Ecology of Freshwater and Estuarine Wetlands

ECOLOGY OF FRESHWATER AND ESTUARINE WETLANDS

Ecology of Freshwater and Estuarine Wetlands

SECOND EDITION

Edited by

DAROLD P. BATZER and REBECCA R. SHARITZ

UNIVERSITY OF CALIFORNIA PRESS

University of California Press, one of the most distinguished university presses in the United States, enriches lives around the world by advancing scholarship in the humanities, social sciences, and natural sciences. Its activities are supported by the UC Press Foundation and by philanthropic contributions from individuals and institutions. For more information, visit www.ucpress.edu.

University of California Press

Oakland, California

© 2014 by The Regents of the University of California

Library of Congress Cataloging-in-Publication Data

Ecology of freshwater and estuarine wetlands / Darold P. Batzer and Rebecca R. Sharitz, editors. — Second edition.

    pages    cm

Includes bibliographical references and index.

ISBN 978-0-520-27858-5 (cloth : alk. paper) —

ISBN 978-0-520-95911-8 (e-book)

    1. Wetland ecology. I. Batzer, Darold P. II. Sharitz, Rebecca R.

QH541.5.M3E266    2014

577.68—dc23            2014011257

Manufactured in China.

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10  9  8  7  6  5  4  3  2  1

The paper used in this publication meets the minimum requirements of ANSI/NISO Z39.48-1992 (R 2002) (Permanence of Paper).

Large cover image: Freshwater marsh in the Mississippi River (Birdsfoot) Delta, Louisiana, with duck potato (Sagittaria lancifolia) in foreground; photo courtesy of Karen McKee.

Small cover images, top to bottom: Muskrat (Ondatra zibethicus); photo courtesy of Michele Rundquist-Franz. Bur-marigold (Bidens laevis); photo courtesy of Andrew Baldwin. Planthopper (Prokelisia sp.); photo courtesy of Steve Pennings. Sea purslane (Sesuvium portulacastrum); photo courtesy of Karen McKee.

CONTENTS

CONTRIBUTORS

PREFACE

1. Ecology of Freshwater and Estuarine Wetlands: An Introduction

REBECCA R. SHARITZ, DAROLD P. BATZER, AND STEVEN C. PENNINGS

What Is a Wetland?

Why Are Wetlands Important?

Characteristics of Selected Wetlands

Wetland Loss and Degradation

What This Book Covers

2. Wetland Soils, Hydrology, and Geomorphology

C. RHETT JACKSON, JAMES A. THOMPSON, AND RANDALL K. KOLKA

Wetland Soils

Hillslope and Wetland Hydrology

Wetland Water Budgets

Hydropatterns

Wetland Hydraulics and Residence Time

Geomorphic Controls on Wetland Hydrology

Effects of Land Use on Wetland Hydrology

3. Abiotic Constraints for Wetland Plants and Animals

IRVING A. MENDELSSOHN, DAROLD P. BATZER, COURTNEY R. HOLT, AND SEAN A. GRAHAM

Hydrology

Salinity

Acid Conditions

4. Wetland Microbial Ecology and Biogeochemistry

PAUL I. BOON, PETER C. POLLARD, AND DARREN RYDER

A Cast of Billions: The Taxonomic and Physiological Diversity of Wetland Microbes

Are Microbes Important to the Ecological Structure and Function of Wetlands?

How Can We Study Wetland Microbes?

Microbial Habitats in Wetlands

Biogeochemistry and Carbon Transformations

Microbes, Plant Decay, and Nutrient Regeneration

Microbial Autotrophy and Wetland Food Webs

Microbial Heterotrophy and Wetland Food Webs: The Microbial Loop

5. Development of Wetland Plant Communities

REBECCA R. SHARITZ AND STEVEN C. PENNINGS

The Importance of Hydrologic Conditions

Plant Community Development

Plant Distributions in Wetlands

Primary Productivity

Limiting Nutrients in Wetlands

6. Wetland Animal Ecology

DAROLD P. BATZER, ROBERT COOPER, AND SCOTT A. WISSINGER

Trophic Ecology

Community Ecology

Focal Wetland Animals

7. Carbon Dynamics and Ecosystem Processes

SCOTT D. BRIDGHAM

The Basic Carbon Cycle: Metrics of Productivity and Carbon Balance

Breakdown and Decomposition of Organic Matter

Anaerobic Carbon Cycling

Why Do Wetlands Accumulate Soil Carbon?

Dissolved Organic Matter Fluxes from Wetlands

The Big Picture

8. United States Wetland Regulation, Policy, and Assessment

C. ANDREW COLE AND D. ERIC SOMERVILLE

Wetland Definitions

Federal Jurisdiction of Wetlands

Wetland Delineation

Wetland Functions and Values

Functional Assessment Methods

9. Wetland Restoration

SUSAN M. GALATOWITSCH AND JOY B. ZEDLER

Adaptive Management and Restoration

Selecting Suitable Targets and Setting Restoration Goals

Restoration Planning

Restoration Methods

Evaluating Progress and Long-Term Stewardship

Restoration Challenges and Emerging Issues

10. Consequences for Wetlands of a Changing Global Environment

ROBERT R. TWILLEY AND MARK M. BRINSON

Assumptions

Effects on Species Composition and Redistribution

Effects on Carbon Balance

Effects on Wetland Ecogeomorphic Types

Management and Policy Options

LITERATURE CITED

INDEX

CONTRIBUTORS

DAROLD P. BATZER Department of Entomology, University of Georgia, Athens, Georgia 30602, USA

PAUL I. BOON Institute for Sustainability and Innovation, Victoria University, Footscray Park campus, P.O. Box 14428, Melbourne City Mail Centre, Victoria 8001, Australia

SCOTT BRIDGHAM Environmental Sciences Institute and Institute of Ecology and Evolution, 5289 University of Oregon, Eugene, Oregon 97403, USA

MARK M. BRINSON (Deceased), Biology Department, Howell Science Complex, N-108, East Carolina University, Greenville, North Carolina, USA

C. ANDREW COLE Department of Landscape Architecture, 329 Stuckeman Family Building, Penn State University, University Park, Pennsylvania 16802, USA

ROBERT COOPER Warnell School of Forest Resources, University of Georgia, Athens, Georgia 30602, USA

SUSAN GALATOWITSCH Department of Fisheries, Wildlife and Conservation Biology, University of Minnesota, 135 Skok Hall, 2003 Upper Buford Circle, St. Paul, Minnesota, 55108, USA

SEAN A. GRAHAM Wetland Biogeochemistry Institute and Department of Oceanography and Coastal Sciences, School of the Coast and Environment, Louisiana State University, Baton Rouge, Louisiana 70803, USA

COURTNEY R. HOLT Department of Entomology, University of Georgia, Athens, Georgia 30602, USA

C. RHETT JACKSON Warnell School of Forest Resources, University of Georgia, Athens, Georgia 30602, USA

RANDALL K. KOLKA USDA Forest Service, Northern Research Station, 1831 Hwy. 169 E., Grand Rapids, MN 55744-3399, USA

IRVING A. MENDELSSOHN Wetland Biogeochemistry Institute and Department of Oceanography and Coastal Sciences, School of the Coast and Environment, Louisiana State University, Baton Rouge, Louisiana 70803, USA

STEVEN C. PENNINGS Department of Biology and Biochemistry, University of Houston, Houston, Texas 77204, USA

