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Support for the “biotic resistance hypothesis,” that species-rich communities are more successful at resisting invasion by exotic species than are species-poor communities, has long been debated. It has been argued that native–exotic richness relationships (NERR) are negative at small spatial scales and positive at large scales, but evidence for the role of spatial scale on NERR has been contradictory. However, no formal quantitative synthesis has previously examined whether NERR is scale-dependent across multiple studies, and previous studies on NERR have not distinguished spatial grain and extent, which may drive very different ecological processes. We used a global systematic review and hierarchical mixed-effects metaanalysis to provide a comprehensive quantitative assessment of the patterns of NERR over a range of spatial grain sizes and spatial extents, based on 204 individual cases of observational (non-experimental) NERRs from 101 publications. We show that NERR was indeed highly scale dependent across studies and increased with the log of grain size. However, mean NERR was not negative at any grain size, although there was high heterogeneity at small grain sizes. We found no clear patterns of NERR across different spatial extents, suggesting that extent plays a less important role in determining NERR than does grain, although there was a complex interaction between extent and grain size. Almost all studies on NERR were conducted in North America, western Europe, and a few other regions, with little information on tropical or Arctic regions. We did find that NERR increased northward in temperate regions and also varied with longitude. We discuss possible explanations for the patterns we found, and caution that our results do not show that invasive species are benign or have no negative consequences for biodiversity preservation. This study represents the first global quantitative analysis of scale-based NERR, and casts doubt on the existence of an “invasion paradox” of negative NERR at small scales and positive correlations at large scales in non-experimental studies.

This study examines the complex feedback mechanisms that regulate a positive relationship between species richness and productivity in a longleaf pine-wiregrass woodland. Across a natural soil moisture gradient spanning wet-mesic to xeric conditions, two large scale manipulations over a 10-yr period were used to determine how limiting resources and fire regulate plant species diversity and productivity at multiple scales. A fully factorial experiment was used to examine productivity and species richness responses to N and water additions. A separate experiment examined standing crop and richness responses to N addition in the presence and absence of fire. Specifically, these manipulations addressed the following questions: (1) How do N and water addition influence annual aboveground net primary productivity of the midstory/overstory and ground cover? (2) How do species richness responses to resource manipulations vary with scale and among functional groups of ground cover species? (3) How does standing crop (including overstory, understory/midstory, and ground cover components) differ between frequently burned and fire excluded plots after a decade without fire? (4) What is the role of fire in regulating species richness responses to N addition? This long-term study across a soil moisture gradient provides empirical evidence that species richness and productivity in longleaf pine woodlands are strongly regulated by soil moisture. After a decade of treatment, there was an overall species richness decline with N addition, an increase in richness of some functional groups with irrigation, and a substantial decline in species richness with fire exclusion. Changes in species richness in response to treatments were scale-dependent, occurring primarily at small scales (≤10 m²). Further, with fire exclusion, standing crop of ground cover decreased with N addition and non-pine understory/midstory increased in wet-mesic sites. Non-pine understory/midstory standing crop increased in xeric sites with fire exclusion, but there was no influence of N addition. This study highlights the complexity of interactions among multiple limiting resources, frequent fire, and characteristics of dominant functional groups that link species richness and productivity.

Mountains are key features of the Earth’s surface and host a substantial proportion of the world’s species. However, the links between the evolution and distribution of biodiversity and the formation of mountains remain poorly understood. Here, we integrate multiple datasets to assess the relationships between species richness in mountains, geology and climate at global and regional scales. Specifically, we analyse how erosion, relief, soil and climate relate to the geographical distribution of terrestrial tetrapods, which include amphibians, birds and mammals. We find that centres of species richness correlate with areas of high temperatures, annual rainfall and topographic relief, supporting previous studies. We unveil additional links between mountain-building processes and biodiversity: species richness correlates with erosion rates and heterogeneity of soil types, with a varying response across continents. These additional links are prominent but under-explored, and probably relate to the interplay between surface uplift, climate change and atmospheric circulation through time. They are also influenced by the location and orientation of mountain ranges in relation to air circulation patterns, and how species diversification, dispersal and refugia respond to climate change. A better understanding of biosphere–lithosphere interactions is needed to understand the patterns and evolution of mountain biodiversity across space and time.

