Jill R. Welter

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.

Sources of nitrogen to the riparian zone of a desert stream: Implications for riparian vegetation and nitrogen retention

Riparian zones effectively remove nitrogen (N) from water flowing through riparian soils, particularly in agricultural watersheds. The mechanism of N removal is still unclear, especially the role of vegetation. Uptake and denitrification are the two most commonly studied mechanisms. Retention of groundwater N by plant uptake is often inferred from measurements of N in net incremental biomass. However, this assumes other sources of N are not contributing to the N demand of plants. The purpose of this work was to investigate the relative importance of three sources of available N to riparian trees in a desert stream—input in stream water during floods, input during baseflow, and mineralization of N from soil organic matter. Two approaches were used; a mass balance approach in which the mass of available N from each source was estimated, and a correlational approach in which indexes of each source were compared to leaf N for individual willow trees. Total N from all sources was 396 kg ha−1 y−1, with 172 kg ha−1 y−1 from mineralization, 214 kg ha−1 y−1 from the stream during baseflow, and 9.6 kg ha−1 y−1 from floods. Leaf N was significantly related to N mineralization rates and flood inputs; it was not related to baseflow inputs. We conclude that mineralization is a major source of available N for willow trees, subsidized by input of N from floods. Baseflow inputs are most likely removed by rapid denitrification at the stream–riparian edge, while higher rates of flood supply exceed the capacity of this “filter.”

Light-mediated thresholds in stream-water nutrient composition in a river network

The elemental composition of solutes transported by rivers reflects combined influences of surrounding watersheds and transformations within stream networks, yet comparatively little is known about downstream changes in effects of watershed loading vs. in-channel processes. In the forested watershed of a river under a mediterranean hydrologic regime, we examined the influence of longitudinal changes in environmental conditions on water-column nutrient composition during summer base flow across a network of sites ranging from strongly heterotrophic headwater streams to larger, more autotrophic sites downstream. Small streams (0.1-10 km2 watershed area) had longitudinally similar nutrient concentration and composition with low (approximately 2) dissolved nitrogen (N) to phosphorus (P) ratios. Abrupt deviations from this pattern were observed in larger streams with watershed areas > 100 km2 where insolation and algal abundance and production rapidly increased. Downstream, phosphorus and silica concentrations decreased by > 50% compared to headwater streams, and dissolved organic carbon and nitrogen increased by approximately 3-6 times. Decreasing dissolved P and increasing dissolved N raised stream-water N:P to 46 at the most downstream sites, suggesting a transition from N limitation in headwaters to potential P limitation in larger channels. We hypothesize that these changes were mediated by increasing algal photosynthesis and N fixation by benthic algal assemblages, which, in response to increasing light availability, strongly altered stream-water nutrient concentration and stoichiometry in larger streams and rivers.

Hydrologic exchange and N uptake by riparian vegetation in an arid-land stream

Abstract Riparian zones can strongly influence the exchange of nutrients between streams and their watersheds. Most riparian studies have been done in mesic watersheds, which differ significantly from arid-land watersheds hydrologically. The goals of our work were to determine the strength and direction of hydrologic linkages between stream and riparian zone, and to estimate the extent of uptake of streamwater N by riparian trees in Sycamore Creek, a Sonoran Desert stream. Br− and 15NH4+ were added simultaneously to the surface stream to trace water and N from stream to riparian zone. Br− concentrations in riparian wells installed downstream of the release point increased during the addition, demonstrating a strong hydrologic linkage from stream to riparian zone. Percentage stream water in wells increased in a downstream direction, suggesting little or no input of water laterally from uplands or vertically from deep groundwater. Leaf and wood samples collected from willow trees downstream of the addition point became significantly labeled with 15N during the addition, indicating uptake of streamwater N. Other tree species did not become labeled, most likely because they were located farther from the stream channel than the willows. Results from our study provide evidence of strong hydrologic linkage between stream and riparian zone and suggest that N demand by riparian vegetation is a potentially significant sink for streamwater N.

