The status and future of essential geodiversity variables.

Rapid environmental change, natural resource overconsumption and increasing concerns about ecological sustainability have led to the development of 'Essential Variables' (EVs). EVs are harmonized data products to inform policy and to enable effective management of natural resources by monitoring global changes. Recent years have seen the instigation of new EVs beyond those established for climate, oceans and biodiversity (ECVs, EOVs and EBVs), including Essential Geodiversity Variables (EGVs). EGVs aim to consistently quantify and monitor heterogeneity of Earth-surface and subsurface abiotic features, including geology, geomorphology, hydrology and pedology. Here we assess the status and future development of EGVs to better incorporate geodiversity into policy and sustainable management of natural resources. Getting EGVs operational requires better consensus on defining geodiversity, investments into a governance structure and open platform for curating the development of EGVs, advances in harmonizing in situ measurements and linking heterogeneous databases, and development of open and accessible computational workflows for global digital mapping using machine-learning techniques. Cross-disciplinary collaboration and partnerships with governmental and private organizations are needed to ensure the successful development and uptake of EGVs across science and policy. This article is part of the Theo Murphy meeting issue 'Geodiversity for science and society'.

The influence of scale-dependent geodiversity on species distribution models in a biodiversity hotspot.

Improving models of species' distributions is essential for conservation, especially in light of global change. Species distribution models (SDMs) often rely on mean environmental conditions, yet species distributions are also a function of environmental heterogeneity and filtering acting at multiple spatial scales. Geodiversity, which we define as the variation of abiotic features and processes of Earth's entire geosphere (inclusive of climate), has potential to improve SDMs and conservation assessments, as they capture multiple abiotic dimensions of species niches, however they have not been sufficiently tested in SDMs. We tested a range of geodiversity variables computed at varying scales using climate and elevation data. We compared predictive performance of MaxEnt SDMs generated using CHELSA bioclimatic variables to those also including geodiversity variables for 31 mammalian species in Colombia. Results show the spatial grain of geodiversity variables affects SDM performance. Some variables consistently exhibited an increasing or decreasing trend in variable importance with spatial grain, showing slight scale-dependence and indicating that some geodiversity variables are more relevant at particular scales for some species. Incorporating geodiversity variables into SDMs, and doing so at the appropriate spatial scales, enhances the ability to model species-environment relationships, thereby contributing to the conservation and management of biodiversity. This article is part of the Theo Murphy meeting issue 'Geodiversity for science and society'.

Diversity-stability relationships across organism groups and ecosystem types become decoupled across spatial scales.

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity-stability relationship [DSR]). At broader spatial extents, regional-scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long-term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ-diversity on metacommunity variability, but ß-diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ-diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial ß-diversity to reduce spatial synchrony.

Life-history responses to temperature and seasonality mediate ectotherm consumer-resource dynamics under climate warming.

Climate warming is altering life cycles of ectotherms by advancing phenology and decreasing generation times. Theoretical models provide powerful tools to investigate these effects of climate warming on consumer-resource population dynamics. Yet, existing theory primarily considers organisms with simplified life histories in constant temperature environments, making it difficult to predict how warming will affect organisms with complex life cycles in seasonal environments. We develop a size-structured consumer-resource model with seasonal temperature dependence, parameterized for a freshwater insect consuming zooplankton. We simulate how climate warming in a seasonal environment could alter a key life-history trait of the consumer, number of generations per year, mediating responses of consumer-resource population sizes and consumer persistence. We find that, with warming, consumer population sizes increase through multiple mechanisms. First, warming decreases generation times by increasing rates of resource ingestion and growth and/or lengthening the growing season. Second, these life-history changes shorten the juvenile stage, increasing the number of emerging adults and population-level reproduction. Unstructured models with similar assumptions found that warming destabilized consumer-resource dynamics. By contrast, our size-structured model predicts stability and consumer persistence. Our study suggests that, in seasonal environments experiencing climate warming, life-history changes that lead to shorter generation times could delay population extinctions.

Consistent stabilizing effects of plant diversity across spatial scales and climatic gradients.

Biodiversity has widely been documented to enhance local community stability but whether such stabilizing effects of biodiversity extend to broader scales remains elusive. Here, we investigated the relationships between biodiversity and community stability in natural plant communities from quadrat (1 m2) to plot (400 m2) and regional (5-214 km2) scales and across broad climatic conditions, using an extensive plant community dataset from the National Ecological Observatory Network. We found that plant diversity provided consistent stabilizing effects on total community abundance across three nested spatial scales and climatic gradients. The strength of the stabilizing effects of biodiversity increased modestly with spatial scale and decreased as precipitation seasonality increased. Our findings illustrate the generality of diversity-stability theory across scales and climatic gradients, which provides a robust framework for understanding ecosystem responses to biodiversity and climate changes.

