All is not loss: plant biodiversity in the anthropocene

Anthropogenic global changes in biodiversity are generally portrayed in terms of massive native species losses or invasions caused by recent human disturbance. Yet these biodiversity changes and others caused directly by human populations and their use of land tend to co-occur as long-term biodiversity change processes in the Anthropocene. Here we explore contemporary anthropogenic global patterns in vascular plant species richness at regional landscape scales by combining spatially explicit models and estimates for native species loss together with gains in exotics caused by species invasions and the introduction of agricultural domesticates and ornamental exotic plants. The patterns thus derived confirm that while native losses are likely significant across at least half of Earth's ice-free land, model predictions indicate that plant species richness has increased overall in most regional landscapes, mostly because species invasions tend to exceed native losses. While global observing systems and models that integrate anthropogenic species loss, introduction and invasion at regional landscape scales remain at an early stage of development, integrating predictions from existing models within a single assessment confirms their vast global extent and significance while revealing novel patterns and their potential drivers. Effective global stewardship of plant biodiversity in the Anthropocene will require integrated frameworks for observing, modeling and forecasting the different forms of anthropogenic biodiversity change processes at regional landscape scales, towards conserving biodiversity within the novel plant communities created and sustained by human systems.     

Disentangling spatiotemporal dynamics in metacommunities through a species-patch network approach

Colonization and extinction at local and regional scales, and gains and losses of patches are important processes in the spatiotemporal dynamics of metacommunities. However, analytical challenges remain in quantifying such spatiotemporal dynamics when species extinction-colonization and patch gain and loss processes act simultaneously. Recent advances in network analysis show great potential in disentangling the roles of colonization, extinction, and patch dynamics in metacommunities. Here, we developed a species-patch network approach to quantify metacommunity dynamics including (i) temporal changes in network structure, and (ii) temporal beta diversity of species-patch links and its components that reflect species extinction-colonization and patch gain and loss. Application of the methods to simulated datasets demonstrated that the approach was informative about metacommunity assembly processes. Based on three empirical datasets, our species-patch network approach provided additional information about metacommunity dynamics through distinguishing the effects of species colonization and extinction at different scales from patch gains and losses and how specific environmental factors related to species-patch network structure. In conclusion, our species-patch network framework provides effective methods for monitoring and revealing long-term metacommunity dynamics by quantifying gains and losses of both species and patches under local and global environmental change.     

Species richness change across spatial scales

Humans have elevated global extinction rates and thus lowered global scale species richness. However, there is no a priori reason to expect that losses of global species richness should always, or even often, trickle down to losses of species richness at regional and local scales, even though this relationship is often assumed. Here, we show that scale can modulate our estimates of species richness change through time in the face of anthropogenic pressures, but not in a unidirectional way. Instead, the magnitude of species richness change through time can increase, decrease, reverse, or be unimodal across spatial scales. Using several case studies, we show different forms of scale‐dependent richness change through time in the face of anthropogenic pressures. For example, Central American corals show a homogenization pattern, where small scale richness is largely unchanged through time, while larger scale richness change is highly negative. Alternatively, birds in North America showed a differentiation effect, where species richness was again largely unchanged through time at small scales, but was more positive at larger scales. Finally, we collated data from a heterogeneous set of studies of different taxa measured through time from sites ranging from small plots to entire continents, and found highly variable patterns that nevertheless imply complex scale‐dependence in several taxa. In summary, understanding how biodiversity is changing in the Anthropocene requires an explicit recognition of the influence of spatial scale, and we conclude with some recommendations for how to better incorporate scale into our estimates of change.     