PETER C. POLLARD Australian Rivers Institute, Griffith University, Nathan campus, Brisbane, Queensland 4111, Australia

DARREN RYDER School of Environmental and Rural Science, University of New England, Armidale, New South Wales 2351, Australia

REBECCA R. SHARITZ University of Georgia, Savannah River Ecology Laboratory, P.O. Drawer E, Aiken, South Carolina 29802, USA

D. ERIC SOMERVILLE US Environmental Protection Agency, Athens, Georgia 30605, USA

JAMES A. THOMPSON West Virginia University, Division of Plant and Soil Sciences, 1090 Agricultural Sciences Building, P.O. Box 6108, Morgantown, West Virginia 26506, USA

ROBERT R. TWILLEY Louisiana Sea Grant College Program, Oceanography and Coastal Sciences, 239 Sea Grant Building, Louisiana State University, Baton Rouge, Louisiana 70803, USA

SCOTT A. WISSINGER Department of Biology, Allegheny College, Meadville, Pennsylvania 16335, USA

JOY B. ZEDLER Botany Department and Arboretum, University of Wisconsin, Madison, Wisconsin 57306, USA

PREFACE

We developed this second edition with two main goals in mind. First, of course, was to update chapters to cover the wealth of new information published in the past several years. Second was to make the text more accessible to those students using it for wetland science courses. To this end we have conducted significant reorganization. In Chapter 1, we now describe up-front the general aspects of different wetland types, to ground students for the more complex material to come. We also reorganized, refocused, and in some cases completely revamped the subject-specific chapters (2–10). This was accomplished by consolidating some chapters, focusing more on the central themes of wetland ecology, and paring more tangential ideas. Some particularly complex, but important, ideas have now been placed into boxes where readers specifically interested in the particular topic can review the material without breaking up the flow of the main themes for more general readers. We have increased the number of figures and photos in this new edition with the aim of making the work more interesting. The text of this second edition is actually somewhat shorter than the first. Despite the focus on students, this book still synthesizes the most recent themes in wetland science, making it a useful reference for all.

Since the first edition, regrettably two of our authors—Robert Wetzel and Mark Brinson—have passed away. We dedicate this second edition to their memories. Fortunately new authors stepped up to cover the material previously in Bob’s and Mark’s chapters. Some of the material formerly covered in Bob’s chapter is now dealt with in the chapter by Paul Boon and co-authors, and some in a completely new chapter by Scott Bridgham. Robert Twilley, a former student of Mark Brinson’s, revised and updated Mark’s climate change chapter, retaining Mark as co-author. Other new lead authors include Sue Galatowitsch, who joined Joy Zedler to revise and update the restoration chapter; and Andy Cole, who joined Eric Somerville to update the new regulations and assessment chapter. Several new co-authors were added to increase the breadth of specific chapters. Overall, we feel that in this second edition we have assembled some of the finest wetland scientists available as authors, and they provide an authoritative look into their areas of wetland expertise.

Darold P. Batzer and Rebecca R. Sharitz

ONE

Ecology of Freshwater and Estuarine Wetlands: An Introduction

REBECCA R. SHARITZ, DAROLD P. BATZER, and STEVEN C. PENNINGS

WHAT IS A WETLAND?

WHY ARE WETLANDS IMPORTANT?

CHARACTERISTICS OF SELECTED WETLANDS

Wetlands with Predominantly Precipitation Inputs

Wetlands with Predominately Groundwater Inputs

Wetlands with Predominately Surface Water Inputs

WETLAND LOSS AND DEGRADATION

WHAT THIS BOOK COVERS

What Is a Wetland?

The study of wetland ecology can entail an issue that rarely needs consideration by terrestrial or aquatic ecologists: the need to define the habitat. What exactly constitutes a wetland may not always be clear. Thus, it seems appropriate to begin by defining the word wetland. The Oxford English Dictionary says, "Wetland (F. wet a. + land sb.)—an area of land that is usually saturated with water, often a marsh or swamp." While covering the basic pairing of the words wet and land, this definition is rather ambiguous. Does usually saturated mean at least half of the time? That would omit many seasonally flooded habitats that most ecologists would consider wetlands. Under this definition, it also seems that lakes or rivers could be considered wetlands. A more refined definition is clearly needed for wetland science or policy.

Because defining wetland is especially important in terms of policy, it is not surprising that governmental agencies began to develop the first comprehensive definitions (see Chapter 8). One influential definition was derived for the U.S. Fish and Wildlife Service (USFWS) (Cowardin et al. 1979):

Wetlands are lands transitional between terrestrial and aquatic systems where the water table is usually at or near the surface or the land is covered by shallow water. Wetlands must have one or more of the following three attributes: (1) at least periodically, the land supports predominately hydrophytes; (2) the substrate is predominantly undrained hydric soil; and (3) the substrate is nonsoil and is saturated with water or covered by shallow water at some time during the growing season of each year.

This USFWS definition emphasizes the importance of hydrology, soils, and vegetation, which you will see is a recurring theme in wetland definitions. The U.S. Army Corps of Engineers (USACE), the primary permitting agency for wetlands of the United States, adopted a slightly different wording (Environmental Laboratory 1987):

The term wetlands means those areas that are inundated or saturated by surface or ground water at a frequency and duration sufficient to support, and that under normal circumstances do support, a prevalence of vegetation typically adapted for life in saturated soil conditions. Wetlands generally include swamps, marshes, bogs, and similar areas.

This definition also incorporates hydrology, soils, and vegetation, but it is more restrictive than the USFWS definition. The USACE definition requires all three features to be present, while the USFWS Cowardin definition indicates that only one of the three conditions needs to occur. Despite its exclusive nature, the USACE definition has been adopted as the authority to define legal (or jurisdictional) wetlands of the United States.

In Canada, a very similar definition for wetland is used by the National Wetlands Working Group (NWWG). This definition also focuses on hydrology, soils, and vegetation, but is more expansive, acknowledging that aquatic processes and biologic factors other than just soils and plants may also be useful for classification (NWWG 1988):

Wetland is defined as land that has the water table at, near, or above the land surface or which is saturated for a long enough period to promote wetland or aquatic processes as indicted by hydric soils, hydrophytic vegetation, and various kinds of biological activity that are adapted to the wet environment.

The NWWG definition is not an official legal standard in Canada but is widely used or adapted by various governmental agencies for setting policy about wetlands (personal communication, Barry Warner, University of Waterloo, Ontario, Canada).

An international definition for wetland was developed for the Ramsar Convention, an intergovernmental treaty regarding wetland conservation initiated in 1971 (which met in Ramsar, Iran). The most recent information (see www.ramsar.org) provides this definition:

Wetlands are areas of marsh, fen, peatland or water, whether natural or artificial, permanent or temporary, with water that is static or flowing, fresh, brackish or salt, including areas of marine water the depth of which at low tide does not exceed six metres.

The emphasis on soils and plant adaptations found in other definitions is absent from this one, possibly because it targets non-scientists who may not be familiar with what constitutes wetland conditions. However, hydrology remains a focal tenet. This definition is currently being applied to identify wetlands in countries in Asia (e.g., Gong et al. 2010).

