Shifting forward: Urban ecology in perspective.

The world has become urban; cities increasingly shape our worldviews, relation to other species, and the large-scale, long-term decisions we make. Cities are nature, but they need to align better with other ecosystems to avoid accelerating climate change and loss of biodiversity. We need a science to guide urban development across the diverse realities of global cities. This need can be met, in part, by shifts in urban ecology and its linkages to related sciences. This perspective is a "synthesis of syntheses", consolidating ideas from the other articles in the Special Section. It re-examines the role of urban ecology, and explores its integration with other disciplines that study cities. We conclude by summarizing the next steps in the ongoing shift in urban ecology, which is fast becoming an integral part of urban studies.

A transformative shift in urban ecology toward a more active and relevant future for the field and for cities.

This paper builds on the expansion of urban ecology from a biologically based discipline-ecology in the city-to an increasingly interdisciplinary field-ecology of the city-to a transdisciplinary, knowledge to action endeavor-an ecology for and with the city. We build on this "prepositional journey" by proposing a transformative shift in urban ecology, and we present a framework for how the field may continue this shift. We conceptualize that urban ecology is in a state of flux, and that this shift is needed to transform urban ecology into a more engaged and action based field, and one that includes a diversity of actors willing to participate in the future of their cities. In this transformative shift, these actors will engage, collaborate, and participate in a continuous spiral of knowledge → action → knowledge spiral and back to knowledge loop, with the goal of co producing sustainable and resilient solutions to myriad urban challenges. Our framework for this transformative shift includes three pathways: (1) a repeating knowledge → action → knowledge spiral of ideas, information, and solutions produced by a diverse community of agents of urban change working together in an "urban sandbox"; (2) incorporation of a social-ecological-technological systems framework in this spiral and expanding the spiral temporally to include the "deep future," where future scenarios are based on a visioning of seemingly unimaginable or plausible future states of cities that are sustainable and resilient; and (3) the expansion of the spiral in space, to include rural areas and places that are not yet cities. The three interrelated pathways that define the transformative shift demonstrate the power of an urban ecology that has moved beyond urban systems science and into a realm where collaborations among diverse knowledges and voices are working together to understand cities and what is urban while producing sustainable solutions to contemporary challenges and envisioning futures of socially, ecologically, and technologically resilient cities. We present case study examples of each of the three pathways that make up this transformative shift in urban ecology and discuss both limitations and opportunities for future research and action with this transdisciplinary broadening of the field.

Forging just ecologies: 25 years of urban long-term ecological research collaboration.

We ask how environmental justice and urban ecology have influenced one another over the past 25 years in the context of the US Long-Term Ecological Research (LTER) program and Baltimore Ecosystem Study (BES) project. BES began after environmental justice emerged through activism and scholarship in the 1980s but spans a period of increasing awareness among ecologists and environmental practitioners. The work in Baltimore provides a detailed example of how ecological research has been affected by a growing understanding of environmental justice. The shift shows how unjust environmental outcomes emerge and are reinforced over time by systemic discrimination and exclusion. We do not comprehensively review the literature on environmental justice in urban ecology but do present four brief cases from the Caribbean, Africa, and Asia, to illustrate the global relevance of the topic. The example cases demonstrate the necessity for continuous engagement with communities in addressing environmental problem solving.

Vegetation cover in relation to socioeconomic factors in a tropical city assessed from sub-meter resolution imagery.

Fine-scale information about urban vegetation and social-ecological relationships is crucial to inform both urban planning and ecological research, and high spatial resolution imagery is a valuable tool for assessing urban areas. However, urban ecology and remote sensing have largely focused on cities in temperate zones. Our goal was to characterize urban vegetation cover with sub-meter (<1 m) resolution aerial imagery, and identify social-ecological relationships of urban vegetation patterns in a tropical city, the San Juan Metropolitan Area, Puerto Rico. Our specific objectives were to (1) map vegetation cover using sub-meter spatial resolution (0.3-m) imagery, (2) quantify the amount of residential and non-residential vegetation, and (3) investigate the relationship between patterns of urban vegetation vs. socioeconomic and environmental factors. We found that 61% of the San Juan Metropolitan Area was green and that our combination of high spatial resolution imagery and object-based classification was highly successful for extracting vegetation cover in a moist tropical city (97% accuracy). In addition, simple spatial pattern analysis allowed us to separate residential from non-residential vegetation with 76% accuracy, and patterns of residential and non-residential vegetation varied greatly across the city. Both socioeconomic (e.g., population density, building age, detached homes) and environmental variables (e.g., topography) were important in explaining variations in vegetation cover in our spatial regression models. However, important socioeconomic drivers found in cities in temperate zones, such as income and home value, were not important in San Juan. Climatic and cultural differences between tropical and temperate cities may result in different social-ecological relationships. Our study provides novel information for local land use planners, highlights the value of high spatial resolution remote sensing data to advance ecological research and urban planning in tropical cities, and emphasizes the need for more studies in tropical cities.

