Trees have overlapping potential niches that extend beyond their realized niches.

Tree species appear to prefer distinct climatic conditions, but the true nature of these preferences is obscured by species interactions and dispersal, which limit species' ranges. We quantified realized and potential thermal niches of 188 North American tree species to conduct a continental-scale test of the architecture of niches. We found strong and consistent evidence that species occurring at thermal extremes occupy less than three-quarters of their potential niches, and species' potential niches overlap at a mean annual temperature of ~12°C. These results clarify the breadth of thermal tolerances of temperate tree species and support the centrifugal organization of thermal niches. Accounting for the nonrealized components of ecological niches will advance theory and prediction in global change ecology.

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.

Operationalizing cultural adaptation to climate change: contemporary examples from United States agriculture.

It has been proposed that climate adaptation research can benefit from an evolutionary approach. But related empirical research is lacking. We advance the evolutionary study of climate adaptation with two case studies from contemporary United States agriculture. First, we define 'cultural adaptation to climate change' as a mechanistic process of population-level cultural change. We argue this definition enables rigorous comparisons, yields testable hypotheses from mathematical theory and distinguishes adaptive change, non-adaptive change and desirable policy outcomes. Next, we develop an operational approach to identify 'cultural adaptation to climate change' based on established empirical criteria. We apply this approach to data on crop choices and the use of cover crops between 2008 and 2021 from the United States. We find evidence that crop choices are adapting to local trends in two separate climate variables in some regions of the USA. But evidence suggests that cover cropping may be adapting more to the economic environment than climatic conditions. Further research is needed to characterize the process of cultural adaptation, particularly the routes and mechanisms of cultural transmission. Furthermore, climate adaptation policy could benefit from research on factors that differentiate regions exhibiting adaptive trends in crop choice from those that do not. This article is part of the theme issue 'Climate change adaptation needs a science of culture'.

Macrobehaviour: behavioural variation across space, time, and taxa.

We explore how integrating behavioural ecology and macroecology can provide fundamental new insight into both fields, with particular relevance for understanding ecological responses to rapid environmental change. We outline the field of macrobehaviour, which aims to unite these disciplines explicitly, and highlight examples of research in this space. Macrobehaviour can be envisaged as a spectrum, where behavioural ecologists and macroecologists use new data and borrow tools and approaches from one another. At the heart of this spectrum, interdisciplinary research considers how selection in the context of large-scale factors can lead to systematic patterns in behavioural variation across space, time, and taxa, and in turn, influence macroecological patterns and processes. Macrobehaviour has the potential to enhance forecasts of future biodiversity change.

Looking back on biodiversity change: lessons for the road ahead.

Estimating biodiversity change across the planet in the context of widespread human modification is a critical challenge. Here, we review how biodiversity has changed in recent decades across scales and taxonomic groups, focusing on four diversity metrics: species richness, temporal turnover, spatial beta-diversity and abundance. At local scales, change across all metrics includes many examples of both increases and declines and tends to be centred around zero, but with higher prevalence of declining trends in beta-diversity (increasing similarity in composition across space or biotic homogenization) and abundance. The exception to this pattern is temporal turnover, with changes in species composition through time observed in most local assemblages. Less is known about change at regional scales, although several studies suggest that increases in richness are more prevalent than declines. Change at the global scale is the hardest to estimate accurately, but most studies suggest extinction rates are probably outpacing speciation rates, although both are elevated. Recognizing this variability is essential to accurately portray how biodiversity change is unfolding, and highlights how much remains unknown about the magnitude and direction of multiple biodiversity metrics at different scales. Reducing these blind spots is essential to allow appropriate management actions to be deployed. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

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.

How does variation in total and relative abundance contribute to gradients of species diversity?

Patterns of biodiversity provide insights into the processes that shape biological communities around the world. Variation in species diversity along biogeographical or ecological gradients, such as latitude or precipitation, can be attributed to variation in different components of biodiversity: changes in the total abundance (i.e., more-individual effects) and changes in the regional species abundance distribution (SAD). Rarefaction curves can provide a tool to partition these sources of variation on diversity, but first must be converted to a common unit of measurement. Here, we partition species diversity gradients into components of the SAD and abundance using the effective number of species (ENS) transformation of the individual-based rarefaction curve. Because the ENS curve is unconstrained by sample size, it can act as a standardized unit of measurement when comparing effect sizes among different components of biodiversity change. We illustrate the utility of the approach using two data sets spanning latitudinal diversity gradients in trees and marine reef fish and find contrasting results. Whereas the diversity gradient of fish was mostly associated with variation in abundance (86%), the tree diversity gradient was mostly associated with variation in the SAD (59%). These results suggest that local fish diversity may be limited by energy through the more-individuals effect, while species pool effects are the larger determinant of tree diversity. We suggest that the framework of the ENS-curve has the potential to quantify the underlying factors influencing most aspects of diversity change.

