Coastal Wetlands: An Integrated Ecosystem Approach

Table of Contents

Cover Image

Copyright Page

Preface

List of Contributors

List of Reviewers

Chapter 1. Coastal Wetlands

1. Introduction

2. A Synthesis of Coastal Wetlands Science

3. Lessons from the Chapters in This Book

Chapter 2. The Morphology and Development of Tropical Coastal Wetlands

1. Introduction

2. Mangrove and Associated Wetlands

3. Sedimentation and the Development of Wetlands

4. Sea-Level Controls on Wetland Development

5. Sea-Level Change and the Diversification of West Indian Mangroves

6. Sea-Level Change and the Evolution of Mangrove Habitats in the IWP

7. Impact of Future Climate and Sea-Level Change

8. Summary and Concluding Remarks

Chapter 3. Temperate Coastal Wetlands

1. Introduction

2. Factors Controlling Sediment Dynamics

3. Factors Controlling Patterns of Vegetation

4. Geographic Variation

5. Human Impact and Climate Change

6. Summary

Chapter 4. Polar Coastal Wetlands

1. Introduction

2. Geology/Geomorphology

3. Oceanography

4. Climate

5. Structure of Coastal Wetlands

6. Vegetation of Polar Coastal Wetlands

7. Fauna of Polar Coastal Wetlands

8. Environmental Hazards

9. Conclusions and Research Priorities

Chapter 5. Intertidal Eco-Geomorphological Dynamics and Hydrodynamic Circulation

Acknowledgments

1. Introduction

2. Intertidal Eco-Geomorphological Evolution

3. Results

4. Discussion

5. Conclusions

Chapter 6. Tidal Courses

Acknowledgments

1. Introduction

2. Proposed Tidal Course Classification

3. Geomorphology of Tidal Courses

4. Course Networks and Drainage Systems

5. Origin of Tidal Courses

6. Course Evolution

7. Summary

Chapter 7. Heat Energy Balance in Coastal Wetlands

Acknowledgments

1. Introduction

2. Mid-Latitudes

3. Low Latitudes

4. High Latitudes

5. Summary

Chapter 8. Hydrodynamics and Modeling of Water Flow in Mangrove Areas

Acknowledgments

1. Introduction

2. Physical Characteristics of Mangrove Topography and Vegetation

3. Peculiar Hydrodynamics in Mangrove Areas

4. Modeling

5. Summary

Chapter 9. Mathematical Modeling of Tidal Flow over Salt Marshes and Tidal Flats with Applications to the Venice Lagoon

