Barbara L. Bedford

PATTERNS IN NUTRIENT AVAILABILITY AND PLANT DIVERSITY OF TEMPERATE NORTH AMERICAN WETLANDS

Few wetland studies from temperate North America have related either species richness or plant community composition to any direct measure of nutrient availability, or examined changes in species composition following experimental nutrient additions. Studies of wetlands in western Europe and of other terrestrial ecosystems in North America frequently show that nutrient enrichment leads to changes in species composition, declines in overall plant species diversity, and loss of rare and uncommon species. We therefore used an extensive literature survey and analysis of data on plant species composition, species richness, productivity or standing crop, and C:N:P stoichiometry in plant tissues and surface soils to draw conclusions about the nature of nutrient limitation in temperate North American bogs, fens, marshes, and swamps, and to infer their potential response to nutrient enrichment. We searched all major bibliographic data bases for studies containing such data (through March 1998) and added relevant d...

The Need to Define Hydrologic Equivalence at the Landscape Scale for Freshwater Wetland Mitigation

Attempts to replace wetlands or define hydrologic equivalence for wetland mitigation must be based on an understanding of the complexity of wetland hydrology and of the relationship of individual wetlands to the landscape. Because mitigation has the potential to re-configure the kinds and spatial distribution of wetland ecosystems over large geographic areas, I advocate a landscape approach to defining hydrologic equivalence. This approach does not depend on specification of hydroperiod or other hydrologic variables for individual wetlands. It relies instead on knowledge of landscape properties that control wetland hydrology and water chemistry. In this paper I develop the conceptual framework for defining hydrologic equivalence for wetland mitigation viewed as a de facto landscape management policy with the potential to reduce the diversity of wetland types within regions. I review modern hydrogeological understanding of where wetlands form in the landscape and identify key hydrologic variables responsible for the formation of specific wetland types. I also review existing evaluations of mitigation projects in several states. On the basis of these reviews, I argue that, in setting regulatory criteria for judging hydrologic equivalence, the scale must be enlarged from the individual wetland project to include the broader landscape. Only this broader view can provide the context within which decision-makers can evaluate the potential cumulative effects of individual mitigation decisions on- broad-scale patterns of wetland diversity. The landscape approach to defining hydrologic equivalence that I advocate is based on the concept of templates for wetland development. These templates are the diversity of settings created in specific landscapes by the complex interactions of hydrogeologic factors and climate. These interactions, in turn, control key hydrologic variables and hydrologically influenced chemical variables that cause specific wetland types to form. Hydrologic equiv- alence then can be defined at the scale of landscapes in terms of the kinds, numbers, relative abundances, and spatial distribution of wetland templates. The approach can be implemented through the identification of landscape goals and profiles based on knowledge of these templates. The profiles would catalog and map the diversity of wetland templates and the diversity of existing wetland types within a given landscape, focusing the attention of decision-makers on broad-scale patterns of loss in wetland types and providing a context within which individual mitigation projects could be evaluated. Landscape goals for main- taining a diversity of wetland templates are suggested.

FENS OF THE UNITED STATES: DISTRIBUTION, CHARACTERISTICS, AND SCIENTIFIC CONNECTION VERSUS LEGAL ISOLATION

The term fen has been variously used by peatland ecologists, ground-water hydrologists, and vegetation scientists. The common denominator among all types of fens is recognition of the importance of ground-water discharge, especially mineral-rich ground water, in determining fen hydrology, chemistry, and vegetation, in contrast to wetlands whose characteristics are determined primarily by precipitation or surface-water inputs. Thus, fens tend to occur where climate and hydrogeologic setting sustain flows to the planrooting zone of mineral-rich ground water. In the United States, these areas include the glaciated Midwest and Northeast, as well as portions of the Appalachian Mountains and mountainous West. Individually and collectively, fens are among the most floristically diverse of all wetland types, supporting a large number of rare and uncommon bryophytes and vascular plant species, as well as uncommon animals including mammals, reptiles, land snails, butterflies, skippers, and dragonflies. Several species listed under the federal Endangered Species Act inhabit or use fens. Fens also help maintain stream water quality through denitrification and phosphorus sorption. Few estimates of loss and current extent exist, but where estimates are available, they indicate extensive loss, fragmentation, and degradation. Cultural eutrophication threatens the biological and functional integrity of remaining fens because, along with mineral-rich water, low availability of nitrogen and phosphorus controls many of their distinctive characteristics. Because they occur where ground water discharges to the surface, fens are isolated from neither ground water nor surface water. However, the majority of fens develop in headwater areas and could be defined as “isolated” for jurisdictional purposes because of their distance from navigable-in-fact waters. If so defined, the critical roles that fens play in maintaining biological diversity and stream water quality are at risk regarding federal jurisdiction over “isolated waters” because of the 2001 U.S. Supreme Court ruling in the case of Solid Waste Agency of Northern Cook County v. U.S. Army Corps of Engineers.