Does N2 fixation amplify the temperature dependence of ecosystem metabolism

Variation in resource supply can cause variation in temperature dependences of metabolic processes (e.g., photosynthesis and respiration). Understanding such divergence is particularly important when using metabolic theory to predict ecosystem responses to climate warming. Few studies, however, have assessed the effect of temperature-resource interactions on metabolic processes, particularly in cases where the supply of limiting resources exhibits temperature dependence. We investigated the responses of biomass accrual, gross primary production (GPP), community respiration (CR), and N2 fixation to warming during biofilm development in a streamside channel experiment. Areal rates of GPP, CR, biomass accrual, and N2 fixation scaled positively with temperature, showing a 32- to 71-fold range across the temperature gradient (approximately 7 degrees-24 degrees C). Areal N2-fixation rates exhibited apparent activation energies (1.5-2.0 eV; 1 eV = approximately 1.6 x 10(-19) J) approximating the activation energy of the nitrogenase reaction. In contrast, mean apparent activation energies for areal rates of GPP (2.1-2.2 eV) and CR (1.6-1.9 eV) were 6.5- and 2.7-fold higher than estimates based on metabolic theory predictions (i.e., 0.32 and 0.65 eV, respectively) and did not significantly differ from the apparent activation energy observed for N2 fixation. Mass-specific activation energies for N2 fixation (1.4-1.6 eV), GPP (0.3-0.5 eV), and CR (no observed temperature relationship) were near or lower than theoretical predictions. We attribute the divergence of areal activation energies from those predicted by metabolic theory to increases in N2 fixation with temperature, leading to amplified temperature dependences of biomass accrual and areal rates of GPP and R. Such interactions between temperature dependences must be incorporated into metabolic models to improve predictions of ecosystem responses to climate change.

Flowpaths as Integrators of Heterogeneity in Streams and Landscapes

Streams are heterogeneous in both space and time. Hydrologic flowpaths along which biogeochemical processing occurs integrate different patches of the stream. Disturbance events (flood and drying) change these patches, alter connectivity, and reinforce spatial heterogeneity. Heterogeneity within patches (surface stream, hyporheic zone, sand bars, and riparian zone) is generated by the interaction of nitrogen (the limiting nutrient) in transport and organisms such as algae and bacteria. These organisms store nitrogen as they grow, alter N forms and concentrations in transport, and in some cases (e.g., denitrification) export it to the atmosphere. Changes in nitrogen in transport can be large, as are community responses to nitrogen availability, thus reinforcing spatial heterogeneity in successional time. Flowpaths connect patches as well and generate changes in recipient patches as a function of nitrogen delivery rate. This is especially evident at patch boundaries. In streams, flow is markedly linear and inexorably downstream in orientation; however, landscapes are drained by coalescing, dendritic networks that intimately connect stream channels with terrestrial flowpaths over and beneath soils. We propose that a unified theory of landscapes will require a focus on spatial linkage, a consideration of both spatial and temporal heterogeneity, and a blurring of distinctions between terrestrial and aquatic elements.

Bridging Food Webs, Ecosystem Metabolism, and Biogeochemistry Using Ecological Stoichiometry Theory

Although aquatic ecologists and biogeochemists are well aware of the crucial importance of ecosystem functions, i.e., how biota drive biogeochemical processes and vice-versa, linking these fields in conceptual models is still uncommon. Attempts to explain the variability in elemental cycling consequently miss an important biological component and thereby impede a comprehensive understanding of the underlying processes governing energy and matter flow and transformation. The fate of multiple chemical elements in ecosystems is strongly linked by biotic demand and uptake; thus, considering elemental stoichiometry is important for both biogeochemical and ecological research. Nonetheless, assessments of ecological stoichiometry (ES) often focus on the elemental content of biota rather than taking a more holistic view by examining both elemental pools and fluxes (e.g., organismal stoichiometry and ecosystem process rates). ES theory holds the promise to be a unifying concept to link across hierarchical scales of patterns and processes in ecology, but this has not been fully achieved. Therefore, we propose connecting the expertise of aquatic ecologists and biogeochemists with ES theory as a common currency to connect food webs, ecosystem metabolism, and biogeochemistry, as they are inherently concatenated by the transfer of carbon, nitrogen, and phosphorous through biotic and abiotic nutrient transformation and fluxes. Several new studies exist that demonstrate the connections between food web ecology, biogeochemistry, and ecosystem metabolism. In addition to a general introduction into the topic, this paper presents examples of how these fields can be combined with a focus on ES. In this review, a series of concepts have guided the discussion: (1) changing biogeochemistry affects trophic interactions and ecosystem processes by altering the elemental ratios of key species and assemblages; (2) changing trophic dynamics influences the transformation and fluxes of matter across environmental boundaries; (3) changing ecosystem metabolism will alter the chemical diversity of the non-living environment. Finally, we propose that using ES to link nutrient cycling, trophic dynamics, and ecosystem metabolism would allow for a more holistic understanding of ecosystem functions in a changing environment.