Synthesis reveals approximately balanced biotic differentiation and homogenization.

It is commonly thought that the biodiversity crisis includes widespread declines in the spatial variation of species composition, called biotic homogenization. Using a typology relating homogenization and differentiation to local and regional diversity changes, we synthesize patterns across 461 metacommunities surveyed for 10 to 91 years, and 64 species checklists (13 to 500+ years). Across all datasets, we found that no change was the most common outcome, but with many instances of homogenization and differentiation. A weak homogenizing trend of a 0.3% increase in species shared among communities/year on average was driven by increased numbers of widespread (high occupancy) species and strongly associated with checklist data that have longer durations and large spatial scales. At smaller spatial and temporal scales, we show that homogenization and differentiation can be driven by changes in the number and spatial distributions of both rare and common species. The multiscale perspective introduced here can help identify scale-dependent drivers underpinning biotic differentiation and homogenization.

Disproportionate declines of formerly abundant species underlie insect loss.

Studies have reported widespread declines in terrestrial insect abundances in recent years1-4, but trends in other biodiversity metrics are less clear-cut5-7. Here we examined long-term trends in 923 terrestrial insect assemblages monitored in 106 studies, and found concomitant declines in abundance and species richness. For studies that were resolved to species level (551 sites in 57 studies), we observed a decline in the number of initially abundant species through time, but not in the number of very rare species. At the population level, we found that species that were most abundant at the start of the time series showed the strongest average declines (corrected for regression-to-the-mean effects). Rarer species were, on average, also declining, but these were offset by increases of other species. Our results suggest that the observed decreases in total insect abundance2 can mostly be explained by widespread declines of formerly abundant species. This counters the common narrative that biodiversity loss is mostly characterized by declines of rare species8,9. Although our results suggest that fundamental changes are occurring in insect assemblages, it is important to recognize that they represent only trends from those locations for which sufficient long-term data are available. Nevertheless, given the importance of abundant species in ecosystems10, their general declines are likely to have broad repercussions for food webs and ecosystem functioning.

Unveiling global species abundance distributions.

Whether most species are rare or have some intermediate abundance is a long-standing question in ecology. Here, we use more than one billion observations from the Global Biodiversity Information Facility to assess global species abundance distributions (gSADs) of 39 taxonomic classes of eukaryotic organisms from 1900 to 2019. We show that, as sampling effort increases through time, the shape of the gSAD is unveiled; that is, the shape of the sampled gSAD changes, revealing the underlying gSAD. The fraction of species unveiled for each class decreases with the total number of species in that class and increases with the number of individuals sampled, with some groups, such as birds, being fully unveiled. The best statistical fit for almost all classes was the Poisson log-normal distribution. This strong evidence for a universal pattern of gSADs across classes suggests that there may be general ecological or evolutionary mechanisms governing the commonness and rarity of life on Earth.

Widespread shifts in body size within populations and assemblages.

Biotic responses to global change include directional shifts in organismal traits. Body size, an integrative trait that determines demographic rates and ecosystem functions, is thought to be shrinking in the Anthropocene. Here, we assessed the prevalence of body size change in six taxon groups across 5025 assemblage time series spanning 1960 to 2020. Using the Price equation to partition this change into within-species body size versus compositional changes, we detected prevailing decreases in body size through time driven primarily by fish, with more variable patterns in other taxa. We found that change in assemblage composition contributes more to body size changes than within-species trends, but both components show substantial variation in magnitude and direction. The biomass of assemblages remains quite stable as decreases in body size trade off with increases in abundance.

Regional occupancy increases for widespread species but decreases for narrowly distributed species in metacommunity time series.