Potential ecological impacts of climate intervention by reflecting sunlight to cool Earth.

As the effects of anthropogenic climate change become more severe, several approaches for deliberate climate intervention to reduce or stabilize Earth's surface temperature have been proposed. Solar radiation modification (SRM) is one potential approach to partially counteract anthropogenic warming by reflecting a small proportion of the incoming solar radiation to increase Earth's albedo. While climate science research has focused on the predicted climate effects of SRM, almost no studies have investigated the impacts that SRM would have on ecological systems. The impacts and risks posed by SRM would vary by implementation scenario, anthropogenic climate effects, geographic region, and by ecosystem, community, population, and organism. Complex interactions among Earth's climate system and living systems would further affect SRM impacts and risks. We focus here on stratospheric aerosol intervention (SAI), a well-studied and relatively feasible SRM scheme that is likely to have a large impact on Earth's surface temperature. We outline current gaps in knowledge about both helpful and harmful predicted effects of SAI on ecological systems. Desired ecological outcomes might also inform development of future SAI implementation scenarios. In addition to filling these knowledge gaps, increased collaboration between ecologists and climate scientists would identify a common set of SAI research goals and improve the communication about potential SAI impacts and risks with the public. Without this collaboration, forecasts of SAI impacts will overlook potential effects on biodiversity and ecosystem services for humanity.

The geodiv r package: Tools for calculating gradient surface metrics.

The geodiv r package calculates gradient surface metrics from imagery and other gridded datasets to provide continuous measures of landscape heterogeneity for landscape pattern analysis. geodiv is the first open-source, command line toolbox for calculating many gradient surface metrics and easily integrates parallel computing for applications with large images or rasters (e.g. remotely sensed data). All functions may be applied either globally to derive a single metric for an entire image or locally to create a texture image over moving windows of a user-defined extent.We present a comprehensive description of the functions available through geodiv. A supplemental vignette provides an example application of geodiv to the fields of landscape ecology and biogeography. geodiv allows users to easily retrieve estimates of spatial heterogeneity for a variety of purposes, enhancing our understanding of how environmental structure influences ecosystem processes. The package works with any continuous imagery and may be widely applied in many fields where estimates of surface complexity are useful.

Foraging strategy mediates ectotherm predator-prey responses to climate warming.

Climate warming and species traits interact to influence predator performance, including individual feeding and growth rates. However, the effects of an important trait-predator foraging strategy-are largely unknown. We investigated the interactions between predator foraging strategy and temperature on two ectotherm predators: an active predator, the backswimmer Notonecta undulata, and a sit-and-wait predator, the damselfly Enallagma annexum. In a series of predator-prey experiments across a temperature gradient, we measured predator feeding rates on an active prey species, zooplankton Daphnia pulex, predator growth rates, and mechanisms that influence predator feeding: body speed of predators and prey (here measured as swimming speed), prey encounter rates, capture success, attack rates, and handling time. Overall, warming led to increased feeding rates for both predators through changes to each component of the predator's functional response. We found that prey swimming speed strongly increased with temperature. The active predator's swimming speed also increased with temperature, and together, the increase in predator and prey swimming speed resulted in twofold higher prey encounter rates for the active predator at warmer temperatures. By contrast, prey encounter rates of the sit-and-wait predator increased fourfold with rising temperatures as a result of increased prey swimming speed. Concurrently, increased prey swimming speed was associated with a decline in the active predator's capture success at high temperatures, whereas the sit-and-wait predator's capture success slightly increased with temperature. We provide some of the first evidence that foraging traits mediate the indirect effects of warming on predator performance. Understanding how traits influence species' responses to warming could clarify how climate change will affect entire functional groups of species.

Towards connecting biodiversity and geodiversity across scales with satellite remote sensing.