Applying the dark diversity concept to plants at the European scale

Large‐scale biodiversity maps are essential to macroecology. However, between‐region comparisons can be more useful if patterns of observed species richness are supplemented by variations in dark diversity – the absent portion of the species pool. We aim to quantify and map plant diversity across Europe by using a measure that accounts for both observed and dark diversity. To do this we need to delimit suitable species pools, and evaluate the potential and limitation of a large‐scale dataset. We used Atlas Florae Europaeae (ca 20% of European plant species mapped within 50 × 50 km grid cells) and defined for each grid cell several species pools by applying various geographical and environmental filters: geographic species pool (number of species within 500 km radius), biogeographic species pool (additionally incorporating species distribution patterns, i.e. dispersion fields), site‐specific species pool (additionally integrating environmental preferences of species based on species co‐occurrence). We integrated dark diversity and observed diversity at a relative scale to calculate the completeness of site diversity: logistic expression of observed and dark diversity. We tested whether our results are robust against regional variation in data availability. We used independent regional databases to test if Atlas Florae Europaeae is a representative subset of total species richness. Environmental filtering was the most influential determinant of species pool size with more species filtered out in southern Europe. Both observed and dark diversity adhered to the well‐known latitudinal gradient, but completeness of site diversity varied throughout Europe with no latitudinal trend. Dark diversity patterns were fairly insensitive to variations in regional sampling intensity. Atlas Florae Europaeae represented well the total variation in plant diversity. In summary, dark diversity and completeness of site diversity add valuable information to broad‐scale diversity patterns since observed diversity is expressed at a relative scale.     

DarkDivNet – A global research collaboration to explore the dark diversity of plant communities

DarkDivNet is a global research collaboration which explores dark diversity — the set of species that are absent from a site despite being suitable under the site conditions and present in the region. Participants of the network survey vascular plant diversity both at local (10  m × 10 m) and regional scales (radius 10 km) using a standardized approach. They also measure simple plant traits and collect soil samples. Observed and dark diversity together form the site‐specific species pool, and the ratio of observed diversity and dark diversity describes community completeness. We shall explore how observed and dark diversity, site‐specific species pool and community completeness vary across natural and anthropogenic gradients. We link plant diversity measures to the information obtained from environmental DNA: soil biota and plant taxa that occurred at the site before. We will refine existing dark diversity methods and use large vegetation databases to infer species habitat suitability. We expand the dark diversity concept from a purely taxonomy‐based approach to include the functional and phylogenetic aspects of diversity. DarkDivNet currently includes 161 planned sampling areas globally, but new participants are welcome. The main vegetation sampling period is scheduled until September 2020, with the first research papers being produced after that.     

Spatial heterogeneity in species composition constrains plant community responses to herbivory and fertilisation

Environmental change can result in substantial shifts in community composition. The associated immigration and extinction events are likely constrained by the spatial distribution of species. Still, studies on environmental change typically quantify biotic responses at single spatial (time series within a single plot) or temporal (spatial beta diversity at single time points) scales, ignoring their potential interdependence. Here, we use data from a global network of grassland experiments to determine how turnover responses to two major forms of environmental change – fertilisation and herbivore loss – are affected by species pool size and spatial compositional heterogeneity. Fertilisation led to higher rates of local extinction, whereas turnover in herbivore exclusion plots was driven by species replacement. Overall, sites with more spatially heterogeneous composition showed significantly higher rates of annual turnover, independent of species pool size and treatment. Taking into account spatial biodiversity aspects will therefore improve our understanding of consequences of global and anthropogenic change on community dynamics.     

Dark diversity reveals importance of biotic resources and competition for plant diversity across habitats

Species richness is the most commonly used metric to quantify biodiversity. However, examining dark diversity, the group of missing species which can potentially inhabit a site, can provide a more thorough understanding of the processes influencing observed biodiversity and help evaluate the restoration potential of local habitats. So far, dark diversity has mainly been studied for specific habitats or large‐scale landscapes, while less attention has been given to variation across broad environmental gradients or as a result of local conditions and biotic interactions. In this study, we investigate the importance of local environmental conditions in determining dark diversity and observed richness in plant communities across broad environmental gradients. Using the ecospace concept, we investigate how these biodiversity measures relate to abiotic gradients (defined as position), availability of biotic resources (defined as expansion), spatiotemporal extent of habitats (defined as continuity), and species interactions through competition. Position variables were important for both observed diversity and dark diversity, some with quadratic relationships, for example, plant richness showing a unimodal response to soil fertility corresponding to the intermediate productivity hypothesis. Interspecific competition represented by community mean Grime C had a negative effect on plant species richness. Besides position‐related variables, organic carbon was the most important variable for dark diversity, indicating that in late‐succession habitats such as forests and shrubs, dark diversity is generally low. The importance of highly competitive species indicates that intermediate disturbance, such as grazing, may facilitate higher species richness and lower dark diversity.     

