Assessing multidimensional sustainability: Lessons from Brazil's social protection programs.

Examining linkages among multiple sustainable development outcomes is key for understanding sustainability transitions. Yet rigorous evidence on social and environmental outcomes of sustainable development policies remains scarce. We conduct a national-level analysis of Brazil's flagship social protection program, Zero Hunger (ZH), which aims to reduce food insecurity and poverty. Using data from rural municipalities across Brazil and quasi-experimental causal inference techniques, we assess relationships between social protection investment and outcomes related to sustainable development goals (SDGs): "no poverty" (SDG 1), "zero hunger" (SDG 2), and "health and well being" (SDG 3). We also assess potential perverse outcomes arising from agricultural development impacting "climate action" (SDG 13) and "life on land" (SDG 15) via clearance of natural vegetation. Despite increasing daily per capita protein and kilocalorie production, summed ZH investment did not alleviate child malnutrition or infant mortality and negligibly influenced multidimensional poverty. Higher investment increased natural vegetation cover in some biomes but increased losses in the Cerrado and especially the Pampa. Effects varied substantially across subprograms. Conditional cash transfer (Bolsa Familia [BF]) was mainly associated with nonbeneficial impacts but increased protein production and improved educational participation in some states. The National Program to Strengthen Family Farming (PRONAF) was typically associated with increased food production (protein and calories), multidimensional poverty alleviation, and changes in natural vegetation. Our results inform policy development by highlighting successful elements of Brazil's ZH program, variable outcomes across divergent food security dimensions, and synergies and trade-offs between sustainable development goals, including environmental protection.

Avian species richness and tropical urbanization gradients: Effects of woodland retention and human disturbance.

Urbanization is a major driver of tropical biodiversity loss. In temperate regions avian species richness-urbanization intensity relationships typically exhibit unimodal patterns, with peak richness at intermediate urbanization levels. In tropical regions, the form of such relationships and the extent to which they are moderated by patches of seminatural habitat are unclear. We address these questions in Bangkok, Thailand (one of the largest and most rapidly expanding tropical mega-cities) and generate conservation recommendations for tropical biodiversity in urban locations. We use repeated point count surveys at a random location, and the largest available woodland patch, in 150 1 km × 1 km grid cells selected along the urbanization gradient. Woodland patches support higher species richness compared with randomized locations (except for non-natives), and avian species richness declines linearly with increasing urbanization. The contrast with unimodal patterns in temperate regions is probably driven by divergent patterns of habitat heterogeneity along tropical and temperate urbanization gradients. Moreover, we provide novel evidence that retaining patches of urban woodland moderates adverse impacts of urbanization on avian species richness. For most species groups, the benefits of woodland increase as urbanization intensifies, despite such woodland patches being very small (mean of 0.38 ha). Avian species richness in woodland patches is maximized, and community composition less similar to that in randomized locations, when woodland patches are larger and visited by fewer people. Assemblages of forest-dependent species (which provide additional ecological functions) have higher richness, and are less similar to those in randomized locations, in patches of woodland with higher tree species richness and biomass. Finally, species richness in randomized sites is greatest when they are closer to woodland patches, and such assemblages more closely resemble those of woodland sites. Our work highlights four strategies for tropical urban bird conservation: (1) conserving woodland patches across the urbanization gradient regardless of patch size, (2) improving the quality of existing woodland by increasing tree biomass and diversity, (3) creating additional woodland that is well distributed throughout the urban area to minimize effects of habitat isolation and (4) reducing human disturbance, especially in areas of the highest habitat quality, while ensuring that the benefits of connecting people to nature are realized in other locations.

Phenological responses in a sycamore-aphid-parasitoid system and consequences for aphid population dynamics: A 20 year case study.