ISSUE: Geodiversity (i.e., the variation in Earth's abiotic processes and features) has strong effects on biodiversity patterns. However, major gaps remain in our understanding of how relationships between biodiversity and geodiversity vary over space and time. Biodiversity data are globally sparse and concentrated in particular regions. In contrast, many forms of geodiversity can be measured continuously across the globe with satellite remote sensing. Satellite remote sensing directly measures environmental variables with grain sizes as small as tens of metres and can therefore elucidate biodiversity-geodiversity relationships across scales. EVIDENCE: We show how one important geodiversity variable, elevation, relates to alpha, beta and gamma taxonomic diversity of trees across spatial scales. We use elevation from NASA's Shuttle Radar Topography Mission (SRTM) and c. 16,000 Forest Inventory and Analysis plots to quantify spatial scaling relationships between biodiversity and geodiversity with generalized linear models (for alpha and gamma diversity) and beta regression (for beta diversity) across five spatial grains ranging from 5 to 100 km. We illustrate different relationships depending on the form of diversity; beta and gamma diversity show the strongest relationship with variation in elevation. CONCLUSION: With the onset of climate change, it is more important than ever to examine geodiversity for its potential to foster biodiversity. Widely available satellite remotely sensed geodiversity data offer an important and expanding suite of measurements for understanding and predicting changes in different forms of biodiversity across scales. Interdisciplinary research teams spanning biodiversity, geoscience and remote sensing are well poised to advance understanding of biodiversity-geodiversity relationships across scales and guide the conservation of nature.

Metabolic asymmetry and the global diversity of marine predators.

Species richness of marine mammals and birds is highest in cold, temperate seas-a conspicuous exception to the general latitudinal gradient of decreasing diversity from the tropics to the poles. We compiled a comprehensive dataset for 998 species of sharks, fish, reptiles, mammals, and birds to identify and quantify inverse latitudinal gradients in diversity, and derived a theory to explain these patterns. We found that richness, phylogenetic diversity, and abundance of marine predators diverge systematically with thermoregulatory strategy and water temperature, reflecting metabolic differences between endotherms and ectotherms that drive trophic and competitive interactions. Spatial patterns of foraging support theoretical predictions, with total prey consumption by mammals increasing by a factor of 80 from the equator to the poles after controlling for productivity.

Asymmetric biotic interactions and abiotic niche differences revealed by a dynamic joint species distribution model.

A species' distribution and abundance are determined by abiotic conditions and biotic interactions with other species in the community. Most species distribution models correlate the occurrence of a single species with environmental variables only, and leave out biotic interactions. To test the importance of biotic interactions on occurrence and abundance, we compared a multivariate spatiotemporal model of the joint abundance of two invasive insects that share a host plant, hemlock woolly adelgid (HWA; Adelges tsugae) and elongate hemlock scale (EHS; Fiorina externa), to independent models that do not account for dependence among co-occurring species. The joint model revealed that HWA responded more strongly to abiotic conditions than EHS. Additionally, HWA appeared to predispose stands to subsequent increase of EHS, but HWA abundance was not strongly dependent on EHS abundance. This study demonstrates how incorporating spatial and temporal dependence into a species distribution model can reveal the dependence of a species' abundance on other species in the community. Accounting for dependence among co-occurring species with a joint distribution model can also improve estimation of the abiotic niche for species affected by interspecific interactions.

Short-term responses to warming vary between native vs. exotic species and with latitude in an early successional plant community.

Climate change is expected to favor exotic plant species over native species, because exotics tend to have wider climatic tolerances and greater phenological plasticity, and also because climate change may intensify enemy release. Here, we examine direct effects of warming (+ 1.8 °C above ambient) on plant abundance and phenology, as well as indirect effects of warming propagated through herbivores, in two heavily invaded plant communities in Michigan, USA, separated by approximately three degrees latitude. At the northern site, warming increased exotic plant abundance by 19% but decreased native plant abundance by 31%, indicating that exotic species may be favored in a warmer world. Warming also resulted in earlier spring green-up (1.65 ± 0.77 days), earlier flowering (2.18 ± 0.92 days), and greater damage by herbivores (twofold increase), affecting exotic and native species equally. Contrary to expectations, native and exotic plants experienced similar amounts of herbivory. Warming did not have strong ecological effects at the southern site, only resulting in a delay of flowering time by 2.42 ± 0.83 days for both native and exotic species. Consistent with the enemy release hypothesis, exotic plants experienced less herbivory than native plants at the southern site. Herbivory was lower under warming for both exotic and native species at the southern site. Thus, climate warming may favor exotic over native plant species, but the response is likely to depend on additional environmental and individual species' traits.

Tropical bird species have less variable body sizes.