Habitat selection behaviour links local and regional scales in aquatic systems

The role of habitat selection behaviour in the assembly of natural communities is an increasingly important theme in ecology. At the same time, ecologists and conservation biologists are keenly interested in scale and how processes at scales from local to regional interact to determine species distributions and patterns of biodiversity. How important is habitat selection in generating observed patterns of distribution and diversity at multiple spatial scales? In theory, habitat selection in response to interacting species can generate both positive and negative covariances among species distributions and create the potential to link processes of community assembly across multiple scales. Here I demonstrate that habitat selection by treefrogs in response to the distribution of fish predators functions at both the regional scale among localities and the local scale among patches within localities, implicating habitat selection as a critical link between local communities and the regional dynamics of metacommunities in complex landscapes.     

Getting biodiversity intactness indices right: ensuring that 'biodiversity' reflects 'diversity'

The 2010 biodiversity target of a ‘significant reduction in the current rate of biodiversity loss’ presents challenges for effective measurement of changes in global/regional biodiversity. A simple ‘biodiversity intactness index’ (BII) is attractive in using available data and expert opinion, but is seen to be only weakly linked to ‘biodiversity’ in its usual sense of ‘variation’. An example illustrates how an improved BII score could result even when there are large species losses. A family of alternative biodiversity ‘representativeness indices’ better reflect variation. They use the same readily available information, plus simple species–area relationships (SAR) and genetic diversity curves. A new genetic‐diversity abundance‐fraction curve, like SAR, is linear in log–log space. The new representativeness indices incorporate, through range‐abundance curves, the abundance fraction estimates normally used for BII. Land use or climate change impacts therefore can reflect partial rather than total biodiversity losses at localities. Estimates of biodiversity gains/losses using these indices enable a novel regional planning‐based approach for addressing the 2010 target.     

Turquoise is the new green: Restoring and enhancing riparian function in the Anthropocene

Riparian ecosystems are hotspots for ecological restoration globally because of the disproportionately high value and diversity of the ecological functions and services which they support and their high level of vulnerability to anthropogenic pressures, including climate change. Degraded riparian ecosystems are associated with many serious anthropogenic problems including increased river bank erosion, water quality decline, increased flood risk and biodiversity loss. Conventional approaches to riparian restoration, however, are frequently too narrow in focus – spatially, temporally, ecologically and socially – to adequately or equitably address the goals to which they aspire. Climate change, along with the intensification of other human pressures, means that static, historically oriented restoration objectives focused solely on prior ecological composition and structure are unlikely to be defensible, achievable or appropriate in the Anthropocene. Conversely, open‐ended restoration strategies lacking clear objectives and targets entail substantial risks such as significant biodiversity losses, especially of native species. A functional approach to planning and prioritising riparian restoration interventions offers an intermediate alternative that is still framed by measurable targets but allows for greater consideration of broader temporal, spatial and cultural influences. Here, we provide an overview of major riparian functions across multiple scales and identify key drivers of, and threats to, these. We also discuss practical approaches to restoring and promoting riparian functions and highlight some key concerns for the development of policy and management of robust riparian restoration in the Anthropocene.     

Magnitude and drivers of plant diversity loss differ between spatial scales in Scania,Sweden 1957–2021

Questions

Changed land use, nitrogen deposition, climate change, and the spread of non-native species have repeatedly been reported as the main drivers of recent floristic changes in northern Europe. However, the relevance of the geographical scale at which floristic changes are observed is less well understood and it has only rarely been possible to quantify biodiversity loss. Therefore, we assessed changes in species richness, species composition and mean ecological indicator values (EIVs) at three nested geographic scales during two different time periods, each ca 30 years, since the mid-1900s.

Location

Two parishes in central Scania, southernmost Sweden.

Methods

We analyzed species presence/absence data from three inventories at ca 30-year intervals over 1957–2021 and three geographic scales (157 m², ca 7 km² and ca 45 km²) to document temporal trends and differences between geographic scales in terms of species richness, species composition and mean EIVs.

Results

We found shifts in species composition across all geographical scales. However, the magnitude of biodiversity loss and the main drivers of these changes were scale-dependent. At the smallest spatial scale, we saw a dramatic loss of plant biodiversity with local species richness in 2021 being only 48% of that of 1960. In contrast, at the larger geographic scales no significant changes in species richness were observed because species losses were compensated for by gains of predominantly non-native species, which made up at least 78% of the new species richness. At the smallest spatial scale, changed land use (ceased grazing/mowing and intensified forestry) appeared as the main driver, while an increasing proportion of non-native species, as well as climatic changes and increasing nitrogen loads appeared relatively more important at larger geographic scales.

