a supplementary tool for conservation efforts.

Sustainable urban nature conservation calls for a rethinking of conventional approaches. Traditionally, conservationists have not incorporated the history of the landscape in management strategies. This study shows that extant vegetation patterns are correlated to past landscapes indicating potential extinction debts. We calculated urban landscape measures for seven time periods (1938&ndash, 2019) and correlated it to three vegetation sampling events (1995, 2012, 2019) using GLM models. We also tested whether urban vegetation was homogenizing. Our results indicated that urban vegetation in our study area is not currently homogenizing but that indigenous forb species richness is declining significantly. Furthermore, long-term studies are essential as the time lags identified for different vegetation sampling periods changed as well as the drivers best predicting these changes. Understanding these dynamics are critical to ensuring sustainable conservation of urban vegetation for future citizens.

1. Over the last decades, biodiversity in agricultural landscapes has declined drastically. Initiatives to enhance biodiversity, such as agri-environment schemes, often have little effect, especially in intensively farmed landscapes. The effectiveness of conservation management may be improved by scheme implementation near high-quality habitats that can act as a source of species. We evaluated up to what distance high-quality habitats (nature reserves and artificially created flower-rich patches) affect the diversity of forbs and pollinators in intensively farmed landscapes of the Netherlands. 2. We surveyed forbs, inflorescences, bees and hover flies and estimated pollination services in transects along ditch banks extending 300 m from four nature reserves forming small islands in landscapes dominated by agriculture. 3. In a separate experiment, we surveyed inflorescences, bees and hover flies in 1500 m long transects on farmland adjacent to five newly introduced flower-rich patches and in five control transects. 4. Species density of forbs declined over the first 75 m and species density and abundance of hover flies declined over the first 125 m beyond the nature reserves. Beyond these distances, no further declines were observed. The effects of flower-rich patches were spatially limited. The species density and abundance of bees and hover flies were significantly enhanced in the flower-rich patch, but only the abundance of hover flies was enhanced up to 50 m beyond the patch. 5. Synthesis and applications. In intensively farmed areas, remnant high-quality habitats sustain more abundant and diverse pollinator and forb communities than the surrounding countryside. They do enhance biodiversity on nearby farmland but increases are spatially restricted (

1.Terrestrial ecosystems are characterised by a strong functional connection between the green (plant-herbivore-based) and brown (detritus-detritivore-based) parts of the food web, which both develop over successional time. However, the interlinked changes in green and brown food web diversity patterns in relation to key ecosystem processes are rarely studied. 2.Here, we demonstrate changes in species richness, diversity and evenness over a wide range of invertebrate green and brown trophic groups during 100 years of primary succession in a salt marsh ecosystem, using a well-calibrated chronosequence. 3.We contrast two hypotheses on the relationship between green and brown food web diversity across succession: i) 'coupled diversity hypothesis', which predicts that all trophic groups covary similarly with the main drivers of successional ecosystem assembly versus ii) the 'decoupled diversity hypothesis', where green and brown trophic groups diversity respond to different drivers during succession. 4.We found that, while species richness for plants and invertebrate herbivores (green web groups) both peaked at intermediate productivity and successional age, the diversity of macro-detritivores, microarthropod microbivores and secondary consumers (brown web groups) continuously increased towards the latest successional stages. These results suggest that green web trophic groups are mainly driven by vegetation parameters, such as the amount of bare soil, vegetation biomass production, and vegetation height, while brown web trophic groups are mostly driven by the production and standing stock of dead organic material and soil development. 5.Our results show that plant diversity cannot simply be used as a proxy for the diversity of all other species groups that drive ecosystem functioning, as brown and green diversity components in our ecosystem responded differently to successional gradients. This article is protected by copyright. All rights reserved.