While human activities are known to elicit rapid turnover in species composition through time, the properties of the species that increase or decrease their spatial occupancy underlying this turnover are less clear. Here, we used an extensive dataset of 238 metacommunity time series of multiple taxa spread across the globe to evaluate whether species that are more widespread (large-ranged species) differed in how they changed their site occupancy over the 10-90 years the metacommunities were monitored relative to species that are more narrowly distributed (small-ranged species). We found that on average, large-ranged species tended to increase in occupancy through time, whereas small-ranged species tended to decrease. These relationships were stronger in marine than in terrestrial and freshwater realms. However, in terrestrial regions, the directional changes in occupancy were less extreme in protected areas. Our findings provide evidence for systematic decreases in occupancy of small-ranged species, and that habitat protection could mitigate these losses in the face of environmental change.

Emerging technologies revolutionise insect ecology and monitoring.

Insects are the most diverse group of animals on Earth, but their small size and high diversity have always made them challenging to study. Recent technological advances have the potential to revolutionise insect ecology and monitoring. We describe the state of the art of four technologies (computer vision, acoustic monitoring, radar, and molecular methods), and assess their advantages, current limitations, and future potential. We discuss how these technologies can adhere to modern standards of data curation and transparency, their implications for citizen science, and their potential for integration among different monitoring programmes and technologies. We argue that they provide unprecedented possibilities for insect ecology and monitoring, but it will be important to foster international standards via collaboration.

Long-term abundance trends of insect taxa are only weakly correlated.

Changes in the abundances of animals, such as with the ongoing concern about insect declines, are often assumed to be general across taxa. However, this assumption is largely untested. Here, we used a database of assemblage-wide long-term insect and arachnid monitoring to compare abundance trends among co-occurring pairs of taxa. We show that 60% of co-occurring taxa qualitatively showed long-term trends in the same direction-either both increasing or both decreasing. However, in terms of magnitude, temporal trends were only weakly correlated (mean freshwater r = 0.05 (±0.03), mean terrestrial r = 0.12 (±0.09)). The strongest correlation was between trends of beetles and those of moths/butterflies (r = 0.26). Overall, even though there is some support for directional similarity in temporal trends, we find that changes in the abundance of one taxon provide little information on the changes of other taxa. No clear candidate for umbrella or indicator taxa emerged from our analysis. We conclude that obtaining a better picture of changes in insect abundances will require monitoring of multiple taxa, which remains uncommon, especially in the terrestrial realm.

InsectChange: a global database of temporal changes in insect and arachnid assemblages.

Insects are the most ubiquitous and diverse group of eukaryotic organisms on Earth, forming a crucial link in terrestrial and freshwater food webs. They have recently become the subject of headlines because of observations of dramatic declines in some places. Although there are hundreds of long-term insect monitoring programs, a global database for long-term data on insect assemblages has so far remained unavailable. In order to facilitate synthetic analyses of insect abundance changes, we compiled a database of long-term (≥10 yr) studies of assemblages of insects (many also including arachnids) in the terrestrial and freshwater realms. We searched the scientific literature and public repositories for data on insect and arachnid monitoring using standardized protocols over a time span of 10 yr or longer, with at least two sampling events. We focused on studies that presented or allowed calculation of total community abundance or biomass. We extracted data from tables, figures, and appendices, and, for data sets that provided raw data, we standardized trapping effort over space and time when necessary. For each site, we extracted provenance details (such as country, state, and continent) as well as information on protection status, land use, and climatic details from publicly available GIS sources. In all, the database contains 1,668 plot-level time series sourced from 165 studies with samples collected between 1925 and 2018. Sixteen data sets provided here were previously unpublished. Studies were separated into those collected in the terrestrial realm (103 studies with a total of 1,053 plots) and those collected in the freshwater realm (62 studies with 615 plots). Most studies were from Europe (48%) and North America (29%), with 34% of the plots located in protected areas. The median monitoring time span was 19 yr, with 12 sampling years. The number of individuals was reported in 129 studies, the total biomass was reported in 13 studies, and both abundance and biomass were reported in 23 studies. This data set is published under a CC-BY license, requiring attribution of the data source. Please cite this paper if the data are used in publications, and respect the licenses of the original sources when using (part of) their data as detailed in Metadata S1: Table 1.