Aim: In urbanized areas, exotic invasions, native extinctions, and the alteration of habitats and natural processes drive homogenization, which is a form of biotic impoverishment This study examines whether urbanization and flooding induce homogenization of herbaceous communities in riparian forests and quantifies the relationships between taxonomic and functional β-diversity. Location: Montreal, Quebec, Canada. Methods: Inventories were conducted in 56 riparian forests. Taxonomic and functional p-diversity were calculated as between-site similarities in species or trait composition for three levels of urbanization and flooding. Differences among the disturbance levels were compared using tests for homogeneity in multivariate dispersions. We quantified the correlation between local species richness, exotic proportion, taxonomic and functional β-diversity. We also partitioned taxonomic p-diversity into species turnover and richness difference. Results: Urbanization led to taxonomic and functional differentiation, while increased flooding led to taxonomic and functional homogenization. We found a significant correlation between taxonomic and functional p-diversity. Changes in β-diversity were associated with species and trait turnover among both urbanization and flood levels, and with changes in species richness. Differentiation was associated with low species richness, and homogenization with high species richness. Exotic invasions tended to favour differentiation, but only at a low urbanization level. Main Conclusions: The effect of urbanization on plant diversity in riparian forests was twofold: first, it directly induced taxonomic and functional differentiation through its effect on species loss and turnover (higher β-diversity at high urbanization level); second, differentiation was indirectly favoured through the reduction in flooding (higher β-diversity at low flood level). Taxonomic and functional p-diversity followed similar patterns, likely because species invasions and extinctions are not random, but are related to species traits. Our results underline the need to move our focus from exotic species to the true underlying factors of biodiversity loss and homogenization, notably land use changes and human disturbances.

To predict the ecological consequences of biodiversity loss, researchers have spent much time and effort quantifying how biological variation affects the magnitude and stability of ecological processes that underlie the functioning of ecosystems. Here we add to this work by looking at how biodiversity jointly impacts two aspects of ecosystem functioning at once: (1) the production of biomass at any single point in time (biomass/area or biomass/volume), and (2) the stability of biomass production through time (the CV of changes in total community biomass through time). While it is often assumed that biodiversity simultaneously enhances both of these aspects of ecosystem functioning, the joint distribution of data describing how species richness regulates productivity and stability has yet to be quantified. Furthermore, analyses have yet to examine how diversity effects on production covary with diversity effects on stability. To overcome these two gaps, we reanalyzed the data from 34 experiments that have manipulated the richness of terrestrial plants or aquatic algae and measured how this aspect of biodiversity affects community biomass at multiple time points. Our reanalysis confirms that biodiversity does indeed simultaneously enhance both the production and stability of biomass in experimental systems, and this is broadly true for terrestrial and aquatic primary producers. However, the strength of diversity effects on biomass production is independent of diversity effects on temporal stability. The independence of effect sizes leads to two important conclusions. First, while it may be generally true that biodiversity enhances both productivity and stability, it is also true that the highest levels of productivity in a diverse community are not associated with the highest levels of stability. Thus, on average, diversity does not maximize the various aspects of ecosystem functioning we might wish to achieve in conservation and management. Second, knowing how biodiversity affects productivity gives no information about how diversity affects stability (or vice versa). Therefore, to predict the ecological changes that occur in ecosystems after extinction, we will need to develop separate mechanistic models for each independent aspect of ecosystem functioning.