Late quaternary biotic homogenization of North American mammalian faunas.

Biotic homogenization-increasing similarity of species composition among ecological communities-has been linked to anthropogenic processes operating over the last century. Fossil evidence, however, suggests that humans have had impacts on ecosystems for millennia. We quantify biotic homogenization of North American mammalian assemblages during the late Pleistocene through Holocene (~30,000 ybp to recent), a timespan encompassing increased evidence of humans on the landscape (~20,000-14,000 ybp). From ~10,000 ybp to recent, assemblages became significantly more homogenous (>100% increase in Jaccard similarity), a pattern that cannot be explained by changes in fossil record sampling. Homogenization was most pronounced among mammals larger than 1 kg and occurred in two phases. The first followed the megafaunal extinction at ~10,000 ybp. The second, more rapid phase began during human population growth and early agricultural intensification (~2,000-1,000 ybp). We show that North American ecosystems were homogenizing for millennia, extending human impacts back ~10,000 years.

Source-sink behavioural dynamics limit institutional evolution in a group-structured society.

Social change in any society entails changes in both behaviours and institutions. We model a group-structured society in which the transmission of individual behaviour occurs in parallel with the selection of group-level institutions. We consider a cooperative behaviour that generates collective benefits for groups but does not spread between individuals on its own. Groups exhibit institutions that increase the diffusion of the behaviour within the group, but also incur a group cost. Groups adopt institutions in proportion to their fitness. Finally, the behaviour may also spread globally. We find that behaviour and institutions can be mutually reinforcing. But the model also generates behavioural source-sink dynamics when behaviour generated in institutionalized groups spreads to non-institutionalized groups and boosts their fitness. Consequently, the global diffusion of group-beneficial behaviour creates a pattern of institutional free-riding that limits the evolution of group-beneficial institutions. Our model suggests that, in a group-structured society, large-scale beneficial social change can be best achieved when the relevant behaviour and institutions remain correlated.

Long-term changes in temperate marine fish assemblages are driven by a small subset of species.

The species composition of plant and animal assemblages across the globe has changed substantially over the past century. How do the dynamics of individual species cause this change? We classified species into seven unique categories of temporal dynamics based on the ordered sequence of presences and absences that each species contributes to an assemblage time series. We applied this framework to 14,434 species trajectories comprising 280 assemblages of temperate marine fishes surveyed annually for 20 or more years. Although 90% of the assemblages diverged in species composition from the baseline year, this compositional change was largely driven by only 8% of the species' trajectories. Quantifying the reorganization of assemblages based on species shared temporal dynamics should facilitate the task of monitoring and restoring biodiversity. We suggest ways in which our framework could provide informative measures of compositional change, as well as leverage future research on pattern and process in ecological systems.

The dimensionality and structure of species trait spaces.

Trait-based ecology aims to understand the processes that generate the overarching diversity of organismal traits and their influence on ecosystem functioning. Achieving this goal requires simplifying this complexity in synthetic axes defining a trait space and to cluster species based on their traits while identifying those with unique combinations of traits. However, so far, we know little about the dimensionality, the robustness to trait omission and the structure of these trait spaces. Here, we propose a unified framework and a synthesis across 30 trait datasets representing a broad variety of taxa, ecosystems and spatial scales to show that a common trade-off between trait space quality and operationality appears between three and six dimensions. The robustness to trait omission is generally low but highly variable among datasets. We also highlight invariant scaling relationships, whatever organismal complexity, between the number of clusters, the number of species in the dominant cluster and the number of unique species with total species richness. When species richness increases, the number of unique species saturates, whereas species tend to disproportionately pack in the richest cluster. Based on these results, we propose some rules of thumb to build species trait spaces and estimate subsequent functional diversity indices.

Long-term trends indicate that invasive plants are pervasive and increasing in eastern national parks.