Amongst many ongoing initiatives to preserve biodiversity, the Millennium Ecosystem Assessment again shows the importance to slow down the loss of biological diversity. However, there is still a gap in the overview of global patterns of species distributions. This paper reviews how remote sensing has been used to assess terrestrial faunal diversity, with emphasis on proxies and methodologies, while exploring prospective challenges for the conservation and sustainable use of biodiversity. We grouped and discussed papers dealing with the faunal taxa mammals, birds, reptiles, amphibians, and invertebrates into five classes of surrogates of animal diversity: (1) habitat suitability, (2) photosynthetic productivity, (3) multi-temporal patterns, (4) structural properties of habitat, and (5) forage quality. It is concluded that the most promising approach for the assessment, monitoring, prediction, and conservation of faunal diversity appears to be the synergy of remote sensing products and auxiliary data with ecological biodiversity models, and a subsequent validation of the results using traditional observation techniques.

Aims - Intercropping increases aboveground and belowground crop productivity, suggesting potential for carbon sequestration. Here we determined whether intercropping affects decomposition of soil organic matter (SOM) and root litter. Methods - We measured in the laboratory and the field the breakdown of SOM, root litter of maize, wheat, or faba bean, litter mixtures, and a standard substrate (compost) in soils from a long term intercropping experiment. Results - Soil organic matter from intercrop plots decomposed faster than SOM from monocrop plots, but compost decomposed at similar rates in different soils. Faster SOM decomposition was associated with lower soil C:N ratio. Root litter mixtures of maize and wheat decomposed as expected from single litters, but litter mixture of maize and faba bean decomposed faster than expected, both in the laboratory and in the field. Root litter decomposed slowly in maize/wheat intercrop soil compared to the two monocropped soils in the laboratory, but the effect was absent in the field. Conclusions - Intercropping increases SOM decomposition, presumably through reduced SOM recalcitrance resulting from lower C:N ratio, higher litter input and better N retention. Depending on the crop combination, also non-additive effects of root litter mixing can enhance organic matter decomposition in intercropping soils.

The importance of low intensity farming for the conservation of biodiversity throughout Europe was acknowledged early in the 1990s when the concept of 'High Nature Value farmlands' (HNVf) was devised. HNVf has subsequently been given high priority within the EU Rural Development Programme. This puts a requirement on each EU Member State not only to identify the extent and condition of HNVf within their borders but also to track trends in HNVf over time. However, the diversity of rural landscapes across the EU, the scarcity of (adequate) datasets on biodiversity, land cover and land use, and the lack of a common methodology for HNVf mapping currently represent obstacles to the implementation of the HNVf concept across Europe. This manuscript provides an overview of the characteristics of HNVf across Europe together with a description of the development of the HNVf concept. Current methodological approaches for the identification and mapping of HNVf across EU-27 and Switzerland are then reviewed, the main limitations of these approaches highlighted and recommendations made as to how the identification, mapping and reporting of HNVf state and trends across Europe can potentially be improved and harmonised. In particular, we propose a new framework that is built on the need for strategic HNVf monitoring based on a hierarchical, bottom-up structure of assessment units, coincident with the EU levels of political decision and devised indicators, and which is linked strongly to a collaborative European network that can provide the integration and exchange of data from different sources and scales under common standards. Such an approach is essential if the scale of the issues facing HNVf landscapes are to be identified and monitored properly at the European level. This would then allow relevant agri-environmental measures to be developed, implemented and evaluated at the scale(s) required to maintain the habitats and species of high nature conservation value that are intimately associated with those landscapes.

