An assessment of urban horticultural soil quality in the United Kingdom and its contribution to carbon storage.

As participation in urban horticulture grows, understanding the quality of urban horticultural soils is of increasing importance. Until now, case studies of individual cities or gardens have limited the potential of such studies to draw generalised conclusions. Here, we present the first national scale assessment of soil quality in allotments, a dominant form of urban horticulture in the United Kingdom. We sampled soils in 200 allotments in 10 urban areas across Great Britain. We assessed a range of soil quality indicators (carbon and nitrogen concentration, C:N ratio, bulk density, carbon density, pH) comparing them to the quality of soils in rural arable and horticultural land. We present the first estimate of nationwide carbon storage on allotments. We found that allotment gardeners consistently employ management practices conducive to high soil quality. Allotment soil quality differed significantly between soil types but in general soils were of a high quality: low bulk density (0.92 g cm-3) and high soil organic carbon concentration and density (58.2 mg g-1 and 58.1 mg cm-3 respectively). Allotment soil organic carbon concentration was 250% higher than in the surrounding arable and horticultural land. Covering only 0.0006% of Great Britain, allotments contribute a disproportionate 0.05-0.14% of nationwide total organic carbon stocks. This national-scale study provides compelling evidence that small-scale urban horticultural production, unlike conventional horticulture, does not degrade soil quality. Indeed, allotments hold a small but previously unaccounted for carbon stock nationally. Urban horticultural land is a vital part of the urban landscape with effectively functioning soils that should be protected. As public demand for urban horticultural land rises and policy-makers from local to trans-national levels of governance advocate for urban food production, our findings demonstrate that urban horticulture can protect or enhance the ecosystem services provided by soils in cities and towns where the majority of people live.

Estimating food production in an urban landscape.

There is increasing interest in urban food production for reasons of food security, environmental sustainability, social and health benefits. In developed nations urban food growing is largely informal and localised, in gardens, allotments and public spaces, but we know little about the magnitude of this production. Here we couple own-grown crop yield data with garden and allotment areal surveys and urban fruit tree occurrence to provide one of the first estimates for current and potential food production in a UK urban setting. Current production is estimated to be sufficient to supply the urban population with fruit and vegetables for about 30 days per year, while the most optimistic model results suggest that existing land cultivated for food could supply over half of the annual demand. Our findings provide a baseline for current production whilst highlighting the potential for change under the scaling up of cultivation on existing land.

Feeding a city - Leicester as a case study of the importance of allotments for horticultural production in the UK.

The process of urbanization has detached a large proportion of the global population from involvement with food production. However, there has been a resurgence in interest in urban agriculture and there is widespread recognition by policy-makers of its potential contribution to food security. Despite this, there is little data on urban agricultural production by non-commercial small-scale growers. We combine citizen science data for self-provisioning crop yields with field-mapping and GIS-based analysis of allotments in Leicester, UK, to provide an estimate of allotment fruit and vegetable production at a city-scale. In addition, we examine city-scale changes in allotment land provision on potential crop production over the past century. The average area of individual allotment plots used to grow crops was 52%. Per unit area yields for the majority of crops grown in allotments were similar to those of UK commercial horticulture. We estimate city-wide allotment production of >1200 t of fruit and vegetables and 200 t of potatoes per annum, equivalent to feeding >8500 people. If the 13% of plots that are completely uncultivated were used this could increase production to >1400 t per annum, feeding ~10,000 people, however this production may not be located in areas where there is greatest need for increased access to fresh fruits and vegetables. The citywide contribution of allotment cultivation peaked in the 1950s when 475 ha of land was allotments, compared to 97 ha currently. This suggests a decline from >45,000 to <10,000 of people fed per annum. We demonstrate that urban allotments make a small but important contribution to the fruit and vegetable diet of a UK city. However, further urban population expansion will exert increasing development pressure on allotment land. Policy-makers should both protect allotments within cities, and embed urban agricultural land within future developments to improve local food security.

Urban meadows as an alternative to short mown grassland: effects of composition and height on biodiversity.