As ecologists, we must realize that political definitions may not cover all habitats that function ecologically as wetlands. For example, mud flats devoid of vegetation, floodplains that primarily flood in winter outside the growing season, and flooded areas of floodplains where anoxic soil conditions do not develop are all probably ecological wetlands but may not fit some legal definitions. In Georgia, for example, we have seen floodplains repeatedly covered by as much as 1 meter of water (Fig. 1.1), yet competent delineators following USACE criteria determined that the majority of the floodplain area did not meet the legal US definition of a wetland (hydric soil was not present). However, the determination that these floodplains were not jurisdictional wetlands did not affect the responses of soil-dwelling arthropods and herbaceous plants that were covered by the water or the functioning of waterfowl or fish that were swimming and feeding in those habitats. Although definitions do serve a purpose, especially for regulation (see Chapter 8), ecologists should not be constrained by them when studying wetlands.

FIGURE 1.1. Flooded riparian wetland of the North Oconee River, Georgia. Although the site floods almost yearly, it does so in winter, out of the growing season. Additionally, hydric soils are not extensive. As such, using current criteria for delineating wetlands in the United States, most of the floodplain is not considered a jurisdictional wetland. Courtesy of Mark Galatowitsch.

Nonetheless, for ecologists seeking a biologically useful definition for wetlands, we recommend the simple, straightforward, yet inclusive, definition put forward by Paul Keddy (2000):

A wetland is an ecosystem that arises when inundation by water produces soils dominated by anaerobic processes and forces the biota, particularly rooted plants, to exhibit adaptations to tolerate flooding.

Why Are Wetlands Important?

Wetlands comprise only about 6% of the earth’s land surface, but ecologically they are disproportionately important. For example, 25% of the plant species in Malaysia occur in only one wetland type, peat swamps (Anderson 1983). Almost 10% of the world’s fish fauna occurs in the Amazon basin (Groombridge and Jenkins 1998). The vast majority of amphibians are linked to wetlands, even if only for reproduction. Wetlands are particularly important habitats for birds, with many species occurring only in association with wetlands. Bird watching and waterfowl hunting are major human activities related to wetlands. Because wetlands support both terrestrial and aquatic biota, they are unusually diverse (Gopal et al. 2000). Those taxa unique to wetlands will contribute significantly to the overall diversity of regions containing numerous wetlands.

Besides supporting the plethora of plants and animals of interest to ecologists and nature enthusiasts, wetlands provide an assortment of ecosystem services of considerable value to all people. Costanza et al. (1997) estimated the economic values of services provided by the world’s ecosystems and found that, on a per-hectare basis, estuaries and freshwater floodplains/swamps were the world’s two most valuable ecosystem types (Table 1.1). The value of these wetlands to people stems primarily from their roles in nutrient cycling, water supplies, disturbance (flood) regulation, and wastewater treatment. However, recreation, food production, and cultural (aesthetic, artistic, educational, spiritual, or scientific) values are also important (Costanza et al. 1997). Many of these services are accomplished by wetland biota (microbes, plants, and animals). In Canada, an assessment of the economic values of the country’s wetlands (see NWWG 1988, Table 10-15) came up with an estimate of almost $10,000,000,000 (1985 Canadian dollars), of which almost half was attributed to recreational values (nature appreciation activities, fishing, and hunting). However, despite the considerable economic value of wetlands, there is a long history of humans destroying or degrading the world’s wetland resources.

TABLE 1.1 Estimated Relative Economic Values per Hectare of Services Provided by the World’s Ecosystems

Characteristics of Selected Wetlands

Throughout the world, the types of plant and animal communities that occur in wetlands are a result of climate, geomorphology and landscape position, soils, water source and chemistry, and numerous environmental factors including disturbance. Wetlands are found on every continent except Antarctica and extend from the tropics to the tundra. Estimates of the extent of the world’s wetlands vary, but generally range from around 7 to 10 million square kilometers (Mitsch and Gosselink 2007). Approximately 30% occur in tropical and 24% in subtropical regions, 12% in temperate areas, and 30% in boreal regions. These estimates do not include large lakes or deepwater coastal systems.

The classification and naming of wetland types is often confusing because names have evolved over centuries in different parts of the world and often reflect regional or continental differences. The wetlands described here reflect a North American perspective and are chosen to be representative of the variety of wetland types rather than totally inclusive of the range of wetland plant and animal communities that occur on this continent. We have grouped them ecologically, according to major sources of water (Fig. 1.2)—(1) precipitation, (2) groundwater, and (3) surface water—with the understanding that sources and amount of water may vary considerably within a wetland type and that most wetlands receive water from more than one source. A more detailed treatment of many North American wetland habitats is available in a companion volume by Batzer and Baldwin (2012).

FIGURE 1.2. The relative contribution of three water sources (precipitation, groundwater discharge, and surface inflow) to wetlands. As there may be much variation in the relative importance of different water sources within many of the wetland types, their locations are approximations and often overlap. Modified from Brinson (1993b).

Wetlands with Predominantly Precipitation Inputs

NORTHERN BOGS

Bogs are freshwater wetlands that occur on acid peat deposits throughout much of the boreal zone of the world (Fig. 1.3). They are frequently part of a larger complex of peatlands that includes fens, with readily apparent gradients of plant species distributions, biogeochemistry, and hydrology (Bridgham et al. 1996; Rochefort et al. 2012; Fens and Related Peatlands below). Bogs are distinguished from fens, however, because they receive water and nutrients exclusively from precipitation (ombrogeneous). Precipitation inputs are greater than evapotranspiration losses, and the slow decomposition of organic material under cold temperatures results in peat accumulation. Bog soils are organic, waterlogged, low in pH, and extremely low in available nutrients for plant growth (Chapter 2). In addition, growing seasons are short. Thus, a specialized and unique flora occurs in this wetland habitat.

FIGURE 1.3. Bog and lake with ericaceous shrubs and conifer trees in northern Michigan. Courtesy of K. E. Francl.

Mosses, primarily Sphagnum, are the most important peat-building plants in bogs. Bogs can be open Sphagnum moss peatlands, Sphagnum-sedge peatlands, Sphagnum-shrub peatlands, or bog forests; these bogs often form a mosaic across the landscape with other peatlands such as fens, which are more influenced by groundwater. Plants often associated with Sphagnum in bogs include various sedges (Carex spp.); cottongrass (Eriophorum vaginatum); and a variety of ericaceous shrubs such as heather (Calluna vulgaris), leatherleaf (Chamaedaphne calyculata), cranberry and blueberry (species of Vaccinium), and Laborador tea (Ledum groenlandicum). Trees such as spruce (Picea spp.) and tamarack (Larix larcina) may occur in bogs, often stunted in growth to reach only a meter or two tall. Many northern peatlands show a considerable overlap of species along the hydrologic and chemical gradients from nutrient-poor bogs to more mineral rich fens (see Fens and Related Peatlands below).

Vegetation development in many bogs may follow the model of hydrarch succession, or terrestrialization, at least to some degree (see Chapter 5). Through the buildup of soil organic matter, vegetation in these peatlands has significant control over habitat development (Moore and Bellamy 1974). Primary production exceeds decomposition of the peat substrates (Clymo et al. 1998), and the accumulation of peat affects hydrologic conditions, chemistry, and plant community composition (Damman 1986; Bridgham et al. 1996; Bauer et al. 2003). As peat builds up, it is often colonized first by shrubs and then by trees. Paleo-ecological evidence from several studies suggests a sequence of plant associations from wet marshes to a Sphagnum bog or wet forest (Walker 1970). Other processes, such as paludification (which occurs when bogs exceed the basin boundaries and encroach on formerly dry land) (e.g., Moore and Bellamy 1974; Bauer et al. 2003; Yu et al. 2003) and fires (Kuhry 1994), complicate the patterns of bog successional development.