International audience; Nutrient-rich water bodies are usually expected to host low species richness at the local scale (water body). Nevertheless, they can support a diverse and sometimes unique biodiversity when diversity is considered at a regional scale. This discrepancy between the two scales is well documented for natural water bodies, but little is known about biodiversity of artificial water bodies, like fish ponds. We hypothesise that nutrient-rich water bodies can collectively host high species richness at the regional scale. Thus, these are important ecosystems for the regional conservation of biodiversity. We investigated 84 fish ponds in the Dombes region, France, with five taxonomic groups: macrophytes, phytoplankton, macroinvertebrates, dragonflies, and amphibians. Species richness patterns were determined for alpha- (single pond), beta- (between ponds), and gamma- (regional pond network) levels. For most studied species groups, richness per fish pond and at the regional level proved to be relatively high in comparison with natural ponds in other landscapes. Contribution of alpha-diversity to regional diversity was highest for dragonflies with 41 %, and lowest for amphibians and macrophytes with 16 and 18 %, respectively. For macroinvertebrate families and phytoplankton genera it was intermediate. Contribution of beta-diversity to regional diversity was similar for all species groups with 22-25 %. Furthermore, some ponds hosted a large number of less frequent species and some endangered species, indicating that the conservation of biodiversity of fish ponds must be established at a regional scale.

Both arthropods and large grazing herbivores are important components and drivers of biodiversity in grassland ecosystems, but a synthesis of how arthropod diversity is affected by large herbivores has been largely missing. To fill this gap, we conducted a literature search, which yielded 141 studies on this topic of which 24 simultaneously investigated plant and arthropod diversity. Using the data from these 24 studies, we compared the responses of plant and arthropod diversity to an increase in grazing intensity. This quantitative assessment showed no overall significant effect of increasing grazing intensity on plant diversity, while arthropod diversity was generally negatively affected. To understand these negative effects, we explored the mechanisms by which large herbivores affect arthropod communities: direct effects, changes in vegetation structure, changes in plant community composition, changes in soil conditions, and cascading effects within the arthropod interaction web. We identify three main factors determining the effects of large herbivores on arthropod diversity: (i) unintentional predation and increased disturbance, (ii) decreases in total resource abundance for arthropods (biomass) and (iii) changes in plant diversity, vegetation structure and abiotic conditions. In general, heterogeneity in vegetation structure and abiotic conditions increases at intermediate grazing intensity, but declines at both low and high grazing intensity. We conclude that large herbivores can only increase arthropod diversity if they cause an increase in (a)biotic heterogeneity, and then only if this increase is large enough to compensate for the loss of total resource abundance and the increased mortality rate. This is expected to occur only at low herbivore densities or with spatio-temporal variation in herbivore densities. As we demonstrate that arthropod diversity is often more negatively affected by grazing than plant diversity, we strongly recommend considering the specific requirements of arthropods when applying grazing management and to include arthropods in monitoring schemes. Conservation strategies aiming at maximizing heterogeneity, including regulation of herbivore densities (through human interventions or top-down control), maintenance of different types of management in close proximity and rotational grazing regimes, are the most promising options to conserve arthropod diversity.

Sown, temporary field margins are a common agri-environment scheme (AES) in the Netherlands. Despite their wide application, though, there has been scarcely any long-term monitoring of the succession of invertebrates. In the field margins of 40 farms, invertebrate diversity and the abundance of three functional groups were assessed in relation to age. The diversity in terms of number of species groups was found to increase with the age of the margins. The abundance of herbivores and detritivores also showed a positive correlation with the age of the margins. However, the abundance of predators decreased with increasing age. Older margins showed a higher total vegetation cover and fewer plant species, also resulting in lower plant species evenness. We suggest several changes to the current AES regulations. For the conservation of invertebrate diversity, longer-lasting field margins are desirable. In addition, old margins are favoured by detritivores, a group that has particular difficulty finding suitable habitats in agricultural landscapes. However, such margins are less favourable from an agricultural perspective, as they appear unsuitable for high abundances of potentially useful predators and the high vegetation cover attracts many potentially harmful herbivores. To circumvent this, the AES might be extended by incorporating hay-making, which would reduce standing biomass and might lead to more predators and fewer herbivores.