Response to Comment on "Meta-analysis reveals declines in terrestrial but increases in freshwater insect abundances".

Desquilbet et al take issue with our data inclusion criteria and make several other dubious claims regarding data processing, analysis, and interpretation. Most of their concerns stem from disagreement on data inclusion criteria and analysis, misunderstanding of our goals, and unrealistic expectations. We maintain that our synthesis provides a state-of-the-art analysis of patterns of trends in insect abundances.

Synchrony matters more than species richness in plant community stability at a global scale.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

Meta-analysis reveals declines in terrestrial but increases in freshwater insect abundances.

Recent case studies showing substantial declines of insect abundances have raised alarm, but how widespread such patterns are remains unclear. We compiled data from 166 long-term surveys of insect assemblages across 1676 sites to investigate trends in insect abundances over time. Overall, we found considerable variation in trends even among adjacent sites but an average decline of terrestrial insect abundance by ~9% per decade and an increase of freshwater insect abundance by ~11% per decade. Both patterns were largely driven by strong trends in North America and some European regions. We found some associations with potential drivers (e.g., land-use drivers), and trends in protected areas tended to be weaker. Our findings provide a more nuanced view of spatiotemporal patterns of insect abundance trends than previously suggested.

Rewilding with large herbivores: Positive direct and delayed effects of carrion on plant and arthropod communities.

Carrion of large animals is an extremely nutrient rich, ephemeral resource that is essential for many species, but is scarce in the anthropogenic Western-European landscape due to legislative restrictions. Rewilding, a novel conservation strategy that aims at restoring natural processes with minimal human intervention, is increasing in popularity and could lead to increased carrion availability in the landscape. It is therefore important to understand the effects of carrion on biodiversity. We investigated the direct and delayed (five months) effects of red deer (Cervus elaphus) carcasses on plants and arthropods in the Oostvaardersplassen, the Netherlands, one of the oldest rewilding sites in Europe. Specifically, we tested whether carrion has a positive direct effect on the abundances and diversity of various arthropod functional groups, as well as a delayed effect on the vegetation and arthropods through the increased nutrient availability. During the active decomposition stage in spring, we, not surprisingly, observed higher abundances of carrion associated species (scavengers and their specialized predators) at the carrion sites than at control sites without carrion, but no higher abundances of predators or detritivores. In late summer, after near-complete decomposition, plant biomass was five times higher, and nutritional plant quality (C:N ratio) was higher at the carrion sites than at the control sites. Arthropod abundance and diversity were also manifold higher, owing to higher numbers of herbivorous and predatory species. Regression analysis showed that abundances of herbivores and detritivores were positively related to plant biomass, and predator abundances were positively related to abundances of herbivores and detritivores, suggesting bottom-up effects propagating through the food chain. Our results show that even in a naturally nutrient-rich ecosystem like the Oostvaardersplassen, carrion can have strong positive effects on local plant biomass and nutritional quality and arthropod abundances, lasting the whole growing season. We found evidence that these effects were first directly caused by the presence of carrion, and later by the enhanced nutrient availability in the soil. This highlights the importance of the indirect pathways by which carrion can structure arthropod communities.

Functional differences stabilize beetle communities by weakening interspecific temporal synchrony.

