Nancy B. Grimm

Global Change and the Ecology of Cities

Urban areas are hot spots that drive environmental change at multiple scales. Material demands of production and human consumption alter land use and cover, biodiversity, and hydrosystems locally to regionally, and urban waste discharge affects local to global biogeochemical cycles and climate. For urbanites, however, global environmental changes are swamped by dramatic changes in the local environment. Urban ecology integrates natural and social sciences to study these radically altered local environments and their regional and global effects. Cities themselves present both the problems and solutions to sustainability challenges of an increasingly urbanized world.

Biogeochemical Hot Spots and Hot Moments at the Interface of Terrestrial and Aquatic Ecosystems

Rates and reactions of biogeochemical processes vary in space and time to produce both hot spots and hot moments of elemental cycling. We define biogeochemical hot spots as patches that show disproportionately high reaction rates relative to the surrounding matrix, whereas hot moments are defined as short periods of time that exhibit disproportionately high reaction rates relative to longer intervening time periods. As has been appreciated by ecologists for decades, hot spot and hot moment activity is often enhanced at terrestrial-aquatic interfaces. Using examples from the carbon (C) and nitrogen (N) cycles, we show that hot spots occur where hydrological flowpaths converge with substrates or other flowpaths containing complementary or missing reactants. Hot moments occur when episodic hydrological flowpaths reactivate and/or mobilize accumulated reactants. By focusing on the delivery of specific missing reactants via hydrologic flowpaths, we can forge a better mechanistic understanding of the factors that create hot spots and hot moments. Such a mechanistic understanding is necessary so that biogeochemical hot spots can be identified at broader spatiotemporal scales and factored into quantitative models. We specifically recommend that resource managers incorporate both natural and artificially created biogeochemical hot spots into their plans for water quality management. Finally, we emphasize the needs for further research to assess the potential importance of hot spot and hot moment phenomena in the cycling of different bioactive elements, improve our ability to predict their occurrence, assess their importance in landscape biogeochemistry, and evaluate their utility as tools for resource management.

Stream denitrification across biomes and its response to anthropogenic nitrate loading

Anthropogenic addition of bioavailable nitrogen to the biosphere is increasing and terrestrial ecosystems are becoming increasingly nitrogen-saturated, causing more bioavailable nitrogen to enter groundwater and surface waters. Large-scale nitrogen budgets show that an average of about 20–25 per cent of the nitrogen added to the biosphere is exported from rivers to the ocean or inland basins, indicating that substantial sinks for nitrogen must exist in the landscape. Streams and rivers may themselves be important sinks for bioavailable nitrogen owing to their hydrological connections with terrestrial systems, high rates of biological activity, and streambed sediment environments that favour microbial denitrification. Here we present data from nitrogen stable isotope tracer experiments across 72 streams and 8 regions representing several biomes. We show that total biotic uptake and denitrification of nitrate increase with stream nitrate concentration, but that the efficiency of biotic uptake and denitrification declines as concentration increases, reducing the proportion of in-stream nitrate that is removed from transport. Our data suggest that the total uptake of nitrate is related to ecosystem photosynthesis and that denitrification is related to ecosystem respiration. In addition, we use a stream network model to demonstrate that excess nitrate in streams elicits a disproportionate increase in the fraction of nitrate that is exported to receiving waters and reduces the relative role of small versus large streams as nitrate sinks.

Integrated approaches to long-term studies of urban ecological systems

This quote captures the spirit of the new urban emphasis in the US Long-Term Ecological Research (LTER) network. We know now that Earth abounds with both subtle and pronounced evidence of the influence of people on natural ecosystems (Russell 1993, Turner and Meyer 1993). Arguably, cities are the most human dominated of all ecosystems. Recent calls for studies on “human-dominated ecosystems” (Vitousek et al. 1997) finally have been heeded, over 60 years after Tansley penned his warning, with the addition of two metropolises (Phoenix and Baltimore) to the LTER network. In this article, we describe an emerging approach to understanding the ecology of urban areas by contrasting these two metropolises, and we present a call to action for ecologists to integrate their science with that of social scientists to achieve a more realistic and useful understanding of the natural world in general and its ecology in particular (Pickett and McDonnell 1993, Ehrlich 1997). We begin by framing a conceptual basis for the study of urban ecological systems: the rationale, contrasting approaches, and special considerations for including human interactions at different scales and in a spatial context. We then discuss the application of our conceptual approach by comparing site conditions and initial research results in Baltimore and Phoenix. We conclude with a summary and synthesis of implications for the integration of social and ecological sciences.

