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This far, quantitative assessments of geodiversity have typically been conducted on the level of alpha geodiversity (i.e., the variety of geofeatures, such as rock types or landforms, in a location) or gamma geodiversity (i.e., sum of all geofeatures across all surveyed locations in a region of interest). These assessments give valuable information of the distribution of geodiversity but do not provide evidence on the uniqueness of the elements of geodiversity (i.e., geofeatures) at different sites. Uniqueness measures can be used to rank locations based on how exceptional they are in terms of geofeatures and when compared with other locations surveyed in a study.In this study, we demonstrate a quantitative approach to distinguish how study sites differ in their geofeature uniqueness. The methodology has been originally proposed for quantifying taxonomic uniqueness in biotic communities but we here extend its use to a geodiversity context. To illustrate the approach, we have identified geological, hydrological and geomorphological geofeatures from three areas (Rekijoki, Oulanka and Paistunturit) located across Finland, and calculated the presence/absence of each geofeature in 500 m grid cells. Founded on these grid-based datasets that cover the three study areas, we have made geodiversity uniqueness calculations. First, we calculated total beta geodiversity, from which local contribution to beta geodiversity (LCBD-g) value for each site can be subsequently derived. The LCBD-g value describes the geofeature uniqueness of each site in comparison to other sites studied in an area. Second, we calculated contributions to beta geodiversity values for each geofeature (geofeature contribution to beta geodiversity, GFCBD), describing the importance of each geofeature for geodiversity variation in the studied sites.Our results show that the uniqueness of geofeatures (LCBD-g values) varies profoundly in each of the studied areas and sites. Also, the importance of each geofeature for geodiversity variation (GFCBD value) exhibits distinct patterns, representing the varying significance of each geofeature for the total geodiversity in the studied areas. The results, especially when displayed as maps, are heuristic and an easily applicable way to flag sites that are unique in their geofeature composition. This information can complement previous approaches (such as geosite or biodiversity assessments) when prioritizing areas for nature conservation.

Geodiversity is an emerging, multi-faceted concept in Earth and environmental sciences. Knowledge on geo-diversity is crucial for understanding functions of natural systems and in guiding sustainable development. Despite the critical nature of geodiversity information, data acquisition and analytical methods have lagged behind the conceptual developments in biosciences. Thus, we propose that geodiversity research could adopt the framework of alpha, beta and gamma concepts widely used in biodiversity research. Especially, geodiversity research would benefit from widening its scope from the evaluation of individual sites towards more holistic geodiversity assessments, where between-site geodiversity is also considered. In this article, we explore the alpha, beta and gamma concepts and how they can be applied in a geodiversity framework. In addition, we scrutinize the statistical methodology related to alpha, beta and gamma geodiversity evaluations, with a special focus on distance metrics for measuring beta geodiversity. As an overview of the process, and to give practical guidelines for the application of the proposed methodology, we present a case study from a UNESCO Global Geopark area. Thus, this study not only develops the geodiversity concept, but also paves the way for simultaneous understanding of both geodiversity and biodiversity within a unified conceptual approach.

It has long been debated why groups such as non-avian dinosaurs became extinct whereas mammals and other lineages survived the Cretaceous/Paleogene mass extinction 66 million years ago. We used Markov networks, ecological niche partitioning, and Earth System models to reconstruct North American food webs and simulate ecospace occupancy before and after the extinction event. We find a shift in latest Cretaceous dinosaur faunas, as medium-sized species counterbalanced a loss of megaherbivores, but dinosaur niches were otherwise stable and static, potentially contributing to their demise. Smaller vertebrates, including mammals, followed a consistent trajectory of increasing trophic impact and relaxation of niche limits beginning in the latest Cretaceous and continuing after the mass extinction. Mammals did not simply proliferate after the extinction event; rather, their earlier ecological diversification might have helped them survive.

Caio Graco-Roza was funded by the Coordination for the Improvement of Higher Education Personnel (CAPES), the Carlos Chagas Filho Research Support Foundation (FAPERJ) and the Ella and Georg Erhnrooth Foundation; Jan Altman by research grants INTER-EXCELLENCE LTAUSA19137 provided by Czech Ministry of Education, Youth and Sports, 20-05840Y of the Czech Science Foundation, and long-term research development project no. RVO 67985939 of the Czech Academy of Sciences; Otso Ovaskainen was funded by Academy of Finland (grant no. 309581), Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme (223257), and the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement no. 856506; ERC-synergy project LIFEPLAN); and Jianjun Wang was funded by CAS Key Research Program of Frontier Sciences (QYZDB-SSW-DQC043) and National Natural Science Foundation of China (91851117). The "sPlot" project was initiated by sDiv, the Synthesis Centre of the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, funded by the German Research Foundation (DFG FZT 118), and is now a platform of iDiv. The study was supported by the TRY initiative on plant traits (). We are also grateful to Jens Kattge and TRY database. TRY is hosted, developed and maintained at the Max Planck Institute for Biogeochemistry (MPI-BGC) in Jena, Germany, in collaboration with the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig. The CESTES database of metacommunities is also an initiative of iDiv led by Alienor Jeliazkov. We thank sDiv for supporting the open science initiative., Aim: Understanding the variation in community composition and species abundances (i.e., β-diversity) is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional variation in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distance. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 148 datasets comprising different types of organisms and environments. Location: Global. Time period: 1990 to present. Major taxa studied: From diatoms to mammals. Method: We measured the strength of the decay using ranked Mantel tests (Mantel r) and the rate of distance decay as the slope of an exponential fit using generalized linear models. We used null models to test whether functional similarity decays faster or slower than expected given the taxonomic decay along the spatial and environmental distance. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm and organismal features. Results: Taxonomic distance decay was stronger than functional distance decay along both spatial and environmental distance. Functional distance decay was random given the taxonomic distance decay. The rate of taxonomic and functional spatial distance decay was fastest in the datasets from mid-latitudes. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distance but a higher rate of decay along environmental distance. Marine ecosystems had the slowest rate of decay along environmental distances. Main conclusions: In general, taxonomic distance decay is a useful tool for biogeographical research because it reflects dispersal-related factors in addition to species responses to climatic and environmental variables. Moreover, functional distance decay might be a cost-effective option for investigating community changes in heterogeneous environments., Coordenacao de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) Fundacao Carlos Chagas Filho de Amparo a Pesquisa do Estado do Rio De Janeiro (FAPERJ), Ella and Georg Erhnrooth Foundation, Ministry of Education, Youth & Sports - Czech Republic LTAUSA19137 Grant Agency of the Czech Republic 20-05840Y Czech Academy of Sciences RVO 67985939, Academy of Finland 309581, Jane and Aatos Erkko Foundation, Research Council of Norway through its Centres of Excellence Funding Scheme 223257, European Research Council (ERC) 856506, CAS Key Research Program of Frontier Sciences QYZDB-SSW-DQC043, National Natural Science Foundation of China (NSFC) 91851117, German Research Foundation (DFG) DFG FZT 118, TRY initiative on plant traits

A multi-faceted approach is needed to better understand how the beta diversity of aquatic assemblages responds to ecological gradients. Using distance-based RDA and variance partitioning, we explored the different components of total beta diversity (replacement and richness difference) based on taxonomic, functional, and phylogenetic data of aquatic macrophytes along environmental and spatial gradients across 49 tropical floodplain lakes in the Pantanal of Mato Grosso in the dry season. We found that taxonomic beta diversity was driven by species replacement, while functional beta diversity was mostly contributed by richness difference and phylogenetic beta diversity was equally contributed by both components. Taxonomic and functional beta diversities were driven by environmental variables, connectivity variables, and directional spatial processes. Phylogenetic beta diversity was mainly affected by the connectivity between the sites and their distance from the river. The relationship between environmental variables and beta diversity indicates that a selection of species with similar functional and phylogenetic characteristics co-occurs at a site. The low connectivity between sites and the increased distance between the lakes and the river suggested dispersal limitation, which was due to the absence of directional exchange of species between lakes. Our findings provided new insights into the beta diversity patterns of aquatic macrophytes and their underlying causes in tropical floodplains.

