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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.

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.

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.

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.

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.

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.

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.

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.

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.

Aim: Geodiversity underpins biodiversity, but the contribution of specific geofeatures or landforms has rarely been explored. In this study, we use multiple vascular plant species diversity measures on alpha, beta and gamma levels to explore the linkage between biodiversity and co‐located landforms (e.g. gullies, dunes and lake shores). We hypothesize that biodiversity will be positively related to geodiversity, which is founded on distinct landforms. Additionally, we propose that different landforms will sustain different amounts of biodiversity and that high alpha and gamma diversity values are related to landform‐driven moisture availability whereas high beta diversity relates especially to landform‐specific microtopographic variation. Location: Rokua UNESCO Global Geopark area, Finland. Taxon: Vascular plants. Methods: We compare vascular plant species richness measures, Shannon’s and Simpson’s diversity indices, rarity‐weighted richness and local contribution to beta diversity at altogether three levels of biodiversity (alpha, beta and gamma) for different landforms. Landform information is compiled from aerial photos, spatial data layers and targeted field surveys. We compare results to control habitat (i.e. sites without any distinct landforms) within the study area. Results: Vascular plant diversity was higher on landforms than in control habitat. There was also notable variation between species diversity of different landforms. Moisture‐rich gullies and river shores were especially diverse at all three levels, whereas aapa mires hosted most unique species composition (highest beta diversity). Beta diversity patterns were rather comparable with alpha and gamma diversity patterns, which contradict our hypothesis. Main conclusions: This study quantitatively established a strong connection between terrestrial plant communities and multiple landforms. Our results highlighted the landform‐controlled variation in soil moisture, microclimate and microtopography in enhancing plant species diversity. Based on the results, we promote the inclusion of landform‐based geodiversity information in conservation management and in further biogeographical studies.

Metacommunity patterns and underlying processes in aquatic organisms have typically been studied within a drainage basin. We examined variation in the composition of six freshwater organismal groups across various drainage basins in Finland. We first modelled spatial structures within each drainage basin using Moran eigenvector maps. Second, we partitioned variation in community structure among three groups of predictors using constrained ordination: (1) local environmental variables, (2) spatial variables, and (3) dummy variable drainage basin identity. Third, we examined turnover and nestedness components of multiple-site beta diversity, and tested the best fit patterns of our datasets using the “elements of metacommunity structure” analysis. Our results showed that basin identity and local environmental variables were significant predictors of community structure, whereas within-basin spatial effects were typically negligible. In half of the organismal groups (diatoms, bryophytes, zooplankton), basin identity was a slightly better predictor of community structure than local environmental variables, whereas the opposite was true for the remaining three organismal groups (insects, macrophytes, fish). Both pure basin and local environmental fractions were, however, significant after accounting for the effects of the other predictor variable sets. All organismal groups exhibited high levels of beta diversity, which was mostly attributable to the turnover component. Our results showed consistent Clementsian-type metacommunity structures, suggesting that subgroups of species responded similarly to environmental factors or drainage basin limits. We conclude that aquatic communities across large scales are mostly determined by environmental and basin effects, which leads to high beta diversity and prevalence of Clementsian community types.

In recent years, there has been an increasing interest in the study of the spatial link between service providing areas (SPA) and service benefiting areas (SBA). Understanding the spatial link between SPAs and SBAs is essential when studying the ecosystem service delivery and the fulfilment of ecosystem service demand. However, far too little attention has been paid to the user movement related ecosystem services and where people should be geographically situated in order to benefit from these services. In the movement related services, benefiting areas are equal to providing areas and the spatial link from residential area to SPA is important. The spatial link is addressed through the concept of accessibility which determines the opportunity to move from the area where beneficiaries are located to areas where ecosystem services are produced. This study presents an accessibility approach to the ecosystem services research. Accessibility analyses offer an opportunity to identify the gap between the ecosystems’ potential to produce services and the actual usage possibilities of such services. We demonstrate the suitability of the method by using outdoor recreation and cultural heritage as examples of cultural ecosystem services that people actively want to reach. Accessibility was calculated using a geographical information system-based least-cost path analysis, which measures travel time by car between residential location and the nearest SPA via road network. The examples highlight that accessibility varies according to the ecosystem service and depends mostly on population distribution and travel possibilities. Our results demonstrate that the density of the analysed ecosystem service opportunities is higher near urban areas than elsewhere. The accessibility of different ecosystem services also depends on how much time people are willing to spend for reaching these services. Our study emphasised that, from a population perspective, accessibility analyses provide a powerful tool for illustrating the utilisation possibilities of spatially distributed ecosystem services. The accessibility approach offers great potential to assess the potential use of SPAs and respond to the need to develop a practical tool for ecosystem service research. It effectively shows, for example, the areas where the risk of overuse of ecosystem services is increased. Knowing about the regional differences in ecosystem service usage also gives background information for the decision-makers for drawing conclusions about how much and where it is sensible to invest in the maintenance of ecosystem services.