Species interactions have a spatiotemporal component driven by environmental cues, which if altered by climate change can drive shifts in community dynamics. There is insufficient understanding of the precise time windows during which inter-annual variation in weather drives phenological shifts and the consequences for mismatches between interacting species and resultant population dynamics-particularly for insects. We use a 20 year study on a tri-trophic system: sycamore Acer pseudoplatanus, two associated aphid species Drepanosiphum platanoidis and Periphyllus testudinaceus and their hymenopteran parasitoids. Using a sliding window approach, we assess climatic drivers of phenology in all three trophic levels. We quantify the magnitude of resultant trophic mismatches between aphids and their plant hosts and parasitoids, and then model the impacts of these mismatches, direct weather effects and density dependence on local-scale aphid population dynamics. Warmer temperatures in mid-March to late-April were associated with advanced sycamore budburst, parasitoid attack and (marginally) D. platanoidis emergence. The precise time window during which spring weather advances phenology varies considerably across each species. Crucially, warmer temperatures in late winter delayed the emergence of both aphid species. Seasonal variation in warming rates thus generates marked shifts in the relative timing of spring events across trophic levels and mismatches in the phenology of interacting species. Despite this, we found no evidence that aphid population growth rates were adversely impacted by the magnitude of mismatch with their host plants or parasitoids, or direct impacts of temperature and precipitation. Strong density dependence effects occurred in both aphid species and probably buffered populations, through density-dependent compensation, from adverse impacts of the marked inter-annual climatic variation that occurred during the study period. These findings explain the resilience of aphid populations to climate change and uncover a key mechanism, warmer winter temperatures delaying insect phenology, by which climate change drives asynchronous shifts between interacting species.

Food availability limits avian reproduction in the city: An experimental study on great tits Parus major.

The altered ecological and environmental conditions in towns and cities strongly affect demographic traits of urban animal populations, for example avian reproductive success is often reduced. Previous work suggests that this is partly driven by low insect availability during the breeding season, but robust experimental evidence that supports this food limitation hypothesis is not yet available. We tested core predictions of the food limitation hypothesis using a controlled experiment that provided supplementary insect food (nutritionally enhanced mealworms supplied daily to meet 40%-50% of each supplemented brood's food requirements) to great tit nestlings in urban and forest habitats. We measured parental provisioning rates and estimated the amount of supplementary food consumed by control and experimental nestlings, and assessed their body size and survival rates. Provisioning rates were similar across habitats and control and supplemented broods, but supplemented (and not control) broods consumed large quantities of supplementary food. As predicted by the food limitation hypothesis we found that nestlings in (a) urban control broods had smaller body size and nestling survival rates than those in forest control broods; (b) forest supplemented and control broods had similar body size and survival rates; (c) urban supplemented nestlings had larger body size and survival rates than those in urban control broods; and crucially (d) urban supplemented broods had similar body size and survival rates to nestlings in forest control broods. Our results provide rare experimental support for the strong negative effects of food limitation during the nestling rearing period on urban birds' breeding success. Furthermore, the fact that supplementary food almost completely eliminated habitat differences in survival rate and nestling body size suggest that urban stressors other than food shortage contributed relatively little to the reduced avian breeding success. Finally, given the impacts of the amount of supplementary food that we provided and taking clutch size differences into account, our results suggest that urban insect populations in our study system would need to be increased by a factor of at least 2.5 for urban and forest great tits to have similar reproductive success.

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.

Quantifying carbon stocks in shifting cultivation landscapes under divergent management scenarios relevant to REDD.

Shifting cultivation dominates many tropical forest regions. It is expanding into old-growth forests, and fallow period duration is rapidly decreasing, limiting secondary forest recovery. Shifting cultivation is thus a major driver of carbon emissions through deforestation and forest degradation, and of biodiversity loss. The impacts of shifting cultivation on carbon stocks have rarely been quantified, and the potential for carbon-based payments for ecosystem services (PES), such as REDD+, to protect carbon in shifting cultivation landscapes is unknown. We present empirical data on aboveground carbon stocks in old-growth forest and shifting cultivation landscapes in northeast India, a hotspot of threatened biodiversity. We then model landscape-level carbon stocks under business-as-usual scenarios, via expansion into the old-growth forest or decreasing fallow periods, and intervention scenarios in which REDD+ is used to either reduce deforestation of primary or secondary forest or increase fallow period duration. We found substantial recovery of carbon stocks as secondary forest regenerates, with a 30-yr fallow storing about one-half the carbon of an old-growth forest. Business-as-usual scenarios led to substantial carbon loss, with an 80% reduction following conversion of old-growth forest to a 30-yr shifting cultivation cycle and, relative to a 30-yr cultivation landscape, a 70% reduction when switching to a 5-yr cultivation cycle. Sparing old-growth forests from deforestation using protected areas and intensifying cropping in the remaining area of shifting cultivation is the most optimal strategy for carbon storage. In areas lacking old-growth forest, substantial carbon stocks accumulate over time by sparing fallows for permanent forest regeneration. Successful implementation of REDD+ in shifting cultivation landscapes can help avert global climate change by protecting forest carbon, with likely co-benefits for biodiversity.

