The role of biotic interactions in shaping distributions and realised assemblages of species: implications for species distribution modelling.

Predicting which species will occur together in the future, and where, remains one of the greatest challenges in ecology, and requires a sound understanding of how the abiotic and biotic environments interact with dispersal processes and history across scales. Biotic interactions and their dynamics influence species' relationships to climate, and this also has important implications for predicting future distributions of species. It is already well accepted that biotic interactions shape species' spatial distributions at local spatial extents, but the role of these interactions beyond local extents (e.g. 10 km(2) to global extents) are usually dismissed as unimportant. In this review we consolidate evidence for how biotic interactions shape species distributions beyond local extents and review methods for integrating biotic interactions into species distribution modelling tools. Drawing upon evidence from contemporary and palaeoecological studies of individual species ranges, functional groups, and species richness patterns, we show that biotic interactions have clearly left their mark on species distributions and realised assemblages of species across all spatial extents. We demonstrate this with examples from within and across trophic groups. A range of species distribution modelling tools is available to quantify species environmental relationships and predict species occurrence, such as: (i) integrating pairwise dependencies, (ii) using integrative predictors, and (iii) hybridising species distribution models (SDMs) with dynamic models. These methods have typically only been applied to interacting pairs of species at a single time, require a priori ecological knowledge about which species interact, and due to data paucity must assume that biotic interactions are constant in space and time. To better inform the future development of these models across spatial scales, we call for accelerated collection of spatially and temporally explicit species data. Ideally, these data should be sampled to reflect variation in the underlying environment across large spatial extents, and at fine spatial resolution. Simplified ecosystems where there are relatively few interacting species and sometimes a wealth of existing ecosystem monitoring data (e.g. arctic, alpine or island habitats) offer settings where the development of modelling tools that account for biotic interactions may be less difficult than elsewhere

Global maps of soil temperature.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km <sup>2</sup> resolution for 0-5 and 5-15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1-km <sup>2</sup> pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (-0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications

Evaluating and communicating simulated wildlife responses to land-use scenarios

Reliable assessments of how human activities affect wildlife populations are essential for effective natural resource management. Agent-based models provide a powerful tool for integration of multiple drivers of ecological systems, but communication of model results is at the same time constrained by the complexity of the model responses. Here, we systematically modify a digital version of a real landscape to produce a set of model landscapes differing in the degree of heterogeneity and test how different landscapes affect abundance and occupancy of six model animal species in four different management scenarios using an agent-based model framework (ALMaSS). ALMaSS is capable of highly detailed modelling of individuals but the outputs can be complex and voluminous. We develop a statistic (the AOR-index) based on the abundance occupancy relationship to simplify presentation of model simulations and facilitate scenario comparisons. Scenario results demonstrate that species respond very differently to a particular land-use scenario and in some cases in opposite directions. The bird and mammal species generally showed larger responses than the invertebrates and changes in occupancy were often smaller than changes in abundance. The species-specific responses are caused by differences in habitat requirements and dispersal abilities, but the importance of such life history traits are affected by landscape dynamics and structure. Hence predictions of species-specific responses to land-use changes in terms of abundance and occupancy are greatly improved by incorporation in a model framework taking spatial and temporal dynamics into account. The use of the AOR-index simplifies the presentation of scenario comparison and provides and objective way to combine impacts across species. Its use, however, still requires management goals in order to evaluate scenario responses

Phenology of high-arctic butterflies and their floral resources: species-specific responses to climate change

Current global warming is particularly pronounced in the Arctic and arthropods are expected to respond rapidly to these changes. Long-term studies of individual arthropod species from the Arctic are, however, virtually absent. We examined butterfly specimens collected from yellow pitfall traps over 14 years (1996-2009) at Zackenberg in high-arctic, north-east Greenland. Specimens were previously sorted to the family level. We identified them to the species level and examined long-term species-specific phenological responses to recent summer warming. Two species were rare in the samples (Polaris fritillary Boloria polaris and Arctic blue Plebejus glandon) and statistical analyses of phenological responses were therefore restricted to the two most abundant species (Arctic fritillary, B. chariclea and Northern clouded yellow Colias hecla). Our analyses demonstrated a trend towards earlier flight seasons in B. chariclea, but not in C. hecla. The timing of onset, peak and end of the flight season in B. chariclea were closely related to snowmelt, July temperature and their interaction, whereas onset, peak and end of the flight season in C. hecla was only related to timing of snowmelt. The duration of the butterfly flight season was significantly positively related to the temporal overlap with floral resources in both butterfly species. We further demonstrate that yellow pitfall traps are a useful alternative to transect walks for butterfly recording in tundra habitats. More phenological studies of Arctic arthropods should be carried out at the species level and ideally be analysed in context with interacting species to assess how ongoing climate change will affect Arctic biodiversity in the near futur