In this paper, we present a novel approach for using ecological heterogeneity in reserve design. We measured five ecological heterogeneity indices (EHI) and we used a database of six biological groups (woody plants, orchids, orthopterans, aquatic and terrestrial herpetofauna and passerine birds) across 30 sites in a Mediterranean reserve (Greece). We found that all the five EHI were significantly related to the overall species richness and to the species richness of woody plants and birds. Two indices, measuring vertical vegetation complexity (1/D) and horizontal heterogeneity of landcover types (SIDI) in terms of Simpson's index, predicted well overall species richness and had significantly higher values inside the complementary reserve networks designed after five of the six biological groups. We compared five methods of forming reserve networks. The method of ecological heterogeneity (selecting those sites with the greatest 1/D and then SIDI) was less efficient (non-significantly) than the species-based methods (scoring and complementary networks) but significantly more efficient than the random method (randomly selected network). We also found that the method of complementary ecological heterogeneity (selecting those sites where each EHI had its maximum value) was not that efficient, as it did not differ significantly from the random method. These results underline the potential of the ecological heterogeneity method as an alternative tool in reserve design. Biodiversity and Conservation

1. Over the last decades, biodiversity in agricultural landscapes has declined drastically. Initiatives to enhance biodiversity, such as agri-environment schemes, often have little effect, especially in intensively farmed landscapes. The effectiveness of conservation management may be improved by scheme implementation near high-quality habitats that can act as a source of species. We evaluated up to what distance high-quality habitats (nature reserves and artificially created flower-rich patches) affect the diversity of forbs and pollinators in intensively farmed landscapes of the Netherlands. 2. We surveyed forbs, inflorescences, bees and hover flies and estimated pollination services in transects along ditch banks extending 300 m from four nature reserves forming small islands in landscapes dominated by agriculture. 3. In a separate experiment, we surveyed inflorescences, bees and hover flies in 1500 m long transects on farmland adjacent to five newly introduced flower-rich patches and in five control transects. 4. Species density of forbs declined over the first 75 m and species density and abundance of hover flies declined over the first 125 m beyond the nature reserves. Beyond these distances, no further declines were observed. The effects of flower-rich patches were spatially limited. The species density and abundance of bees and hover flies were significantly enhanced in the flower-rich patch, but only the abundance of hover flies was enhanced up to 50 m beyond the patch. 5. Synthesis and applications. In intensively farmed areas, remnant high-quality habitats sustain more abundant and diverse pollinator and forb communities than the surrounding countryside. They do enhance biodiversity on nearby farmland but increases are spatially restricted (

International audience; The importance of the conservation of all three fundamental levels of biodiversity (ecosystems, species and genes) has been widely acknowledged, but only in recent years it has become technically feasible to consider intraspecific diversity, i.e. the genetic component to biodiversity. In order to facilitate the assessment of biodiversity, considerable efforts have been made towards identifying surrogates because the efficient evaluation of regional biodiversity would help in designating important areas for nature conservation at larger spatial scales. However, we know little about the fundamental relationships among the three levels of biodiversity, which impedes the formulation of a general, widely applicable concept of biodiversity conservation through surrogates. Here, we present the set-up of an international, interdisciplinary project, INTRABioDiv (http://www.intrabiodiv.eu), which studied vascular plant biodiversity at a large scale, i.e. across the European Alps and the Carpathians. Our assessment comprises species richness (high-mountain flora), genetic variation (amplified fragment length polymorphisms, AFLPs) and environmental diversity (modelled potential habitat diversity). Our primary aims were to test for correlations between intra- and interspecific diversity and to identify possible environmental surrogates to describe biodiversity in the two study regions. To the best of our knowledge, INTRABioDiv represents the first multispecies study on intraspecific, molecular-genetic variation in relation with species and habitat diversity. Here, we outline the theoretical background, our sampling scheme, the technical approaches and the feasibility of a concentrated and standardized sampling effort. We further show exemplary results. Our three data sets will be made freely available and will provide a playground for further hypothesis testing in conservation, ecology or evolution open to the scientific community. (C) 2008 Rubel Foundation, ETH Zurich. Published by Elsevier GmbH. All rights reserved.

