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Abstract Urbanization is a leading cause of global biodiversity loss, yet cities can provide resources required by many species throughout the year. In recognition of this, cities around the world are adopting strategies to increase biodiversity. These efforts would benefit from a robust understanding of how natural and enhanced features in urbanized areas influence various taxa. We explored seasonal and spatial patterns in occupancy and taxonomic richness of birds and pollinators among office parks in Santa Clara County, California, USA, where natural features and commercial landscaping have generated variation in conditions across scales. We surveyed birds and insect pollinators, estimated multi‐species occupancy and species richness, and found that spatial scale (local, neighborhood, and landscape scale), season, and urban sensitivity were all important for understanding how communities occupied sites. Features at the landscape (distance to streams or baylands) and local scale (tree canopy, shrub, or impervious cover) were the strongest predictors of avian occupancy in all seasons. Pollinator richness was influenced by local tree canopy and impervious cover in spring, and distance to baylands in early and late summer. We then predicted the relative contributions of different spatial scales to annual bird species richness by simulating “good” and “poor” quality sites based on influential covariates returned by the previous models. Shifting from poor to good quality conditions locally increased annual avian richness by up to 6.8 species with no predicted effect on the quality of the neighborhood. Conversely, sites of poor local and neighborhood scale quality in good‐quality landscapes were predicted to harbor 11.5 more species than sites of good local‐ and neighborhood‐scale quality in poor‐quality landscapes. Finally, more urban‐sensitive bird species were gained at good quality sites relative to urban tolerant species, suggesting that urban natural features at the local and landscape scales disproportionately benefited them.

Abstract Green spaces in urban areas—like remnant habitat, parks, constructed wetlands, and street trees—supply multiple benefits. Many studies show green spaces in and near urban areas play important roles harbouring biodiversity and promoting human well‐being. On the other hand, evidence suggests that greater human population density enables compact, low‐carbon cities that spare habitat conversion at the fringes of expanding urban areas, while also allowing more walkable and livable cities. How then can urban areas have abundant green spaces as well as density? In this paper, we review the empirical evidence for the relationships between urban density, nature, and sustainability. We also present a quantitative analysis of data on urban tree canopy cover and open space for United States large urbanized areas, as well as an analysis of non‐US Functional Urban Areas in OECD countries. We found that there is a negative correlation between population density and these green spaces. For Functional Urban Areas in the OECD, a 10% increase in density is associated with a 2.9% decline in tree cover. We argue that there are competing trade‐offs between the benefits of density for sustainability and the benefits of nature for human well‐being. Planners must decide an appropriate density by choosing where to be on this trade‐off curve, taking into account city‐specific urban planning goals and context. However, while the negative correlation between population density and tree cover is modest at the level of US urbanized areas (R2 = 0.22), it is weak at the US Census block level (R2 = 0.05), showing that there are significant brightspots, neighbourhoods that manage to have more tree canopy than would be expected based upon their level of density. We then describe techniques for how urban planners and designers can create more brightspots, identifying a typology of urban forms and listing green interventions appropriate for each form. We also analyse policies that enable these green interventions illustrating them with the case studies of Curitiba and Singapore. We conclude that while there are tensions between density and urban green spaces, an urban world that is both green and dense is possible, if society chooses to take advantage of the available green interventions and create it. Read the free Plain Language Summary for this article on the Journal blog.

Abstract Conserving urban biodiversity will rely on updating our planning paradigms. This work synthesizes urban ecological research and identifies seven elements—patch size, connections, matrix quality, habitat diversity, native vegetation, special resources, and stewardship and management—as key factors associated with urban biodiversity. It then distils these findings into a science-based framework for building urban biodiversity. Beyond providing a framework, it demonstrates how to use this approach to integrate conservation into urban planning and provides an example where this framework has been applied. Such a landscape ecological approach will provide a science-driven approach to advance urban biodiversity and promote equitable access to urban nature.

