James B. Heffernan

RIPARIAN ZONES INCREASE REGIONAL SPECIES RICHNESS BY HARBORING DIFFERENT, NOT MORE, SPECIES

Riparian zones are habitats of critical conservation concern worldwide, as they are known to filter agricultural contaminants, buffer landscapes against erosion, and provide habitat for high numbers of species. Here we test the generality of the notion that riparian habitats harbor more species than adjacent upland habitats. Using previously pub- lished data collected from seven continents and including taxa ranging from Antarctic soil invertebrates to tropical rain forest lianas and primates, we show that riparian habitats do not harbor higher numbers of species, but rather support significantly different species pools altogether. In this way, riparian habitats increase regional ( g-) richness across the globe by .50%, on average. Thus conservation planners can easily increase the number of species protected in a regional portfolio by simply including a river within terrestrial biodiversity reserves. Our analysis also suggests numerous possible improvements for future studies of species richness gradients across riparian and upland habitats. First, ,15% of the studies in our analysis included estimates of more than one taxonomic group of interest. Second, within a given taxonomic group, studies employed variable methodologies and sampling areas in pursuit of richness and turnover estimates. Future analyses of species richness patterns in watersheds should aim to include a more comprehensive suite of taxonomic groups and should measure richness at multiple spatial scales.

HORIZONS IN STREAM BIOGEOCHEMISTRY: FLOWPATHS TO PROGRESS

Over the past 50 years, conceptual developments in stream ecology and ecosystem ecology have converged, thanks to biogeochemistry and the recognition that in situ processing on one hand and spatial translation of materials, processes, and influence along flowpaths on the other, unite to generate a holistic picture of ecosystem functioning at the landscape level. Early emphases in stream biogeochemistry involved organic carbon dynamics and whole-ecosystem budgets. These approaches were holistic but cumbersome and laborious and ignored several crucial issues, such as variation in organic matter quality. Nutrient-spiraling approaches rectified this shortcoming and provided a flowpath-specific technique for resolving the dynamics of both inorganic and organic materials and for comparing streams of different sizes and flow rates. The ability of nutrient-spiraling approaches to deal with multiple elements and fluctuating flows, including floods, remains elusive, however. There are several opportunities for stre...

Macrosystems ecology: understanding ecological patterns and processes at continental scales

Macrosystems ecology is the study of diverse ecological phenomena at the scale of regions to continents and their interactions with phenomena at other scales. This emerging subdiscipline addresses ecological questions and environmental problems at these broad scales. Here, we describe this new field, show how it relates to modern ecological study, and highlight opportunities that stem from taking a macrosystems perspective. We present a hierarchical framework for investigating macrosystems at any level of ecological organization and in relation to broader and finer scales. Building on well-established theory and concepts from other subdisci- plines of ecology, we identify feedbacks, linkages among distant regions, and interactions that cross scales of space and time as the most likely sources of unexpected and novel behaviors in macrosystems. We present three examples that highlight the importance of this multiscaled systems perspective for understanding the ecology of regions to continents.

Ecological homogenization of urban USA

A visually apparent but scientifically untested outcome of land-use change is homogenization across urban areas, where neighborhoods in different parts of the country have similar patterns of roads, residential lots, commercial areas, and aquatic features. We hypothesize that this homogenization extends to ecological structure and also to ecosystem functions such as carbon dynamics and microclimate, with continental-scale implications. Further, we suggest that understanding urban homogenization will provide the basis for understanding the impacts of urban land-use change from local to continental scales. Here, we show how multi-scale, multi-disciplinary datasets from six metropolitan areas that cover the major climatic regions of the US (Phoenix, AZ; Miami, FL; Baltimore, MD; Boston, MA; Minneapolis–St Paul, MN; and Los Angeles, CA) can be used to determine how household and neighborhood characteristics correlate with land-management practices, land-cover composition, and landscape structure and ecosystem functions at local, regional, and continental scales.