Impact of urbanization on abundance and phenology of caterpillars and consequences for breeding in an insectivorous bird.

Urbanization can have marked effects on plant and animal populations' phenology, population size, predator-prey, interactions and reproductive success. These aspects are rarely studied simultaneously in a single system, and some are rarely investigated, e.g., how insect phenology responds to urban development. Here, we study a tri-trophic system of trees, phytophagous insects (caterpillars), and insectivorous birds (Great Tits) to assess how urbanization influences (1) the phenology of each component of this system, (2) insect abundance, and (3) avian reproductive success. We use data from two urban and two forest sites in Hungary, central Europe, collected over four consecutive years. Despite a trend of earlier leaf emergence in urban sites, there is no evidence for an earlier peak in caterpillar abundance. Thus, contrary to the frequently stated prediction in the literature, the earlier breeding of urban bird populations is not associated with an earlier peak in caterpillar availability. Despite this the seasonal dynamics of caterpillar biomass exhibited striking differences between habitat types with a single clear peak in forests, and several much smaller peaks in urban sites. Caterpillar biomass was higher in forests than urban areas across the entire sampling period, and between 8.5 and 24 times higher during the first brood's chick-rearing period. This higher biomass was not associated with taller trees in forest sites, or with tree species identity, and occurred despite most of our focal trees being native to the study area. Urban Great Tits laid smaller clutches, experienced more frequent nestling mortality from starvation, reared fewer offspring to fledging age, and their fledglings had lower body mass. Our study strongly indicates that food limitation is responsible for lower avian reproductive success in cities, which is driven by reduced availability of the preferred nestling diet, i.e., caterpillars, rather than phenological shifts in the timing of peak food availability.

'Green' on the ground but not in the air: Pro-environmental attitudes are related to household behaviours but not discretionary air travel.

The rise in greenhouse gas emissions from air travel could be reduced by individuals voluntarily abstaining from, or reducing, flights for leisure and recreational purposes. In theory, we might expect that people with pro-environmental value orientations and concerns about the risks of climate change, and those who engage in more pro-environmental household behaviours, would also be more likely to abstain from such voluntary air travel, or at least to fly less far. Analysis of two large datasets from the United Kingdom, weighted to be representative of the whole population, tested these associations. Using zero-inflated Poisson regression models, we found that, after accounting for potential confounders, there was no association between individuals' environmental attitudes, concern over climate change, or their routine pro-environmental household behaviours, and either their propensity to take non-work related flights, or the distances flown by those who do so. These findings contrasted with those for pro-environmental household behaviours, where associations with environmental attitudes and concern were observed. Our results offer little encouragement for policies aiming to reduce discretionary air travel through pro-environmental advocacy, or through 'spill-over' from interventions to improve environmental impacts of household routines.

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.

Meeting sustainable development goals via robotics and autonomous systems.

Robotics and autonomous systems are reshaping the world, changing healthcare, food production and biodiversity management. While they will play a fundamental role in delivering the UN Sustainable Development Goals, associated opportunities and threats are yet to be considered systematically. We report on a horizon scan evaluating robotics and autonomous systems impact on all Sustainable Development Goals, involving 102 experts from around the world. Robotics and autonomous systems are likely to transform how the Sustainable Development Goals are achieved, through replacing and supporting human activities, fostering innovation, enhancing remote access and improving monitoring. Emerging threats relate to reinforcing inequalities, exacerbating environmental change, diverting resources from tried-and-tested solutions and reducing freedom and privacy through inadequate governance. Although predicting future impacts of robotics and autonomous systems on the Sustainable Development Goals is difficult, thoroughly examining technological developments early is essential to prevent unintended detrimental consequences. Additionally, robotics and autonomous systems should be considered explicitly when developing future iterations of the Sustainable Development Goals to avoid reversing progress or exacerbating inequalities.