Our existence on Earth is founded on a vital nature, which supports human physical and mental health. However, nature is often depicted only through biodiversity, whereas geodiversity-the diversity of non-living nature-has so far been neglected. Geodiversity consists of assemblages, structures, and systems of geological, geomorphological, soil, and hydrological components that fundamentally underlie biodiversity. Biodiversity can support overall human health only with the foundation of geodiversity. Landscape characteristics, such as varying topography or bodies of water, promote aesthetic and sensory experiences and are also a product of geodiversity. In this Personal View, we introduce the concept of geodiversity as a driver for planetary health, describe its functions and services, and outline the intricate relationships between geodiversity, biodiversity, and human health. We also propose an agenda for acknowledging the importance of geodiversity in health-related research and decision making. Geodiversity is an emerging topic with untapped potential for ensuring ecosystem functionality and good living conditions for people in a time of changing environments.

Unravelling patterns and mechanisms of biogeographical transitions is crucial if we are to understand compositional gradientsat large spatial extents, but no studies have thus far examined breakpoints in community composition of freshwater plants acrosscontinents. Using a dataset of almost 500 observations of lake plant community composition from six continents, we examined,for the first time, if such breakpoints in geographical space exist for freshwater plants and how well a suite of ecological factors(including climatic and local environmental variables) can explain transitions in community composition from the subtropicsto the poles. Our combination of multivariate regression tree (MRT) analysis and k-means partitioning suggests that the mostabrupt breakpoint exists between temperate to boreal regions on the one hand and freshwater plant communities harbouringmainly subtropical or Mediterranean assemblages on the other. The spatially structured variation in current climatic conditionsis the most likely candidate for controlling these latitudinal patterns, although one cannot rule out joint effects of eco-evolutionary constraints in the harsher high-latitude environments and post-glacial migration lags after Pleistocene Ice Ages. Overall, ourstudy supports the foundations of global regionalisation for freshwater plants and anticipates further biogeographical researchon freshwater plant communities once datasets have been harmonised for conducting large-scale spatial analyses.

1. To comprehensively understand the impact of anthropogenic activities on biodiversity, we must understand how biodiversity has changed over time and what are the underlying processes. A growing body of evidence has shown that beta diversity reveals more about temporal changes in biodiversity compared with alpha diversity. Temporal beta diversity indicates, for example, degrees of change in species composition at single locations through time. 2. We examined whether freshwater plant communities showed different patterns in temporal beta diversity in relation to concomitant changes in environmental conditions across decades. To do this, we used presence–absence data of lake plants for five decades (1940s–2010s) from southern Finland and calculated temporal beta diversity indices (TBI) for each lake between pairs of decades to the whole community, hydrophytes and helophytes. To get insights into possible processes behind the observed trends, we decomposed TBIs into beta diversity contributed by either temporal losses or temporal gains of species. We related TBIs and their loss and gain components to lake landscape position and changes in environmental variables. 3. Based on comparisons of TBIs between the survey decade pairs, the temporal change in aquatic plant communities was modest through decades. Hydrophyte assemblages have changed more than helophyte assemblages. The main changes in temporal beta diversity occurred from the 1940s to the 1970s, when the gain of new species was the dominant process in the lakes throughout the landscape. Following that period, there was only modest changes, but from the 2000s to the 2010s, the dominant process was the loss of species. Temporal changes in environmental conditions played a key role in explaining the TBI. 4. Our results showed that relying on only two survey points in time can result in limited knowledge of the ecological phenomenon under study and, for example, an exceptional year in terms of weather conditions can hinder detecting overall long‐term trends in compositional changes. Therefore, future studies should try to combine data from several decades to overcome the typical limitations of temporal information.

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.

Studying the geographical distribution of species can reveal conditions and processes that may drive species presence and abundance. Organism distribution has frequently been explained by climate, but the relative role of local environmental predictors is not fully understood. Moreover, in the freshwater realm, intrinsic differences existing between different categories of water bodies can lead to significant differences in species–environment relationships. Here, we tested the relative importance of broad‐scale climate and local environmental predictors in explaining plant species distributions in freshwater lakes and streams. We built species distribution models to investigate which predictors best explain aquatic plant distribution in two categories of water bodies. We used species inventories and records of three climate and eight local environmental predictors for 150 lakes and 150 streams in Finland. We found that sets of predictors that explain the distribution of macrophyte species are unique depending on if species are in a lake or a stream. Overall, air temperature and ecosystem size were essential to predict aquatic plant species presence in both water body categories. Broad‐scale climate predictors were always very important in explaining species distribution, while local environmental conditions such as water chemistry were of variable influence, depending on species and water body category. These results are probably due to high spatial and temporal variability and range of water physico‐chemical parameters, especially in streams. Nonetheless, despite a lower relative importance than climatic factors, local environmental predictors also strongly affected species distributions. Our findings highlight that incorporating local environmental conditions to species distribution models in addition to climate predictors is necessary to improve predictions, particularly for distribution of stream flora. Considering the species‐specific responses of aquatic plants to their environment, studying species individually with species distribution models represents a useful analysis.

Highlights •We examined how eutrophication affects macroinvertebrate biodiversity in 33 lakes. •Three facets of alpha diversity declined with increasing of eutrophication. •Functional diversity indices performed best in portraying anthropogenic disturbances. •Environmental filtering dominated over spatial factors in explaining multifaceted alpha diversity. Abstract The accelerated eutrophication of freshwater lakes has become an environmental problem worldwide. Increasing numbers of studies highlight the need to incorporate functional and phylogenetic information of species into bioassessment programms, but it is still poorly understood how eutrophication affects multiple diversity facets of freshwater communities. Here, we assessed the responses of taxonomic, phylogenetic and functional diversity of benthic macroinvertebrates to water eutrophication in 33 lakes in the Yangtze River floodplain in China. Our results showed that macroinvertebrate assemblage structure was significantly different among four lake groups (river-connected, macrophyte-dominated, macrophyte-algae transition, and algae-dominated). Three taxonomic, two phylogenetic and two functional diversity indices were significantly different among the lake groups. Except for the increasing trend of Lambda+, these metrics showed a clear decreasing trend with increasing levels of eutrophication, with highest values detected in river-connected and macrophyte-dominated lakes, followed by macrophyte-algae transition lakes and algal-dominated lakes. Although differing in the number and identity of key environmental and spatial variables among the explanatory models of different diversity indices, environmental factors (eutrophication-related water quality variables) played more important role than spatial factors in structuring all three facets of alpha diversity. The predominant role of environmental filtering can be attributed to the strong eutrophication gradient across the studied lakes. Among the three diversity facets, functional diversity indices performed best in portraying anthropogenic disturbances, with variations in these indices being solely explained by environmental factors. Spatial factors were mostly weak or negligible in accounting for the variation in functional diversity indices, implying that trait-based indices are robust in portraying anthropogenic eutrophication in floodplain lakes. However, variation in some taxonomic and phylogenetic diversity indices were also affected by spatial factors, indicating that conservation practitioners and environmental managers should use these metrics with caution when providing solutions for addressing eutrophication in floodplain lakes.