While invasive plant distributions are relatively well known in the eastern United States, temporal changes in species distributions and interactions among species have received little attention. Managers are therefore left to make management decisions without knowing which species pose the greatest threats based on their ability to spread, persist and outcompete other invasive species. To fill this gap, we used the U.S. National Park Service's Inventory and Monitoring Program data collected from over 1,400 permanent forest plots spanning 12 yr and covering 39 eastern national parks to analyze invasive plant trends. We analyzed trends in abundance at multiple scales, including plot frequency, quadrat frequency, and average quadrat cover. We examined trends overall, by functional group, and by species. We detected considerably more increasing than decreasing trends in invasive plant abundance. In fact, 80% of the parks in our study had at least one significant increasing trend in invasive abundance over time. Where detected, significant negative trends tended to be herbaceous or graminoid species. However, these declines were often countered by roughly equivalent increases in invasive shrubs over the same time period, and we only detected overall declines in invasive abundance in two parks in our study. Present in over 30% of plots and responsible for the steepest and greatest number of significant increases, Japanese stiltgrass (Microstegium vimineum) was the most aggressive invader in our study and is a high management priority. Invasive shrubs, especially Japanese barberry (Berberis thunbergii), Japanese honeysuckle (Lonicera japonica), multiflora rose (Rosa multiflora), and wineberry (Rubus phoenicolasius), also increased across multiple parks, and sometimes at the expense of Japanese stiltgrass. Given the added risks to human health from tick-borne diseases, invasive shrubs are a high management priority. While these findings provide critical information to managers for species prioritization, they also demonstrate the incredible management challenge that invasive plants pose in protected areas, particularly since we documented few overall declines in invasive abundance. As parks work to overcome deferred maintenance of infrastructure, our findings suggest that deferred management of natural resources, particularly invasive species, requires similar attention and long-term commitment to reverse these widespread increasing invasive trends.

A multiscale framework for disentangling the roles of evenness, density, and aggregation on diversity gradients.

Disentangling the drivers of diversity gradients can be challenging. The Measurement of Biodiversity (MoB) framework decomposes scale-dependent changes in species diversity into three components of community structure: species abundance distribution (SAD), total community abundance, and within-species spatial aggregation. Here we extend MoB from categorical treatment comparisons to quantify variation along continuous geographic or environmental gradients. Our approach requires sites along a gradient, each consisting of georeferenced plots of abundance-based species composition data. We demonstrate our method using a case study of ants sampled along an elevational gradient of 28 sites in a mixed deciduous forest of the Great Smoky Mountains National Park, USA. MoB analysis revealed that decreases in ant species richness along the elevational gradient were associated with decreasing evenness and total number of species, which counteracted the modest increase in richness associated with decreasing spatial aggregation along the gradient. Total community abundance had a negligible effect on richness at all but the finest spatial grains, SAD effects increased in importance with sampling effort, and the aggregation effect had the strongest effect at coarser spatial grains. These results do not support the more-individuals hypothesis, but they are consistent with a hypothesis of stronger environmental filtering at coarser spatial grains. Our extension of MoB has the potential to elucidate how components of community structure contribute to changes in diversity along environmental gradients and should be useful for a variety of assemblage-level data collected along gradients.

The commonness of rarity: Global and future distribution of rarity across land plants.

A key feature of life's diversity is that some species are common but many more are rare. Nonetheless, at global scales, we do not know what fraction of biodiversity consists of rare species. Here, we present the largest compilation of global plant diversity to quantify the fraction of Earth's plant biodiversity that are rare. A large fraction, ~36.5% of Earth's ~435,000 plant species, are exceedingly rare. Sampling biases and prominent models, such as neutral theory and the k-niche model, cannot account for the observed prevalence of rarity. Our results indicate that (i) climatically more stable regions have harbored rare species and hence a large fraction of Earth's plant species via reduced extinction risk but that (ii) climate change and human land use are now disproportionately impacting rare species. Estimates of global species abundance distributions have important implications for risk assessments and conservation planning in this era of rapid global change.

A balance of winners and losers in the Anthropocene.

Scientists disagree about the nature of biodiversity change. While there is evidence for widespread declines from population surveys, assemblage surveys reveal a mix of declines and increases. These conflicting conclusions may be caused by the use of different metrics: assemblage metrics may average out drastic changes in individual populations. Alternatively, differences may arise from data sources: populations monitored individually, versus whole-assemblage monitoring. To test these hypotheses, we estimated population change metrics using assemblage data. For a set of 23 241 populations, 16 009 species, in 158 assemblages, we detected significantly accelerating extinction and colonisation rates, with both rates being approximately balanced. Most populations (85%) did not show significant trends in abundance, and those that did were balanced between winners (8%) and losers (7%). Thus, population metrics estimated with assemblage data are commensurate with assemblage metrics and reveal sustained and increasing species turnover.