Conclusion

Our results highlight the precarious situation for biodiversity in the region and at the same time the fundamental importance of geographic scale in studies of biodiversity change. Both the magnitude and drivers of changes may differ depending on the geographic scale and must be considered also when previously published studies are interpreted.     

Dispersal and neutral sampling mediate contingent effects of disturbance on plant beta‐diversity: a meta‐analysis

A major challenge in ecology, conservation and global‐change biology is to understand why biodiversity responds differently to similar environmental changes. Contingent biodiversity responses may depend on how disturbance and dispersal interact to alter variation in community composition (β‐diversity) and assembly mechanisms. However, quantitative syntheses of these patterns and processes across studies are lacking. Using null‐models and meta‐analyses of 22 factorial experiments in herbaceous plant communities across Europe and North America, we show that disturbance diversifies communities when dispersal is limited, but homogenises communities when combined with increased immigration from the species pool. In contrast to the hypothesis that disturbance and dispersal mediate the strength of niche assembly, both processes altered β‐diversity through neutral‐sampling effects on numbers of individuals and species in communities. Our synthesis suggests that stochastic effects of disturbance and dispersal on community assembly play an important, but underappreciated, role in mediating biotic homogenisation and biodiversity responses to environmental change.     

Plant species diversity and grazing in the Scandinavian mountains - patterns and processes at different spatial scales

There is a long tradition of grazing by semi‐domestic reindeer and sheep in alpine and sub‐alpine Scandinavian habitats, but present management regimes are questioned from a conservation point of view. In this review we discuss plant diversity patterns in the Scandinavian mountains in a global, regional and local perspective. The main objective was to identify processes that influence diversity at different spatial scales with a particular focus on grazing. In a global perspective the species pool of the Scandinavian mountains is limited. partly reflecting the general latitudinal decline of species but also historical and ecological factors operating after the latest glaciation. At the local scale, both productivity and disturbance are primary factors structuring diversity, but abiotic factors such as soil pH, snow distribution and temperature are also important. Although evidence is scarce, grazing favours local species richness in productive habitats, whereas species richness decreases with grazing when productivity is low. Regional patterns of plant diversity is set by, 1) the species pool. 2) the heterogeneity and fragmentation of communities, and 3) local diversity of each plant community. We suggest that local shifts in community composition depend both on the local grazing frequency and the return‐time of the plant community after a grazing session. In addition, an increasing number of grazing‐modified local patches homogenises the vegetation and is likely to reduce the regional plant diversity. The time scale of local shifts in community composition depends on plant colonisation and persistence, From a mechanistic point of view, diversity patterns at a regional scale also depend on the regional dynamics of single species. Colonisation is usually a slow and irregular process in alpine environments, whereas the capacity for extended local persistence is generally high. Although the poor knowledge of plant regional dynamics restricts our understanding of how grazing influences plant diversity, we conclude that grazing is a key process for maintaining biodiversity in the Scandinavian mountains.     

Dark diversity: shedding light on absent species

Ecological theory and nature conservation have traditionally relied solely on observed local diversity. In this review, we recommend including those species that are absent from an ecosystem but which belong to its species pool; that is, all species in the region that can potentially inhabit those particular ecological conditions. We call the set of absent species 'dark diversity'. Relating local and dark diversities enables biodiversity comparisons between regions, ecosystems and taxonomic groups, and the evaluation of the roles of local and regional processes in ecological communities. Dark diversity can also be used to counteract biodiversity loss and to estimate the restoration potential of ecosystems. We illustrate the dark diversity concept by globally mapping plant dark diversity and the local:dark diversity ratio.     