Current global environmental change calls for comprehensive and complementing approaches for biodiversity conservation. According to recent research, consideration of the diversity of Earth's abiotic features (i.e. geodiversity) could provide new insights and applications into the investigation and management of biodiversity. However, methods to map and quantify geodiversity at local scale have not been developed although this scale is important for conservation planning. Here, we introduce a field methodology for observing plot-scale geodiversity, pilot the method in an Arctic–alpine tundra environment, provide empirical evidence on the plot-scale biodiversity–geodiversity relationship and give guidance for practitioners on the implementation of the method. The field method is based on observation of geofeatures, that is, elements of geology, geomorphology and hydrology, from a given area surrounding a location of species observations. As a result, the method provides novel information on the variation of abiotic nature for biodiversity research and management. The method was piloted in northern Norway and Finland by observing geofeatures from 76 sites at three scales (5, 10 and 25 m radii). To explore the relationship between measures of biodiversity and geodiversity, the occurrence of vascular plant species was recorded from 2 m × 2 m plots at the same sites. According to the results, vascular plant species richness was positively correlated with the richness of geofeatures (Rs = 0.18–0.59). The connection was strongest in habitats characterized by deciduous shrubs. The method has a high potential for observing geofeatures without extensive geological or geomorphological training or field survey experience and could be applied by conservation practitioners. Synthesis and applications. Consideration of geodiversity in understanding, analysing and conserving biodiversity could facilitate environmental management and ensure the long-term sustainability of ecosystem functions. With the developed method, it is possible to cost-efficiently observe the elements of geodiversity that are useful in ecology and biodiversity conservation. Our approach can be adapted in different ecosystems and biodiversity investigations. The method can be adjusted depending on the abiotic conditions, expertise of the observer(s) and the equipment available. publishedVersion

1) Isoëtes sabatina is the rarest aquatic quillwort in Europe. Although recently found (2013) in Lake Bracciano (central Italy), the species is just one step away from extinction with an estimated population not exceeding 400 individuals and a spatial range of a few hundred square metres. 2) Lake Bracciano is a deep, oligo-mesotrophic Mediterranean volcanic lake that has been subjected to human activities. From January to October 2017, the lake experienced a dramatic water level decrease (up to −1.50 m), which significantly affected the littoral zone and the habitat of I. sabatina. 3) To improve the chances of survival of I. sabatina, the first eco-taxonomic investigation on this species was carried out to describe its genetic distinctness, physical and chemical requirements and companion species. 4) The phylogenetic position of I. sabatina was investigated by applying standard DNA barcoding methods. Simultaneously, during summer 2019, the physical and chemical features of water and sediments of the I. sabatina population and five small Alpine lakes colonized by Isoëtes echinospora – a supposed close relative – were characterized. These data were then compared with the available data on the trophic requirements of the target obligate aquatic Isoëtes, together with Isoëtes lacustris and Isoëtes malinverniana. 5) The present survey confirmed the taxonomic and ecological distinctness of I. sabatina – providing the first evidence of genetic differentiation from I. echinospora. Isoëtes sabatina grows in waters with temperature, conductivity and total alkalinity up to 30°C, 561 μS cm−1 and 3.45 meq L−1, respectively. 6) The edaphic requirements of I. sabatina confirm its outstanding conservation value, and this study offers a basic understanding of how to prevent its extinction. Now, all possible actions must be taken immediately to save this species.

Lateral hydrological connectivity (LHC) is a key process in maintaining aquatic biodiversity in river floodplain ecosystems. However, the effects of LHC loss on aquatic biodiversity are rarely studied. Here, we evaluated, for the first time, the responses of multiple facets (i.e., taxonomic, functional and phylogenetic) of alpha and beta diversity of freshwater molluscs to the LHC loss in 23 floodplain lakes in the Yangtze River Basin in China. Our results showed that taxonomic and functional alpha diversities were all significantly higher in connected lakes (CLs) than in disconnected lakes (DLs), whereas phylogenetic alpha diversity (Δ

It has been predicted that spatial beta diversity shows a decreasing trend in the Anthropocene due to increasing human impact, causing biotic homogenisation. We aimed to discover if vascular aquatic macrophyte communities show different spatial patterns in beta diversity in relation to land use and environmental characteristics in different decades from 1940s to 2010s. We aimed to discover if spatial structures differ between species‐, phylogeny‐ and functional‐based beta diversity. We used presence–absence data of aquatic macrophytes from five decades from small boreal lakes. We utilized generalised dissimilarity modelling to analyse spatial patterns in beta diversity in relation to environmental gradients. We found that lake elevation and pH were the most important variables in each decade, while land use was not particularly important in shaping beta diversity patterns. We did not find signs of a decreasing trend in spatial beta diversity in our study area during the past 70 yr. We did not find signs of either biotic homogenisation or biotic differentiation (taxonomic, phylogenetic or functional). Vascular aquatic macrophyte communities showed only slightly different beta diversity patterns in relation to human impact across decades. The patterns of different facets of beta diversity diverged only slightly from each other. Lake position in the landscape, reflecting both natural connectivity and lake characteristics, explained the patterns found in beta diversity, probably because our study area has faced only modest changes in land use from 1940s to 2010s when compared globally. Our study highlights the fact that biotic homogenisation is not an unambiguous process acting similarly at all spatial and temporal scales or in different environments and different organism groups.

Aim: Conserving freshwater biodiversity in a rapidly changing world requires updated planning schemes and research efforts. Geodiversity — the diversity of Earth surface forms, materials and processes — and biodiversity are interlinked at a fundamental level. This relationship is being considered in a growing number of studies, yet research from freshwater environments is scarce. We used geodiversity (rock‐type, soil‐type and geomorphological richness), local and climatic variables to explore whether geodiversity can be used as a surrogate for aquatic plant species richness in lakes and rivers. Location: Finland. Taxon: Aquatic plants. Methods: We compared geodiversity variables (measured within 1‐km² grid cells) to well‐studied local (e.g. area, alkalinity) and climate (e.g. growing degree‐days) variables, and examined the patterns between habitat types (lakes and rivers) and among all taxa and major functional groups (helophytes and hydrophytes). We modelled lake (n = 145) and river (n = 146) plant species richness with generalized linear models, and further partitioned variation to measure the independent and shared contributions of the geodiversity, climate and local environmental variable groups. As a complementary analysis, and to identify single important variables explaining variation in aquatic plant species richness, we utilized boosted regression trees. Results: We found a positive relationship between aquatic plant species richness and catchment geodiversity variation with recurring patterns across two different freshwater habitat types and two aquatic plant functional groups. Higher variation in geodiversity (measured at landscape scale) supported higher freshwater biodiversity (measured at the local scale) of lakes and rivers. Main conclusions: Geodiversity can be a useful addition to biodiversity modelling, and it should be considered in conservation schemes and monitoring efforts, further supporting the principle of conserving nature’s stage. Yet, differences between habitats and functional groups suggest that more habitat‐specific approaches and multiple biodiversity measures should be considered. Our study is an important signpost guiding further studies on the biodiversity–geodiversity relationship in freshwater ecosystems.