There are increasing calls to provide greenspace in urban areas, yet the ecological quality, as well as quantity, of greenspace is important. Short mown grassland designed for recreational use is the dominant form of urban greenspace in temperate regions but requires considerable maintenance and typically provides limited habitat value for most taxa. Alternatives are increasingly proposed, but the biodiversity potential of these is not well understood. In a replicated experiment across six public urban greenspaces, we used nine different perennial meadow plantings to quantify the relative roles of floristic diversity and height of sown meadows on the richness and composition of three taxonomic groups: plants, invertebrates, and soil microbes. We found that all meadow treatments were colonized by plant species not sown in the plots, suggesting that establishing sown meadows does not preclude further locally determined grassland development if management is appropriate. Colonizing species were rarer in taller and more diverse plots, indicating competition may limit invasion rates. Urban meadow treatments contained invertebrate and microbial communities that differed from mown grassland. Invertebrate taxa responded to changes in both height and richness of meadow vegetation, but most orders were more abundant where vegetation height was longer than mown grassland. Order richness also increased in longer vegetation and Coleoptera family richness increased with plant diversity in summer. Microbial community composition seems sensitive to plant species composition at the soil surface (0-10 cm), but in deeper soils (11-20 cm) community variation was most responsive to plant height, with bacteria and fungi responding differently. In addition to improving local residents' site satisfaction, native perennial meadow plantings can produce biologically diverse grasslands that support richer and more abundant invertebrate communities, and restructured plant, invertebrate, and soil microbial communities compared with short mown grassland. Our results suggest that diversification of urban greenspace by planting urban meadows in place of some mown amenity grassland is likely to generate substantial biodiversity benefits, with a mosaic of meadow types likely to maximize such benefits.

Optimising UK urban road verge contributions to biodiversity and ecosystem services with cost-effective management.

Urban road verges can contain significant biodiversity, contribute to structural connectivity between other urban greenspaces, and due to their proximity to road traffic are well placed to provide ecosystem services. Using the UK as a case study we review and critically evaluate a broad range of evidence to assess how this considerable potential can be enhanced despite financial, contractual and public opinion constraints. Reduced mowing frequency and other alterations would enhance biodiversity, aesthetics and pollination services, whilst delivering costs savings and potentially being publically acceptable. Retaining mature trees and planting additional ones is favourable to residents and would enhance biodiversity, pollution and climate regulation, carbon storage, and stormwater management. Optimising these services requires improved selection of tree species, and creating a more diverse tree stock. Due to establishment costs additional tree planting and maintenance could benefit from payment for ecosystem service schemes. Verges could also provide areas for cultivation of biofuels and possibly food production. Maximising the contribution of verges to urban biodiversity and ecosystem services is economical and becoming an increasingly urgent priority as the road network expands and other urban greenspace is lost, requiring enhancement of existing greenspace to facilitate sustainable urban development.

What personal and environmental factors determine frequency of urban greenspace use?

For many people, urban greenspaces are the only places where they encounter the natural world. This is concerning as there is growing evidence demonstrating that human well-being is enhanced by exposure to nature. There is, therefore, a compelling argument to increase how frequently people use urban greenspaces. This may be achieved in two complementary ways by encouraging: (I) non-users to start visiting urban greenspaces; (II) existing users to visit more often. Here we examine the factors that influence frequency of greenspace visitation in the city of Sheffield, England. We demonstrate that people who visit a site least frequently state lower self-reported psychological well-being. We hypothesised that a combination of socio-demographic characteristics of the participants, and the biophysical attributes of the greenspaces that they were visiting, would be important in influencing visit frequency. However, socio-demographic characteristics (income, age, gender) were not found to be predictors. In contrast, some biophysical attributes of greenspaces were significantly related to use frequency. Frequent use was more likely when the time taken to reach a greenspace was shorter and for sites with a higher index of greenspace neglect, but were unrelated to tree cover or bird species richness. We related these results to the motivations that people provide for their visits. Infrequent users were more likely to state motivations associated with the quality of the space, while frequent users gave motivations pertaining to physical, repeated activities. This suggests that there may be no simple way to manage greenspaces to maximise their use across user cohorts as the motivations for visits are very different.

Quantifying preferences for the natural world using monetary and nonmonetary assessments of value.