Sphagnum has the ability to acidify its environment, probably through the production of organic acids located on its cell walls (Clymo and Hayward 1982). The acid environment retards bacterial action and reduces decomposition rates, enabling peat accumulation. Sphagnum also maintains waterlogging in the substrate. Its compact growth habit and overlapping, rolled leaves form a wick that draws up water by capillarity. Many bog plants are adapted to waterlogged anaerobic (low oxygen) environments by aerenchyma production, reduced oxygen consumption, and a leakage of oxygen from the roots to the rhizosphere. Many bog plants also have adaptations to the low available nutrient supply; these include evergreenness, sclerophylly (thickening of the plant epidermis to minimize grazing), the uptake of amino acids as a nitrogen source, and high root biomass (Bridgham et al. 1996). In addition, carnivorous plants such as pitcher plants (Sarracenia spp.) and sundews (Drosera spp.) have the ability to trap and digest insects. Some bog plants, such as the sweet gale (Myrica gale) and alders (Alnus spp.) also carry out symbiotic bacterial nitrogen fixation in nodules on their roots.

POCOSINS

Pocosins are evergreen shrub bogs (Fig. 1.4; Richardson 2012) restricted to the southeastern US Atlantic Coastal Plain, chiefly in North Carolina. They occur on waterlogged, acidic, nutrient-poor sandy or peaty soils (Bridgham and Richardson 1993; Richardson 2003) and are located primarily on flat topographic plateaus of the outer Coastal Plain. Their source of water is precipitation, and most of their water loss is through evapotranspiration during the summer and fall, although surface runoff also occurs, especially during winter and spring (Richardson 2012).

FIGURE 1.4. Short pocosin with ericaceous shrubs and pond pine (Pinus serotina) in eastern North Carolina. Courtesy of R. K. Peet.

Evergreen shrub and tree species dominate in pocosins, and the composition and stature of the vegetation is related to depth of the peat and nutrient availability. On deep peat accumulations (>1 m), roots do not penetrate into the underlying mineral soils, and ombrotrophic shrub bogs develop (Otte 1981). Ericaceous shrubs in these communities, called short pocosin, include titi (Cyrilla racemiflora), fetterbush (Lyonia lucida), and honeycup (Zenobia pulverulenta) as well as vines, particularly greenbriar (Smilax spp.). A sparse and often stunted canopy of pond pine (Pinus serotina) and loblolly bay (Gordonia lasianthus) may be present. Where organic substrates are shallower (approximately 50–100 cm), roots can penetrate into the underlying mineral soil and the vegetation grows somewhat taller. Additional species in these tall pocosins may include red maple (Acer rubrum), black gum (Nyssa sylvatica), and sweetbay (Magnolia virginiana). Fire during drought may also be an important factor in pocosin community development. Shallow peat burns allow the regeneration of pocosin species, although if the depth of the peat is reduced, roots of the recovering plants may be able to reach mineral soils. Severe burns that destroy peat substrates lead to development of a nonpocosin community, such as a marsh.

Pocosins temporarily hold water, especially during winter and early spring, and then slowly release it to adjacent wetlands. Since they occur in close proximity to estuaries, this slow release of freshwater may stabilize the salinity of regional estuaries (Daniel 1981). Draining and development activities may greatly change these hydrologic outputs. Richardson and McCarthy (1994) reported that peat mining resulted in an increased runoff of approximately 30%. Furthermore, drainage and agricultural conversion have increased the turbidity and levels of phosphate, nitrate, and ammonia in adjacent estuaries (Sharitz and Gresham 1998).

CAROLINA BAYS

Carolina bays are elliptical depressional wetlands (Fig. 1.5) that occur throughout the southeastern US Atlantic Coastal Plain from New Jersey to northern Florida (Kirkman et al. 2012). These wetlands range in size from greater than 3,600 hectares to less than 1 hectare. Early estimates of their number were as high as 500,000 (Prouty 1952), although it is more likely that only 10,000 to 20,000 remain (Richardson and Gibbons 1993). Carolina bays characteristically have no natural drainage, either into them or out of them, and overland water flows are minimal. Precipitation is their predominant source of water, and these shallow basins range from nearly permanently inundated to frequently dry, depending on their depth and local rainfall patterns. Most have highly variable hydroperiods (Sharitz 2003) and tend to be wetter in the winter and drier in the summer. Many small bays typically dry completely during most summers and refill during fall and winter rains. Soils in the basins of Carolina bays range from highly organic to predominantly mineral, and most are underlain with sand and impervious clay layers that retard vertical water movement. This sandy clay hardpan is often assumed to limit interactions between surface and groundwater and result in a perched water table in the basin, although a few studies have shown some connection with shallow groundwater (Lide et al. 1995; Chmielewski 1996; Pyzoha et al. 2008). The water in most bays is nutrient poor and acidic (with a pH range of 3.4 to 6.7) (Newman and Schalles 1990).

FIGURE 1.5. A 5-hectare Carolina bay showing characteristic elliptical shape. Aquatic herbaceous vegetation in the center is surrounded by bald cypress trees on the rim. Courtesy of L. S. Lee.

Plant communities in Carolina bays are influenced by soils and hydroperiod, and at least 11 vegetation types have been described (Schafale and Weakley 1990; Bennett and Nelson 1991). Pocosin communities, pond cypress (Taxodium ascendens) savannas, and pond cypress ponds are more common in bays on the lower Coastal Plain, which tend to have soils that are more organic. Herbaceous depression meadows are found on mineral soils in bays of the upper Coastal Plain, and forested habitats occur throughout. Various submersed and floating-leaved species such as bladderworts (Utricularia spp.), water lily (Nymphaea odorata), and water shield (Brasenia schreberi) are often common in bay ponds. Depression meadows are dominated by graminoids, including grasses such as Panicum, Leersia, and Dicanthelium; sedges such as Carex; and rushes including Juncus and Rhynchospora as well as a variety of other herbaceous plants. Forested bays may contain pond cypress as well as broadleaved trees such as swamp tupelo (Nyssa biflora), red maple, and sweetgum (Liquidambar styraciflua). Pocosin vegetation that occurs in bays is dominated by evergreen shrubs similar to that of the larger regional pocosins. Across this range of plant communities, Carolina bays have high plant species richness and contribute greatly to the regional biodiversity. The seed banks of some Carolina bays, especially depression meadows, are highly species rich (Kirkman and Sharitz 1994; Collins and Battaglia 2001; Mulhouse et al. 2005).

Rich zooplankton communities have been reported from Carolina bays (Mahoney et al. 1990), and a wide variety of aquatic and semiaquatic insects live in these habitats (Taylor et al. 1999). These seasonal wetlands are critical breeding habitat for numerous species of amphibians, some of which are entirely dependent on these ecosystems (Gibbons and Semlitsch 1991) and may be found in huge numbers during the breeding season (Pechmann et al. 1991; Gibbons et al. 2006).

Many Carolina bays, especially the smaller ones, have been drained and converted to agriculture or other uses, and the great majority of those remaining have drainage ditches (Bennett and Nelson 1991). Since 2001, one of their most serious threats has come from US Supreme Court decisions that held that isolated non-navigable waters are not necessarily protected under the Clean Water Act (Downing et al. 2003; Sharitz 2003; see Chapter 8).