Preserving and restoring wildlife in urban areas benefits both urban ecosystems and the well-being of urban residents. While urban wildlife conservation is a rapidly developing field, the majority of conservation research has been performed in wildland areas. Understanding the applicability of wildland science to urban populations and the relative importance of factors limiting species persistence are of critical importance to identifying prescriptive management strategies for restoring wildlife to urban parks. We evaluated how habitat fragmentation, habitat quality and mortality threats influence species occupancy and persistence in urban parks. We chose California quail Callipepla californica as a representative species with potential to respond to urban conservation. We used publicly available eBird data to construct occupancy models of quail in urban parks across their native range, and present an application using focal parks interested in exploring quail reintroduction. Urban parks had a 0.23 +/- 0.02 probability of quail occupancy, with greater occupancy in larger parks that were less isolated from potential source populations, had higher shrub cover and had lower impervious cover. Less isolated parks had higher colonization rates, while larger parks had lower extinction rates. These results align with findings across urban ecology showing greater biodiversity in larger and more highly connected habitat patches. A case study highlighted that interventions to increase effective park size and improve connectivity would be most influential for two highly urban focal parks, while changes to internal land cover would have a relatively small impact. Low joint extinction probability in the parks (0.010 +/- 0.013) indicated reintroduced populations could persist for some time. Synthesis and applications. We show how eBird data can be harnessed to evaluate the responsiveness of wildlife to urban parks of variable size, connectivity and habitat quality, highlighting what management actions are most needed. Using California quail as an example, we found park size, park isolation and presence of coyotes are all important drivers of whether quail can colonize and persist in parks. Our results suggest reintroducing quail to parks could be successful provided parks are large enough to support quail, and management actions are taken to enhance regional connectivity or periodic assisted colonization is used to supplement local populations.

Urban nature—such as greenness and parks—can alleviate distress and provide space for safe recreation during the COVID-19 pandemic. However, nature is often less available in low-income populations and communities of colour—the same communities hardest hit by COVID-19. In analyses of two datasets, we quantified inequity in greenness and park proximity across all urbanized areas in the United States and linked greenness and park access to COVID-19 case rates for ZIP codes in 17 states. Areas with majority persons of colour had both higher case rates and less greenness. Furthermore, when controlling for sociodemographic variables, an increase of 0.1 in the Normalized Difference Vegetation Index was associated with a 4.1% decrease in COVID-19 incidence rates (95% confidence interval: 0.9–6.8%). Across the United States, block groups with lower income and majority persons of colour are less green and have fewer parks. Our results demonstrate that the communities most impacted by COVID-19 also have the least nature nearby. Given that urban nature is associated with both human health and biodiversity, these results have far-reaching implications both during and beyond the pandemic. Access to green space has been a critical, and contentious, issue for neighbourhood inequality and health outcomes. This Analysis looks at how the COVID-19 pandemic interacts with availability of nature for urban residents.

Urban tree cover provides benefits to human health and well-being, but previous studies suggest that tree cover is often inequitably distributed. Here, we use National Agriculture Imagery Program digital ortho photographs to survey the tree cover inequality for Census blocks in US large urbanized areas, home to 167 million people across 5,723 municipalities and other Census-designated places. We compared tree cover to summer land surface temperature, as measured using Landsat imagery. In 92% of the urbanized areas surveyed, low-income blocks have less tree cover than high-income blocks. On average, low-income blocks have 15.2% less tree cover and are 1.5⁰C hotter than high-income blocks. The greatest difference between low- and high-income blocks was found in urbanized areas in the Northeast of the United States, where low-income blocks in some urbanized areas have 30% less tree cover and are 4.0⁰C hotter. Even after controlling for population density and built-up intensity, the positive association between income and tree cover is significant, as is the positive association between proportion non-Hispanic white and tree cover. We estimate, after controlling for population density, that low-income blocks have 62 million fewer trees than high-income blocks, equal to a compensatory value of $56 billion ($1,349/person). An investment in tree planting and natural regeneration of $17.6 billion would be needed to close the tree cover disparity, benefitting 42 million people in low-income blocks.

Cities are both embedded within and ecologically linked to their surrounding landscapes. Although urbanization poses a substantial threat to biodiversity, cities also support many species, some of which have larger populations, faster growth rates, and higher productivity in cities than outside of them. Despite this fact, surprisingly little attention has been paid to the potentially beneficial links between cities and their surroundings. We identify five pathways by which cities can benefit regional ecosystems by releasing species from threats in the larger landscape, increasing regional habitat heterogeneity and genetic diversity, acting as migratory stopovers, preadapting species to climate change, and enhancing public engagement and environmental stewardship. Increasing recognition of these pathways could help cities identify effective strategies for supporting regional biodiversity conservation and could provide a science-based platform for incorporating biodiversity alongside other urban greening goals.

In response to the widely recognized negative impacts of urbanization on biodiversity, many cities are reimagining urban design to provide better biodiversity support. Some cities have developed urban biodiversity plans, primarily focused on improving biodiversity support and ecosystem function within the built environment through habitat restoration and other types of urban greening projects. The biophilic cities movement seeks to reframe nature as essential infrastructure for cities, seamlessly integrating city and nature to provide abundant, accessible nature for all residents and corresponding health and well-being outcomes. Urban biodiversity planning and biophilic cities have significant synergies in their goals and the means necessary to achieve them. In this paper, we identify three key ways by which the urban biodiversity planning process can support biophilic cities objectives: engaging the local community; identifying science-based, quantitative goals; and setting priorities for action. Urban biodiversity planning provides evidence-based guidance, tools, and techniques needed to design locally appropriate, pragmatic habitat enhancements that support biodiversity, ecological health, and human health and well-being. Developing these multi-functional, multi-benefit strategies that increase the abundance of biodiverse nature in cities has the potential at the same time to deepen and enrich our biophilic experience in daily life.