TEMPORAL SUCCESSION IN A DESERT STREAM ECOSYSTEM FOLLOWING FLASH FLOODING

Recovery of a desert stream after an intense flash flooding event is described as a model of temporal succession in lotic ecosystems. A late summer flood in Sycamore Creek, Arizona, virtually eliminated algae and reduced invertebrate standing crop by 98%. Physical and morphometric conditions typical of the preflood period were restored in 2 d and the biota recovered in 2—3 wk. Algal communities responded rapidly and achieved a standing crop of nearly 100 g/m2 in 2 wk. Community composition was dominated by diatoms early in succession and by filamentous greens and blue—greens later. Macroinvertebrates also recolonized denuded substrates rapidly, largely by immigration of aerial adults and subsequent oviposition. Growth and development were rapid and several generations of the dominant mayfly and dipteran taxa were completed during the 1st mo of recovery. Invertebrate dry biomass reached 7.3 g/m2 in 1 mo. Gross primary production (Pg) measured as O2 increased in a similar asymptotic fashion and reached 6.6 g°m—2°d—1 in 30 d. Pg exceeded community respiration (R) after day 5 and Pg/R averaged 1.46 for the remainder of the 2—mo sequence. This ecosystem is thus autotrophic and exports organic matter downstream and by drying, laterally. Uptake of nitrate and phosphorus were proportional to net primary production and exhibited a marked downstream decline in concentration during both light and dark periods. Temporal trajectories of various community and ecosystem attributes are compared with those suggested by Odum (1969) to be diagnostic of successional status. Agreement was poor in attributes which are especially modified in open, frequently disturbed ecosystems such as streams.

Towards an ecological understanding of biological nitrogen fixation

N limitation to primary production and other ecosystem processes is widespread. To understand the causes and distribution of N limitation, we must understand the controls of biological N fixation. The physiology of this process is reasonably well characterized, but our understanding of ecological controls is sparse, except in a few cultivated ecosystems. We review information on the ecological controls of N fixation in free-living cyanobacteria, vascular plant symbioses, and heterotrophic bacteria, with a view toward developing improved conceptual and simulation models of ecological controls of biological N fixation.A model (Howarth et al. 1999) of cyanobacterial fixation in lakes (where N fixation generally increases substantially when N:P ratios are low) versus estuaries (where planktonic N fixation is rare regardless of N:P ratios) concludes that an interaction of trace-element limitation and zooplankton grazing could constrain cyanobacteria in estuaries and so sustain N limitation. Similarly. a model of symbiotic N fixation on land (Vitousek & Field 1999) suggests that shade intolerance, P limitation, and grazing on N-rich plant tissues could suppress symbiotic N fixers in late-successional forest ecosystems. This congruence of results raises the question – why do late-successional tropical forests often contain many potentially N-fixing canopy legumes, while N fixers are absent from most late-successional temperate and boreal forests? We suggest that relatively high N availability in lowland tropical forests permits legumes to maintain an N-demanding lifestyle (McKey 1994) without always being required to pay the costs of fixing N.Overall, both the few simulation models and the more-numerous conceptual models of ecological controls of biological N fixation suggest that there are substantial common features across N-fixing organisms and ecosystems. Despite the many groups of organisms capable of fixing N, and the very different ecosystems in which the process is important, we suggest that these common controls provide a foundation for the development of regional and global models that incorporate ecological controls of biological N fixation.