CYPRESS DOMES

Cypress swamps found in nearly circular isolated depressions throughout the karst landscape of Florida are called cypress domes because of the dome-like appearance of the tree canopy (Fig. 1.6). Trees are usually taller and grow faster in the centers of the depressions than at the edges (Ewel and Wickenheiser 1988). Trees on the perimeters of domes are also more susceptible to fire mortality (Watts et al. 2012). These depressions are formed by the dissolution of underlying limestone, and most are less than 10 hectares in size (Ewel 1998). Most of the water is received from precipitation, although surface inflows may also occur. Water may also move from these depression ponds into shallow groundwater (Heimburg 1984). Pond cypress and swamp tupelo are the dominant species, with slash pine (Pinus elliottii) co-dominant in partly drained cypress domes (Mitsch and Ewel 1979).

FIGURE 1.6. Cypress dome wetland with pond cypress ( Taxodium ascendens ) trees in Florida. Note that the tallest trees are in the center, which is usually the deepest part of the wetland.

These wetlands play a major role in maintaining the region’s biodiversity. Many are significant amphibian breeding grounds. In addition, since they hold water for long periods, cypress domes help prevent flooding of local areas and aid in groundwater discharge. Nearly all cypress domes in northern Florida have been harvested, although in many the trees have regenerated. The most detrimental human impact is caused by development and conversion to residential and commercial sites. Drainage of cypress domes also causes oxidation of the organic soils, land subsidence, and an increase in fire susceptibility (Ewel 1998).

PRAIRIE POTHOLES

One of the most important areas of freshwater wetlands in the world is the prairie pothole complex of North America (Galatowitsch 2012). These shallow depressional wetlands (Fig. 1.7) are found in Minnesota, Iowa, and the Dakotas in the United States, and in Alberta, Saskatchewan, and Manitoba in Canada. Although individual pothole marshes are usually small, they are regionally abundant. Between 4 and 10 million potholes are estimated to occur in Canada (Adams 1988), and about 2.3 million existed in the 1960s in North and South Dakota (Kantrud et al. 1989), although many had been drained even by that time. Using assumptions from these estimates regarding abundance and size, van der Valk and Pederson (2003) suggested that these wetlands covered approximately 63,000 square kilometers prior to drainage.

FIGURE 1.7. Prairie pothole marsh wetland in the upper Midwest. From Tiner et al. (2002).

Precipitation is the primary source of water for prairie potholes. Annual precipitation can vary significantly from year to year, with periods of severe drought alternating with periods of above-normal precipitation. Because of the small size of their catchments, changes in annual precipitation can result in major changes in these potholes’ annual water levels. Almost all prairie potholes also have some connection to groundwater (Winter 1989). They can be groundwater recharge sites, groundwater discharge sites, or groundwater flow-through wetlands. Because of these groundwater connections, prairie potholes are interlinked wetland complexes. In addition, during wet years when the catchments fill, water may overflow on the surface from one basin to another. Such ephemeral surface-water connections may provide opportunity for seed dispersal or movement of aquatic animals among prairie potholes.

Herbaceous marshes containing robust perennial plants along with submersed species characterize prairie potholes throughout most of their range. Often, stands of emergent vegetation dominated primarily by one species such as cattail (Typha spp.) will have high primary productivity (van der Valk and Davis 1978b). Periodic drawing down and refilling of these shallow basins results in changes in vegetation types and species that can be predicted from a knowledge of the seed bank, the potential dispersal of plant propagules, and the conditions under which different species will germinate and become established (van der Valk 1981). The vegetation cycle results in four distinct stages: a dry marsh stage, a regenerating marsh stage, a degenerating marsh stage, and a lake stage (van der Valk and Davis 1978b). During droughts when the substrate is exposed, perennial emergent species (e.g., cattails, perennial sedges Scirpus spp., bur-reed Sparganium eurycarpum) and annual mud flat species (e.g., smartweed Polygonum spp., annual sedges Cyperus spp., beggarticks Bidens cernua) become established from the seed bank or from propagules dispersed into the basins. This dry marsh stage is followed by a wet marsh community when rainfall returns to normal and basins refill. The annual species that required exposed substrate for germination disappear, leaving the perennial emergents. Submersed species that can germinate under water (e.g., pondweed Potamogeton spp., water nymph Najas flexilis, water milfoil Myriophyllum sibiricum) also appear. Wet marsh may persist for several years, but eventually the emergent vegetation begins to decline, perhaps because of the failure of some emergent species to continue to reproduce vegetatively (van der Valk and Davis 1978a), or destruction by muskrats. This degenerating marsh may become a pond or shallow lake in which the dominant vegetation is comprised primarily of freefloating and submersed plants. When drought once again exposes the marsh bottom, the cycle repeats (see Chapter 6 for animal responses to this cycle).

Prairie potholes are especially important ecologically and economically because they are the major waterfowl breeding area in North America. An estimated 50% to 80% of North America’s game waterfowl species are produced in this region (Batt et al. 1989). Successful breeding requires availability of a variety of wetlands and wetland plants (for food and habitat) because no single wetland basin provides for all their reproductive needs throughout the breeding season (Swanson and Duebbert 1989). The existence of large numbers of small wetlands allows the birds to disperse across the landscape, thereby lowering their vulnerability to predation and diseases and increasing the likelihood of successful reproduction and brood rearing (Kantrud et al. 1989).

About half the original prairie potholes in the Dakotas have been destroyed, mostly by agriculture, and more than 99% of Iowa’s original marshes have been lost (Tiner 1984, 2003). Drainage of potholes significantly reduces their water-storage capacity, and destruction of natural vegetation buffers around remaining wetlands has significantly reduced valuable waterfowl nesting and rearing areas (Tiner 2003). Like all depressional wetlands, prairie pothole protection through federal regulations has been weakened as a result of recent Supreme Court decisions (see Chapter 8).

PLAYA AND RAINWATER BASIN/SANDHILLS WETLANDS

Playas are shallow recharge wetlands found in semiarid prairie areas of the southern Great Plains (Smith et al. 2012). They range in area from less than 1 hectare to greater than 250 hectares and average 6.3 hectares (Guthery and Bryant 1982). There are probably more than 30,000 of these small circular depressions, which are thought to result from a combination of dissolution of subsurface materials and wind action (Haukos and Smith 2003). Playas receive most of their water from rainfall and local runoff (including irrigation water), and it is rare for them to be connected to groundwater sources (Haukos and Smith 1994). These wetlands are usually dry in late winter, early spring, and late summer; multiple wet-dry cycles during a single growing season are common.

Playas are considered to be keystone ecosystems serving as biological refugia and critical sites of biodiversity in this semiarid and intensive agricultural region (Smith and Haukos 2002). More than 340 plant species have been recorded in playas (Haukos and Smith 1997), although most of these species are also commonly found in other wetland and terrestrial habitats. Because of their rapidly changing environmental conditions, the flora is dominated by annuals and short-lived perennials. In a survey of 224 playa wetlands, Smith and Haukos (2002) found that only 38% of plant species present in the early growing season were still present late in the season. Thus, the vegetation is influenced by the composition of the seed bank and the environmental conditions that regulate germination and seedling growth. Because the land surrounding most playas is cultivated, annual and exotic plant species are common invaders (Smith and Haukos 2002). Playas also support a broad array of bird species and are vital overwintering, migration, and breeding habitats for waterfowl in the region (Haukos and Smith 2003). They are recognized as important for invertebrates and amphibians as well.