Integrating solutions to adapt cities for climate change.

Record climate extremes are reducing urban liveability, compounding inequality, and threatening infrastructure. Adaptation measures that integrate technological, nature-based, and social solutions can provide multiple co-benefits to address complex socioecological issues in cities while increasing resilience to potential impacts. However, there remain many challenges to developing and implementing integrated solutions. In this Viewpoint, we consider the value of integrating across the three solution sets, the challenges and potential enablers for integrating solution sets, and present examples of challenges and adopted solutions in three cities with different urban contexts and climates (Freiburg, Germany; Durban, South Africa; and Singapore). We conclude with a discussion of research directions and provide a road map to identify the actions that enable successful implementation of integrated climate solutions. We highlight the need for more systematic research that targets enabling environments for integration; achieving integrated solutions in different contexts to avoid maladaptation; simultaneously improving liveability, sustainability, and equality; and replicating via transfer and scale-up of local solutions. Cities in systematically disadvantaged countries (sometimes referred to as the Global South) are central to future urban development and must be prioritised. Helping decision makers and communities understand the potential opportunities associated with integrated solutions for climate change will encourage urgent and deliberate strides towards adapting cities to the dynamic climate reality.

Independent colonization of multiple urban centres by a formerly forest specialist bird species.

Urban areas are expanding rapidly, but a few native species have successfully colonized them. The processes underlying such colonization events are poorly understood. Using the blackbird Turdus merula, a former forest specialist that is now one of the most common urban birds in its range, we provide the first assessment of two contrasting urban colonization models. First, that urbanization occurred independently. Second, that following initial urbanization, urban-adapted individuals colonized other urban areas in a leapfrog manner. Previous analyses of spatial patterns in the timing of blackbird urbanization, and experimental introductions of urban and rural blackbirds to uncolonized cities, suggest that the leapfrog model is likely to apply. We found that, across the western Palaearctic, urban blackbird populations contain less genetic diversity than rural ones, urban populations are more strongly differentiated from each other than from rural populations and assignment tests support a rural source population for most urban individuals. In combination, these results provide much stronger support for the independent urbanization model than the leapfrog one. If the former model predominates, colonization of multiple urban centres will be particularly difficult when urbanization requires genetic adaptations, having implications for urban species diversity.

Spatial scale, abundance and the species-energy relationship in British birds.

1. The spatial scale of analysis may influence the nature, strength and underlying drivers of macroecological patterns, one of the most frequently discussed of which is the relationship between species richness and environmental energy availability. 2. It has been suggested that species-energy relationships are hump-shaped at fine spatial grains and consistently positive at larger regional grains. The exact nature of this scale dependency is, however, the subject of much debate as relatively few studies have investigated species-energy relationships for the same assemblage across a range of spatial grains. Here, we contrast species-energy relationships for the British breeding avifauna at spatial grains of 1 km x 1 km, 2 km x 2 km and 10 km x 10 km plots, while maintaining a constant spatial extent. 3. Analyses were principally conducted using data on observed species richness. While survey work may fail to detect some species, observed species richness and that estimated using nonparametric techniques were strongly positively correlated with each other, and thus exhibit very similar spatial patterns. Moreover, the forms of species-energy relationships using observed and estimated species richness were statistically indistinguishable from each other. 4. Positive decelerating species-energy relationships arise at all three spatial grains. There is little evidence that the explanatory power of these relationships varies with spatial scale. However, ratios of regional (large-scale) to local (small-scale) species richness decrease with increasing energy availability, indicating that local richness responds to energy with a steeper gradient than does regional richness. Local assemblages thus sample a greater proportion of regional richness at higher energy levels, suggesting that spatial turnover of species richness is lower in high-energy regions. Similarly, a crude measure of temporal turnover, the ratio of cumulative species richness over a 4-year period to species richness in a single year, is lower in high-energy regions. These negative relationships between turnover and energy appear to be causal as both total and mean occupancy per species increases with energy. 5. While total density in 1 km x 1 km plots correlates positively with energy availability, such relationships are very weak for mean density per species. This suggests that the observed association between total abundance and species richness may not be mediated by population extinction rates, as predicted by the more individuals hypothesis. 6. The sampling mechanism suggests that species-energy relationships arise as high-energy areas support a greater number of individuals, and that random allocation of these individuals to local areas from a regional assemblage will generate species-energy relationships. While randomized local species-energy relationships are linear and positive, predicted richness is consistently greater than that observed. The mismatch between the observed and randomized species-energy relationships probably arises as a consequence of the aggregated nature of species distributions. The sampling mechanism, together with species spatial aggregation driven by limited habitat availability, may thus explain the species-energy relationship observed at this spatial scale.

