Kate Lajtha

Stable isotopes in ecology and environmental science

Contributors. Abbreviations. Introduction. 1. Stable isotope chemistry and measurement: a primer. Elizabeth W. Sulzman. Introduction. What isotopes are, what makes them distinct. Properties of ecologically useful stable isotopes. Technological advances and current trends in the ecological use of isotopes. Acknowledgments. References. 2. Sources of variation in the stable isotopic composition of plants. John D. Marshall, J. Renee Brooks, and Kate Lajtha. Introduction. Carbon isotopes. Nitrogen isotopes. Hydrogen and oxygen isotopes. Conclusions. References. 3. Natural 15N- and 13C-abundance as indicators of forest nitrogen status and soil carbon dynamics. Charles T. Garten, Jr, Paul J. Hanson, Donald E. Todd, Jr, Bonnie B. Lau, and Deanne J. Brice. Introduction. Significance of 15N-abundance to soil carbon sequestration. Vertical changes in soil 13C-abundance and soil carbon dynamics. Conclusions. Acknowledgments. References. 4. Soil nitrogen isotope composition. R. Dave Evans. Introduction. Sources of variation in soil 15N. Patterns of soil nitrogen isotope composition. Conclusions. References. 5. Isotopic study of the biology of modern and fossil vertebrates. Paul L. Koch. Introduction. Vertebrate tissues in the fossil record. Controls on the isotopic composition of vertebrate tissues. Preservation of biogenic isotope compositions by vertebrate fossils. Paleobiological applications. Conclusions. A post-script on workshops and literature resources. References. 6. Isotopic tracking of migrant wildlife. Keith A. Hobson. Introduction. Basic principles. Marine systems. Terrestrial systems (excluding deuterium). Using deuterium patterns in precipitation. Conclusions. References. 7. Natural abundance of 15N in marine planktonic ecosystems. Joseph P. Montoya. Introduction. Background. Isotopic variation in marine nitrogen. Source delineation and isotope budgets. Animal fractionation and food web processes. Isotopic transients in marine systems. Compound-specific nitrogen isotope analyses. Conclusions. Acknowledgment. References. 8. Stable isotope studies in marine chemoautotrophically based ecosystems: An update. Cindy Lee Van Dover. Introduction. Isotopic tracing of carbon at methane seeps. Whale falls. Hydrothermal vents. Conclusions. References. 9. Stable isotope ratios as tracers in marine food webs: An update. Robert H. Michener and Les Kaufman. Introduction. Methods of assessing food webs. Phytoplankton and particulate organic carbon. Phytoplankton and particulate organic nitrogen. Marine food webs. Stable isotopes in marine conservation biology. Conclusions. Acknowledgments. References. 10. Stable isotope tracing of temporal and spatial variability in organic matter sources to freshwater ecosystems. Jacques C. Finlay and Carol Kendall. Introduction. Overview of river food webs and stable isotope approaches. Stable isotope ratios of organic matter sources in stream ecosystems. C, N, and S isotopic variability and its applications in river ecology. Conclusions. Acknowledgments. References. 11. Stable isotope tracers in watershed hydrology. Kevin J. McGuire and Jeff McDonnell. Introduction. Basic concepts in watershed hydrology. Why are stable isotopes needed?. General concepts in isotope hydrology. Applications of isotope hydrology in watershed and ecosystem studies. Conclusions. Acknowledgments. References. 12. Tracing anthropogenic inputs of nitrogen to ecosystems. Carol Kendall, Emily M. Elliott, and Scott D. Wankel. Introduction. Isotopic compositions of major N sources to ecosystems. Processes affecting the isotopic composition of DIN. Separating mixing of sources from the effects of cycling. Applications to different environmental settings. What sources of agricultural and urban sources of nitrate can be distinguished using isotopes?. Other tools for tracing anthropogenic contaminants. Conclusions. References. 13. Modeling the dynamics of stable-isotope ratios for ecosystem biogeochemistry. William S. Currie. Introduction. Designing consistent model-data linkages and comparisons. Principles and techniques of stable isotope modeling. Conclusions. Acknowledgments. References. 14. Compound-specific stable isotope analysis in ecology and paleoecology. Richard P. Evershed, Ian D. Bull, Lorna T. Corr, Zoe M. Crossman, Bart E. van Dongen, Claire Evans, Susan Jim, Hazel Mottram, Anna J. Mukherjee, and Richard D. Pancost. Introduction. Why use compound-specific stable isotopes?. Analytical considerations in compound-specific stable isotope analysis. Applications of compound-specific stable isotope approaches in ecology and paleoecology. Conclusions. References. Index