Urban nature can alleviate distress and provide space for safe recreation during the COVID-19 pandemic. However, nature is often less available in low-income and communities of color—the same communities hardest hit by COVID-19. We quantified nature inequality across all urbanized areas in the US and linked nature access to COVID-19 case rates for ZIP Codes in 17 states. Areas with majority persons of color had both higher case rates and less greenness. Furthermore, when controlling for socio-demographic variables, an increase of 0.1 in Normalized Difference Vegetation Index (NDVI) was associated with a 4.1% decrease in COVID-19 incidence rates (95% confidence interval: 0.9-6.8%). Across the US, block groups with lower-income and majority persons of color are less green and have fewer parks. Thus, communities most impacted by COVID-19 also have the least nature nearby. Given urban nature is associated with both human health and biodiversity, these results have far-reaching implications both during and beyond the pandemic.

Investing in nature and nature-based solutions is an important pathway to address the current ecological crisis. The loss of biodiversity, with around one million animal and plant species threatened with extinction, is putting at risk our economies, livelihoods, food security, health, and quality of life worldwide. Diverse and innovative measures are needed to reverse this decline and to restore healthy ecosystems. The magnitude of the challenge calls for the collaboration and contribution of all actors, including non-state actors, such as businesses, local authorities, civil society organisations, and individual citizens. Sports and urban biodiversity identifies the opportunities for sports federations and sport venue owners and operators to develop the seven elements essential for supporting nature in cities. Case studies from Europe, North America, Australia and South Africa capture the biodiversity actions taken by a variety of sports, from Olympic city planning to local projects at stadiums and practice facilities. These examples also reveal ways in which improvements to urban biodiversity elements can offer a better sporting experience for event participants and spectators.

Ecological resilience is a powerful heuristic for ecosystem management in the context of rapid environmental change. Significant efforts are underway to improve the resilience of biodiversity and ecological function to extreme events and directional change across all types of landscapes, from intact natural systems to highly modified landscapes such as cities and agricultural regions. However, identifying management strategies likely to promote ecological resilience remains a challenge. In this article, we present seven core dimensions to guide long-term and large-scale resilience planning in highly modified landscapes, with the objective of providing a structure and shared vocabulary for recognizing opportunities and actions likely to increase resilience across the whole landscape. We illustrate application of our approach to landscape-scale ecosystem management through case studies from two highly modified California landscapes, Silicon Valley and the Sacramento–San Joaquin Delta. We propose that resilience-based management is best implemented at large spatial scales and through collaborative, cross-sector partnerships.

Understanding spatial distributions of invasive plant species at early infestation stages is critical for assessing the dynamics and underlying factors of invasions. Recent progress in very high resolution remote sensing is facilitating this task by providing high spatial detail over whole-site extents that are prohibitive to comprehensive ground surveys. This study assessed the opportunities and constraints to characterize landscape distribution of the invasive grass medusahead (Elymus caput-medusae) in a ∼36.8 ha grassland in California, United States from 0.15m-resolution visible/near-infrared aerial imagery at the stage of late spring phenological contrast with dominant grasses. We compared several object-based unsupervised, single-run supervised and hierarchical approaches to classify medusahead using spectral, textural, and contextual variables. Fuzzy accuracy assessment indicated that 44–100% of test medusahead samples were matched by its classified extents from different methods, while 63–83% of test samples classified as medusahead had this class as an acceptable candidate. Main sources of error included spectral similarity between medusahead and other green species and mixing of medusahead with other vegetation at variable densities. Adding texture attributes to spectral variables increased the accuracy of most classification methods, corroborating the informative value of local patterns under limited spectral data. The highest accuracy across different metrics was shown by the supervised single-run support vector machine with seven vegetation classes and Bayesian algorithms with three vegetation classes; however, their medusahead allocations showed some “spillover” effects due to misclassifications with other green vegetation. This issue was addressed by more complex hierarchical approaches, though their final accuracy did not exceed the best single-run methods. However, the comparison of classified medusahead extents with field segments of its patches overlapping with survey transects indicated that most methods tended to miss and/or over-estimate the length of the smallest patches and under-estimate the largest ones due to classification errors. Overall, the study outcomes support the potential of cost-effective, very high-resolution sensing for the site-scale detection of infestation hotspots that can be customized to plant phenological schedules. However, more accurate medusahead patch delineation in mixed-cover grasslands would benefit from testing hyperspectral data and using our study’s framework to inform and constrain the candidate vegetation classes in heterogeneous locations.

