Alistair G. Auffret

The spatial and temporal components of functional connectivity in fragmented landscapes.

Connectivity is key for understanding how ecological systems respond to the challenges of land-use change and habitat fragmentation. Structural and functional connectivity are both established concepts in ecology, but the temporal component of connectivity deserves more attention. Whereas functional connectivity is often associated with spatial patterns (spatial functional connectivity), temporal functional connectivity relates to the persistence of organisms in time, in the same place. Both temporal and spatial processes determine biodiversity responses to changes in landscape structure, and it is therefore necessary that all aspects of connectivity are considered together. In this perspective, we use a case study to outline why we believe that both the spatial and temporal components of functional connectivity are important for understanding biodiversity patterns in the present-day landscape, and how they can also help us to make better-informed decisions about conserving and restoring landscapes and improving resilience to future change.

Plant functional connectivity – integrating landscape structure and effective dispersal

Summary 1.Dispersal is essential for species to survive the threats of habitat destruction and climate change. Combining descriptions of dispersal ability with those of landscape structure, the concept of functional connectivity has been popular for understanding and predicting species’ spatial responses to environmental change. 2.Following recent advances, the functional connectivity concept is now able to move beyond landscape structure to consider more explicitly how other external factors such as climate and resources affect species movement. We argue that these factors, in addition to a consideration of the complete dispersal process, are critical for an accurate understanding of functional connectivity for plant species in response to environmental change. 3.We use recent advances in dispersal, landscape and molecular ecology to describe how a range of external factors can influence effective dispersal in plant species, and how the resulting functional connectivity can be assessed. 4.Synthesis. We define plant functional connectivity as the effective dispersal of propagules or pollen among habitat patches in a landscape. Plant functional connectivity is determined by a combination of landscape structure, interactions between plant, environment and dispersal vectors, and the successful establishment of individuals. We hope that this consolidation of recent research will help focus future connectivity research and conservation. This article is protected by copyright. All rights reserved.

Humans as Long-Distance Dispersers of Rural Plant Communities

Humans are known for their capacity to disperse organisms long distances. Long-distance dispersal can be important for species threatened by habitat destruction, but research into human-mediated dispersal is often focused upon few and/or invasive species. Here we use citizen science to identify the capacity for humans to disperse seeds on their clothes and footwear from a known species pool in a valuable habitat, allowing for an assessment of the fraction and types of species dispersed by humans in an alternative context. We collected material from volunteers cutting 48 species-rich meadows throughout Sweden. We counted 24 354 seeds of 197 species, representing 34% of the available species pool, including several rare and protected species. However, 71 species (36%) are considered invasive elsewhere in the world. Trait analysis showed that seeds with hooks or other appendages were more likely to be dispersed by humans, as well as those with a persistent seed bank. More activity in a meadow resulted in more dispersal, both in terms of species and representation of the source communities. Average potential dispersal distances were measured at 13 km. We consider humans capable seed dispersers, transporting a significant proportion of the plant communities in which they are active, just like more traditional vectors such as livestock. When rural populations were larger, people might have been regular and effective seed dispersers, and the net rural-urban migration resulting in a reduction in humans in the landscape may have exacerbated the dispersal failure evident in declining plant populations today. With the fragmentation of habitat and changes in land use resulting from agricultural change, and the increased mobility of humans worldwide, the dispersal role of humans may have shifted from providers of regular local and landscape dispersal to providers of much rarer long-distance and regional dispersal, and international invasion.