The temporal stability of communities is essential for the maintenance of ecosystem functioning across trophic levels. The stabilizing effect of biodiversity is, among other factors, modulated by the level of synchrony in population fluctuations among the species in the community. What drives community synchrony, however, remains largely unclear. Community synchrony can be affected by external drivers such as disturbances, but also by the properties of the community. Species with different ecological strategies should fluctuate less synchronously than more similar species; thus, an increase in diversity of ecological strategies should decrease synchrony, and increase the stability of the community. Here, using an exceptionally large data set of ground beetle trappings in Dutch heathlands (~370,000 individuals in 19 plots, each sampled between 9 and 36 yr), we assess the drivers of community stability and synchrony, and their relationship with disturbance, species richness, and functional diversity (FD). We found no effect of disturbance (fire and topsoil removal) on community stability or synchrony, probably because of unpredictable patterns of increase or decrease of the populations. Community synchrony was overall positive, giving more support for independent and positive correlation between species than for compensatory dynamics. Synchrony decreased with increasing FD, but not with species richness. Supporting this, we found that the more species pairs differ in their traits, the less synchronously their populations fluctuate, where 74% of all pairs showed no significant correlation. Significant positive synchrony (19% of species pairs) was concentrated among pairs with low trait dissimilarity, and the 7% of pairs with significant negative temporal correlation showed no relation with pairwise functional dissimilarity. The stabilizing effect of FD via decreased synchrony supports largely untested theoretical expectations that an increased diversity of functional strategies in a community will have a stabilizing effect on community abundance. We hypothesize that because competition is low in this community, the stabilizing effect of FD reflects interspecific variation in responses to environmental fluctuations rather than competition.

Rewilding complex ecosystems.

The practice of rewilding has been both promoted and criticized in recent years. Benefits include flexibility to react to environmental change and the promotion of opportunities for society to reconnect with nature. Criticisms include the lack of a clear conceptualization of rewilding, insufficient knowledge about possible outcomes, and the perception that rewilding excludes people from landscapes. Here, we present a framework for rewilding that addresses these concerns. We suggest that rewilding efforts should target trophic complexity, natural disturbances, and dispersal as interacting processes that can improve ecosystem resilience and maintain biodiversity. We propose a structured approach to rewilding projects that includes assessment of the contributions of nature to people and the social-ecological constraints on restoration.

Larval and phenological traits predict insect community response to mowing regime manipulations.

For the restoration of biodiversity in agricultural grasslands, it is essential to understand how management acts as an ecological filter on the resident species. Mowing constitutes such a filter: only species that possess functional traits enabling them to withstand its consequences can persist in the community. We investigated how the timing of mowing modulates this filtering effect for insects. We predicted that two traits drive species responses. Species with larval development within the meadow vegetation will suffer more from mowing than species whose larvae develop in or on the ground, or outside the meadows, while species with a later phenology should benefit from later mowing. We conducted a five-year experiment, replicated at 12 sites across the Swiss lowlands, applying three different mowing regimes to low-intensity hay meadows: (1) first cut of the year not earlier than 15 June (control regime); (2) the first cut delayed until 15 July; and (3) leaving an uncut grass refuge on 10-20% of the meadow area (after earliest first cut on 15 June). Before the first cut in years 4 or 5, we sampled larvae of Lepidoptera and sawflies, and adults of moths, parasitoid wasps, wild bees, hoverflies, ground beetles, and rove beetles. Overall, before the first cut of the year, abundances of species with vegetation-dwelling larvae were higher in meadows with delayed mowing or an uncut grass refuge, with some taxon-specific variation. In contrast, species whose larval development is independent of the meadow vegetation showed no differences in abundance between mowing regimes. Species richness did not differ among regimes. For species with vegetation-dwelling larvae, a fourth-corner analysis showed an association between early phenology and the control regime. No associations were found for the other functional groups. Our results show that slight modifications of mowing regimes, easily implementable in agri-environmental policy schemes, can boost invertebrate abundance, potentially benefitting insectivorous vertebrates.

Risks and opportunities of trophic rewilding for arthropod communities.