Socioeconomics drive urban plant diversity

Spatial variation in plant diversity has been attributed to heterogeneity in resource availability for many ecosystems. However, urbanization has resulted in entire landscapes that are now occupied by plant communities wholly created by humans, in which diversity may reflect social, economic, and cultural influences in addition to those recognized by traditional ecological theory. Here we use data from a probability-based survey to explore the variation in plant diversity across a large metropolitan area using spatial statistical analyses that incorporate biotic, abiotic, and human variables. Our prediction for the city was that land use, along with distance from urban center, would replace the dominantly geomorphic controls on spatial variation in plant diversity in the surrounding undeveloped Sonoran desert. However, in addition to elevation and current and former land use, family income and housing age best explained the observed variation in plant diversity across the city. We conclude that a functional relationship, which we term the “luxury effect,” may link human resource abundance (wealth) and plant diversity in urban ecosystems. This connection may be influenced by education, institutional control, and culture, and merits further study.

Ecosystem Expansion and Contraction in Streams Desert streams vary in both space and time and fluctuate dramatically in size

St re<l ms are hydrologically diverse and dynamic ecosystems. Flow may vary between extremes, from high-discharge floods to periods when surface water is absent. Although much is known about the role of floods in shaping ecological processes, far less is known about the biological and chemical changes that occu r du ring periods of water loss in stream ecosystems (Boulton and Suter 1986, Stanley and Fisher 1992). Nowhere is this lack of knowledge more apparent than in desert streams; these lotic ecosystems exist in a setting defined by water limitation, and periods of declining or absent flow are common. However, water loss is by no means unique to desert streams, because intermittent streams are found in many different environments. Moreover, escalating demands on a finite water supply arc increasing the likelihood of drying in streams and rivers worldwide. Irrigation, impoundment, diversion, and groundwater abstraction reduce streamflow in mesic and xeric regions alike. In arid and semiarid areas, large rivers that are devoid of water are common, and in more mesic locales, profligate water use decreases the total amount of surface water present

Exchange between interstitial and surface water: implications for stream metabolism and nutrient cycling

Metabolism of a Sonoran Desert stream was investigated by both enclosure and whole system oxygen techniques. We used recirculating chambers to estimate surface sediment metabolism and measured deep sediment respiration in isolated sediment cores. Metabolism of the stream ecosystem was determined for a 30-m reach as dark and light oxygen change with and without black plastic sheeting that darkened the stream and prevented diffusion. Average ecosystem respiration for two dates in August (440 mg O2 m-2 h-1) exceeded respiration of either the surface sediment community (155 Mg O2 m-2 h-1) or the hyporheic community (170 mg O2 m-2 h-1) alone. Deep sediments show substantial oxygen and nitrate uptake when isolated. In the stream, this low nitrate interstitial water is exchanged with surface water. Metabolism of the isolated surface community suggests a highly productive and autotrophic system, yet gross production is balanced or exceeded by community respiration when ecosystem boundaries include the hyporheic zone. Thus, despite high rates of gross primary production (600–1200 mg O2 m-2 h-1), desert streams may be heterotrophic (PG < R) during summer.

The changing landscape: ecosystem responses to urbanization and pollution across climatic and societal gradients

Urbanization, an important driver of climate change and pollution, alters both biotic and abiotic ecosystem properties within, surrounding, and even at great distances from urban areas. As a result, research challenges and environmental problems must be tackled at local, regional, and global scales. Ecosystem responses to land change are complex and interacting, occurring on all spatial and temporal scales as a consequence of connectivity of resources, energy, and information among social, physical, and biological systems. We propose six hypotheses about local to continental effects of urbanization and pollution, and an operational research approach to test them. This approach focuses on analysis of “megapolitan” areas that have emerged across North America, but also includes diverse wildland-to-urban gradients and spatially continuous coverage of land change. Concerted and coordinated monitoring of land change and accompanying ecosystem responses, coupled with simulation models, will permit robust foreca...