Erratum to: Cumulative effects on wetland landscapes: links to wetland restoration in the United States and southern Canada

The cumulative effects of human actions on wetland ecosystems motivate current efforts at wetland restoration. They also have created in part the context within which restorations are undertaken. Using modern hydrogeological understanding of wetland-landscape linkages, I argue that restorations should begin with a cumulative impact analysis for the entire region in which the restoration is proposed. The analysis, however, should not focus merely on number of hectares of wetlands lost or degraded. It should be based on the concept of templates for wetland development. These templates are the diversity of settings created in specific landscapes by the complex interactions of hydrogeologic factors and climate. They control key hydrologic variables and hydrologically influenced chemical variables that cause specific wetland types to form and to be maintained through time. They also determine in large part the biogeochemical cycling characteristics specific to different types of wetlands. They thus account for both the biological and functional diversity of wetlands. A cumulative impact assessment for restoration purposes should identify the kinds, numbers, relative abundances, and spatial distribution of wetland templates in a region—both past and present. These past and present profiles of the wetland landscape can be used to make decisions regarding the type and location of restorations. Matching type and location to the appropriate hydrogeologic setting will maximize the probability of success for individual projects. Regional wetland diversity can be restored if individual restoration decisions about wetland type and location are made in light of the diversity of templates in past and present regional profiles.

Evaluating cumulative effects on wetland functions: A conceptual overview and generic framework

This article outlines conceptual and methodological issues that must be confronted in developing a sound scientific basis for investigating cumulative effects on freshwater wetlands. We are particularly concerned with: (1) effects expressed at temporal and spatial scales beyond those of the individual disturbance, specific project, or single wetland, that is, effects occurring at the watershed or regional landscape level; and (2) the scientific (technical) component of the overall assessment process. Our aim is to lay the foundation for a research program to develop methods to quantify cumulative effects of wetland loss or degradation on the functioning of interacting systems of wetlands. Toward that goal we: (1) define the concept of cumulative effects in terms that permit scientific investigation of effects; (2) distinguish the scientific component of cumulative impact analysis from other aspects of the assessment process; (3) define critical scientific issues in assessing cumulative effects on wetlands; and (4) set up a hypothetical and generic structure for measuring cumulative effects on the functioning of wetlands as landscape systems.We provide a generic framework for evaluating cumulative effects on three basic wetland landscape functions: flood storage, water quality, and life support. Critical scientific issues include appropriate delineations of scales, identification of threshold responses, and the influence on different functions of wetland size, shape, and position in the landscape.The contribution of a particular wetland to landscape function within watersheds or regions will be determined by its intrinsic characteristics, e.g., size, morphometry, type, percent organic matter in the sediments, and hydrologic regime, and by extrinsic factors, i.e., the wetlands context in the landscape mosaic. Any cumulative effects evaluation must take into account the relationship between these intrinsic and extrinsic attributes and overall landscape function. We use the magnitude of exchanges among component wetlands in a watershed or larger landscape as the basis for defining the geographic boundaries of the assessment. The time scales of recovery for processes controlling particular wetland functions determine temporal boundaries. Landscape-level measures are proposed for each function.

Developing the scientific basis for assessing cumulative effects of wetland loss and degradation on landscape functions: Status, perspectives, and prospects

The incongruity between the regional and national scales at which wetland losses are occurring, and the project-specific scale at which wetlands are regulated and studied, has become obvious. This article presents a synthesis of recent efforts by the US Environmental Protection Agency and the Ecosystems Research Center at Cornell University to bring wetland science and regulation into alignment with the reality of the cumulative effects of wetland loss and degradation on entire landscapes and regions. The synthesis is drawn from the other articles in this volume, the workshop that initiated them, and the scientific literature. It summarizes the status of our present scientific understanding, discusses means by which to actualize the existing potential for matching the scales of research and regulation with the scales at which effects are observed, and provides guidelines for building a stronger scientific base for landscape-level assessments of cumulative effects. It also provides the outlines for a synoptic and qualitative approach to cumulative effects assessment based on a reexamination of the generic assessment framework we proposed elsewhere in this volume.The primary conclusion to be drawn from the articles and the workshop is that a sound scientific basis for regulation will not come merely from acquiring more information on more variables. It will come from recognizing that a perceptual shift to larger temporal, spatial, and organizational scales is overdue. The shift in scale will dictate different—not necessarily more—variables to be measured in future wetland research and considered in wetland regulation.