Trophic rewilding is a restoration strategy focusing on the restoration of trophic interactions to promote self-regulating, biodiverse ecosystems. It has been proposed as an alternative to traditional conservation management in abandoned or defaunated areas. Arthropods constitute the most species-rich group of eukaryotic organisms, but are rarely considered in rewilding. Here, we first present an overview of direct and indirect pathways by which large herbivores and predators affect arthropod communities. We then review the published evidence of the impacts of rewilding with large herbivores on arthropods, including grey literature. We find that systematic monitoring is rare and that a comparison with a relevant control treatment is usually lacking. Nevertheless, the available data suggest that when the important process of top-down control of large-herbivore populations is missing, arthropod diversity tends to decrease. To ensure that rewilding is supportive of biodiversity conservation, we propose that if natural processes can only partially be restored, substitutes for missing processes are applied. We also propose that boundaries of acceptable outcomes of rewilding actions should be defined a priori, particularly concerning biodiversity conservation, and that action is taken when these boundaries are transgressed. To evaluate the success of rewilding for biodiversity, monitoring of arthropod communities should be a key instrument.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Cross-taxa generalities in the relationship between population abundance and ambient temperatures.

Identifying patterns in the effects of temperature on species' population abundances could help develop a general framework for predicting the consequences of climate change across different communities and realms. We used long-term population time series data from terrestrial, freshwater, and marine species communities within central Europe to compare the effects of temperature on abundance across a broad range of taxonomic groups. We asked whether there was an average relationship between temperatures in different seasons and annual abundances of species in a community, and whether species attributes (temperature range of distribution, range size, habitat breadth, dispersal ability, body size, and lifespan) explained interspecific variation in the relationship between temperature and abundance. We found that, on average, warmer winter temperatures were associated with greater abundances in terrestrial communities (ground beetles, spiders, and birds) but not always in aquatic communities (freshwater and marine invertebrates and fish). The abundances of species with large geographical ranges, larger body sizes, and longer lifespans tended to be less related to temperature. Our results suggest that climate change may have, in general, positive effects on species' abundances within many terrestrial communities in central Europe while the effects are less predictable in aquatic communities.

Cross-realm assessment of climate change impacts on species' abundance trends.

Climate change, land-use change, pollution and exploitation are among the main drivers of species' population trends; however, their relative importance is much debated. We used a unique collection of over 1,000 local population time series in 22 communities across terrestrial, freshwater and marine realms within central Europe to compare the impacts of long-term temperature change and other environmental drivers from 1980 onwards. To disentangle different drivers, we related species' population trends to species- and driver-specific attributes, such as temperature and habitat preference or pollution tolerance. We found a consistent impact of temperature change on the local abundances of terrestrial species. Populations of warm-dwelling species increased more than those of cold-dwelling species. In contrast, impacts of temperature change on aquatic species' abundances were variable. Effects of temperature preference were more consistent in terrestrial communities than effects of habitat preference, suggesting that the impacts of temperature change have become widespread for recent changes in abundance within many terrestrial communities of central Europe.

Herbivore exclusion promotes a more stochastic plant community assembly in a natural grassland.

Both bottom-up (e.g., nutrients) and top-down (e.g., herbivory) forces structure plant communities, but it remains unclear how they affect the relative importance of stochastic and deterministic processes in plant community assembly. Moreover, different-sized herbivores have been shown to have contrasting effects on community structure and function, but their effects on the processes governing community assembly (i.e., how they generate the impacts on structure) remain largely unknown. We evaluated the influence of bottom-up and top-down forces on the relative importance of deterministic and stochastic processes during plant community assembly. We used the data of a 7-yr factorial experiment manipulating nutrient availability (ambient and increased) and the presence of vertebrate herbivores (>1 kg) of different body size in a floodplain grassland in The Netherlands. We used a null model that describes a community composition expected by chance (i.e., stochastic assembly) and compared the plant community composition in the different treatments with this null model (the larger the difference, the more deterministically assembled). Our results showed that herbivore exclusion promoted a more stochastic plant community assembly, whereas increased nutrients played a relatively minor role in determining the relative importance of stochasticity in community assembly. Large herbivores facilitated intermediate-sized mammal herbivores, resulting in synergistic effects of enhanced grazing pressure and a more deterministic and convergent plant community assembly. We conclude that herbivores can act as strong deterministic forces during community assembly in natural systems. Our results also reveal that although large- and intermediate-sized mammal herbivores often have contrasting effects on many community and ecosystem properties, they can also synergistically homogenize plant communities.