Given that funds for biodiversity conservation are limited, there is a need to understand people's preferences for its different components. To date, such preferences have largely been measured in monetary terms. However, how people value biodiversity may differ from economic theory, and there is little consensus over whether monetary metrics are always appropriate or the degree to which other methods offer alternative and complementary perspectives on value. We used a choice experiment to compare monetary amounts recreational visitors to urban green spaces were willing to pay for biodiversity enhancement (increases in species richness for birds, plants, and aquatic macroinvertebrates) with self-reported psychological gains in well-being derived from visiting the same sites. Willingness-to-pay (WTP) estimates were significant and positive, and respondents reported high gains in well-being across 3 axes derived from environmental psychology theories (reflection, attachment, continuity with past). The 2 metrics were broadly congruent. Participants with above-median self-reported well-being scores were willing to pay significantly higher amounts for enhancing species richness than those with below-median scores, regardless of taxon. The socio-economic and demographic background of participants played little role in determining either their well-being or the probability of choosing a paying option within the choice experiment. Site-level environmental characteristics were only somewhat related to WTP, but showed strong associations with self-reported well-being. Both approaches are likely to reflect a combination of the environmental properties of a site and unobserved individual preference heterogeneity for the natural world. Our results suggest that either metric will deliver mutually consistent results in an assessment of environmental preferences, although which approach is preferable depends on why one wishes to measure values for the natural world.

Experimentally testing the accuracy of an extinction estimator: Solow's optimal linear estimation model.

Mathematical methods for inferring time to extinction have been widely applied but poorly tested. Optimal linear estimation (also called the 'Weibull' or 'Weibull extreme value' model) infers time to extinction from a temporal distribution of species sightings. Previous studies have suggested optimal linear estimation provides accurate estimates of extinction time for some species; however, an in-depth test of the technique is lacking. The use of data from wild populations to gauge the error associated with estimations is often limited by very approximate estimates of the actual extinction date and poor sighting records. Microcosms provide a system in which the accuracy of estimations can be tested against known extinction dates, whilst incorporating a variety of extinction rates created by changing environmental conditions, species identity and species richness. We present the first use of experimental microcosm data to exhaustively test the accuracy of one sighting-based method of inferring time of extinction under a range of search efforts, search regimes, sighting frequencies and extinction rates. Our results show that the accuracy of optimal linear estimation can be affected by both observer-controlled parameters, such as change in search effort, and inherent features of the system, such as species identity. Whilst optimal linear estimation provides generally accurate and precise estimates, the technique is susceptible to both overestimation and underestimation of extinction date. Microcosm experiments provide a framework within which the accuracy of extinction predictors can be clearly gauged. Variables such as search effort, search regularity and species identity can significantly affect the accuracy of estimates and should be taken into account when testing extinction predictors in the future.

Interactions between assembly order and temperature can alter both short- and long-term community composition.

Both the order in which species arrive in a community, and environmental conditions, such as temperature, are known to affect community structure. Little is known, however, about the potential for, and occurrence of, interactions between assembly history and the environment. Of particular, interest may be the interaction between temperature and community assembly dynamics, especially in the light of predicted global climatic change and the fundamental processes that are governed, through metabolic rate, by an individual's environmental temperature. We present, to our knowledge, the first experimental exploration of how the influence of assembly history, temperature, and the interaction between the two alters the structure of communities of competitors, using small-scale protist microcosm communities where temperature and assembly order were manipulated factorially. In our experiment, the most important driver of long-term abundance was temperature but long-lasting assembly order effects influenced the relationship between temperature and abundance. Any advantage of early colonization proved to be short-lived, and there was rarely any long-term advantage to colonizing a habitat before other species. The results presented here suggest that environmental conditions shape community composition, but that occasionally temperature could interact with the stochastic nature of community assembly to significantly alter future community composition, especially where temperature change has been large. This could have important implications for the dynamics of both rare and invasive species.

Environmental impact propagated by cross-system subsidy: chronic stream pollution controls riparian spider populations.

Resource subsidies between habitats are common and create the potential for the propagation of environmental impacts across system boundaries. However, recent understanding of the potential for subsidy-mediated cross-system impact propagations is limited and primarily based on passive flows of nutrients and detritus or short-term effects. Here, we assess the effects of sustained alterations in aquatic insect emergence (active subsidy pathway), due to chronic stream pollution, for riparian spiders. The sustained reduction in aquatic insect densities at the polluted reaches resulted in a marked decline in web spider population density and a shift in spider community composition. Our results provide the first evidence that stream pollution can control populations and community structure of terrestrial predators via sustained alterations in aquatic subsidies, emphasizing the role of subtle trophic linkages in the transmission of environmental impacts across ecosystem boundaries.

Fit, efficiency, and biology: some thoughts on judging food web models.