Species distributions are structured by regional and local determinants, which operate at multiple spatial and temporal scales. The purpose of our work was to distinguish the relative roles of local variables, climate, geographical location and post glaciation condition (i.e., delineation between supra- and subaquatic lakes during the post-glacial Ancylus Lake) in explaining variation in macrophyte community composition of all taxa, helophytes and hydrophytes. In addition, we investigated how these four explanatory variable groups affected macrophyte strategy groups based on Grime’s classification. Using partial linear regression and variation partitioning, we found that macrophyte communities are primarily filtered by local determinants together with regional characteristics at the studied spatial scale. We further evidenced that post glaciation condition indirectly influenced on local water quality variables, which in turn directly contributed to the macrophyte communities. We thus suggest that regional determinants interact with local-scale abiotic factors in explaining macrophyte community patterns and examining only regional or local factors is not sufficient for understanding how aquatic macrophyte communities are structured locally and regionally.

Geographical information systems (GIS)-based approaches support the traditional empirical assessment and monitoring measures to indicate large-scale environmental alterations caused by global change. We examined the relationship between cover patterns of emergent aquatic plants derived from the CORINE land cover database, and environmental gradients (i.e. lake, climate and land cover and soil variables) at four different buffer scales (50 m, 100 m, 300 m and 500 m) in 408 boreal lakes in Finland. Our main statistical methods were hierarchical partitioning (HP), generalized linear models (GLM) and variation partitioning (VP). Based on correlations and HP, the importance of different buffer zones varied between environmental variables in explaining the vegetation cover. Our study also demonstrated that landscape productivity variables (i.e. Normalized Difference Vegetation Index and Tasseled Cap Greenness) strongly structured the vegetation cover. These productivity variables proved to be proxies for agricultural land and nutrient leaching from terrestrial areas to water, but they were also associated with shading of riparian vegetation. In the VP, land cover and soil variable groups influenced emergent vegetation cover the most, followed by lake and, lastly, by climate variables. Our work emphasized the importance of spatial scale in structuring freshwater assemblages that should be taken into account in ecological research and bioassessment. The use of a single spatial scale may lead to flawed results on the relevance of various environmental gradients on freshwater assemblages. Moreover, our study showed that satellite-based productivity variables can be powerful proxies for the amount of nutrients eventually entering aquatic ecosystems from adjacent terrestrial areas. The GIS-based approach used here provided an ecologically sound and cost-efficient early-warning estimation of vegetation overgrowth in boreal lakes. This approach is applicable to other regions with similar land cover data sets.

Metacommunity paradigms are increasingly studied to explain how environmental control and spatial patterns determine variation in community composition. However, the relative importance of these patterns on biological assemblages among different habitats is not well known. We investigated the relative roles of local, catchment and spatial variables based on overland and watercourse distances in explaining the variation of community structure of lake and river macrophytes in two large river basins at two spatial extents (within and across river basins). Partial redundancy analysis was used to explore the share of variability in macrophyte communities attributable to local environmental conditions, catchment land cover and space (generated with Principle Coordinates of Neighbour Matrices). We found that local variables had the highest effect on both lake and river macrophyte communities, followed by catchment variables. Space had no or only marginal influence on the community structure regardless of used distance measure. Total phosphorus, conductivity and turbidity of the local variables contributed most for lake macrophytes, whereas pH and color had largest independent contribution for variation in river macrophytes. Size of catchment area and proportion of lakes and agriculture were the most important catchment variables in both habitats. The strong importance of environmental control suggests that both lake and river macrophyte communities are structured by species sorting. This finding gives support to the validity of assessment systems based on the European Water Framework Directive.