Ecologists have often predicted that species' niche breadths should decline towards the Equator. Dan Janzen arrived at this prediction based on climatic constraints, while Robert MacArthur argued that a latitudinal gradient in resource specialization drives the pattern. This idea has some support when it comes to thermal niches, but has rarely been explored for other niche dimensions. Body size is linked to niche dimensions related to diet, competition and environmental tolerance in vertebrates. We identified 68 pairs of tropical and nontropical sister bird species using a comprehensive phylogeny and used the VertNet specimen database to ask whether tropical birds have lower intraspecific body-size variation than their nontropical sister species. Our results show that tropical species have less intraspecific variability in body mass ([Formula: see text]; p = 0.009). Variation in body-size variability was poorly explained by both abiotic and biotic drivers; thus the mechanisms underlying the pattern are still unclear. The lower variation in body size of tropical bird species may have evolved in response to more stable climates and resource environments.

Incorporating Context Dependency of Species Interactions in Species Distribution Models.

SYNOPSIS: Species distribution models typically use correlative approaches that characterize the species-environment relationship using occurrence or abundance data for a single species. However, species distributions are determined by both abiotic conditions and biotic interactions with other species in the community. Therefore, climate change is expected to impact species through direct effects on their physiology and indirect effects propagated through their resources, predators, competitors, or mutualists. Furthermore, the sign and strength of species interactions can change according to abiotic conditions, resulting in context-dependent species interactions that may change across space or with climate change. Here, we incorporated the context dependency of species interactions into a dynamic species distribution model. We developed a multi-species model that uses a time-series of observational survey data to evaluate how abiotic conditions and species interactions affect the dynamics of three rocky intertidal species. The model further distinguishes between the direct effects of abiotic conditions on abundance and the indirect effects propagated through interactions with other species. We apply the model to keystone predation by the sea star Pisaster ochraceus on the mussel Mytilus californianus and the barnacle Balanus glandula in the rocky intertidal zone of the Pacific coast, USA. Our method indicated that biotic interactions between P. ochraceus and B. glandula affected B. glandula dynamics across >1000 km of coastline. Consistent with patterns from keystone predation, the growth rate of B. glandula varied according to the abundance of P. ochraceus in the previous year. The data and the model did not indicate that the strength of keystone predation by P. ochraceus varied with a mean annual upwelling index. Balanus glandula cover increased following years with high phytoplankton abundance measured as mean annual chlorophyll-a. M. californianus exhibited the same pattern to a lesser degree, although this pattern was not significant. This work bridges the disciplines of biogeography and community ecology to develop tools to better understand the direct and indirect effects of abiotic conditions on ecological communities.

Searching for Biotic Multipliers of Climate Change.

SYNOPSIS: As climates change, biologists need to prioritize which species to understand, predict, and protect. One way is to identify key species that are both sensitive to climate change and that disproportionately affect communities and ecosystems. These "biotic multipliers" provide efficient targets for research and conservation. Here, we propose eight mechanistic hypotheses related to impact and sensitivity that suggest that top consumers might often act as biotic multipliers of climate change. For impact, top consumers often affect communities and ecosystems through strong top-down effects. For sensitivity, metabolic theory and data suggest that photosynthesis and respiration differ in temperature responses, potentially increasing the sensitivity of consumers relative to plants. Larger-bodied organisms are typically more thermally sensitive than smaller ones, suggesting how large top consumers might be more sensitive than their smaller prey. In addition, traits related to predation are more sensitive than defensive traits to temperature. Top consumers might also be more sensitive because they often lag behind prey in phenological responses. The combination of low population sizes and demographic traits of top consumers could make them more sensitive to disturbances like climate change, which could slow their recovery. As top consumers are positioned at the top of the food chain, many small effects can accumulate from other trophic levels to affect top consumers. Finally, top consumers also often disperse more frequently and farther than prey, potentially leading to greater sensitivity to climate-induced changes in ranges and subsequent impacts on invaded communities. Overall, we expect that large, ectothermic top consumers and mobile predators might frequently be biotic multipliers of climate change. However, this prediction depends on the particular features of species, habitats, and ecosystems. In specific cases, herbivores, plants, or pathogens might be more sensitive than top consumers or have greater community impacts. To predict biotic multipliers, we need to compare sensitivities and impacts across trophic groups in a broader range of ecosystems as well as perform experiments that uncouple proposed mechanisms. Overall, the biotic multiplier concept offers an alternative prioritization scheme for research and conservation that includes impacts on communities and ecosystems.

Coastal foredune evolution: the relative influence of vegetation and sand supply in the US Pacific Northwest.