In the era of the Anthropocene, environmental change is accelerating biodiversity loss across ecosystems on Earth, among which freshwaters are likely the most threatened. Different biodiversity facets in the freshwater realm suffer from various environmental changes that jeopardize the ecosystem functions and services important for humankind. In this work we examine how environmental changes (e.g., climate change, eutrophication, or invasive species) affect trait-based, functional, and phylogenetic diversity of biological communities. We first developed a simple conceptual model of the possible relationships between environmental change and these three diversity facets in freshwaters and, secondly, systematically reviewed articles where these relationships had been investigated in different freshwater ecosystems. Finally, we highlighted research gaps from the perspectives of organisms, ecosystems, stressors, and geographical locations. Our conceptual model suggested that both natural factors and global change operating at various spatial scales influence freshwater community structure and ecosystem functioning. The relationships between biodiversity and environmental change depend on geographical region, organism group, spatial scale, and environmental change gradient length. The systematic review revealed that environmental change impacts biodiversity patterns in freshwaters, but there is no single type of biodiversity response to the observed global changes. Natural stressors had different, even contradictory, effects (i.e., multiple, negative, and positive) on biodiversity compared with anthropogenic stressors. Anthropogenic stressors more often decreased biodiversity, although eutrophication and climate change affected freshwater ecosystems in a complex, more multi-dimensional way. The research gaps we identified were related, for example, to the low number of community-based biodiversity studies, the lack of information on true phylogenies for all freshwater organism groups, the missing evaluations whether species traits are phylogenetically conserved, and the geographical biases in research (i.e., absence of studies from Africa, Southern Asia, and Russia). We hope that our review will stimulate more research on the less well-known facets and topics of biodiversity loss in highly vulnerable freshwater ecosystems.

1. Ecogeographical rules refer to recurring patterns in nature, including the latitudinal diversity gradient (LDG), Rapoport's rule and Bergmann's rule, amongst others. In the present study, the existence of these rules was examined for diving beetles (Coleoptera: Dytiscidae), a family of aquatic predatory beetles. 2. Assemblage‐level data were analysed for diving beetles, focusing on species richness, local contribution to beta diversity (LCBD), mean range size and mean body size across the biogeographical provinces of Northern Europe. First, each of these variables was correlated with latitude, and then variation in each variable was modelled using actual environmental variables in boosted regression tree analysis. 3. Species richness was found to decrease with latitude, LCBD increased with latitude, mean range size did not show a significant relationship with latitude, and mean body size decreased with latitude. The latter finding was in contrast to Bergmann's rule. The actual environmental variables best predicting variation in these four response variables varied among the models, although they generally included temperature‐related and land use variables as the most influential ones. 4. The results obtained in the present study suggest that diving beetles conformed to the LDG, did not follow Rapoport's rule, and showed a reversed latitudinal gradient in the context of Bergmann's rule. In addition, species‐poor provinces harboured ecologically most unique faunas, suggesting that species richness and LCBD are complementary measures of biodiversity. 5. Even though general support was not found for most of the ecogeographical rules examined, the findings of the present study are interesting because they suggest that aquatic ectothermic invertebrates may show patterns different from those originally described for terrestrial endothermic vertebrates.

AimBiogeographical regionalisations are actively studied in different ecosystems, because they increase our understanding on fundamental broad□scale patterns and can help us in the establishment of conservation areas. Thus, we studied how well existing freshwater ecoregions describe geographical delineation for inland water plants and which ecogeographical gradients explain them.LocationNorth America, excluding Mexico and remote islands.TaxonFreshwater vascular plants of all taxa and different functional groups.MethodsUsing newly available fine–grained data on freshwater plant distributions, we calculated internal homogeneity and cross–boundary heterogeneity among neighbouring ecoregions. We further integrated measures of community dissimilarity to assess whether the degree of within–ecoregion homogeneity and distinctness are driven by their relationships to species replacements and richness differences, and explored how a complex suite of ecogeographical mechanisms and plant life forms affect ecoregion delineation using spatially explicit regression routines.ResultsWe found a clear geographical patterning of ecoregion robustness for North American freshwater plants, with their communities being more internally homogeneous and more similar to one another in polar and subtropical inland waters. Surprisingly, the degree of internal homogeneity and ecoregion distinctness were almost equally driven by species replacements and richness differences. Considering different life forms, ecoregion delineation performed best for emergent and floating–leaved plants. Finally, within–ecoregion homogeneity and distinctness were best explained by annual mean temperature and terrain ruggedness, respectively, with mean water alkalinity, ecoregion area and Late Quaternary Ice Age legacies having supplementary effects.Main conclusionsOur findings emphasise that geographical regionalisations founded on a particular organismal group are not applicable for all taxa. Our study is a promising starting point for further investigations of geographical delineations for different freshwater taxa. These updated regionalisations can then be used for conserving different biotas in freshwaters, which are currently among the most threatened ecosystems in the world.Statement of significanceIn biogeographical regionalisation biota is categorized to meaningful geographical units, such as ecoregions. However, ecoregions delineated for a particular group of organisms may not be applicable to another assemblages. We studied how ecoregions founded on fish are suitable for freshwater plants across North America. Our findings suggest that these ecoregions did not show consistent robustness for freshwater plants in North America. This study is a promising starting point for further investigations of geographical delineations for different freshwater taxa, having also value in conservation planning and management.

Understanding the variation in community composition and species abundances, i.e., β-diversity, is at the heart of community ecology. A common approach to examine β-diversity is to evaluate directional turnover in community composition by measuring the decay in the similarity among pairs of communities along spatial or environmental distances. We provide the first global synthesis of taxonomic and functional distance decay along spatial and environmental distance by analysing 149 datasets comprising different types of organisms and environments. We modelled an exponential distance decay for each dataset using generalized linear models and extracted r2 and slope to analyse the strength and the rate of the decay. We studied whether taxonomic or functional similarity has stronger decay across the spatial and environmental distances. We also unveiled the factors driving the rate of decay across the datasets, including latitude, spatial extent, realm, and organismal features. Taxonomic distance decay was stronger along spatial and environmental distances compared with functional distance decay. The rate of taxonomic spatial distance decay was the fastest in the datasets from mid-latitudes while the rate of functional decay increased with latitude. Overall, datasets covering larger spatial extents showed a lower rate of decay along spatial distances but a higher rate of decay along environmental distances. Marine ecosystems had the slowest rate of decay. This synthesis is an important step towards a more holistic understanding of patterns and drivers of taxonomic and functional β-diversity.

A variety of ways can be used to measure geodiversity, but no consensus on quantifying it exists to date. Most quantifications have thus far focused on the abiotic diversity of individual sites, which offers only limited views on variation of abiotic nature. It is, therefore, important to examine the between-site geodiversity, i.e., the dissimilarities of geofeatures (elements of geodiversity) between different sites. For instance, it would be interesting to recognize not only the most geodiverse sites of an area, but also the sites that have the most unique compositions of geofeatures. This extended geodiversity information could be further applied in nature conservation or land-use planning. We propose that geodiversity research would benefit from adopting the alpha, beta and gamma concepts of species diversity research to provide a more holistic framework for geodiversity assessments. In particular, the inclusion of distance metrics for measuring beta biodiversity could open new perspectives in evaluating beta geodiversity. The integration of these diversity concepts into geodiversity research would also allow a better joint understanding of biotic and abiotic diversity.