We investigated how land use at multiple scales affects functional macrophytes groups and ecological status index in the boreal region. We employed a variance partitioning analysis to quantify the relative role of lake characteristics, multiple-scaled land use (catchment, buffer zones of 100, 300 and 500 m), and space in explaining the composition and richness of functional macrophyte groups (emergent and submerged macrophytes and hydrophytes) and ecological status of macrophytes in 110 Finnish lakes. Partial redundancy analysis (community composition) and partial linear regression (richness and status index) revealed that macrophyte community composition, richness, and status index were mostly explained by the pure effect of lake characteristics, which dominated over space for most macrophyte variables. Land use adjacent to shoreline had a higher effect on emergent macrophytes and status index compared to the land use of the whole catchment. Our findings suggest that emergent macrophytes can indicate changes in water quality and hydro-morphology originated from the close vicinity of the littoral zone. Ecological quality assessment based on emergent macrophytes only is probably not sufficient, but including emergent species in the assessments is recommended, especially in the species-poor boreal region.

We assessed the relative roles of natural covariates, human disturbance (water quality and catchment land use) together with geography in driving variation in aquatic macrophyte community composition, richness and status among 101 lakes in southern and central Finland. In addition to all species together, we studied different growth forms (i.e. emergent and submerged macrophytes and aquatic bryophytes) separately. Partial redundancy analysis (taxonomic composition) and partial least-squares regression (species richness and status index) were employed to display the share of variability in macrophyte assemblages that was attributable to the environmental factors (both natural and human-affected) and the spatial filters generated through principal coordinates of neighbor matrices (PCNM). Macrophyte community composition, richness and status were explained by natural covariates, together with joint effects of human disturbance variables and space. The contributions of pure fractions of human disturbance and space were mostly modest, albeit variable among macrophyte groups and status indices. Alkalinity, historical distributions, colour, dynamic ratio and lake area were most important natural covariates for macrophytes. Of those variables influenced by human, macrophytes were mostly explained by conductivity, total phosphorus, turbidity and chlorophyll- a . Our results demonstrate, as expected, that macrophytes are dominantly affected by local environmental variables, whereas dispersal-related processes seem not to be important at regional extent. Response of macrophyte growth forms to environment and space, however, varied significantly. Community composition and richness of emergent macrophytes showed congruent response to natural covariates and human disturbance. Aquatic bryophytes, which are rarely studied along lake macrophytes, responded stronger than other growth forms to human disturbance. Contrary to our expectations, ecological indices were not affected by dispersal-related processes, but were mainly explained by natural covariates. This study is the first to investigate spatial patterns in aquatic macrophytes derived bioassessment. Geographical structuring of environmental variables and regional extent negatively affected indices, suggesting that ecological status assessment needs further development.

Aim Spatial extent is inherently related to the potential roles of the main mechanisms structuring metacommunities. We examined the effects of varying spatial extent (ecological province, region and subregion) on the environmental and spatial components of variation in lake macrophyte communities. We also studied these effects separately for three macrophyte functional groups. Location The US state of Minnesota. Methods We examined average and heterogeneity differences in macrophyte community composition and environmental variation among the subregions of Minnesota using canonical analysis of principal coordinates (CAP) and homogeneity of multivariate dispersion (PERMDISP), respectively. We further used partial redundancy analysis (pRDA) to decompose variation in macrophyte community composition between environmental variables and spatial location at each spatial extent and geographical region. Spatial variables were derived using principal coordinates of neighbour matrices (PCNM) analysis. Results CAP and PERMDISP analyses showed that the subregions differed both in average community composition and in the heterogeneity of community composition for all macrophyte taxa, for emergent and submerged macrophytes, but not for non-rooted macrophytes. We did not, however, find significant differences in overall environmental heterogeneity among the subregions. Variation partitioning using pRDAs showed that species sorting is more important than spatial processes for macrophytes, although these patterns were relatively weak. There was, however, much regional specificity, with the environmental and spatial fractions of community composition varying widely at different spatial extents, among different geographical regions and among functional groups. Contrary to our initial expectations, we did not find increasing spatial structuring and decreasing environmental control with increasing spatial extent. Main conclusions Our findings indicate that, in macrophyte metacommunities, the relative contribution of spatial processes and environmental control varies rather unpredictably with spatial extent and geographical region. Our findings are thus of importance in advancing metacommunity ecology by showing that drawing wide-ranging conclusions based on a single spatial extent or a single geographical region may be unwise.