All flora and fauna occupying playas must be adapted to the fluctuating environmental conditions, and any alteration of the hydroperiod may have drastic effects on species persistence. Unfortunately, many playas have been affected by sedimentation as a result of cultivation and erosion of the surrounding landscape. This has caused a dramatic decrease in playa hydroperiod and altered floral and faunal communities (Haukos and Smith 1994).

In Nebraska, aeolian forces from winds have created depressional wetlands in the Rainwater Basin in the south-central part of the state and in the Sandhills region of the northern and central areas. The Rainwater Basin wetlands depend on precipitation and overland runoff for their water supply (Frankforter 1996), and most are marshes, wet meadows, or ponds (Tiner 2003). Lakes and marshes in the Sandhills region are interconnected with the regional groundwater (LaBaugh 1986b; Winter 1986). Both Rainwater Basin and Sandhills wetlands have been identified as wetlands of international importance to waterfowl and other wildlife. Millions of waterfowl in the Central Flyway use these wetlands during spring migration (Gersib 1991); an abundance of aquatic invertebrates and fish provide a food source for these migratory birds. Agricultural activities, such as drainage and groundwater pumping, have been major causes of loss or degradation of these wetlands. At least 66% of the original area of Rainwater Basin wetlands has been lost (LaGrange 2001), as have more than 30% of the original Sandhills wetlands (Erickson and Leslie 1987).

VERNAL/SEASONAL POOLS

Vernal pools, broadly defined as ephemeral depressional wetlands that flood during the spring months of the year but that dry during the summer, are distributed throughout the world (Zedler 2003). These wetlands are largely collectors of rainfall and snowmelt water, although groundwater inputs may occur in some. Depending on climate, geology, hydrology, and other factors, vernal pools may be dominated by trees and shrubs, by marsh and wet meadow species, or by aquatic plants, or they may be devoid of vegetation (Tiner 2003). In the United States, true vernal pools are particularly abundant on the Pacific Coast. In the glaciated landscapes of the North and Northeast, seasonal woodland pools are very abundant and are often called vernal pools, but in fact they can be flooded in seasons other than spring, depending on precipitation patterns for a given year (Calhoun et al. 2012). Both types—west coast vernal pools and eastern woodland vernal pools—have received considerable attention because of their importance as habitats for rare plants and amphibians.

West coast vernal pools fill from winter rains characteristic of the region’s Mediterranean climate, and then dry to extreme desiccating soil conditions during the dry summers (Zedler 2003). The isolated nature and unpredictable flooding of these wetlands promote endemism, thereby creating unique flora and fauna and making these vernal pools vital sites for the conservation of biodiversity (Tiner 2003). The flora of vernal pools in California contains numerous federally listed threatened and endangered species as well as state-listed endangered and rare species. In the past, west coast vernal pools were used for grazing and other forms of agriculture. More recently, population growth and corresponding urbanization in California have greatly reduced the extent of these ecosystems, and the largest remaining complexes are found in the open lands of military facilities.

Seasonal woodland pools occur throughout forested regions of the eastern United States and southeastern Canada (Calhoun et al. 2012). These habitats are typically inundated during the spring and early summer and then dry out in late summer or autumn (but, as mentioned, can be flooded longer in wet years). The flora may consist primarily of the surrounding forest trees and shrub species as well as various grasses, depending in part on hydroperiod and canopy openness; vegetation characteristics are highly variable (Palik et al. 2001). Because predatory fish are not present, these seasonal wetlands can be extremely productive sites for macroinvertebrate and amphibian reproduction, and several salamander species are entirely dependent on seasonal woodland pools for breeding (Gibbs 1993; Semlitsch and Bodie 1998; Kenney and Burne 2000). While the seasonal pool breeders require such habitats for reproduction and growth of larvae, adult salamanders and frogs spend their lives in the surrounding woodland. Thus the protection of seasonal pools and the surrounding forest is important for the conservation of biodiversity (Semlitsch and Bodie 1998; Kenney and Burne 2000; Calhoun et al. 2012). Unfortunately, since these pools are usually very small, they are often destroyed by development activities.

THE EVERGLADES

The Everglades (Fig. 1.8) is perhaps one of the most well recognized wetlands in the world, its notoriety derived from the wealth of its biotic heritage as well as the magnitude of factors that threaten its resources (Gunderson and Loftus 1993; Gaiser et al. 2012). Occurring in the subtropical southern part of the Florida peninsula, the Everglades historically covered a vast area of about 1.2 million hectares; about half has now been drained for agriculture and development (Davis et al. 1994). The bedrock substrate underlying most of the Everglades is limestone, of marine and freshwater origin. The soil substrate is predominantly peat, formed during the last 5,000 years (Gleason and Stone 1994) and often interspersed with light-colored calcitic soils called marl. It is the largest and most important freshwater subtropical peatland in North America (Koch and Reddy 1992; Gaiser et al. 2012).

FIGURE 1.8. Everglades freshwater marsh and tree island complex in southern Florida. Courtesy of South Florida Water Management District.

Precipitation is the main route by which water enters the Everglades ecosystem (Duever et al. 1994). Thus waters of the historic Everglades were probably very low in dissolved nitrogen and phosphorus (oligotrophic), but relatively high in calcium and bicarbonate (Flora and Rosendahl 1982). Approximately 60% of the rain falls between June and September, produced primarily by localized thunderstorms and, at times, tropical cyclones. The hydroperiod is quite variable, with water levels declining slowly during the winter and droughts common during the dry spring months when evapotranspiration is high. Lake Okeechobee, to the north, is linked hydrologically with the Everglades by groundwater connections and, during high water periods, by overland flow. The topography of this region is very flat, and water moves southward through the Everglades marshes at velocities ranging from approximately 0 to 1 cm/sec (Rosendahl and Rose 1982). This slow southerly flow of water inspired Marjorie Stoneman Douglas (1947) to call the Everglades a River of Grass.

Major freshwater wetland plant communities include marshes and wet prairies, forested communities (tree islands), and ponds and sloughs with little emergent vegetation (Gunderson 1994) (Fig. 1.8). These communities are arrayed in a mosaic influenced by hydrologic and substrate conditions. Sawgrass (Cladium jamaicense [C. mariscus ssp. jamaicense]) is the most common and widespread species in the Everglades marshes. A robust, rhizomatous, perennial sedge rather than a grass, as its name implies, sawgrass is well adapted to the low nutrient conditions as well as to flooding and also burning during droughts. Throughout much of the Everglades, sawgrass marshes are interspersed with shallow sloughs that contain spikerush (Eleocharis spp.) and floating-leaved aquatic plants such as water lily, yellow pond-lily (Nuphar lutea), and submerged aquatics including bladderworts. The submerged portions of most aquatic macrophytes in the Everglades are covered with periphyton (a community of many species of microalgae, including calcareous species), which serves as a food web base, as well as oxygenating the water column and building calcite mud sediment (Browder et al. 1994; Gaiser et al. 2005a, 2005b, 2012).

Wet prairies of graminoid species develop on peat or marl (limestone) substrates, and each soil type has distinct plant communities. Those on peat commonly are dominated by species of spikerush, beakrush, or maidencane (Panicum hemitomon). Prairies on marl substrates are dominated by sawgrass and muhly grass (Muhlenbergia spp.). Tree islands are clumps of bayhead/swamp forest taller than the surrounding marsh. Canopy species include redbay (Persea borbonia) and sweetbay as well as dahoon holly (Ilex cassine) and pond apple (Annona glabra), with a dense layer of shrubs underneath. Found throughout many parts of the Everglades marshes, these tree islands are often in the shape of an elongated teardrop with the long axis parallel to the main direction of flow (Gaiser et al. 2012).