Scale-dependent diversity effects of seed dispersal by a wild herbivore in fragmented grasslands

Dispersal limitation between habitat fragments is a known driver of landscape-scale biodiversity loss. In Europe, agricultural intensification during the twentieth century resulted in losses of both grassland habitat and traditional grassland seed dispersal vectors such as livestock. During the same period, populations of large wild herbivores have increased in the landscape. Usually studied in woodland ecosystems, these animals are found to disperse seeds from grasslands and other open habitats. We studied endozoochorous seed dispersal by roe deer (Capreolus capreolus) in fragmented grasslands and grassland remnants, comparing dispersed subcommunities of plant species to those in the established vegetation and the seed bank. A total of 652 seedlings of 67 species emerged from 219 samples of roe deer dung. This included many grassland species, and several local grassland specialists. Dispersal had potentially different effects on diversity at different spatial scales. Almost all sites received seeds of species not observed in the vegetation or seed bank at that site, suggesting that local diversity might not be dispersal limited. This pattern was less evident at the landscape scale, where fewer new species were introduced. Nonetheless, long-distance dispersal by large wild herbivores might still provide connectivity between fragmented habitats within a landscape in the areas in which they are active. Finally, as only a subset of the available species were found to disperse in space as well as time, the danger of future biodiversity loss might still exist in many isolated grassland habitats.

Spatial scale and specialization affect how biogeography and functional traits predict long‐term patterns of community turnover

Summary Immigration, extirpation and persistence of individual populations of species are key processes determining community responses to environmental change. However, they are difficult to study over long time periods without corresponding historical and modern-day species occurrences. We used historical and present-day plant species occurrence data from two different spatial scales (resolutions) to investigate the plant community turnover during the 20th century in a Baltic Sea archipelago. Patterns of turnover were analysed in relation to plant functional traits relating to dispersal and competition/persistence, as well as biogeographical variables. Turnover was largely driven by interactions between functional traits and measures of area, connectivity and distance to mainland. However, the combinations of traits and biogeographical variables that were most important for predicting immigration and extirpation differed between data sets, and between species associated with grassland management and the entire species pool. Taller plants were more likely to persist regardless of scale and biogeography, reflecting the grazing abandonment that occurred in the study area. Interactions between dispersal traits and biogeography were related to immigrations when the entire species pool was considered. However, increased dispersal potential, a smaller island size and increasing distance to mainland combined to promote extirpations in management-associated species. A perennial life span and seed banking contributed to species persistence. At the larger spatial scale, trait-driven turnover was not mediated by the biogeographical context. We showed that it is important to consider functional traits, biogeographical variables and their interactions when analysing community turnover over time. Furthermore, we found that the understanding of how combinations of traits and biogeography predict turnover depends on the source and spatial scale of the available data, and the species pool analysed.

Extinction risk from climate change is reduced by microclimatic buffering

Protecting biodiversity against the impacts of climate change requires effective conservation strategies that safeguard species at risk of extinction1. Microrefugia allowed populations to survive adverse climatic conditions in the past2,3, but their potential to reduce extinction risk from anthropogenic warming is poorly understood3–5, hindering our capacity to develop robust in situ measures to adapt conservation to climate change6. Here, we show that microclimatic heterogeneity has strongly buffered species against regional extirpations linked to recent climate change. Using more than five million distribution records for 430 climate-threatened and range-declining species, population losses across England are found to be reduced in areas where topography generated greater variation in the microclimate. The buffering effect of topographic microclimates was strongest for those species adversely affected by warming and in areas that experienced the highest levels of warming: in such conditions, extirpation risk was reduced by 22% for plants and by 9% for insects. Our results indicate the critical role of topographic variation in creating microrefugia, and provide empirical evidence that microclimatic heterogeneity can substantially reduce extinction risk from climate change.Topographic variations result in microclimatic heterogeneity that can substantially reduce extinction risk from climate change, according to a study of 430 climate-threatened and range-declining species in England.

Methodological bias in the seed bank flora holds significant implications for understanding seed bank community functions.