Community ecology has had a strong focus on single snapshots of species compositional variation in time. However, environmental change often occurs slowly at relatively broad spatio-temporal scales, which requires historically explicit assessments of long-term metacommunity dynamics, such as the order of species arrival during community assembly (i.e., priority effects), a theme that merits further empirical quantification. In this study, we applied the Bayesian inference scheme of Hierarchical Modeling of Species Communities together with information on functional traits and evolutionary dependencies to efficiently explore the question of how ecological communities are organized in space and time. To do this, we used a comprehensive time-series dataset from boreal lake plants and adopted the perspective that more sound conclusions on metacommunity dynamics can be gained from studies that consider a historically integrative approach over long timeframes. Our findings revealed that historical contingency via priority effects can profoundly shape community assembly under the influence of environmental change across decades (here, from the 1940s to the 2010s). Similarly, our results supported the existence of both positive and negative species-to-species associations in lake plants, suggesting that functional divergence can switch the inhibition–facilitation balance at the metacommunity level. Perhaps more importantly, this proof-of-concept study supports the notion that community ecology should include a historical perspective and suggests that ignoring priority effects may risk our ability to identify the true magnitude of change in present-day biotic communities.

Highlights • Few broad-scale patterns are well-studied for aquatic macrophytes. • Local environmental predictors were important for freshwater plants at broad-scales. • Several knowledge gaps on macroecology of freshwater plants were identified. • Lack of lotic studies and databases of species traits and phylogeny were found. • Additional macroecological investigations on freshwater plants are clearly needed. Broad-scale studies of species distributions and diversity have contributed to the emergence of general macroecological rules. These rules are typically founded on research using well-known terrestrial taxa as models and it is thus uncertain whether aquatic macrophytes follow these macroecological rules. Our purpose is to draw together available information from broad-scale research on aquatic macrophytes growing in lakes, ponds, wetlands, rivers and streams. We summarize how different macroecological rules fit the patterns shown by freshwater plants at various spatial scales. Finally, we outline future actions which should be taken to advance macroecological research on freshwater plants. Our review suggested that some macroecological patterns are relatively well-evidenced for aquatic macrophytes, whereas little information exists for others. We found, for example, that the species richness-latitude relationship follows a unimodal pattern, and species turnover prevails over species nestedness, whereas higher nestedness-related richness differences are found in low beta diversity regions. Contrary to terrestrial plants, climate or history seem not to be dominant determinants explaining these broad-scale patterns; instead local explanatory variables (e.g., water quality, such as alkalinity and nutrients, and hydromorphology) are often important for freshwater plants. We identified several knowledge gaps related, for example, to a smaller number of studies in lotic habitats, compared with lentic habitats, lack of spatially-adequate aquatic plant studies, deficiency of comprehensive species traits databases for aquatic macrophytes, and absence of a true phylogeny comprising most freshwater plant lineages. We hope this review will encourage the undertaking of additional macroecological investigations on freshwater plants across broad spatial and temporal scales.

Aim: Range size conservatism suggests that closely-related species maintain geographic ranges of similar extent. However, consensus regarding this suggestion has not been reached. To shed more light on this phenomenon, we studied freshwater plant range size conservatism, range overlap and environmental niche conservatism using congeneric species comparison in two continents. In addition, we investigated whether a phylogenetic signal is found in the range sizes of aquatic plants. Location: Europe and North America. Taxon: Freshwater plants. Methods: Across spatial resolutions (50 km², 100 km² and 200 km²), we applied Spearman correlations among 347 and 730 pairs of congeneric species in Europe and North America, respectively, and 63 pairs shared between them. In addition, Spearman correlations were used to evaluate how the degree of spatial overlap influences range sizes and which environmental variables explain variation in range sizes. Brownian evolutionary model was used to assess the phylogenetic signal in species range sizes. Results: We found no evidence of range size conservatism across spatial resolutions for any species and species shared between the continents. In addition, range sizes of more closely related species did not overlap geographically more than those of distantly related ones and no support for environmental niche conservatism was evidenced. Main conclusions: We found that aquatic plants show no range size conservatism in the Northern Hemisphere. This means that it is challenging to define different range sizes of freshwater plants through species traits. Furthermore, we are unable to predict unknown distributions of extant aquatic plant species based on known distributional attributes of closely related species. However, our findings suggest that the interpretations of previous investigations on the range sizes of aquatic plants remain valid due to lack of range size conservatism. These practical implications encourage studying range size conservatism across realms and regions, especially for understudied organismal groups.

Highlights • New empirical studies are repeatedly demonstrating the importance of rare species. • However, very little is still known about species rarity in the aquatic realm. • Continental maps suggest variable latitudinal trends in freshwater plants. • Human footprint, past and present-day climates predict rarity hotspots. • Current centers of freshwater plant rarity might shrink within a century. Patterns of species rarity have long fascinated ecologists, yet most of what we know about the natural world stems from studies of common species. A large proportion of freshwater plant species has small range sizes and are therefore considered rare. However, little is known about the mechanisms and geographical distribution of rarity in the aquatic realm and to what extent diversity of rare species in freshwater plants follows their terrestrial counterparts. Here, we present the first in–depth analysis of geographical patterns, potential deterministic ecogeographical factors and projected scenarios of freshwater vascular plant rarity using 50 × 50 km grid cells across Europe (41°N–71°N) and North America (25°N–78°N). Our results suggest that diversity of rare species shows different patterns in relation to latitude on the two continents, and that hotspots of rarity concentrate in a relatively small proportion of the European and North American land surface, especially in mountainous as well as in climatically rare and stable areas. Interestingly, we found no differences among alternative rarity definitions and measures when delineating areas with notably high diversity of rare species. Our findings also indicate that few variables, namely a combination of current climate, Late Quaternary climate–change velocity and human footprint, are able to accurately predict the location of continental centers of rare species diversity. However, these relationships are not geographically homogeneous, and the underlying factors likely act synergistically. Perhaps more importantly, we provide empirical evidence that current centers of rare species diversity are characterized by higher anthropogenic impacts and might shrink disproportionately within this century as the climate changes. Our reported distributional patterns of species rarity align with the known trends in species richness of other freshwater organisms and may help conservation planners make informed decisions mitigating the effects of climate change and other anthropogenic impacts on biodiversity.