Lakes in the era of global change: moving beyond single-lake thinking in maintaining biodiversity and ecosystem services

The Anthropocene presents formidable threats to freshwater ecosystems. Lakes are especially vulnerable and important at the same time. They cover only a small area worldwide but harbour high levels of biodiversity and contribute disproportionately to ecosystem services. Lakes differ with respect to their general type (e.g. land-locked, drainage, floodplain and large lakes) and position in the landscape (e.g. highland versus lowland lakes), which contribute to the dynamics of these systems. Lakes should be generally viewed as ‘meta-systems’, whereby biodiversity is strongly affected by species dispersal, and ecosystem dynamics are contributed by the flow of matter and substances among locations in a broader waterscape context. Lake connectivity in the waterscape and position in the landscape determine the degree to which a lake is prone to invasion by non-native species and accumulation of harmful substances. Highly connected lakes low in the landscape accumulate nutrients and pollutants originating from ecosystems higher in the landscape. The monitoring and restoration of lake biodiversity and ecosystem services should consider the fact that a high degree of dynamism is present at local, regional and global scales. However, local and regional monitoring may be plagued by the unpredictability of ecological phenomena, hindering adaptive management of lakes. Although monitoring data are increasingly becoming available to study responses of lakes to global change, we still lack suitable integration of models for entire waterscapes. Research across disciplinary boundaries is needed to address the challenges that lakes face in the Anthropocene because they may play an increasingly important role in harbouring unique aquatic biota as well as providing ecosystem goods and services in the future.     

Global patterns of stream detritivore distribution: implications for biodiversity loss in changing climates

Aim We tested the hypothesis that shredder detritivores, a key trophic guild in stream ecosystems, are more diverse at higher latitudes, which has important ecological implications in the face of potential biodiversity losses that are expected as a result of climate change. We also explored the dependence of local shredder diversity on the regional species pool across latitudes, and examined the influence of environmental factors on shredder diversity. Location World‐wide (156 sites from 17 regions located in all inhabited continents at latitudes ranging from 67° N to 41° S). Methods We used linear regression to examine the latitudinal variation in shredder diversity at different spatial scales: alpha (α), gamma (γ) and beta (β) diversity. We also explored the effect of γ‐diversity on α‐diversity across latitudes with regression analysis, and the possible influence of local environmental factors on shredder diversity with simple correlations. Results Alpha diversity increased with latitude, while γ‐ and β‐diversity showed no clear latitudinal pattern. Temperate sites showed a linear relationship between γ‐ and α‐diversity; in contrast, tropical sites showed evidence of local species saturation, which may explain why the latitudinal gradient in α‐diversity is not accompanied by a gradient in γ‐diversity. Alpha diversity was related to several local habitat characteristics, but γ‐ and β‐diversity were not related to any of the environmental factors measured. Main conclusions Our results indicate that global patterns of shredder diversity are complex and depend on spatial scale. However, we can draw several conclusions that have important ecological implications. Alpha diversity is limited at tropical sites by local factors, implying a higher risk of loss of key species or the whole shredder guild (the latter implying the loss of trophic diversity). Even if regional species pools are not particularly species poor in the tropics, colonization from adjacent sites may be limited. Moreover, many shredder species belong to cool‐adapted taxa that may be close to their thermal maxima in the tropics, which makes them more vulnerable to climate warming. Our results suggest that tropical streams require specific scientific attention and conservation efforts to prevent loss of shredder biodiversity and serious alteration of ecosystem processes.     

Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver–response relationships to develop conceptual models across ecosystems

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed ‘beta diversity’) is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.     

Dark diversity in dry calcareous grasslands is determined by dispersal ability and stress‐tolerance

Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat‐specific regional species pool may be absent from local communities and represent the ‘dark diversity’ of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat‐specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing). We analysed a total of 1223 vegetation plots (1 × 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co‐occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S‐strategy type scores were used to characterise low stress‐tolerance. Levels of small‐scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions. We conclude that, even if temperate grasslands have high levels of small‐scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress‐tolerance.     

How pervasive is biotic homogenization in human‐modified tropical forest landscapes?

Land‐cover change and ecosystem degradation may lead to biotic homogenization, yet our understanding of this phenomenon over large spatial scales and different biotic groups remains weak. We used a multi‐taxa dataset from 335 sites and 36 heterogeneous landscapes in the Brazilian Amazon to examine the potential for landscape‐scale processes to modulate the cumulative effects of local disturbances. Biotic homogenization was high in production areas but much less in disturbed and regenerating forests, where high levels of among‐site and among‐landscape β‐diversity appeared to attenuate species loss at larger scales. We found consistently high levels of β‐diversity among landscapes for all land cover classes, providing support for landscape‐scale divergence in species composition. Our findings support concerns that β‐diversity has been underestimated as a driver of biodiversity change and underscore the importance of maintaining a distributed network of reserves, including remaining areas of undisturbed primary forest, but also disturbed and regenerating forests, to conserve regional biota.