Acknowledgments

1. Introduction

2. Wetting and Drying, and the Dynamics of Very Shallow Flows

3. Wind and Wind Waves

4. Salt Marsh Vegetation

5. Salt Marshes and Tidal Flats Morphodynamics

6. Conclusions

Chapter 10. Geomorphology and Sedimentology of Tidal Flats

Acknowledgments

1. Introduction

2. Basic Conditions for the Formation of Tidal Flats

3. Zonation in Sedimentation and Flat Surface Morphology

4. Factors and Processes

5. Summary

Chapter 11. Intertidal Flats

Acknowledgments

1. Introduction

2. The Depositional Habitat

3. The Functional Role of Biota

4. Future Shock: Climate Change and Ecosystem Function

Chapter 12. Biogeochemical Dynamics of Coastal Tidal Flats

Acknowledgments

1. Introduction

2. Transport Processes on Intertidal Flats

3. Microbial Processes

4. Nitrogen Cycle

5. Phosphorus Cycle

6. Silicon Cycle

7. Concluding Remarks

Chapter 13. Productivity and Biogeochemical Cycling in Seagrass Ecosystems

Acknowledgments

1. Introduction

2. Sediment Biogeochemistry – Modified by Seagrasses

3. Human Pressures and Effects on Biogeochemistry

4. Future Perspectives and Conclusions

Chapter 14. Tidal Salt Marshes

1. Introduction

2. Geographical Distribution

3. Why Salt Marshes Exist?

4. Geomorphology

5. Morphodynamics

6. Sedimentology

7. Concluding Discussion

Chapter 15. Ecosystem Structure of Tidal Saline Marshes

Acknowledgments

1. Introduction

2. Saline Marsh Communities

3. Interaction Among Communities

Chapter 16. Salt Marsh Biogeochemistry – An Overview

Acknowledgments

1. Introduction

2. Carbon

3. Nitrogen

4. Iron and Sulfur

5. Phosphorus

6. Marshes in Transition and Directions for Future Work

Chapter 17. The Role of Freshwater Flows on Salt Marsh Growth and Development

1. Introduction

2. Freshwater Routes in Salt Marshes

3. Associated Processes

4. Hydrological Impacts in Salt Marshes

5. Techniques for the Study of Marsh Hydrology

6. Implications of Freshwater Flows for Salt Marsh Management

7. Implications of Freshwater Flows for Salt Marsh Creation

8. The Ecohydrological Approach in Salt Marsh Studies

9. The Way Ahead – Problems and Challenges

Chapter 18. Tidal Freshwater Wetlands

1. Introduction

2. Hydrogeomorphic Setting

3. Biodiversity

4. Primary Production and Nutrient Cycling

5. Threats and Future Prospectus

Chapter 19. Biogeochemistry of Tidal Freshwater Wetlands

Acknowledgments

1. Introduction

2. Carbon Biogeochemistry

3. Processes Governing Organic Carbon Metabolism

4. Nitrogen Biogeochemistry

5. Phosphorus Biogeochemistry

6. Silicon Biogeochemistry

7. Biogeochemical Effects of Sea-Level Rise

8. Concluding Comments

Chapter 20. Geomorphology and Sedimentology of Mangroves

Acknowledgments

1. Introduction

2. Mangrove Environmental Settings

3. Tidal Range and Sea-Level Control

4. Sedimentation in Mangroves

5. Mangroves as Sea-Level Indicators

6. Storms/Tsunamis

7. Inundation Changes Affecting Mangroves

8. Conclusions

Chapter 21. Geomorphology and Sedimentology of Mangroves and Salt Marshes

1. Driving Forces Determining Main Morphology and Vegetation Types in the Coastal Zone

2. Depositional Environment and Substrate Formation for the Development of Mangroves and Salt Marshes

3. Influence of Sea Level and Climate Oscillations on Local Geobotanical Features

4. Major Factors Leading to the Development of Salt Marshes and Mangroves at the Amazon Coastal Region: An Integrated Analysis

5. Influence of Geomorphology and Inundation Regime of Geobotanical Units on their Sediment Biogeochemistry

Chapter 22. Paradigm Shifts in Mangrove Biology

1. Introduction

2. Shifts in Established Paradigms

3. Conclusions

Chapter 23. Ecogeomorphic Models of Nutrient Biogeochemistry for Mangrove Wetlands