The Effects of Arbuscular Mycorrhizal Fungal Colonisation on Nutrient Status, Growth, Productivity, and Canker Resistance of Apple (Malus pumila).

We assess whether arbuscular mycorrhizal fungi (AMF) improve growth, nutritional status, phenology, flower and fruit production, and disease resistance in woody perennial crops using apple (Malus pumila) as a study system. In a fully factorial experiment, young trees were grown for 3 years with or without AMF (Funneliformis mosseae and Rhizophagus irregularis), and with industrial standard fertiliser applications or restricted fertiliser (10% of standard). We use two commercial scions (Dabinett and Michelin) and rootstocks (MM111 and MM106). Industrial standard fertiliser applications reduced AMF colonisation and root biomass, potentially increasing drought sensitivity. Mycorrhizal status was influenced by above ground genotypes (scion type) but not rootstocks, indicating strong interactions between above and below ground plant tissue. The AMF inoculation significantly increased resistance to Neonectria ditissima, a globally economically significant fungal pathogen of apple orchards, but did not consistently alter leaf nutrients, growth, phenology or fruit and flower production. This study significantly advances understanding of AMF benefits to woody perennial crops, especially increased disease resistance which we show is not due to improved tree nutrition or drought alleviation. Breeding programmes and standard management practises can limit the potential for these benefits.

Tritrophic phenological match-mismatch in space and time.

Increasing temperatures associated with climate change may generate phenological mismatches that disrupt previously synchronous trophic interactions. Most work on mismatch has focused on temporal trends, whereas spatial variation in the degree of trophic synchrony has largely been neglected, even though the degree to which mismatch varies in space has implications for meso-scale population dynamics and evolution. Here we quantify latitudinal trends in phenological mismatch, using phenological data on an oak-caterpillar-bird system from across the UK. Increasing latitude delays phenology of all species, but more so for oak, resulting in a shorter interval between leaf emergence and peak caterpillar biomass at northern locations. Asynchrony found between peak caterpillar biomass and peak nestling demand of blue tits, great tits and pied flycatchers increases in earlier (warm) springs. There is no evidence of spatial variation in the timing of peak nestling demand relative to peak caterpillar biomass for any species. Phenological mismatch alone is thus unlikely to explain spatial variation in population trends. Given projections of continued spring warming, we predict that temperate forest birds will become increasingly mismatched with peak caterpillar timing. Latitudinal invariance in the direction of mismatch may act as a double-edged sword that presents no opportunities for spatial buffering from the effects of mismatch on population size, but generates spatially consistent directional selection on timing, which could facilitate rapid evolutionary change.

Contrasting effects of visiting urban green-space and the countryside on biodiversity knowledge and conservation support.

Conservation policy frequently assumes that increasing people's exposure to green-space enhances their knowledge of the natural world and desire to protect it. Urban development is, however, considered to be driving declining connectedness to nature. Despite this the evidence base supporting the assumption that visiting green-spaces promotes biodiversity knowledge and conservation support, and the impacts of urbanization on these relationships, is surprisingly limited. Using data from door-to-door surveys of nearly 300 residents in three pairs of small and large urban areas in England we demonstrate that people who visit green-space more regularly have higher biodiversity knowledge and support for conservation (measured using scales of pro-environmental behavior). Crucially these relationships only arise when considering visits to the countryside and not the frequency of visits to urban green-space. These patterns are robust to a suite of confounding variables including nature orientated motivations for visiting green-space, socio-economic and demographic factors, garden-use and engagement with natural history programs. Despite this the correlations that we uncover cannot unambiguously demonstrate that visiting the countryside improves biodiversity knowledge and conservation support. We consider it likely, however, that two mechanisms operate through a positive feedback loop i.e. increased visits to green-space promote an interest in and knowledge of biodiversity and support for conservation, which in turn further increase the desire to visit green-space and experience nature. The intensity of urbanization around peoples' homes, but not city size, is negatively associated with their frequency of countryside visits and biodiversity knowledge. Designing less intensely urbanized cities with good access to the countryside, combined with conservation policies that promote access to the countryside thus seems likely to maximize urban residents' biodiversity knowledge and support for conservation.