Restoration of wetland vegetation with transplanted wetland soil: An experimental study

Restoration of drained wetlands requires the re-establishment of a native wetland plant community. This can be difficult in areas where long-term drainage has eliminated wetland vegetation and significantly reduced the number of viable wetland plant seeds in the seed bank. This study of U.S. Fish and Wildlife Service wetland restoration sites in northern New York examines the effectiveness of transplanting wetland soil from small remnant wetlands in the drainage ditches to the area that becomes shallow marsh following reflooding. The results of two experiments are reported, including a small-scale study of transplantation techniques using small plots with treatments and controls established by hand, and a large-scale application of soil transplantation and site-preparation techniques using heavy equipment to establish large plots across entire wetland basins. In the small-scale study, the transplant plots had significantly lower wetland index values, indicating greater dominance of wetland plants, after one growing season but not after two. Transplant plots also had more wetland plant species and more wetland plant cover than natural control plots, and these differences persisted through the second growing season. Litter removal and soil disturbance also lowered the wetland index values and increased wetland plant species number and cover, but only for the first growing season. In the large-scale study, soil transplantation significantly increased both the number of species and the amount of cover of wetland plants and of plants valuable as wildlife food sources. Mowing and plowing treatments increased wetland plant establishment, but much less than soil transplantation, and plowing significantly increased the establishment of cattail (Typha spp.), an undesirable invasive species in small wildlife marshes. Soil transplantation should be a particularly effective technique for improving wetland plant establishment and limiting cattail encroachment in areas disturbed by dike construction.

Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.

Wetland dicots and monocots differ in colonization by arbuscular mycorrhizal fungi and dark septate endophytes

As an initial step towards evaluating whether mycorrhizas influence composition and diversity in calcareous fen plant communities, we surveyed root colonization by arbuscular mycorrhizal fungi (AMF) and dark septate endophytic fungi (DSE) in 67 plant species in three different fens in central New York State (USA). We found colonization by AMF and DSE in most plant species at all three sites, with the type and extent of colonization differing between monocots and dicots. On average, AMF colonization was higher in dicots (58±3%, mean±SE) than in monocots (13±4%) but DSE colonization followed the opposite trend (24±3% in monocots and 9±1% in dicots). In sedges and cattails, two monocot families that are often abundant in fens and other wetlands, AMF colonization was usually very low (<10%) in five species and completely absent in seven others. However, DSE colonization in these species was frequently observed. Responses of wetland plants to AMF and DSE are poorly understood, but in the fen communities surveyed, dicots appear to be in a better position to respond to AMF than many of these more abundant monocots (e.g., sedges and cattails). In contrast, these monocots may be more likely to respond to DSE. Future work directed towards understanding the response of these wetland plants to AMF and DSE should provide insight into the roles these fungal symbionts play in influencing diversity in fen plant communities.

PATHWAYS OF NUTRIENT LOADING AND IMPACTS ON PLANT DIVERSITY IN A NEW YORK PEATLAND

Nutrient loading is a subtle, yet serious threat to the preservation of high diversity wetlands such as peatlands. Pathways of nutrient loading and impacts on plant diversity in a small peatland in New York State, USA were determined by collecting and analyzing a suite of hydrogeological, hydro-chemical, soil, and vegetation data. Piezometer clusters within an intensive network constituted hydro-chemical sampling points and focal points for randomly selected vegetation quadrats and soil-coring locations. Hydrogeological data and nutrient analyses showed that P and K loading occurred chiefly by means of overland flow from an adjacent farm field, whereas N loading occurred predominantly through ground-water flow from the farm field. Redundancy analysis and polynomial regression showed that nutrients, particularly total P in peat, total K in peat, extractable NH4-N, and NO3-N flux in ground water, were strongly negatively correlated with plant diversity measures at the site. No other environmental variables except vegetation measures associated with eutrophication demonstrated such a strong relationship with plant diversity. Nitrate loading over 4 mg m−2 day−1 was associated with low plant diversity, and Ca fluxes between 80 and 130 mg m−2 day−1 were associated with high plant diversity. Areas in the site with particularly low vascular plant and bryophyte species richness and Shannon-Wiener diversity (H′) occurred adjacent to the farm field and near a hillside spring. High H′ and species richness of vascular plants and bryophytes occurred in areas that were further removed from agriculture, contained no highly dominant vegetation, and were situated directly along the ground-water flow paths of springs. These areas were characterized by relatively constant water levels and consistent, yet moderate fluxes of base cations and nutrients. Overall, this study demonstrates that knowledge of site hydrogeology is crucial for determining potential pathways of nutrient loading and for developing relationships between nutrient inflows and wetland plant diversity.