Regional nitrogen budgets and riverine N & P fluxes for the drainages to the North Atlantic Ocean: Natural and human influences

We present estimates of total nitrogen and total phosphorus fluxes in rivers to the North Atlantic Ocean from 14 regions in North America, South America, Europe, and Africa which collectively comprise the drainage basins to the North Atlantic. The Amazon basin dominates the overall phosphorus flux and has the highest phosphorus flux per area. The total nitrogen flux from the Amazon is also large, contributing 3.3 Tg yr-1 out of a total for the entire North Atlantic region of 13.1 Tg yr-1. On a per area basis, however, the largest nitrogen fluxes are found in the highly disturbed watersheds around the North Sea, in northwestern Europe, and in the northeastern U.S., all of which have riverine nitrogen fluxes greater than 1,000 kg N km-2 yr-1.

Couplings of watersheds and coastal waters: Sources and consequences of nutrient enrichment in Waquoit Bay, Massachusetts

Human activities on coastal watersheds provide the major sources of nutrients entering shallow coastal ecosystems. Nutrient loadings from watersheds are the most widespread factor that alters structure and function of receiving aquatic ecosystems. To investigate this coupling of land to marine systems, we are studying a series of subwatersheds of Waquoit Bay that differ in degree of urbanization and hence are exposed to widely different nutrient loading rates. The subwatersheds differ in the number of septic tanks and the relative acreage of forests. In the area of our study, groundwater is the major mechanism that transports nutrients to coastal waters. Although there is some attenuation of nutrient concentrations within the aquifer or at the sediment-water interface, in urbanized areas there are significant increases in the nutrient content of groundwater arriving at the shore’s edge. The groundwater seeps or flows through the sediment-water boundary, and sufficient groundwater-borne nutrients (nitrogen in particular) traverse the sediment-water boundary to cause significant changes in the aquatic ecosystem. These loading-dependent alterations include increased nutrients in water, greater primary production by phytoplankton, and increased macroaglal biomass and growth (mediated by a suite of physiological responses to abundance of nutrients). The increased macroalgal biomass dominates the bay ecosystem through second- or third-order effects such as alterations of nutrient status of water columns and increasing frequency of anoxic events. The increases in seaweeds have decreased the areas covered by eelgrass habitats. The change in habitat type, plus the increased frequency of anoxic events, change the composition of the benthic fauna. The data make evident the importance of bottom-up control in shallow coastal food webs. The coupling of land to sea by groundwater-borne nutrient transport is mediated by a complex series of steps; the cascade of processes make it unlikely to find a one-to-one relation between land use and conditions in the aquatic ecosystem. Study of the process and synthesis by appropriate models may provide a way to deal with the complexities of the coupling.

NITROGEN LOADING FROM COASTAL WATERSHEDS TO RECEIVING ESTUARIES: NEW METHOD AND APPLICATION

In this paper we develop a model to estimate nitrogen loading to watersheds and receiving waters, and then apply the model to gain insight about sources, losses, and transport of nitrogen in groundwater moving through a coastal watershed. The model is developed from data of the Waquoit Bay Land Margin Ecosystems Research project (WBLMER), and from syntheses of published information. The WBLMER nitrogen loading model first estimates inputs by atmospheric deposition, fertilizer use, and wastewater to surfaces of the major types of land use (natural vegetation, turf, agricultural land, residential areas, and impervious surfaces) within the landscape. Then, the model estimates losses of nitrogen in the various compartments of the watershed ecosystem. For atmospheric and fertilizer nitrogen, the model allows losses in vegetation and soils, in the vadose zone, and in the aquifer. For wastewater nitrogen, the model allows losses in septic systems and effluent plumes, and it adds further losses that occur during diffuse transport within aquifers. The calculation of losses is done separately for each major type of land cover, because the processes and loss rates involved differ for different tesserae of the land cover mosaic. If groundwater flows into a freshwater body, the model adds a loss of nitrogen for traversing the freshwater body and then subjects the surviving nitrogen to losses in the aquifer. The WBLMER model is developed for Waquoit Bay, but with inputs for local conditions it is applicable to other rural to suburban watersheds underlain by unconsolidated sandy sediments. Model calculations suggest that the atmosphere contributes 56%, fertilizer 14%, and wastewater 27% of the nitrogen delivered to the surface of the watershed of Waquoit Bay. Losses within the watershed amount to 89% of atmospheric nitrogen, 79% of fertilizer nitrogen, and 65% of wastewater nitrogen. The net result of inputs to the watershed surface and losses within the watershed is that wastewater becomes the largest source (48%) of nitrogen loads to receiving estuaries, followed by atmospheric deposition (30%) and fer- tilizer use (15%). The nitrogen load to estuaries of Waquoit Bay is transported primarily through land parcels covered by residential areas (39%, mainly via wastewater), natural vegetation (21%, by atmospheric deposition), and turf (16%, by atmospheric deposition and fertilizers). Other land covers were involved in lesser throughputs of nitrogen. The model results have implications for management of coastal landscapes and water quality. Most attention should be given to wastewater disposal within the watershed, par- ticularly within 200 m of the shore. Rules regarding setbacks of septic system location relative to shore and nitrogen retention ability of septic systems, will be useful in control of wastewater nitrogen loading. Installation of multiple conventional leaching fields or septic systems in high-flow parcels could be one way to increase nitrogen retention. Control of fertilizer use can help to a modest degree, particularly for optional uses such as lawns situated near shore. Conservation of parcels of accreting natural vegetation should be given high priority, because these environments effectively intercept atmospheric deposition. Areas upgradient from freshwater bodies should be given low priority in plans to control nitrogen loading, because ponds intercept much of the nitrogen transported from upgradient.