Documenting the patterns of biological diversity on Earth has always been a central challenge in macroecology and biogeography. However, for the diverse group of freshwater plants, such research program is still in its in- fancy. Here, we examined global variation in taxonomic, functional and phylogenetic beta diversity patterns of lake macrophytes using regional data from six continents. A data set of ca. 480 lake macrophyte community ob- servations, together with climatic, geographical and environmental variables, was compiled across 16 regions worldwide. We (a) built the very first phylogeny comprising most freshwater plant lineages; (b) exploited a wide array of functional traits that are important to macrophyte autoecology or that relate to lake ecosystem functioning; (c) assessed if different large-scale beta diversity patterns show a clear latitudinal gradient from the equator to the poles using null models; and (d) employed evolutionary and regression models to first identify the degree to which the studied functional traits show a phylogenetic signal, and then to estimate community- environment relationships at multiple spatial scales. Our results supported the notion that ecological niches evolved independently of phylogeny in macrophyte lineages worldwide. We also showed that taxonomic and phylogenetic beta diversity followed the typical global trend with higher diversity in the tropics. In addition, we were able to confirm that species, multi-trait and lineage compositions were first and foremost structured by climatic conditions at relatively broad spatial scales. Perhaps more importantly, we showed that large-scale processes along latitudinal and elevational gradients have left a strong footprint in the current diversity patterns and community-environment relationships in lake macrophytes. Overall, our results stress the need for an inte- grative approach to macroecology, biogeography and conservation biology, combining multiple diversity facets at different spatial scales. JGG appreciates financial support from the Spanish Ministry of Economy and Industry [project METAPONDS, grant CGL2017- 84176R], the Junta de Castilla y León [grant LE004G18] and from the Fundación Biodiversidad (Spanish Ministry for Ecological Transi- tion and Demographic Challenge). BAL was supported by National Research, Development and Innovation Fund [grant NKFIH, OTKA PD120775] and by the Bolyai János Research Scholarship of the Hun- garian Academy of Sciences. S.K. was supported by NWO Veni [grant 86312012]. Sampling of the coastal Brazilian lakes was financed by NWO [grant W84-549]; The National Geographic Society [grant 7864-5]; and CNPq [grants 480122, 490409, 311427]. We thank the SALGA team, especially Gissell Lacerot, Nestor Mazzeo, Vera Huszar, David da Motta Marques, and Erik Jeppesen for organizing and exe- cuting the SALGA field sampling campaign and Bruno Irgang† and Eduardo Alonso Paz for helping with identification. We thank Minne- sota and Wisconsin Departments of Natural Resources for collecting the macrophyte data. We are grateful to Carol Reschke for her work in combining and performing quality control for the Minnesota mac- rophyte data used in the analysis. This is contribution no. 607 of the Natural Resources Research Inst. of the Univ. of Minnesota Duluth. Provision of New Zealand macrophyte data was possible via NIWA SSIF funding. JGG appreciates financial support from the Spanish Ministry of Economy and Industry [project METAPONDS, grant CGL2017- 84176R], the Junta de Castilla y León [grant LE004G18] and from the Fundación Biodiversidad (Spanish Ministry for Ecological Transi- tion and Demographic Challenge). BAL was supported by National Research, Development and Innovation Fund [grant NKFIH, OTKA PD120775] and by the Bolyai János Research Scholarship of the Hun- garian Academy of Sciences. S.K. was supported by NWO Veni [grant 86312012]. Sampling of the coastal Brazilian lakes was financed by NWO [grant W84-549]; The National Geographic Society [grant 7864-5]; and CNPq [grants 480122, 490409, 311427]. We thank the SALGA team, especially Gissell Lacerot, Nestor Mazzeo, Vera Huszar, David da Motta Marques, and Erik Jeppesen for organizing and exe- cuting the SALGA field sampling campaign and Bruno Irgang† and Eduardo Alonso Paz for helping with identification. We thank Minne- sota and Wisconsin Departments of Natural Resources for collecting the macrophyte data. We are grateful to Carol Reschke for her work in combining and performing quality control for the Minnesota mac- rophyte data used in the analysis. This is contribution no. 607 of the Natural Resources Research Inst. of the Univ. of Minnesota Duluth. Provision of New Zealand macrophyte data was possible via NIWA SSIF funding.

Functional homogenisation occurs across many areas and organism groups, thereby seriously affecting biodiversity loss and ecosystem functioning. In this study, we examined how functional features of aquatic macrophytes have changed during a 70-year period at community and species levels in a boreal lake district. At the community level, we examined if aquatic macrophyte communities showed different spatial patterns in functional composition and functional richness in relation to main environmental drivers between the time periods. We also observed each species in functional space to assess if species with certain sets of traits have become more common or rare in the 70-year study period. We found changes in the relationship between functional community composition and the environment. The aquatic macrophyte communities showed different patterns in functional composition between the two time periods, and the main environmental drivers for these changes were partly different. Temporal changes in functional richness were only partially linked to concomitant changes in the environment, while stable factors were more important. Species’ functional traits were not associated with commonness or rarity patterns. Our findings revealed that functional homogenisation has not occurred across these boreal lakes, ranging from small oligotrophic forest lakes to larger lakes affected by human impacts.

Biotic interactions are fundamental drivers governing biodiversity locally, yet their effects on geographical variation in community composition (i.e. incidence‐based) and community structure (i.e. abundance‐based) at regional scales remain controversial. Ecologists have only recently started to integrate different types of biotic interactions into community assembly in a spatial context, a theme that merits further empirical quantification. Here, we applied partial correlation networks to infer the strength of spatial dependencies between pairs of organismal groups and mapped the imprints of biotic interactions on the assembly of pond metacommunities. To do this, we used a comprehensive empirical dataset from Mediterranean landscapes and adopted the perspective that community assembly is best represented as a network of interacting organismal groups. Our results revealed that the co‐variation among the beta diversities of multiple organismal groups is primarily driven by biotic interactions and, to a lesser extent, by the abiotic environment. These results suggest that ignoring biotic interactions may undermine our understanding of assembly mechanisms in spatially extensive areas and decrease the accuracy and performance of predictive models. We further found strong spatial dependencies in our analyses which can be interpreted as functional relationships among several pairs of organismal groups (e.g. macrophytes–macroinvertebrates, fish–zooplankton). Perhaps more importantly, our results support the notion that biotic interactions make crucial contributions to the species sorting paradigm of metacommunity theory and raise the question of whether these biologically‐driven signals have been equally underappreciated in other aquatic and terrestrial ecosystems. Although more research is still required to empirically capture the importance of biotic interactions across ecosystems and at different spatial resolutions and extents, our findings may allow decision makers to better foresee the main consequences of human‐driven impacts on inland waters, particularly those associated with the addition or removal of key species.

1. The latitudinal diversity gradient hypothesis suggests that species richness should be highest at low latitudes, whereas Rapoport’s rule states that largest ranges ought to be found for species at high latitudes. However, there is no consensus over these patterns and their underlying drivers in the freshwater realm. 2. We investigated species richness and mean range size of freshwater plants in 50 × 50 km grid cells across Europe (40°N–71°N) and North America (25°N–78°N), supplemented with data based on 1° latitudinal bands for mean range size. We were especially interested to find out whether there are similarities and differences in these ecogeographical patterns and their underlying drivers between the continents due to their contrasting historical characteristics, spatial extent and topography. 3. First, we used partial regression to reveal whether species richness and mean range size of freshwater plants have a linear or quadratic relationship with latitude. Second, we employed variation partitioning based on partial regression to model relationships between plant species richness and mean range size and four explanatory variable groups (i.e. environmental features, current climate, historical climate and geographical location). Third, we utilized boosted regression tree analysis to further investigate species richness and mean range size of freshwater plants in relation to a set of explanatory variables. 4. Our results revealed that species richness showed relatively similar patterns in relation to latitude between the continents. Similarly, mean range size trends were alike in North America whether we used 50 × 50 km grid cell data or 1° latitudinal bands. Instead, different patterns in mean range size emerged between the used datasets in Europe. For both species richness and mean range size, current climate (with different individual predictor variables) was the main driver in both the continents, but historical effects had a small influence on the response variables. 5. Synthesis. Our findings indicated that major ecogeographical rules can strongly vary for the same taxonomic group across broad scales between continents. It is also premature to rely solely on well‐known terrestrial taxonomic groups when drawing generalizations about ecogeographical rules.