Major parts of the Everglades wetlands now have been drained for agriculture and urbanization, and the water flows in the remaining areas are fragmented by canals, large water conservation areas, and roads and drainage ditches and pipes. Today, only 0.62 million hectares of the original Everglades remain (Davis et al. 1994). Increased phosphorus loading from agricultural runoff in the northern parts of the remaining Everglades may be promoting an increase in southern cattail (Typha domingensis) and declines in sawgrass and periphyton, which prosper in nutrient-poor waters (Newman et al. 1996; Noe et al. 2001; Childers et al. 2003; Gaiser et al. 2005a). Many exotic invasive plant species also threaten the plant communities of the Everglades. Among the most aggressive and difficult to control are melaleuca (Melaleuca quinquenervia), a pioneering Australian tree species; Brazilian peppertree (Schinus terebinthifolius); old world climbing fern (Lygodium microphyllum), which overtops and smothers Everglades tree islands; and torpedo grass (Panicum repens), which has replaced large areas of marsh plants in Lake Okeechobee (White 1994). Beginning in the late 1990s, a major restoration of the Everglades was initiated by the USAC E and the state of Florida to try to restore some of the hydrologic integrity of the Everglades (Comprehensive Everglades Restoration Plan 2000; Gaiser et al. 2012).

Wetlands with Predominately Groundwater Inputs

SPRING-FED WETLANDS

Groundwater can emerge to the surface in isolated locations called springs. On flat areas or at the base of slopes, small wetlands can develop around these points of discharge (Fig. 1.9). Most water discharge is cold, but if the Earth’s magma is in close proximity to the groundwater aquifer, hot springs (>50°C) can develop. While spring-fed wetlands occur in many different landscapes, they are particularly important ecologically in arid and semi-arid areas, such as the Great Basin of the western United States and Canada (Keleher and Sada 2012); they are one of the few wetland habitats occurring in such areas, and the lack of other water sources in these areas make these spring-fed wetlands true oases for many animals from the surrounding landscape. Spring-fed wetlands are also common in karstic (limestone) landscapes such as Florida.

FIGURE 1.9. A cold spring-fed wetland in Utah. Courtesy of M. J. Keleher.

The near-constant supply of water for spring-fed wetlands and the thermal and chemical characteristics of the water make the flora and fauna of spring-fed wetlands unique. In hot springs, unique species of bacteria, algae, plants, and animals tolerant of high temperatures occur. In cold springs, the water becomes flushed with oxygen after it emerges and the water consistently flows, and thus numerous fishes can inhabit these wetlands (Keleher and Sada 2012), and invertebrates more typically occurring in cool streams (e.g., Plecoptera stoneflies) and not in traditional wetlands may thrive (Batzer and Ruhí 2013). The emerging water contains high levels of dissolved minerals, such as calcium and phosphorus, and thus spring-fed wetlands may support a diversity of emergent (e.g., Ranunculus) and submersed plants (e.g., Potamogeton), as well as copious growth of filamentous algae (e.g., Chara) (Fig. 1.9) characteristic of nutrient-rich waters. Because of ample calcium, various snails, which need calcium for shell development, can reach very high densities in spring-fed wetlands. Spring-fed wetlands are typically geographically isolated from other types of wetland and from each other, and thus the flora and fauna of the habitats show a high degree of endemism (i.e., species that only occur in a restricted area).

Because endemic species are so prevalent, an inordinate number of threatened or endangered species occur in spring-fed wetlands. For example, several species of fish and snails in the Great Basin are associated with a small number (sometimes only one) of spring-fed wetlands (Keleher and Sada 2012). In Florida, endangered manatees (Trichechus manatus) often migrate to spring-fed wetlands in winter as a thermal refuge (Laist et al. 2013). Because of their reliance on groundwater, one of the biggest human threats to spring-fed wetlands is excessive groundwater extraction that could decrease or even eliminate natural discharge into these wetlands.

FENS AND RELATED PEATLANDS

Peatlands are wetland ecosystems that accumulate carbon because primary plant productivity exceeds decomposition and dead organic material builds up as peat. Most of the global peatland area is found in boreal and subarctic zones of the Northern Hemisphere (Gorham 1991; Rochefort et al. 2012), although peatlands also exist in southern locations (pocosins, southern Appalachian fens). The terminology applied to peatlands is often confusing. Throughout the world, they have been known by many different names, such as bog, fen, moor, mire, marsh, swamp, and heath (Bedford and Godwin 2003).

Peatlands are generally classified into ombrogenous (rain fed) and geogenous (receiving water from the regional water table or other outside sources). Ombrogenous peatlands vegetated largely with Sphagnum mosses and geogenous peatlands vegetated mostly with graminoid species (such as grasses, sedges, and rushes) are commonly called bogs and fens, respectively. Geogenous fens may be further divided into (1) limnogeneous peatlands, which develop along lakes and slow-flowing streams; (2) topogeneous peatlands, which develop in topographic depressions, with a portion of their water derived from the regional ground water table; and (3) soligeneous peatlands, which are affected by water from outside sources percolating through or over surface peat (Bridgham et al. 1996). There is strong overlap among all these peatland categories in environmental conditions, such as soil or water pH, and in plant species. Thus, Bridgham et al. (1996) recommend that the term peatland be used for all these systems to reduce confusion. They suggested that the terms bog and fen be used only in a broad sense, with bogs referring to acidic low alkalinity peatlands that are typically dominated by Sphagnum mosses, various species of ericaceous shrubs, and/or conifers such as spruces or pines. Similarly, fens should refer broadly to somewhat less acidic, more alkaline peatlands dominated by graminoid species, brown mosses, taller shrubs, and coniferous and/or deciduous trees.

The defining characteristic of all types of fens is the importance of groundwater inputs in determining their hydrology, chemistry, and vegetation (Bedford and Godwin 2003). Thus, fens occur where climate and the hydrologic and geologic setting sustains flows of mineral-rich groundwater to the plant rooting zone. They may develop on slopes, in depressions, or on flats. The relatively constant supply of groundwater maintains saturated conditions most of the time, and the water chemistry reflects the mineralogy of the surrounding and underlying substrates. Fens may be slightly acidic (poor fens), circumneutral (rich fens), or strongly alkaline (extremely rich fens). In an extensive survey of North American fens, Bedford and Godwin (2003) reported pH ranging from 3.5 to 8.4.

Peatland development is a dynamic process (e.g., Heinselman 1970; Kuhry 1994; Wheeler and Proctor 2000; Yu et al. 2001, 2003; Bauer et al. 2003; Rochefort et al. 2012) that is controlled by multiple biotic and abiotic factors. Climate, physiography, and peat accumulation all play a role. Terrestrialization (filling in of shallow lakes) and paludification (encroachment of bogs over formerly dry land) are the major processes in peatland development.