WETLANDS AS AN ALTERNATIVE STABLE STATE IN DESERT STREAMS

Historically, desert drainages of the American southwest supported productive riverine wetlands (ciénegas). Region-wide erosion of ciénegas during the late 19th and early 20th century dramatically reduced the abundance of these ecosystems, but recent reestablishment of wetlands in Sycamore Creek, Arizona, USA, provides an opportunity to evaluate the mechanisms underlying wetland development. A simple model demonstrates that density-dependent stabilization of channel substrate by vegetation results in the existence of alternative stable states in desert streams. A two-year (October 2004-September 2006) field survey of herbaceous cover and biomass at 26 sites located along Sycamore Creek is used to test the underlying assumption of this model that vegetation cover loss during floods is density dependent, as well as the prediction that the distribution of vegetation abundance should shift toward bimodality in response to floods. Observations of nonlinear, negative relationships between herbaceous biomass prior to flood events and the proportion of persistent vegetation cover were consistent with the alternative stable state model. In further support of the alternative-state hypothesis, vegetation cover diverged from an approximately normal distribution toward a distinctly bimodal distribution during the monsoon flood season of 2006. These results represent the first empirically supported example of alternative-state behavior in stream ecosystems. Identification of alternative stable states in desert streams supports recent hypotheses concerning the importance of strong abiotic-disturbance regimes and biogeomorphic mechanisms in multiple-state ecosystems.

Assessing the homogenization of urban land management with an application to US residential lawn care

Significance This paper offers conceptual and empirical contributions to sustainability science in general and urban-ecological studies in particular. We present a new analytical framework for classifying socioecological measures along a homogenization–differentiation spectrum. This simple 2 × 2 matrix highlights the multiscale nature of the processes and outcomes of interest. Our application of the conceptual framework produces needed empirical insights into the extent to which land management appears to be homogenizing in differing biophysical settings. Results suggest that US lawn care behaviors are more differentiated in practice than in theory. Thus even if the biophysical outcomes of urbanization are homogenizing, managing the associated sustainability implications may require a multiscale, differentiated approach. Changes in land use, land cover, and land management present some of the greatest potential global environmental challenges of the 21st century. Urbanization, one of the principal drivers of these transformations, is commonly thought to be generating land changes that are increasingly similar. An implication of this multiscale homogenization hypothesis is that the ecosystem structure and function and human behaviors associated with urbanization should be more similar in certain kinds of urbanized locations across biogeophysical gradients than across urbanization gradients in places with similar biogeophysical characteristics. This paper introduces an analytical framework for testing this hypothesis, and applies the framework to the case of residential lawn care. This set of land management behaviors are often assumed—not demonstrated—to exhibit homogeneity. Multivariate analyses are conducted on telephone survey responses from a geographically stratified random sample of homeowners (n = 9,480), equally distributed across six US metropolitan areas. Two behaviors are examined: lawn fertilizing and irrigating. Limited support for strong homogenization is found at two scales (i.e., multi- and single-city; 2 of 36 cases), but significant support is found for homogenization at only one scale (22 cases) or at neither scale (12 cases). These results suggest that US lawn care behaviors are more differentiated in practice than in theory. Thus, even if the biophysical outcomes of urbanization are homogenizing, managing the associated sustainability implications may require a multiscale, differentiated approach because the underlying social practices appear relatively varied. The analytical approach introduced here should also be productive for other facets of urban-ecological homogenization.

Unintended Consequences of Urbanization for Aquatic Ecosystems: A Case Study from the Arizona Desert

ABSTRACT Many changes wrought during the construction of “designer ecosystems” are intended to ensure—and often succeed in ensuring—that a city can provide ecosystem goods and services; but other changes have unintended impacts on the ecology of the city, impairing its ability to provide these critical functions. Indian Bend Wash, an urbanizing watershed in the Central Arizona–Phoenix (CAP) ecosystem, provides an excellent case study of how human alteration of land cover, stream channel structure, and hydrology affect ecosystem processes, both intentionally and unintentionally. The construction of canals created new flowpaths that cut across historic stream channels, and the creation of artificial lakes produced sinks for fine sediments and hotspots for nitrogen processing. Further hydrologic manipulations, such as groundwater pumping, linked surface flows to the aquifer and replaced ephemeral washes with perennial waters. These alterations of hydrologic structure are typical by-products of urban growth in arid and semiarid regions and create distinct spatial and temporal patterns of nitrogen availability.