Taxonomy and remote sensing of leaf mass per area (LMA) in humid tropical forests.

Leaf mass per area (LMA) is a trait of central importance to plant physiology and ecosystem function, but LMA patterns in the upper canopies of humid tropical forests have proved elusive due to tall species and high diversity. We collected top-of-canopy leaf samples from 2873 individuals in 57 sites spread across the Neotropics, Australasia, and Caribbean and Pacific Islands to quantify environmental and taxonomic drivers of LMA variation, and to advance remote-sensing measures of LMA. We uncovered strong taxonomic organization of LMA, with species accounting for 70% of the global variance and up to 62% of the variation within a forest stand. Climate, growth habit, and site conditions are secondary contributors (1-23%) to the observed LMA patterns. Intraspecific variation in LMA averages 16%, which is a fraction of the variation observed between species. We then used spectroscopic remote sensing (400-2500 nm) to estimate LMA with an absolute uncertainty of 14-15 g/m2 (r2 = 0.85), or approximately 10% of the global mean. With radiative transfer modeling, we demonstrated the scalability of spectroscopic remote sensing of LMA to the canopy level. Our study indicates that remotely sensed patterns of LMA will be driven by taxonomic variation against a backdrop of environmental controls expressed at site and regional levels.

The potential for species conservation in tropical secondary forests.

In the wake of widespread loss of old-growth forests throughout the tropics, secondary forests will likely play a growing role in the conservation of forest biodiversity. We considered a complex hierarchy of factors that interact in space and time to determine the conservation potential of tropical secondary forests. Beyond the characteristics of local forest patches, spatial and temporal landscape dynamics influence the establishment, species composition, and persistence of secondary forests. Prospects for conservation of old-growth species in secondary forests are maximized in regions where the ratio of secondary to old-growth forest area is relatively low, older secondary forests have persisted, anthropogenic disturbance after abandonment is relatively low, seed-dispersing fauna are present, and old-growth forests are close to abandoned sites. The conservation value of a secondary forest is expected to increase over time, as species arriving from remaining old-growth forest patches accumulate. Many studies are poorly replicated, which limits robust assessments of the number and abundance of old-growth species present in secondary forests. Older secondary forests are not often studied and few long-term studies are conducted in secondary forests. Available data indicate that both old-growth and second-growth forests are important to the persistence of forest species in tropical, human-modified landscapes.

A link between hurricane-induced tree sprouting, high stem density and short canopy in tropical dry forest.

The physiognomy of Caribbean dry forest is shorter, denser and contains a greater proportion of multi-stemmed trees than other neotropical dry forests. Our previous research, conducted after Hurricane Georges in 1998, has shown that dry forest trees sprout near the base following hurricane disturbance, even if the trees have not incurred structural damage. However, for these hurricane-induced sprouts to contribute to the physiognomy of the forest, they must grow and survive. We followed sprout dynamics and stem mortality on 1,407 stems from 1998, after Hurricane Georges, until 2005. The number of surviving sprouts and the proportion of sprouting stems decreased during the 7-year period, but the sprouting rate was still 3-fold higher and the proportion of sprouting stems 5-fold higher than before the hurricane. Mortality of non-sprouting stems (15.4%) was about the same as for sprouting stems (13.9%) after 7 years. The mean length of the dominant sprout surpassed 1.6 m by 2005, with over 13% of the dominant sprouts reaching subcanopy height. Sprout growth and survival varied among species. These results demonstrate that, despite some thinning, hurricane-induced sprouts survive and grow and that the unique physiognomic characteristic of Caribbean dry forests is related to hurricane disturbance.