Biophysical feedbacks between vegetation and sediment are important for forming and modifying landscape features and their ecosystem services. These feedbacks are especially important where landscape features differ in their provision of ecosystem services. For example, the shape of coastal foredunes, a product of both physical and biological forces, determines their ability to protect communities from rising seas and changing patterns of storminess. Here we assessed how sand supply and changes in vegetation over interannual (3 year) and decadal (21 year) scales influenced foredune shape along 100 km of coastline in the US Pacific Northwest. Across 21 years, vegetation switched from one congeneric non-native beachgrass to another (Ammophila arenaria to A. breviligulata) while sand supply rates were positive. At interannual timescales, sand supply rates explained the majority of change in foredune height (64-69%) and width (56-80%). However, at decadal scales, change in vegetation explained the majority of the change in foredune width (62-68%), whereas sand supply rates explained most of the change in foredune height (88-90%). In areas with lower shoreline change rates (±2 m yr(-1)), the change in vegetation explained the majority of decadal changes in foredune width (56-57%) and height (59-76%). Foredune shape directly impacts coastal protection, thus our findings are pertinent to coastal management given pressures of development and climate change.

Invasive congeners differ in successional impacts across space and time.

Invasive species can alter the succession of ecological communities because they are often adapted to the disturbed conditions that initiate succession. The extent to which this occurs may depend on how widely they are distributed across environmental gradients and how long they persist over the course of succession. We focus on plant communities of the USA Pacific Northwest coastal dunes, where disturbance is characterized by changes in sediment supply, and the plant community is dominated by two introduced grasses--the long-established Ammophila arenaria and the currently invading A. breviligulata. Previous studies showed that A. breviligulata has replaced A. arenaria and reduced community diversity. We hypothesize that this is largely due to A. breviligulata occupying a wider distribution across spatial environmental gradients and persisting in later-successional habitat than A. arenaria. We used multi-decadal chronosequences and a resurvey study spanning 2 decades to characterize distributions of both species across space and time, and investigated how these distributions were associated with changes in the plant community. The invading A. breviligulata persisted longer and occupied a wider spatial distribution across the dune, and this corresponded with a reduction in plant species richness and native cover. Furthermore, backdunes previously dominated by A. arenaria switched to being dominated by A. breviligulata, forest, or developed land over a 23-yr period. Ammophila breviligulata likely invades by displacing A. arenaria, and reduces plant diversity by maintaining its dominance into later successional backdunes. Our results suggest distinct roles in succession, with A. arenaria playing a more classically facilitative role and A. breviligulata a more inhibitory role. Differential abilities of closely-related invasive species to persist through time and occupy heterogeneous environments allows for distinct impacts on communities during succession.

Climate change and the past, present, and future of biotic interactions.

Biotic interactions drive key ecological and evolutionary processes and mediate ecosystem responses to climate change. The direction, frequency, and intensity of biotic interactions can in turn be altered by climate change. Understanding the complex interplay between climate and biotic interactions is thus essential for fully anticipating how ecosystems will respond to the fast rates of current warming, which are unprecedented since the end of the last glacial period. We highlight episodes of climate change that have disrupted ecosystems and trophic interactions over time scales ranging from years to millennia by changing species' relative abundances and geographic ranges, causing extinctions, and creating transient and novel communities dominated by generalist species and interactions. These patterns emerge repeatedly across disparate temporal and spatial scales, suggesting the possibility of similar underlying processes. Based on these findings, we identify knowledge gaps and fruitful areas for research that will further our understanding of the effects of climate change on ecosystems.

Moving forward: dispersal and species interactions determine biotic responses to climate change.

We need accurate predictions about how climate change will alter species distributions and abundances around the world. Most predictions assume simplistic dispersal scenarios and ignore biotic interactions. We argue for incorporating the complexities of dispersal and species interactions. Range expansions depend not just on mean dispersal, but also on the shape of the dispersal kernel and the population's growth rate. We show how models using species-specific dispersal can produce more accurate predictions than models applying all-or-nothing dispersal scenarios. Models that additionally include species interactions can generate distinct outcomes. For example, species interactions can slow climate tracking and produce more extinctions than models assuming no interactions. We conclude that (1) just knowing mean dispersal is insufficient to predict biotic responses to climate change, and (2) considering interspecific dispersal variation and species interactions jointly will be necessary to anticipate future changes to biological diversity. We advocate for collecting key information on interspecific dispersal differences and strong biotic interactions so that we can build the more robust predictive models that will be necessary to inform conservation efforts as climates continue to change.