Diel phosphorus variation and the stoichiometry of ecosystem metabolism in a large spring‐fed river

Elemental cycles are coupled directly and indirectly to ecosystem metabolism at multiple time scales. Understanding coupling in lotic ecosystems has recently advanced through simultaneous high-frequency measurements of multiple solutes. Using hourly in situ measurements of soluble reactive phosphorus (SRP), specific conductance (SpC), and dissolved oxygen (DO), we estimated phosphorus (P) retention pathways and dynamics in a large (discharge, Q ≈ 7.5 m3/s) spring-fed river (Ichetucknee River, Florida, USA). Across eight multi-day deployments, highly regular diel SRP variation of 3–9 μg P/L (mean ∼50 μg P/L) was strongly correlated with DO variation, suggesting photosynthetic control directly via assimilation, and/or indirectly via geochemical reactions. Consistent afternoon SRP maxima and midnight minima suggest peak removal lags gross primary production (GPP) by ∼8 hours. Two overlapping processes were evident, one dominant with maximum removal near midnight, the other smaller with maximum removal near m...

Algal blooms and the nitrogen-enrichment hypothesis in Florida springs: evidence, alternatives, and adaptive management

Contradictions between system-specific evidence and broader paradigms to explain ecosystem behavior present a challenge for natural resource management. In Florida (U.S.A.) springs, increasing nitrate (NO3-) concentrations have been implicated as the cause of algal overgrowth via alleviation of N-limitation. As such, policy and management efforts have centered heavily on reduction of nitrogen (N) loads. While the N-limitation hypothesis appears well founded on broadly supported aquatic eutrophication models, several observations from Florida springs are inconsistent with this hypothesis in its present simplified form. First, NO3- concentration is not correlated with algal abundance across the broad population of springs and is weakly negatively correlated with primary productivity. Second, within individual spring runs, algal mats are largely confined to the headwater reaches within 250 m of spring vents, while elevated NO3- concentrations persist for several kilometers or more. Third, historic observations suggest that establishment of macroalgal mats often lags behind observed increases in NO3- by more than a decade. Fourth, although microcosm experiments indicate high thresholds for N-limitation of algae, experiments in situ have demonstrated only minimal response to N enrichment. These muted responses may reflect large nutrient fluxes in springs, which were sufficient to satisfy present demand even at historic concentrations. New analyses of existing data indicate that dissolved oxygen (DO) has declined dramatically in many Florida springs over the past 30 years, and that DO and grazer abundance are better predictors of algal abundance in springs than are nutrient concentrations. Although a precautionary N-reduction strategy for Florida springs is warranted given demonstrable effects of nutrient enrichment in a broad suite of aquatic systems worldwide, the DO-grazer hypothesis and other potential mechanisms merit increased scientific scrutiny. This case study illustrates the importance of an adaptive approach that explicitly evaluates paradigms as hypotheses and actively seeks alternative explanations.

Reciprocal Biotic Control on Hydrology, Nutrient Gradients, and Landform in the Greater Everglades

Restoration can be viewed as the process of reestablishing both exogenous drivers and internal feedbacks that maintain ecosystems in a desirable state. Correcting exogenous and abiotic drivers is clearly necessary, but may be insufficient to achieve desired outcomes in systems with self-organizing biotic feedbacks that substantially influence ecological stability and timing of responses. Evidence from a broad suite of systems demonstrates the prevalence of biotic control over key ecosystem attributes such as hydroperiod, nutrient gradients, and landform that are most commonly conceived of as exogenously controlled. While a general theory to predict conditions under which biotic controls exert such strong feedbacks is still nascent, it appears clear that the Greater Everglades/South Florida landscape has a high density of such effects. The authors focus on three examples of biotic control over abiotic processes: hydroperiod and discharge controls exerted by peat accretion in the ridge-slough landscape; phosphorus (P) gradients that emerge, at least in part, from interactions between accelerated peat accretion rates, vegetation structure and fauna; and reinforcing feedbacks among land elevation, aquatic respiration, and carbonate dissolution that produce local and landscape basin structure. The authors propose that the unifying theme of biogeomorphic landforms in South Florida is low extant topographic variability, which allows reciprocal biotic modification of local site conditions via mechanisms of peat accretion (including via effects of landscape P redistribution on primary production) or limestone dissolution. Coupling these local positive feedbacks, which drive patch expansion, with inhibitory or negative feedbacks on site suitability at distance, which serve to constrain patch expansion, provide the mechanistic basis for landscape pattern formation. The spatial attributes (range and isotropy) of the distal negative feedback, in particular, control pattern geometry; elucidating the mechanisms and properties of these distal feedbacks is critical to restoration planning.