SummaryRainfall is a key determinant of production and composition in arid and semi‐arid systems. Long‐term studies relating composition and water availability primarily focus on current‐year precipitation patterns, though mounting evidence highlights the importance of previous‐year rainfall particularly in grasslands dominated by perennial species. The extent to which lagged precipitation effects occur in annual grasslands, however, remains largely unexplored.We pair a long‐term study with two manipulative experiments to identify patterns and mechanisms of lagged precipitation effects in annual grasslands. The long‐term study captured variation in functional group (exotic annual forbs and grasses) abundance and precipitation across 8 years at three northern California grassland sites. We then tested whether lagged rainfall effects were created through seed production and litter (residual dry matter, RDM) by manipulating rainfall and litter, respectively.Rainfall from the previous‐year growing season (both seasonal and total rainfall) shifted functional group abundance. High lagged rainfall was associated with increased grass and decreased forb abundance the following year. Current‐year seasonal rainfall also influenced species composition, with winter rain increasing forb and decreasing grass abundance. Lagged precipitation effects were generally stronger for forbs than for grasses. Our experimental studies provided evidence for two mechanisms that contributed to lagged effects in annual grasslands. Higher rainfall increased seed production for grasses, which translated to more germinable seed the following year. Higher rainfall also increased biomass production and RDM, which benefited grasses and reduced forb abundance.Synthesis. Our results highlight the importance of previous‐year precipitation in structuring annual community composition and suggest two important biotic pathways, seed rain and RDM, that regulate lagged community responses to rainfall. Incorporating lagged effects into models of grassland diversity and productivity could improve predictions of climate change impacts in annual grasslands.

Questions: Californian grasslands have a long history of invasion, starting with the introduction of exotic forage species a century ago, and followed by newer waves of invaders. Both exotic and invasive species produce large amounts of litter, but the importance of litter accumulation on the growth and expansion of these species has not been rigorously assessed. We addressed the following questions: (1) do litter type (exotic or invasive species) and litter quantity affect exotic and invasive plant performance; (2) at which plant life stages does litter accumulation effects occur; and (3) is litter accumulation impacting native species recruitment and diversity?Location: Semi-natural grasslands, northern California.Methods: We created litter quantity gradients (0, 62.5, 125, 250 or 500 g.m(-2)) of three litter types (exotic or invasive species litter or a mixture of both). We evaluated litter effects on the germination, recruitment, production of flowers and biomass production of the exotic Avena fatua, the invasive Elymus caput-medusae and a mixture of native species. We used litter depth as a proxy for litter accumulation and measured litter impacts on light availability at ground level, soilmoisture, soil andmicrobial C and N.Results: We found a significant litter depth impact on all species, but no litter type effects. Litter accumulation principally reduced light availability at ground level and positively affected exotic and invasive species at different life-history stages. Above-ground biomass production of the exotic species Avena increased with increased litter depth, suggesting a litter accumulation feedback. The invasive species Elymus responded to increased litter with increased seed production, rather than biomass production. In contrast, litter negatively affected the recruitment and diversity of native species.Conclusion: Overall, our findings point to cascading effects in invasion with positive litter accumulation feedbacks of an exotic species simultaneously reducing native diversity and promoting seed production of a more recent invasive arrival. These results have strong consequences for invasion management and suggest that litter removal practices, such as burning and grazing, would better succeed at limiting the growth and spread of invasive species than herbicide application or seeding.

Summary Successful colonization by invasive species depends on both the ability to disperse seeds to a site and an ability to establish once seeds have arrived. While seed and establishment limitation are known to jointly influence colonization, decomposing establishment limitation into density‐dependent and density‐independent components has remained challenging. Here, we couple theoretical models of recruitment with a multispecies invasion experiment conducted within a natural gradient of soil moisture and productivity to assess how variation in establishment limitation shapes outcomes for invasion. Recruitment was affected by both density‐dependent and density‐independent sources of establishment limitation in three of four species. Soil moisture stress and productivity both increased density‐independent mortality in one species, whereas density‐dependent mortality increased in locations with favourable soil moisture. Synthesis. Successful establishment of invading species can be limited by both density‐dependent and density‐independent mechanisms. In particular, the strength of density‐independent limitation may depend on natural gradients in abiotic factors. The varying strengths of establishment limitation suggest that patterns of invasion are likely to be uneven both in space and in time. Understanding how intraspecific competitive constraints and density‐independent limitation vary with abiotic gradients can assist with predicting when invasions are likely to occur, information that can be harnessed in the development of better methods for control.