Species traits and the form of individual species-energy relationships.

Environmental energy availability explains much of the spatial variation in species richness at regional scales. While numerous mechanisms that may drive such total species-energy relationships have been identified, knowledge of their relative contributions is scant. Here, we adopt a novel approach to identify these drivers that exploits the composite nature of species richness, i.e. its summation from individual species distributions. We construct individual species-energy relationships (ISERs) for each species in the British breeding avifauna using both solar (temperature) and productive energy metrics (normalized difference vegetation index) as measures of environmental energy availability. We use the slopes of these relationships and the resultant change in deviance, relative to a null model, as measures of their strength and use them as response variables in multiple regressions that use ecological traits as predictors. The commonest species exhibit the strongest ISERs, which is counter to the prediction derived from the more individuals hypothesis. There is no evidence that predatory species have stronger ISERs, which is incompatible with the suggestion that high levels of energy availability increase the length of the food chain allowing larger numbers of predators to exist. We find some evidence that species with narrow niche breadths have stronger ISERs, thus providing one of the few pieces of supportive evidence that high-energy availability promotes species richness by increasing the occurrence of specialist species that use a narrow range of resources.

The species-area-energy relationship.

Area and available energy are major determinants of species richness. Although scale dependency of the relationship between energy availability and species richness (the species-energy relationship) has been documented, the exact relationship between the species-area and the species-energy relationship has not been studied explicitly. Here we show, using two extensive data sets on avian distributions in different biogeographic regions, that there is a negative interaction between energy availability and area in their effect on species richness. The slope of the species-area relationship is lower in areas with higher levels of available energy, and the slope of the species-energy relationship is lower for larger areas. This three-dimensional species-area-energy relationship can be understood in terms of probabilistic processes affecting the proportions of sites occupied by individual species. According to this theory, high environmental energy elevates species' occupancies, which depress the slope of the species-area curve.

Dissecting the species-energy relationship.

Environmental energy availability can explain much of the spatial variation in species richness. Such species-energy relationships encompass a diverse range of forms, and there is intense debate concerning which of these predominate, and the factors promoting this diversity. Despite this there has been relatively little investigation of whether the form, and relative strength, of species-energy relationships varies with (i) the currency of energy availability that is used, and (ii) the ecological characteristics of the constituent species. Such investigations can, however, shed light on the causal mechanisms underlying species-energy relationships. We illustrate this using the British breeding avifauna. The strength of the species-energy relationship is dependent on the energy metric used, with species richness being more closely correlated with temperature than the Normalized Difference Vegetation Index, which is a strong correlate of net primary productivity. We find little evidence, however, for the thermoregulatory load hypothesis that high temperatures enable individuals to invest in growth and reproduction, rather than thermoregulation, increasing population sizes that buffer species from extinction. High levels of productive energy may also elevate population size, which is related to extinction risk by a negative decelerating function. Therefore, the rarest species should exhibit the strongest species-energy relationship. We find evidence to the contrary, together with little support for suggestions that high-energy availability elevates species richness by increasing the numbers of specialists or predators.

Birds and people in Europe.

At a regional scale, species richness and human population size are frequently positively correlated across space. Such patterns may arise because both species richness and human density increase with energy availability. If the species-energy relationship is generated through the 'more individuals' hypothesis, then the prediction is that areas with high human densities will also support greater numbers of individuals from other taxa. We use the unique data available for the breeding birds in Europe to test this prediction. Overall regional densities of bird species are higher in areas with more people; species of conservation concern exhibit the same pattern. Avian density also increases faster with human density than does avian biomass, indicating that areas with a higher human density have a higher proportion of small-bodied individuals. The analyses also underline the low numbers of breeding birds in Europe relative to humans, with a median of just three individual birds per person, and 4 g of bird for every kilogram of human.