Understanding the factors that regulate the functioning of our ecosystems in response to environmental changes can help to maintain the stable provisioning of ecosystem services to mankind. This is especially relevant given the increased variability of environmental conditions due to human activities. In particular, maintaining a stable production and plant biomass during the growing season (intra‐annual stability) despite pervasive and directional changes in temperature and precipitation through time can help to secure food supply to wild animals, livestock, and humans. Here, we conducted a 29‐year field observational study in a temperate grassland to explore how the intra‐annual stability of primary productivity is influenced by biotic and abiotic variables through time. We found that intra‐annual precipitation variability in the growing season indirectly influenced the community intra‐annual biomass stability by its negative effect on intra‐annual species asynchrony. While the intra‐annual temperature variability in the growing season indirectly altered community intra‐annual biomass stability through affecting the intra‐annual species richness. At the same time, although the intra‐annual biomass stability of the dominant species and the dominant functional group were insensitive to climate variability, they also promoted the stable community biomass to a certain extent. Our results indicate that ongoing intra‐annual climate variability affects community intra‐annual biomass stability in the temperate grassland, which has important theoretical significance for us to take active measures to deal with climate change. Intra‐annual growing season precipitation variability and temperature variability indirectly influenced the community intra‐annual biomass stability by negative effect on intra‐annual species asynchrony and intra‐annual species richness, respectively. ; Intra‐annual biomass stability of dominant species and dominant functional group were insensitive to climate variability, they also promoted the stable community biomass to a certain extent.

Networks of protected areas are fundamental for biodiversity conservation, but many factors determine their conservation efficiency. In particular, on top of other human-driven disturbances, invasions by non-native species can cause habitat and biodiversity loss. Jointly understanding what drives patterns of plant diversity and of non-native species in protected areas is therefore a priority. We tested whether the richness and composition of native and non-native plant species within a network of protected areas follow similar patterns across spatial scales. Specifically, we addressed three questions: (a) what is the degree of congruence in species richness between native and non-native species? (b) do changes in the composition of non-native species across ecological gradients reflect a similar turnover of native species along the same gradients ? (c) what are the main environmental and human disturbance drivers controlling species richness in these two groups of species? Species richness and composition of native and non-native plant species were compared at two spatial scales: the plot scale (10 m × 10 m) and the Protected Area scale (PA). In addition, we fit Generalized Linear Models to identify the most important drivers of native and non-native species richness at each scale, focusing on environmental conditions (climate, topography) and on the main sources of human disturbance in the area (land use and roads). We found a significant positive correlation between the turnover of native and non-native species composition at both plot and PA scales, whereas their species richness was only correlated at the larger PA scale. The lack of congruence between the richness of native and non-native species at the plot scale was likely driven by differential responses to fine scale environmental factors, with non-natives favoring drier climates and milder slopes (climate and slope). In addition, more non-native species were found closer to road-ways in the reserve network. In contrast, the congruence in the richness of native and non-native species at the broader PA scale was mainly driven by the common influence of PA area, but also by similar responses of the two groups of species to climatic heterogeneity. Thus, our study highlights the strong spatial dependence of the relationship between native and non-native species richness and of their responses to environmental variation. Taken together, our results suggest that within the study region the introduction and establishment of non-native species would be more likely in warmer and dryer areas, with high native species richness at large spatial scale but intermediate levels of anthropogenic disturbances and mild slope inclinations and elevation at fine scale. Such an exhaustive understanding of the factors that influence the spread of non-native species, especially in networks of protected areas is crucial to inform conservation managers on how to control or curb non-native species.