Revealing the processes that determine who eats whom, and thereby the structure of food webs, is a long running challenge in ecological research. Recent advances include development of new methods for measuring fit of models to observed food web data, and thereby testing which are the 'best' food web models. The best model could be considered the most efficient with relatively few parameters and high explanatory power. Another recent advance involves adding some additional biology to food web models in the form of foraging theory based on maximisation of energy intake as the predictor of species' diets in food webs. While it is interesting to compare efficiency among food web models, we believe that such comparisons at least should be interpreted with caution, since they do not account for any differences in motivation, formulation, and potential that might also exist among models. Furthermore, we see an important but somewhat neglected role for experimental tests of models of food web structure.

Urban domestic gardens (XIV): the characteristics of gardens in five cities.

Domestic gardens make substantial contributions to the provision of green space in urban areas. However, the ecological functions provided by such gardens depend critically on their configuration and composition. Here, we present the first detailed analysis of variation in the composition of urban gardens, in relation to housing characteristics and the nature of the surrounding landscape, across different cities in the United Kingdom. In all five cities studied (Belfast, Cardiff, Edinburgh, Leicester, and Oxford), garden size had an overwhelming influence on garden composition. Larger gardens supported more of the land-use types recorded, in greater extents, and were more likely to contain particular features, including tall trees and mature shrubs, areas of unmown grass and uncultivated land, vegetable patches, ponds, and composting sites. The proportional contribution of non-vegetated land-uses decreased as garden area increased. House age was less significant in determining the land-use within gardens, although older houses, which were more likely to be found further from the urban edge of the city, contained fewer hedges and greater areas of vegetation canopy >2 m in height. Current UK government planning recommendations will ultimately reduce the area of individual gardens and are thus predicted to result in fewer tall trees and, in particular, less vegetation canopy >2 m. This might be detrimental from ecological, aesthetic, social, and economic stand points.

Size, foraging, and food web structure.

Understanding what structures ecological communities is vital to answering questions about extinctions, environmental change, trophic cascades, and ecosystem functioning. Optimal foraging theory was conceived to increase such understanding by providing a framework with which to predict species interactions and resulting community structure. Here, we use an optimal foraging model and allometries of foraging variables to predict the structure of real food webs. The qualitative structure of the resulting model provides a more mechanistic basis for the phenomenological rules of previous models. Quantitative analyses show that the model predicts up to 65% of the links in real food webs. The deterministic nature of the model allows analysis of the model's successes and failures in predicting particular interactions. Predacious and herbivorous feeding interactions are better predicted than pathogenic, parasitoid, and parasitic interactions. Results also indicate that accurate prediction and modeling of some food webs will require incorporating traits other than body size and diet choice models specific to different types of feeding interaction. The model results support the hypothesis that individual behavior, subject to natural selection, determines individual diets and that food web structure is the sum of these individual decisions.

Psychological benefits of greenspace increase with biodiversity.

The world's human population is becoming concentrated into cities, giving rise to concerns that it is becoming increasingly isolated from nature. Urban public greenspaces form the arena of many people's daily contact with nature and such contact has measurable physical and psychological benefits. Here we show that these psychological benefits increase with the species richness of urban greenspaces. Moreover, we demonstrate that greenspace users can more or less accurately perceive species richness depending on the taxonomic group in question. These results indicate that successful management of urban greenspaces should emphasize biological complexity to enhance human well-being in addition to biodiversity conservation.

Daytime noise predicts nocturnal singing in urban robins.

Ambient noise interferes with the propagation of acoustic signals through the environment from sender to receiver. Over the past few centuries, urbanization and the development of busy transport networks have led to dramatic increases in the levels of ambient noise with which animal acoustic communications must compete. Here we show that urban European robins Erithacus rubecula, highly territorial birds reliant on vocal communication, reduce acoustic interference by singing during the night in areas that are noisy during the day. The effect of ambient light pollution, to which nocturnal singing in urban birds is frequently attributed, is much weaker than that of daytime noise.

Foraging biology predicts food web complexity.

Food webs, the networks of feeding links between species, are central to our understanding of ecosystem structure, stability, and function. One of the key aspects of food web structure is complexity, or connectance, the number of links expressed as a proportion of the total possible number of links. Connectance (complexity) is linked to the stability of webs and is a key parameter in recent models of other aspects of web structure. However, there is still no fundamental biological explanation for connectance in food webs. Here, we propose that constraints on diet breadth, driven by optimal foraging, provide such an explanation. We show that a simple diet breadth model predicts highly constrained values of connectance as an emergent consequence of individual foraging behavior. When combined with features of real food web data, such as taxonomic and trophic aggregation and cumulative sampling of diets, the model predicts well the levels of connectance and scaling of connectance with species richness, seen in real food webs. This result is a previously undescribed synthesis of foraging theory and food web theory, in which network properties emerge from the behavior of individuals and, as such, provides a mechanistic explanation of connectance currently lacking in food web models.