Sources of nitrate in rivers draining sixteen watersheds in the northeastern U.S.: Isotopic constraints

The feasibility of using nitrogen and oxygenisotope ratios of nitrate (NO3−) forelucidating sources and transformations ofriverine nitrate was evaluated in a comparativestudy of 16 watersheds in the northeastern U.S.A. Stream water was sampled repeatedly at theoutlets of the watersheds between January andDecember 1999 for determining concentrations,δ15N values, and δ18Ovalues of riverine nitrate.In conjunction with information about land useand nitrogen fluxes,δ15Nnitrate andδ18Onitrate values providedmainly information about sources of riverinenitrate. In predominantly forested watersheds,riverine nitrate had mean concentrations ofless than 0.4 mg NO3−-N L−1,δ15Nnitrate values of lessthan +5‰, and δ18Onitratevalues between +12 and +19‰. This indicatesthat riverine nitrate was almost exclusivelyderived from soil nitrification processes withpotentially minor nitrate contributions fromatmospheric deposition in some catchments. Inwatersheds with significant agricultural andurban land use, concentrations of riverinenitrate were as high as 2.6 mg NO3−-NL−1 with δ15Nnitratevalues between +5 and +8‰ andδ18Onitrate values generallybelow +15‰. Correlations between nitrateconcentrations, δ15Nnitratevalues, and N fluxes suggest that nitrate inwaste water constituted a major, and nitrate inmanure a minor additional source of riverinenitrate. Atmospheric nitrate deposition ornitrate-containing fertilizers were not asignificant source of riverine nitrate inwatersheds with significant agricultural andurban land use. Although complementary studiesindicate that in-stream denitrification wassignificant in all rivers, the isotopiccomposition of riverine nitrate sampled at theoutlet of the 16 watersheds did not provideevidence for denitrification in the form ofelevated δ15Nnitrate andδ18Onitrate values. Relativelylow isotopic enrichment factors for nitrogenand oxygen during in-stream denitrification andcontinuous admixture of nitrate from theabove-described sources are thought to beresponsible for this finding.

Where did all the nitrogen go? Fate of nitrogen inputs to large watersheds in the northeastern U.S.A.

To assess the fate of the large amounts of nitrogen (N) brought into the environment by human activities, we constructed N budgets for sixteen large watersheds (475 to 70,189 km2) in the northeastern U.S.A. These watersheds are mainly forested (48–87%), but vary widely with respect to land use and population density. We combined published data and empirical and process models to set up a complete N budget for these sixteen watersheds. Atmospheric deposition, fertilizer application, net feed and food inputs, biological fixation, river discharge, wood accumulation and export, changes in soil N, and denitrification losses in the landscape and in rivers were considered for the period 1988 to 1992. For the whole area, on average 3420 kg of N is imported annually per km2 of land. Atmospheric N deposition, N2 fixation by plants, and N imported in commercial products (fertilizers, food and feed) contributed to the input in roughly equal contributions. We quantified the fate of these inputs by independent estimates of storage and loss terms, except for denitrification from land, which was estimated from the difference between all inputs and all other storage and loss terms. Of the total storage and losses in the watersheds, about half of the N is lost in gaseous form (51%, largely by denitrification). Additional N is lost in riverine export (20%), in food exports (6%), and in wood exports (5%). Change in storage of N in the watersheds in soil organic matter (9%) and wood (9%) accounts for the remainder of the sinks. The presence of appreciable changes in total N storage on land, which we probably under-rather than overestimated, shows that the N budget is not in steady state, so that drainage and denitrification exports of N may well increase further in the future.