The vegetation of fens (Fig. 1.10) is generally dominated by bryophytes, sedges, grasses, dicotyledonous herbs, and coniferous trees. Vegetation of poor fens resembles that of bogs, with Sphagnum mosses and ericaceous shrubs. Rich fens, however, are dominated by sedges and brown mosses (mostly of the Amblestegiaceae family), with many distinctive species of dicotyledonous herbs (Bedford and Godwin 2003). Tussock cottongrass, sedges (genera of the Cyperaceae family, especially Carex) and shrubs such as heather, leatherleaf, Laborador tea, and cranberry and blueberry may occur. In forested peatlands, trees such as spruce and tamarack may be found, often in stunted condition. Individually and collectively, fens are among the most floristically diverse of all wetland types, supporting rare and uncommon bryophytes and vascular plant species (Bedford and Godwin 2003).

FIGURE 1.10. Fen peatland in Canada. Courtesy of K. E. Francl.

In a description of the Lake Agassiz peatlands of northern Minnesota, Heinselman (1970) described seven vegetation associations: (1) a rich swamp forest dominated by northern red cedar (Thuja occidentalis) with a shrub layer of alder (Alnus incana) and hummocks of Sphagnum, (2) a poor swamp forest dominated by tamarack with an understory of bog birch (Betula pumila) and Sphagnum hummocks, (3) a cedar string bog-and-fen complex with ridges of northern red cedar and treeless hollows of sedges (mostly Carex spp.) between them, (4) a larch string bog-and-fen complex in which tamarack dominated the ridges, (5) a black spruce (Picea marina) forest with a carpet of feathermoss (Pleurozium) and other mosses, (6) a Sphagnum-black spruce-leatherleaf bog forest of stunted black spruce and a heavy evergreen shrub layer over Sphagnum mosses, and (7) a Sphagnum-leatherleaf-laurel-spruce heath in which a low shrub layer including laurel (Kalmia spp.) and stunted spruce grow over a continuous blanket of Sphagnum mosses.

Peatlands are important wetland ecosystems for several reasons, including their vast extent across northern boreal regions. They are one of the largest terrestrial carbon reservoirs. Northern peatlands have accumulated about 400 to 500 Gt (1 Gt = 10¹⁵ g) of carbon during the Holocene (Gorham 1991; Clymo et al. 1998; Roulet 2000; Vitt et al. 2000; see Chapter 7). Their extent, high-latitude location, and the large size of their carbon pool raise concerns that they may become significant sources for atmospheric carbon under a changing climate (Moore et al. 1998; Schindler 1998; see Chapter 10).

ATLANTIC WHITE CEDAR SWAMPS

These forests, dominated by Chamaecyparis thyoides, occur within a wide climatic range along the Atlantic and Gulf of Mexico coastline areas of the United States from Maine to Mississippi. Throughout their range, however, they are uncommon, having decreased historically in area and in biological diversity (Laderman 1989). They are most abundant in the southeastern New Jersey Pine Barrens, in the Dismal Swamp of Virginia and North Carolina, and along several river systems in northwest Florida (Sheffield et al. 1998).

Atlantic white cedar swamps may occur in isolated basins, along lake shorelines, shoreward of coastal tidal marshes, on river floodplains, or on slopes (Laderman 1989). Hydrologic conditions can vary considerably among these forests, although flooding typically occurs in late winter and early spring, sometimes for extended periods. White cedar swamps occurring in basins, with precipitation as the major source of water, are usually oligotrophic. Those in other locations, however, may receive significant groundwater and are more nutrient rich (Laderman 1989). Most occur on peat soils of relatively low pH (2.5–6.7) (Day 1984; Whigham and Richardson 1988; Laderman 1989; Ehrenfeld and Schneider 1991). Under these conditions, C. thyoides may grow in dense, almost monospecific stands. Other canopy species commonly associated with Atlantic white cedar are red maple, black gum, and sweetbay, and pines such as loblolly (Pinus taeda), white (P. strobus), or pitch (P. rigida).

Relatively open conditions are necessary for the healthy growth of Chamaecyparis seedlings (Laderman 1989), and several studies have indicated poor seedling recruitment under the closed canopy in dense stands (Motzkin et al. 1993; Stoltzfus and Good 1998). This suggests that natural biotic processes in which Chamaecyparis seedlings are replaced by more shade-tolerant species such as red maple may play a role in the decline of these forests. From age structure analyses and paleo-ecological investigations of an old-growth Atlantic white cedar swamp, Motzkin et al. (1993) determined that extensive Chamaecyparis establishment occurred during distinct episodes following disturbance events. Prior to European settlement, fires frequently destroyed existing Atlantic white cedar stands but allowed for subsequent regeneration from seed stored in the upper soil horizon or from surviving trees. Thus abiotic factors associated with disturbances have also been important in influencing the development of Atlantic white cedar swamps (Motzkin et al. 1993).

Much of the historic decrease in C. thyoides stands is attributable to selective logging for their valuable lumber, however, and to conversion to agricultural, industrial, or commercial uses (Motzkin et al. 1993). Up to 50% of the Atlantic white cedar area of North Carolina was cut between 1870 and 1890, for example (Frost 1987). Herbivory of seedlings and saplings by deer often has an impact on stand regeneration, and deer browse can destroy young stands (Laderman 1989). Swamps that have frequent, lowlevel disturbances such as suburban runoff and the presence of roads generally have few Chamaecyparis seedlings and lack soil conditions conducive to their growth (Ehrenfeld and Schneider 1991). It is likely, however, that adjacent land uses are more important than regional land-use patterns in affecting the runoff of sediments and contaminants into these swamps (Laidig and Zampella 1999).

Wetlands with Predominately Surface Water Inputs

SOUTHERN DEEPWATER SWAMPS

Deepwater swamps, primarily baldcypress-water tupelo (Taxodium distichum–Nyssa aquatica) forests (Fig. 1.11), are freshwater ecosystems that have standing water for most or all of the year (Penfound 1952). They are generally found along rivers and streams of the Atlantic Coastal Plain from Delaware to Florida, along the Gulf Coastal Plain to southeastern Texas, and up the Mississippi River to southern Illinois (Conner and Buford 1998). Other southern deepwater swamps include cypress domes (see the Cypress Domes section). On river floodplains, these baldcypress-water tupelo swamps are found in meander scrolls created as the rivers change course (ridge and swale topography), oxbow lakes created as meanders become separated from the main river channel, and sloughs that are areas of ponded water in meanders and backwater swamps (Brinson et al. 1981).

FIGURE 1.11. Southern deepwater swamp with baldcypress (Taxodium distichum) trees in Congaree National Park, South Carolina. Courtesy of R. R . Sharitz.

The major hydrologic inputs to these deepwater swamps are overflow from flooding rivers and runoff from surrounding uplands. In addition, larger and deeper topographic features may impound water from rainfall and may be connected with the regional groundwater. Even though deepwater swamps are usually flooded, water levels may vary seasonally and annually. High water levels typically coincide with winter-spring rains and melting snow runoff. Low levels occur in the summer from high evapotranspiration and low rainfall (Wharton and Brinson 1979). During extreme droughts, even deepwater swamp forests may lack surface water for extended periods (Mancil 1969). Some peat development is characteristic of these deepwater ecosystems because of slow decomposition rates (Conner and Buford 1998). Since many deepwater swamps are found along the floodplains of rivers, their soils generally have adequate nutrients, and these forests are relatively productive. However, anaerobic conditions associated with continuous flooding may limit the availability of several nutrients to plants and reduce their productivity (Wharton et al. 1982; Megonigal et al. 1997).

Southern deepwater swamps have unique plant communities that either depend on or adapt