Similarities and differences in intrapopulation trait correlations of co-occurring tree species: consistent water-use relationships amid widely different correlation patterns.

PREMISE OF THE STUDY: General relationships among functional traits have been identified across species, but the forces shaping these relationships remain largely unknown. Adopting an approach from evolutionary biology, we studied similarities and differences in intrapopulation trait correlations among locally co-occurring tree species to assess the roles of constraints, phylogeny, and the environmental niche in shaping multivariate phenotypes. We tested the hypotheses (1) that intrapopulation correlations among functional traits are largely shaped by fundamental trade-offs or constraints and (2) that differences among species reflect adaptation to their environmental niches. METHODS: We compared pairwise correlations and correlation matrices of 17 key functional traits within and among temperate tree species. These traits describe three well-established trade-off dimensions characterizing interspecific relationships among physiological functions: resource acquisition and conservation; sap transport and mechanical support; and branch architecture. KEY RESULTS: Six trait pairs are consistently correlated within populations. Of these, only one involves dimensionally independent traits: LMA-δ13 C. For all other traits, intrapopulation functional trait correlations are weak, are species-specific, and differ from interspecific correlations. Species intrapopulation correlation matrices are related to neither phylogeny nor environmental niche. CONCLUSIONS: The results (1) suggest that the functional design of these species is centered on efficient water use, (2) highlight flexibility in plant functional design across species, and (3) suggest that intrapopulation, local interspecific, and global interspecific correlations are shaped by processes acting at each of these scales.

Embracing scale-dependence to achieve a deeper understanding of biodiversity and its change across communities.

Because biodiversity is multidimensional and scale-dependent, it is challenging to estimate its change. However, it is unclear (1) how much scale-dependence matters for empirical studies, and (2) if it does matter, how exactly we should quantify biodiversity change. To address the first question, we analysed studies with comparisons among multiple assemblages, and found that rarefaction curves frequently crossed, implying reversals in the ranking of species richness across spatial scales. Moreover, the most frequently measured aspect of diversity - species richness - was poorly correlated with other measures of diversity. Second, we collated studies that included spatial scale in their estimates of biodiversity change in response to ecological drivers and found frequent and strong scale-dependence, including nearly 10% of studies which showed that biodiversity changes switched directions across scales. Having established the complexity of empirical biodiversity comparisons, we describe a synthesis of methods based on rarefaction curves that allow more explicit analyses of spatial and sampling effects on biodiversity comparisons. We use a case study of nutrient additions in experimental ponds to illustrate how this multi-dimensional and multi-scale perspective informs the responses of biodiversity to ecological drivers.

Predictors of specialist avifaunal decline in coastal marshes.

Coastal marshes are one of the world's most productive ecosystems. Consequently, they have been heavily used by humans for centuries, resulting in ecosystem loss. Direct human modifications such as road crossings and ditches and climatic stressors such as sea-level rise and extreme storm events have the potential to further degrade the quantity and quality of marsh along coastlines. We used an 18-year marsh-bird database to generate population trends for 5 avian species (Rallus crepitans, Tringa semipalmata semipalmata, Ammodramus nelsonii subvirgatus, Ammodramus caudacutus, and Ammodramus maritimus) that breed almost exclusively in tidal marshes, and are potentially vulnerable to marsh degradation and loss as a result of anthropogenic change. We generated community and species trends across 3 spatial scales and explored possible drivers of the changes we observed, including marsh ditching, tidal restriction through road crossings, local rates of sea-level rise, and potential for extreme flooding events. The specialist community showed negative trends in tidally restricted marshes (-2.4% annually from 1998 to 2012) but was stable in unrestricted marshes across the same period. At the species level, we found negative population trends in 3 of the 5 specialist species, ranging from -4.2% to 9.0% annually. We suggest that tidal restriction may accelerate degradation of tidal marsh resilience to sea-level rise by limiting sediment supply necessary for marsh accretion, resulting in specialist habitat loss in tidally restricted marshes. Based on our findings, we predict a collapse of the global population of Saltmarsh Sparrows (A. caudacutus) within the next 50 years and suggest that immediate conservation action is needed to prevent extinction of this species. We also suggest mitigation actions to restore sediment supply to coastal marshes to help sustain this ecosystem into the future.