The Biogeochemistry of Phosphorus Cycling and Phosphorus Availability Along a Desert Soil Chronosequence

The biogeochemistry ofthe weathering, landscape movements, and chemical transformations of phosphorus and its availability to plants were examined in a chrono- sequence of soils developed from quartz monzonite alluvium in southern New Mexico. Total P in the soil profile decreased with increasing soil age and was removed from the ecosystem as readily as the most easily leachable base cations. Although Ca-bound forms of P decreased with increasing soil age, Ca-P remained the single largest fraction of total P in all soils. In contrast, Fe- and Al-bound P was a very small percent of total P in all soils. There was little evidence for the stabilization of P by soil organic matter within this ecosystem; both soil organic P and microbial P represented very small pools of total soil P. Phosphorus availability, measured by in situ resin bags, was not well correlated with soil age or total soil P, and P concentrations in shrub tissues did not reflect changes in forms or total amounts of soil P. The biogeochemical cycle of P in this system differs sharply from that in a more mesic, forested system, where fixation by iron and aluminum oxides and biologic activity play more dominant roles in the conservation of P within the ecosystem.

The effect of water and nitrogen amendments on photosynthesis, leaf demography, and resource-use efficiency in Larrea tridentata, a desert evergreen shrub

SummaryIn the Chihuahuan Desert of southern New Mexico, both water and nitrogen limit the primary productivity of Larrea tridentata, a xerophytic evergreen shrub. Net photosynthesis was positively correlated to leaf N, but only in plants that received supplemental water. Nutrient-use efficiency, defined as photosynthetic carbon gain per unit N invested in leaf tissue, declined with increasing leaf N. However, water-use efficiency, defined as the ratio of photosynthesis to transpiration, increased with increasing leaf N, and thus these two measures of resource-use efficiency were inversely correlated. Resorption efficiency was not significantly altered over the nutrient gradient, nor was it affected by irrigation treatments. Leaf longevity decreased significantly with fertilization although the absolute magnitude of this decrease was fairly small, in part due to a large background of insect-induced mortality. Age-specific gas exchange measurements support the hypothesis that leaf aging represents a redistribution of resources, rather than actual deterioration or declining resource-use efficiency.

Carbon cycling in soil

As yet, nobody knows what effects climate change will have on soil carbon reserves, or how those changes will affect the global carbon cycle. Soils are the primary terrestrial repository for carbon, so minor changes in the balance between belowground carbon storage and release could have major impacts on greenhouse gases. Soil fauna, roots, fungi, and microbes interact with mineral and organic matter to process soil carbon. Studies have been hampered by the difficulty of observing processes beneath the earths surface, but advances in science and technology are improving our ability to understand belowground ecosystems.

Biogeochemistry of unpolluted forested watersheds in the Oregon Cascades: temporal patterns of precipitation and stream nitrogen fluxes

We analyzed long-term organic and inorganic nitrogen inputs and outputs in precipitation and streamwater in six watersheds at the H.J. Andrews Experimental Forest in the central Cascade Mountains of Oregon. Total bulk N deposition, averaging 1.6 to 2.0 kg N ha−1 yr−1, is low compared to other sites in the United States and little influenced by anthropogenic N sources. Streamwater N export is also low, averaging <1 kg ha−1 yr−1. DON is the predominant form of N exported from all watersheds, followed by PON, NH4-N, and NO3-N. Total annual stream discharge was a positive predictor of annual DON output in all six watersheds, suggesting that DON export is related to regional precipitation. In contrast, annual discharge was a positive predictor of annual NO3-N output in one watershed, annual NH4-N output in three watersheds, and annual PON output in three watersheds. Of the four forms of N, only DON had consistent seasonal concentration patterns in all watersheds. Peak streamwater DON concentrations occurred in November-December after the onset of fall rains but before the peak in the hydrograph, probably due to flushing of products of decomposition that had built up during the dry summer. Multiple biotic controls on the more labile nitrate and ammonium concentrations in streams may obscure temporal DIN flux patterns from the terrestrial environment. Results from this study underscore the value of using several watersheds from a single climatic zone to make inferences about controls on stream N chemistry; analysis of a single watershed may preclude identification of geographically extensive mechanisms controlling N dynamics.