Persistent seed banks are a key plant regeneration strategy, buffering environmental variation to allow population and species persistence. Understanding seed bank functioning within herb layer dynamics is therefore important. However, rather than assessing emergence from the seed bank in herb layer gaps, most studies evaluate the seed bank functioning via a greenhouse census. We hypothesise that greenhouse data may not reflect seed bank-driven emergence in disturbance gaps due to methodological differences. Failure in detecting (specialist) species may then introduce methodological bias into the ecological interpretation of seed bank functions using greenhouse data. The persistent seed bank was surveyed in 40 semi-natural grassland plots across a fragmented landscape, quantifying seedling emergence in both the greenhouse and in disturbance gaps. Given the suspected interpretational bias, we tested whether each census uncovers similar seed bank responses to fragmentation. Seed bank characteristics were similar between censuses. Census type affected seed bank composition, with >25% of species retrieved better by either census type, dependent on functional traits including seed longevity, production and size. Habitat specialists emerged more in disturbance gaps than in the greenhouse, while the opposite was true for ruderal species. Both censuses uncovered fragmentation-induced seed bank patterns. Low surface area sampling, larger depth of sampling and germination conditions cause underrepresentation of the habitat-specialised part of the persistent seed bank flora during greenhouse censuses. Methodological bias introduced in the recorded seed bank data may consequently have significant implications for the ecological interpretation of seed bank community functions based on greenhouse data.

HistMapR: Rapid digitization of historical land‐use maps in R

1. Habitat destruction and degradation represent serious threats to biodiversity, and quantification of land-use change over time is important for understanding the consequences of these changes to organisms and ecosystem service provision. 2. Comparing land use between maps from different time periods allows estimation of the magnitude of habitat change in an area. However, digitizing historical maps manually is time-consuming and analyses of change are usually carried out at small spatial extents or at low resolutions. 3. HistMapR contains a number of functions that can be used to semi-automatically digitize historical land use according to a maps colours, as defined by the RGB bands of the raster image. We test the method on different historical land-use map series and compare results to manual digitizations. 4. Digitization is fast, and agreement with manually digitized maps of around 80–90% meets common targets for image classification. We hope that the ability to quickly classify large areas of historical land use will promote the inclusion of land-use change into analyses of biodiversity, species distributions and ecosystem services.

Land uplift creates important meadow habitat and a potential original niche for grassland species

Semi-natural grasslands have been severely affected by agricultural land-use change. However, the isostatic land adjustment following deglaciation in the Northern Hemisphere means that new land is continually being created in coastal areas. We modelled isostatic adjustment during the last 4000 years in a region of the Baltic coast to estimate the emergence of potential grassland habitat. We also compared the α and β diversity of existing managed and abandoned coastal meadows, and assessed their contribution to biodiversity at landscape scales. We estimated that half the 7866 km2 of emerging land had the potential to become coastal meadow habitat, which is an order of magnitude larger than the total area of all valuable semi-natural grassland in the study region today. The small area of managed coastal habitat remaining was found to have a disproportionate influence on the richness of threatened species at landscape scales, but our results also show that continued management is essential for the maintenance of grassland biodiversity. Our combination of approaches identifies uplifted coastal meadows as an additional original niche for grassland plant species, while highlighting that low-intensity disturbance through grassland management is essential for the maintenance of diversity at multiple scales.

Super-regional land-use change and effects on the grassland specialist flora

Habitat loss through land-use change is the most pressing threat to biodiversity worldwide. European semi-natural grasslands have suffered an ongoing decline since the early twentieth century, but we have limited knowledge of how grassland loss has affected biodiversity across large spatial scales. We quantify land-use change over 50–70 years across a 175,000 km2 super-region in southern Sweden, identifying a widespread loss of open cover and a homogenisation of landscape structure, although these patterns vary considerably depending on the historical composition of the landscape. Analysing species inventories from 46,796 semi-natural grasslands, our results indicate that habitat loss and degradation have resulted in a decline in grassland specialist plant species. Local factors are the best predictors of specialist richness, but the historical landscape predicts present-day richness better than the contemporary landscape. This supports the widespread existence of time-lagged biodiversity responses, indicating that further species losses could occur in the future.Biodiversity declines can be difficult to attribute to habitat loss at large spatial scales. Here, the authors document land-use change in Sweden at high spatial resolution over 70 years, showing that habitat loss at local and landscape scales is associated with reduced grassland biodiversity.