Aims Saprophytic fungi are important agents of soil mineralization and carbon cycling. Their community structure is known to be affected by soil conditions such as organic matter and pH. However, the effect of plant species, whose roots provide the litter input into the soil, on the saprophytic fungal community is largely unknown. Methods We examined the saprophytic fungi in a grassland biodiversity experiment with eight plant species belonging to two functional groups (grasses and forbs), combining DNA extraction from plant roots, next-generation sequencing and literature research. Results We found that saprophyte richness increased with plant species richness, but plant functional group richness was the best predictor. Plant functional group was also the main factor driving fungal saprophytic community structure. This effect was correlated with differences in root lignin content and C:N ratio between grasses and forbs. In monocultures, root traits and plant functional group type explained 16% of the variation in community structure. The saprophyte taxa detected in mixed plant communities were to a large extent subsets of those found in monocultures. Conclusions Our work shows that the richness and community structure of the root-associated saprophytic fungi can largely be predicted by plant functional groups and their associated root traits. This means that the effects of plant diversity on ecosystem functions such as litter decomposition may also be predictable using information on plant functional groups in grasslands.

Tropical rainforests (TRFs) harbour almost half of the world's vascular plant species diversity while covering only about 6-7% of land. However, why species richness varies amongst the Earth's major TRF regions remains poorly understood. Here we investigate the evolutionary processes shaping continental species richness disparities of the pantropical, epiphytic and mostly TRF-dwelling orchid mega-genus Bulbophyllum (c. 1948 spp. in total) using diversification analyses based on a time-calibrated molecular phylogeny (including c. 45-50% spp. each from Madagascar, Africa, Neotropics, and 8.4% from the Asia-Pacific region), coupled with ecological niche modelling (ENM) of geographic distributions under present and past (Last Glacial Maximum; LGM) conditions. Our results suggest an early-to-late Miocene scenario of 'out-of-Asia-Pacific' origin and progressive, dispersal-mediated diversification in Madagascar, Africa and the Neotropics, respectively. Species richness disparities amongst these four TRF lineages are best explained by a time-for-speciation (i.e. clade age) effect rather than differences in net diversification or diversity-dependent diversification due to present or past spatial-bioclimatic limits. For each well-sampled lineage (Madagascar, Africa, Neotropics), we inferred high rates of speciation and extinction over time (i.e. high species turnover), yet with the origin of most extant species falling into the Quaternary. In contrast to predictions of classical 'glacial refuge' theories, all four lineages experienced dramatic range expansions during the LGM. As the Madagascan, African and Neotropical lineages display constant-rate evolution since their origin (early-to-mid-Miocene), Quaternary environmental change might be a less important cause of their high species turnover than intrinsic features generally conferring rapid population turnover in tropical orchids (e.g., epiphytism, specialization on pollinators and mycorrhizal fungi, wind dispersal). Nonetheless, climate-induced range fluctuations during the Quaternary could still have played an influential role in the origination and extinction of Bulbophyllum species in those three, if not in all four TRF regions.

Understanding the relationship of soil carbon storage and species diversity in grasslands can provide insights into managing these ecosystems. We studied relationships among soil C and plant species richness within ~ 9700 ha of grasslands in Colorado, US. Using 141 grassland transects, we tested how soil C was related to plant species richness, grassland type, soil texture, and prairie dog presence. Soil C was significantly, positively related to plant species richness, while native perennial graminoid species richness exhibited an even stronger positive relationship. However, the relationship of soil C and plant richness was not found in all three grassland types studied, but instead was unique to the most common grassland type, mixed grass prairie, and absent from both xeric tallgrass and mesic tallgrass prairie. The presence of a single indicator species, Andropogon gerardii , showed a significant, positive relationship with soil carbon. Our best possible model explained 45% of the variance in soil C using species richness, grassland type, and their interaction. Surprisingly, soil C was negatively related to soil clay, suggesting that surface clays amplify evaporation and water runoff rather than protecting soil organic matter from decomposition. Soil C was negatively related to prairie dog presence, suggesting that prairie dogs do not enhance soil carbon sequestration; in fact, prairie dog occupied sites had significantly lower soil C, likely related to loss of topsoil from prairie dog colonies. Our results suggest that management for species richness provides the co-benefit of soil C storage, and high clay and prairie dog disturbance compromises both.