We investigated whether environmental filtering or dispersal-related factors mostly drive helophyte and hydrophyte species richness and community composition in 93 lakes situated in Baikal Siberia. Using partial linear regression and partial redundancy analysis, we studied (1) what are the relative roles of environmental variables, dispersal variables, spatial processes and region identity (i.e., river basins) in explaining variation in the species richness and species composition of helophytes and hydrophytes across 93 Siberian lakes, and (2) what are the differences in the most important explanatory variables driving community variation in helophytes versus hydrophytes? We found that, for both species richness and species composition, environmental variables clearly explained most variation for both plant groups, followed by region identity and dispersal-related variables. Spatial variables were significant only for the species composition of hydrophytes. Nutrient-salinity index, a proxy for habitat trophic-salinity status, was by far the most significant environmental determinant of helophytes and hydrophytes. Our results indicate that environmental factors explained the most variation in both species richness and species composition of helophytes and hydrophytes. Nevertheless, dispersal-related variables (i.e. spatial and dispersal) were also influential but less important than environmental factors. Furthermore, the dispersal-related variables were more important for hydrophytes than for helophytes. Most brackish permanent lakes were mostly located in the steppe biomes of southern Transbaikalia. This characteristic along with the oldest age, the largest distances to both river and settlements and the lowest temperatures in the study region distinguished them from freshwater, drained and more nutrient-rich floodplain lakes.

Documenting the patterns of biological diversity on Earth has always been a central challenge in macroecology and biogeography. However, we are only beginning to generate an understanding of the global patterns and determinants of macrophyte diversity. Here, we studied large-scale variation and community-environment relationships of lake macrophytes along climatic and geographical gradients using regional data from six continents. We applied statistical routines typically used in the context of metacommunity studies to provide novel insights into macrophyte community compositional patterns within regions worldwide. We found that lake macrophyte metacommunities followed clumped species replacement structures, suggesting that two or more species groups were responding similarly to the environment within regions. Underlying such general convergence, our results also provided evidence that community-environment relationships were largely context-dependent, stressing that no single mechanism is enough to account for the complex nature of compositional variation. Surprisingly, we found no general relationships between functional or phylogenetic composition and main metacommunity types, suggesting that linking multi-trait and evolutionary information to the elements of metacommunity structure is not straightforward. Our findings highlight that global conservation initiatives and biodiversity protection need to capture environmental variation at the metacommunity level, and acknowledge the highly context-dependent patterns in the community-environment relationships of lake macrophytes. Overall, we emphasize the need to embrace the potential complexity of ecological inferences in metacommunity organization across the globe.

Mapping of ecosystem services (ESs) provide valuable information on the geographical variation of ESs and their relation to overall diversity. Although the relationship between biodiversity and ESs has been intensively explored, little is known how geodiversity (i.e., variety of geological, geomorphological and soil features) is associated with different ESs. We studied 1) the spatial variation of geodiversity and biodiversity in relation to six ESs (i.e., forest carbon budget, potential supply of groundwater, milk and meat production, crop production, amount of free-time residences and nationally valuable landscapes) using variation partitioning (VP), and 2) the spatial overlap between geodiversity and biodiversity and ESs using generalized additive models (GAM) in 1006 intensively surveyed grid cells of 100 km2 located across Finland. In the VP, biodiversity independently explained more of the variation than geodiversity for majority of the ESs. However, shared explanation ability of biodiversity and geodiversity was considerable for majority of ESs (forest carbon budget: 41.3%, crop production: 15.0%, free-time residences: 15.2% and valuable landscapes: 7.3%), often exceeding that of both independent contributions. GAMs indicated that increase in both biodiversity and geodiversity enhances forest carbon budget (D2 = 66.8% and 12.4%, respectively), potential production of groundwater (8.3% and 0.1%), crop production (35.7% and 8.9%), free-time residences (40.0% and 7.9%) and valuable landscapes (11.6% and 6.9%). However, the positive relationship between diversity and ESs levelled off for many of the ESs. Our findings suggest that geodiversity is an important complementing factor in explaining spatial variation of the ESs in high-latitude regions. We also found dominantly synergic effects between abiotic diversity and ESs. Thus, our study results highlight the need to more deeply incorporate abiotic diversity into ESs research. Environmental conservation and management would benefit from the more comprehensive integration of geodiversity to ESs research along with the changing environmental conditions of future decades.

Different components of global change (e.g., climate change, land use, pollution, and introduced species) continue to alter biodiversity worldwide. As northern regions are still relatively undisturbed and will likely face clear increases in temperature in the near-future, we examined the signs of biodiversity change due to anthropogenic stressors using a systematic review of previous studies. Our aim was to map where, in which way, and owing to which stressor biodiversity in northern regions has changed. We made a systematic literature search covering the years between 2000 and 2015 to obtain a comprehensive selection of recent research. As species richness was clearly the most commonly used indicator of biodiversity, we only concentrated on this aspect of biodiversity. We compared different biological groups, regions, and ecosystems. In the majority of the cases, anthropogenic stressors had decreased species richness, or had no effects on it, while increasing or multiple effects of stressors on species richness were less common. Freshwater ecosystems were most sensitive to anthropogenic stressors, as species richness often decreased owing to these stressors. The effects of land use on richness were covered relatively widely in the selected set of articles, but the effects of other components of global change on species richness require further attention. Despite the fact that pollution was not as commonly studied stressor as land use, it was the most harmful stressor type affecting species richness. Geographically, most studies were located in boreal Canada or Fennoscandia, while no studies were executed in vast circumpolar areas where the temperature rise has been greatest and the projected climate change is likely to be fast. Overall, we could find an alarmingly small set of studies that described the effects of actual anthropogenic stressors in real-life circumstances in northern high latitudes.

The biodiversity of aquatic ecosystems is under threat and there is an urgent need to quantify the various facets of biodiversity to assess the conservation value of freshwater ecosystems. The effects of taxonomic relatedness have so far not been taken into account in biodiversity assessments of lake macrophytes. We therefore tested the response of species richness and average taxonomic distinctness (AvTD) of aquatic macrophytes along environmental gradients using linear regression models and Bayesian Information Criterion variable selection method. We selected data from four regions, each with 50–60 lakes, situated in northern Europe (Finland and Sweden) and northern America (Minnesota and Wisconsin). We separately studied all macrophyte species, hydrophytes and helophytes. Species richness and AvTD of aquatic macrophytes were generally negatively related in all regions, although it was not statistically significant. Both biodiversity measures responded to environmental gradients to various degrees among the studied macrophyte groups and regions. Species richness was best explained by alkalinity and lake area in Finland, by elevation, annual mean temperature and total phosphorus in Minnesota, and by alkalinity in Wisconsin. AvTD was best explained by alkalinity, annual mean temperature and total phosphorus in Finland and by alkalinity in Wisconsin. Very weak relationships were found in Sweden. Our findings strongly suggest that complementary indices are needed to indicate more comprehensively the effects of environmental conditions on freshwater biodiversity. Species richness was found to be a better measure than AvTD to account for conservation value in freshwaters. However, further research is required to evaluate the usefulness of AvTD to indicate conservation value (e.g. randomisation tests), because alternative measures are clearly needed for those freshwater taxa lacking complete information on true phylogenetic diversity.