We studied the relations of aquatic macrophyte communities, ecological quality metrics and water quality to lake characteristics and to land use defined at multiple spatial scales (three lake marginal zones (LMZs) of contrasting width and the whole topographic catchment) in 110 boreal lakes in Finland using ordination and generalised additive modelling. Variation in macrophyte community composition of our lakes was related to water quality, land use, latitude and lake morphology. Relationships of macrophyte status metrics – the proportion of type-specific species (TT50), Percent Model Affinity (PMA) and Trophic Index (TI) – to land use were scale dependent. The proportion of agricultural and urban land use adjacent to the lake shoreline (LMZs of 300 m and 500 m) had a more significant effect on the metrics compared to land use at the catchment scale. The scale-related pattern in the effect of land use was not congruent among metrics and the metrics also showed different indications of deterioration. Lower sensitivity to the measured disturbance of the pressure sensitive metric TI compared to the other metrics (TT50 and PMA) based on general type-specific species composition was presumably due to a lack of helophytes in the species pool used, and due to invariable indicator values across lake types. Moreover, TI varied more with hydromorphology related natural gradients than TT50 and PMA. This could be due to lack of habitat for eutrophication sensitive submerged indicator species in shallow lakes. Our study demonstrates that choosing the most influential catchment scale for the evaluation and mitigation of anthropogenic impacts on lacustrine biota, may be significant in conserving and restoring the ecological quality of freshwaters, as required by, e.g. , the European Water Framework Directive.

Land use planning was modernised in Finland at the turn of the millennium. Simultaneously, river basin planning based on the EU Water Framework Directive (WFD) markedly changed the environmental goals for water bodies. The WFD’s main objective is to achieve good ecological status in all water bodies by the year 2015. To meet this objective, traditional water protection measures are planned for preventing and reducing eutrophication in water bodies suffering from diffuse pollution, which is the major obstacle reaching a good ecological status in Finnish waterbodies. The need for an alternative and broader selection of water protection methods is nevertheless evident due to the low efficiency and high costs of traditional means. We have studied the potential to integrate water protection objectives into land use planning and found several interfaces between river basin planning and land use planning in Finland. The emphasis is on the prevention of diffuse pollution originating from different land use practices. Our study highlights the linkages between the two planning processes and proposes ideas for further integration by drawing on the experiences gained from two pilot studies focusing mainly on water protection measures, which provide support to achieving the ecological objectives of WFD.

Aim Aquatic–terrestrial ecotones are vulnerable to climate change, and degradation of the emergent aquatic macrophyte zone would have severe ecological consequences for freshwater, wetland and terrestrial ecosystems. Our aim was to uncover future changes in boreal emergent aquatic macrophyte zones by modelling the occurrence and percentage cover of emergent aquatic vegetation under different climate scenarios in Finland by the 2050s. Location Finland, northern Europe. Methods Data derived from different GIS sources were used to estimate future emergent aquatic macrophyte distributions in all catchments in Finland (848 in total). We used generalized additive models (GAM) with a full stepwise selection algorithm and Akaike information criterion to explore the main environmental determinates (climate and geomorphology) of emergent aquatic macrophyte distributions, which were derived from the national subclass of CORINE land-cover classification. The accuracy of the distribution models (GAMs) was cross-validated, using percentage of explained deviance and the area under the curve derived from the receiver-operating characteristic plots. Results Our results indicated that emergent aquatic macrophytes will expand their distributions northwards from the current catchments and percentage cover will increase in all of the catchments in all climate scenarios. Growing degree-days was the primary determinant affecting distributions of emergent aquatic macrophytes. Inclusion of geomorphological variables clearly improved model performance in both model exercises compared with pure climate variables. Main conclusions Emergent aquatic macrophyte distributions will expand due to climate change. Many emergent aquatic plant species have already expanded their distributions during the past decades, and this process will continue in the years 2051–80. Emergent aquatic macrophytes pose an increasing overgrowth risk for sensitive macrophyte species in boreal freshwater ecosystems, which should be acknowledged in management and conservation actions. We conclude that predictions based on GIS data can provide useful ‘first-filter’ estimates of changes in aquatic–terrestrial ecotones.