Species-energy relationships at the macroecological scale: a review of the mechanisms.

Correlations between the amount of energy received by an assemblage and the number of species that it contains are very general, and at the macro-scale such species-energy relationships typically follow a monotonically increasing curve. Whilst the ecological literature contains frequent reports of such relationships, debate on their causal mechanisms is limited and typically focuses on the role of energy availability in controlling the number of individuals in an assemblage. Assemblages from high-energy areas may contain more individuals enabling species to maintain larger, more viable populations, whose lower extinction risk elevates species richness. Other mechanisms have, however, also been suggested. Here we identify and clarify nine principal mechanisms that may generate positive species-energy relationships at the macro-scale. We critically assess their assumptions and applicability over a range of spatial scales, derive predictions for each and assess the evidence that supports or refutes them. Our synthesis demonstrates that all mechanisms share at least one of their predictions with an alternative mechanism. Some previous studies of species-energy relationships appear not to have recognised the extent of shared predictions, and this may detract from their contribution to the debate on causal mechanisms. The combination of predictions and assumptions made by each mechanism is, however, unique, suggesting that, in principle, conclusive tests are possible. Sufficient testing of all mechanisms has yet to be conducted, and no single mechanism currently has unequivocal support. Each may contribute to species-energy relationships in some circumstances, but some mechanisms are unlikely to act simultaneously. Moreover, a limited number appear particularly likely to contribute frequently to species-energy relationships at the macro-scale. The increased population size, niche position and diversification rate mechanisms are particularly noteworthy in this context.

Body size in ecological networks.

Body size determines a host of species traits that can affect the structure and dynamics of food webs, and other ecological networks, across multiple scales of organization. Measuring body size provides a relatively simple means of encapsulating and condensing a large amount of the biological information embedded within an ecological network. Recently, important advances have been made by incorporating body size into theoretical models that explore food web stability, the patterning of energy fluxes, and responses to perturbations. Because metabolic constraints underpin body-size scaling relationships, metabolic theory offers a potentially useful new framework within which to develop novel models to describe the structure and functioning of ecological networks and to assess the probable consequences of biodiversity change.

Interspecific abundance-occupancy relationships and the effects of disturbance: a test using microcosms.

The effects of disturbance on interspecific relationships between abundance and occupancy are tested using the results of a factorial experiment carried out with microcosm communities of protists. A positive relationship was documented whether marked disturbance was present or not; the pattern was neither a product of disturbance, nor was it destroyed by disturbance. Coefficients of determination, and the slopes and intercepts of abundance-occupancy relationships did not appear to change systematically with treatment. This robustness of the relationship may reflect the tendency for species to maintain approximately the same relative abundances and levels of occupancy in a broad range of circumstances.

The effects of between-habitat dispersal rate on protist communities and metacommunities in microcosms at two spatial scales.

The effect of manipulation of between-habitat dispersal rates in multiple patch systems was examined experimentally using protist communities in laboratory microcosms. Replicate "landscapes" of eight microcosms (patches) at two spatial scales (patch sizes) were inoculated with 13 species of protists. Dispersal was carried out by transferring a small random sample of medium and protists from one randomly selected microcosm to another within a landscape. Four dispersal rates (24, 6, 2 and 0 transfers very 3 days) were used, and the microcosms were sampled after 6 and 12 weeks. Patch size had a consistent effect on within-path (community) and within-landscape (metacommunity) diversity, both being lower in small patch systems. Higher dispersal rates had a slight effect on community and metacommunity diversity after 12 weeks, with a tendency for higher dispersal to slightly offset the rate of loss of species. Both dispersal and patch size had effects on the abundance of many individual species, though in a variety of ways. The individual species results suggest that extinction is selective with respect to both patch size and dispersal rate treatments, and may be influenced by species interactions. It seems likely that in metacommunity systems of this sort, rather than mainland-island systems, the potential effect of between-patch dispersal rate in rescuing and recolonizing where local extinctions occur may be much reduced by the effect of selective extinction, relative to that expected under the assumption of random extinction.