1. Comparative analyses of diversity variation among and between regions allow testing of alternative explanatory models and ideas. Here, we explore the relationships between the tree alpha-diversity of small rain forest plots in Africa and in Amazonia and climatic variables, to test the explanatory power of climate and the consistency of relationships between the two continents. 2. Our analysis included 1003 African plots and 512 Amazonian plots. All are located in old-growth primary non-flooded forest under 900 m altitude. Tree alpha-diversity is estimated using Fisher's alpha calculated for trees with diameter at breast height >= 10 cm. Mean diversity values are lower in Africa by a factor of two. 3. Climate-diversity analyses are based on data aggregated for grid cells of 2.5 x 2.5 km. The highest Fisher's alpha values are found in Amazonian forests with no climatic analogue in our African data set. When the analysis is restricted to pixels of directly comparable climate, the mean diversity of African forests is still much lower than that in Amazonia. Only in regions of low mean annual rainfall and temperature is mean diversity in African forests comparable with, or superior to, the diversity in Amazonia. 4. The climatic variables best correlated with the tree alpha-diversity are largely different in the African and Amazonian data, or correlate with African and Amazonian diversity in opposite directions. 5. These differences in the relationship between local/landscape-scale alpha-diversity and climate variables between the two continents point to the possible significance of an array of factors including: macro-scale climate differences between the two regions, overall size of the respective species pools, past climate variation, other forms of long-term and short-term environmental variation, and edaphics. We speculate that the lower alpha-diversity of African lowland rain forests reported here may be in part a function of the smaller regional species pool of tree species adapted to warm, wet conditions. 6. Our results point to the importance of controlling for variation in plot size and for gross differences in regional climates when undertaking comparative analyses between regions of how local diversity of forest varies in relation to other putative controlling factors.

Red list species densities of birds (maximally 22 km¿2), and angiosperms (maximally 39 km¿2) were used as biodiversity indicators in 21 larger complexes of wetlands across the Netherlands. Their covariability with a range of indicators of human land use was assessed, including population, road and visitor density, area covered by agriculture, open water, forest and residential housing. Data were collected on the wetland complexes as well as for a perimeter with 10 km radius. In a principal components analysis (PCA) with all land use variables, it was found that the population-density-related complex of urbanisation, fragmentation (by roads), and intensity of fertilizer use together explained most of the variability present (i.e. the first PCA axis explained 50%), whilst land use within these complexes was second with an additional 19% and waterside recreation third with 12%. Red list bird species density did not correlate with that of angiosperms, nor with any of the indicators used. For the 13 complexes on organic peatland, we observed an increase in maximum red list angiosperm species density with the proportion of open marshland (P 0.55), which, in turn, was negatively and closely correlated with the first PCA axis reflecting an urbanisation gradient across the Netherlands

Many semi-natural landscape elements, the so-called green veining, are disappearing from the intensively used agricultural landscapes of Europe. In order to develop or restore biodiversity in these networks, it is necessary to quantify the relation between biodiversity and amount, spatial arrangement and management intensity of green veining elements. In this review, we investigate whether biodiversity increases with the amount of green veining in an agricultural landscape following the species–area relationship, and whether a certain level of biodiversity can be reached at lower densities of green veining if green veining elements are better connected (higher spatial connectivity) or if they are managed less intensively (lower management intensity). We reviewed studies on aboveground biodiversity in green veining structures in 39 scientific papers on field and experimental studies within Europe. More of these studies focussed on management intensity than on amount or spatial configuration of green veining. Also more studies focussed on the spatial scale of individual landscape elements than on the farm or landscape scale, which may be caused by the large number of studies focussing on plant or invertebrate species. Species living at larger spatial scales, e.g. mammals and birds were not often studied at the level of green veining elements as they also use agricultural fields as part of their habitat. We could not verify the species–area relation for green veining, nor the effect of amount, spatial configuration or management intensity on this relation, because only few studies quantified the found effects and no studies were found on the effect of management intensity or spatial configuration on the species–area curve in green veining. We addressed the most important challenges for future field and model research in order to fill the identified gaps in knowledge.