Context: ‘Conserving Nature’s stage’ has been advanced as an important conservation principle because of known links between biodiversity and abiotic environmental diversity, especially in sensitive high-latitude environments and at the landscape scale. However these links have not been examined across gradients of human impact on the landscape. Objectives: To (1) analyze the relationships between land-use intensity and both landscape-scale biodiversity and geodiversity, and (2) assess the contributions of geodiversity, climate and spatial variables to explaining vascular plant species richness in landscapes of low, moderate and high human impact. Methods: We used generalized additive models (GAMs) to analyze relationships between land-use intensity and both geodiversity (geological, geomorphological and hydrological richness) and plant species richness in 6191 1-km² grid squares across Finland. We used linear regression-based variation partitioning (VP) to assess contributions of climate, geodiversity and spatial variable groups to accounting for spatial variation in species richness. Results: In GAMs, geodiversity correlated negatively, and plant species richness positively, with land-use intensity. Both relationships were non-linear. In VP, geodiversity best accounted for species richness in areas of moderate to high human impact. These overall contributions were mainly due to variation explained jointly with climate, which dominated the models. Independent geodiversity contributions were highest in pristine environments, but low throughout. Conclusions: Human action increases biodiversity but may reduce geodiversity, at landscape scale in high-latitude environments. Better understanding of the connections between biodiversity and abiotic environment along changing land-use gradients is essential in developing sustainable measures to conserve biodiversity under global change.

Different species' niche breadths in relation to ecological gradients are infrequently examined within the same study and, moreover, species niche breadths have rarely been averaged to account for variation in entire ecological communities. We investigated how average environmental niche breadths (climate, water quality and climate-water quality niches) in aquatic macrophyte communities are related to ecological gradients (latitude, longitude, altitude, species richness and lake area) among four distinct regions (Finland, Sweden and US states of Minnesota and Wisconsin) on two continents. We found that correlations between the three different measures of average niche breadths and ecological gradients varied considerably among the study regions, with average climate and average water quality niche breadth models often showing opposite trends. However, consistent patterns were also found, such as widening of average climate niche breadths and narrowing of average water quality niche breadths of aquatic macrophytes along increasing latitudinal and altitudinal gradients. This result suggests that macrophyte species are generalists in relation to temperature variations at higher latitudes and altitudes, whereas species in southern, lowland lakes are more specialised. In contrast, aquatic macrophytes growing in more southern nutrient-rich lakes were generalists in relation to water quality, while specialist species are adapted to low-productivity conditions and are found in highland lakes. Our results emphasise that species niche breadths should not be studied using only coarse-scale data of species distributions and corresponding environmental conditions, but that investigations on different kinds of niche breadths (e.g., climate vs. local niches) also require finer resolution data at broad spatial extents.

Metacommunity ecology has broadened considerably with the recognition that measuring beta diversity beyond the purely taxonomic viewpoint may improve our understanding of the dispersal- and niche-based mechanisms across biological communities. In that perspective, we applied a novel multidimensional approach including taxonomic, functional and phylogenetic data to enhance our basic understanding of macrophyte metacommunity dynamics. For each beta diversity metric, we calculated the mean overall value and tested whether the mean value was different from that expected by chance using null models. We also employed evolutionary and spatially constrained models to first identify the degree to which the studied functional traits showed a phylogenetic signal, and then to estimate the relative importance of spatial and environmental effects on metacommunity structure. We first found that most individual ponds were inhabited by species that were merely random draws from the taxonomic and phylogenetic species pool available in the study region. Contrary to our expectations, not all measured traits were conserved along the phylogeny. We also showed that trait and phylogenetic dimensions strongly increased the amount of variation in beta diversity that can be explained by degree of environmental filtering and dispersal limitation. This suggests that accounting for functional traits and phylogeny in metacommunity ecology helps to explain idiosyncratic patterns of variation in macrophyte species distribution. Importantly, phylogenetic and functional analyses identified the influence of underlying mechanisms that would otherwise be missed in an analysis of taxonomic turnover. Together, these results let us conclude that macrophyte species have labile functional traits adapted to dispersal-based processes and some evolutionary trade-offs that drive community assembly via species sorting. Overall, our exploration of different facets of beta diversity showed how functional and phylogenetic information may be used with species-level data to test community assembly hypotheses that are more ecologically meaningful than assessments of environmental patterns based on the purely taxonomic viewpoint.

AIM: Ecogeographical patterns have been widely studied in endothermic vertebrates, but relatively few studies have simultaneously examined patterns and causes of gradients in species richness, range size and body size in ectothermic insects. We examined patterns in species richness, mean range size and mean body size of ground beetle assemblages across the biogeographical provinces of Northern Europe, a region that was mostly covered by ice sheets during the latest Ice Age and that presents strong contemporary climatic gradients. LOCATION: Northern Europe. METHODS: We used literature information on the occurrence of ground beetles, and analysed patterns in species richness, mean range size and mean body size across the provinces using generalized linear models and boosted regression tree (BRT) analysis. RESULTS: We found a strongly decreasing gradient in species richness with increasing latitude, a strongly unimodal range size‐latitude relationship, and a weak unimodal body size‐latitude relationship in entire ground beetle assemblages. These gradients also varied among four major genera, suggesting that the overall patterns result from the nuances of smaller clades of ground beetles. The relative importance of contemporary environmental drivers also varied between species richness, mean range size and mean body size in BRT analysis. While species richness increased with mean annual temperature, mean range size showed an opposite relationship. Mean body size was most clearly associated with the precipitation of the driest month. MAIN CONCLUSIONS: Our findings showed that the latitudinal species richness gradient was strong, and it was closely related to concomitant variation in temperature, whereas variations in mean range size and mean body size were more complex. These findings suggest that the causes for range size and body size variation in insects may be complex, requiring additional insights from studies conducted at local, regional and continental scales.

Ecosystem services are fundamental to the well-being and health of people. Despite the growing awareness of the positive impacts of ecosystem services on human health, researchers have often ignored many ecosystem functions that are disadvantageous to humans. These negative facets of ecosystems are called ecosystem disservices. The central focus of this study was to test the applicability of Geographic Information Systems-based spatial accessibility analysis in mapping the potential risk of ecosystem disservice at a national scale. We used tick exposure as an example of a disservice. Worldwide, ticks (genus Ixodes) are the primary vectors of several dangerous diseases which pose threats to people. As the probability of encountering infectious ticks has increased during the last decades, new spatial information on high-risk tick exposure areas are needed. To evaluate exposure risk, we developed a tick probability map based on tick observations and environmental variables in Finland. First, we analyzed what kind of threat ticks pose to populations in residential areas and around free-time residences. Second, we studied if the movement of people (here school children) in the everyday environment increased tick exposure risk. We calculated the shortest school route for all children by using spatial accessibility analysis. Our results showed that taking the movement of people into consideration through the accessibility analysis, we can get a more realistic picture of tick exposure risk. Further, we gained a better overview of the number of children at higher exposure risk. This kind of information is crucial for pre-assessment and identification of public health strategies for control and minimizing tick-borne diseases. In general, the accessibility approach provided a good overview of areas where the greatest tick exposure was present and produced valuable information to support decision-making. The method enabled new insights into the assessment of exposure to ecosystem disservices.