A new classifi cation system compliant with the EU's Water Framework Directive (WFD) was employed to assess the ecological status of six small forest lakes differently affected by catchment forestry. The classifi ca- tion is based on metrics involving four biological quality elements (phytoplankton, aquatic macrophytes, benthic macroinvertebrates and fi sh) and their nationally defi ned reference and class boundary values. The classifi cation evidence obtained from single metrics and quality elements, on the "One-out, All-out" principle, which establishes status according to the weakest measure, was compared with the integrated classifi cation system used in the Finnish national status assessments. The national classifi cation system harmonizes the individual measures by scoring and expresses overall status class as a median score across quality elements. In addition, because the amount of refer- ence data for this lake type was limited, palaeolimnological analyses were conducted to determine lake-specifi c changes during past decades. The results demonstrated that individual metrics and quality elements may result in highly variable classifi cations, and consequently pessimistic ecological estimates, if the One-out, All-out principle is applied. It is concluded that the integration of evidence from multiple quality elements evens out their variability and results in more realistic status assessments in small humic boreal forest lakes.

Regional faunas are structured by historical, spatial and environmental factors. We studied large-scale variation in four ecologically different beetle groups (Coleoptera: Dytiscidae, Carabidae, Hydrophiloidea, Cerambycidae) along climate, land cover and geographical gradients, examined faunal breakpoints in relation to environmental variables, and investigated the best fit pattern of assemblage variation (i.e. randomness, checkerboards, nestedness, evenly spaced, Gleasonian, Clementsian). We applied statistical methods typically used in the analysis of local ecological communities to provide novel insights into faunal compositional patterns at large spatial grain and geographical extent. We found that spatially structured variation in climate and land cover accounted for most variation in each beetle group in partial redundancy analyses, whereas the individual effect of each explanatory variable group was generally much less important in accounting for variation in provincial species composition. We also found that climate variables were most strongly associated with faunal breakpoints, with temperature-related variables alone accounting for about 20% of variation at the first node of multivariate regression tree for each beetle group. The existence of faunal breakpoints was also shown by the 'elements of faunal structure' analyses, which suggested Clementsian gradients across the provinces, that is, that there were two or more clear groups of species responding similarly to the underlying ecological gradients. The four beetle groups showed highly similar biogeographical patterns across our study area. The fact that temperature was related to faunal breakpoints in the species composition of each beetle group suggests that climate sets a strong filter to the distributions of species at this combination of spatial grain and spatial extent. This finding held true despite the ecological differences among the four beetle groups, ranging from fully aquatic to fully terrestrial and from herbivorous to predaceous species. The existence of Clementsian gradients may be a common phenomenon at large scales, and it is likely to be caused by crossing multiple species pools determined by climatic and historical factors on the distributions of species.

Documenting the patterns and potential associated processes of ancient biotas has always been a central challenge in palaeontology. Over recent decades, intense debate has focused on the organization of dinosaur-dominated communities, yet no general consensus has been reached on how these communities were organized in a spatial context. Here, we used analytical routines typically applied in metacommunity ecology to provide novel insights into dinosaurian distributions across the latest Cretaceous of North America. To do this, we combined fossil occurrences with functional, phylogenetic and palaeoenvironmental modelling, and adopted the perspective that more reasonable conclusions on palaeoecological reconstructions can be gained from studies that consider the organization of biotas along ecological gradients at multiple spatial scales. Our results showed that dinosaurs were restricted in range to different parts of the Hell Creek Formation, prompting the recognition of discrete and compartmentalized faunal areas during the Maastrichtian at fine-grained scales, whereas taxa with the broadest ranges included those with narrower distributions when combining data from various geological formations across the Western Interior of North America. Although groups of dinosaurs had coincident range boundaries, their communities responded to multiple ecologically-important gradients when compensating for differences in sampling effort. Metacommunity structures of both ornithischians and theropods were correlated with climatic barriers and potential trophic relationships between herbivores and carnivores, thereby suggesting that dinosaurian faunas were shaped by physiological constraints, limited food resources abundance, and a combination of bottom-up and top-down forces across multiple spatial grains and extents.