Acknowledgments

1. Introduction

2. Ecogeomorphology of Mangroves (Model 1)

3. A Multigradient Model (Model 2)

4. Geochemical Model (Model 3)

5. Soil Biogeochemistry Model (NUMAN, Model 4)

6. Mass Balance Exchange (Model 5)

7. Contrasting Coastal Settings and Biogeochemical Models

Chapter 24. Seagrass Restoration

Acknowledgment

1. Introduction

2. Regional Activities

3. Policy Issues Relevant to Mitigation

4. Conclusions

Chapter 25. Tidal Marsh Creation

1. Introduction

2. Principles and Techniques of Tidal Marsh Creation

3. Evaluating Functional Equivalence of Created Tidal Marshes

4. Summary

Chapter 26. Salt Marsh Restoration

1. Introduction

2. Setting Objectives

3. Planning for the Future

4. Addressing Causes and not Symptoms

5. Managing Disturbance

6. Conflicting Priorities

7. Discussion

8. Conclusions

Chapter 27. Managed Realignment

Acknowledgments

1. Introduction

2. Location, Drivers, and Constraints to Managed Realignment

3. Site Evolution

4. Challenges in Managed Realignment Research

Chapter 28. Methods and Criteria for Successful Mangrove Forest Restoration

1. Introduction

2. General Site Selection for Restoration

3. Specific Site Selection for Restoration

4. Establishing Success Criteria

5. Monitoring and Reporting Success

6. Functionality of Restored Mangrove Forests

7. Summary

Chapter 29. Evaluation of Restored Tidal Freshwater Wetlands

Acknowledgments

1. Introduction

2. Characteristics of Restored TFW

3. Success Evaluation of Restored Wetlands

4. Ecosystem Attributes Measured at Restored TFW

5. Criteria for Successful Restoration of TFW

6. Case Study: Evaluation of Restored TFW of the Anacostia River, Washington, DC, USA

7. Conclusions and Recommendations

Chapter 30. Surface Elevation Models

1. Introduction

2. Measuring Processes that Affect Wetland Elevation

3. Types of Models

4. Future Directions for Model Improvement

5. Conclusions

Chapter 31. Salt Marsh–Mangrove Interactions in Australasia and the Americas

1. Introduction

2. Distribution/Geomorphic Settings – Where do Mangrove and Salt Marsh Coexist?

3. Long-Term Dynamics

4. Recent Interactions

5. Stressors Controlling Delimitation of Mangrove

6. Conclusions

Chapter 32. Wetland Landscape Spatial Models

Acknowledgments

1. Introduction

2. Physical Models

3. Hydraulic Modeling

4. Hydrodynamic Modeling

5. Ecological Models

6. Individual-Based Modeling

7. Eco-Geomorphological Modeling

8. Ecosystem-Level Modeling

9. Desktop Dynamic Modeling

10. Conclusions

Subject Index

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British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Library of Congress Cataloging-in-Publication Data

Coastal wetlands : an integrated ecosystem approach/edited by Gerardo M. E. Perillo … [et al.]. — 1st ed. p. cm.

Includes bibliographical references and index.

ISBN 978-0-444-53103-2

1. Estuarine restoration. 2. Coastal zone management. 3. Wetland ecology.

4. Coasts. I. Perillo, G. M. E. (Gerardo M. E.)

QH541.5.E8C63 2009

577.69—dc22

2008046508

ISBN: 978-0-444-53103-2

For information on all Elsevier publications visit our website at elsevierdirect.com

Printed and bound in The Netherlands

09 10 11 12 10 9 8 7 6 5 4 3 2 1

Preface

Gerardo M.E. Perillo, Eric Wolanski, Donald R. Cahoon and Mark M. Brinson

This document is based on work partially supported by the U.S. National Science Foundation under Grants No. BSR-8702333-06, DEB-9211772, DEB-9411974, DEB-0080381 and DEB-0621014 and to SCOR under Grant No. OCE-0608600. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the authors and do not necessarily reflect the views of the U.S. National Science Foundation (NSF).

Why coastal wetlands? What is so important about them that a whole book is required to try to review and explain their large variety of properties? Of all the coastal habitats, wetlands are the least depicted in the tourist brochures because they lack those paradisiacal long, white sandy beaches backed by palm trees or expensive resort hotels close to transparent blue waters. In fact, most coastal wetlands are quite muddy and are more likely to be inhabited by crabs and worms than by charismatic fish, birds and mammals. Hence, most inhabitants of our world either have never thought about coastal wetlands or may consider them a nuisance, not realizing that their seafood dinner likely had its origin as a detrital food web in a salt marsh or mangrove swamp. Bahía Blanca (Argentina) inhabitants are a classical example: a city of over 300,000 people living within 10 km of a 2,300-km ² wetland, the largest of Argentina, but fewer than 40% have any idea that they are so close to the sea and a short distance of places that are globally unique (Perillo and Iribarne, 2003, in Chapter 6).

Similarly, there are many other coastlines dominated by wetlands, yet they are only seen as areas to exploit in an unsustainable fashion. For example, mangroves have served local communities for generations in many Asian tropical countries for harvesting wood and fish in contrast to their wholesale replacement for rice cultivation and shrimp farming.

Even though management guidelines have been available for decades, the negative consequences of uninformed exploitation have resulted in poor or even total lack of management criteria by most governments at all levels. Even local stakeholders fail to act in their own best interest without consideration of the ecosystem goods and services that the nearby wetlands provide.

Coastal wetlands best develop along passive-margin coasts with low-gradient coastal plains and wide continental shelves. The combination of low hydraulic energy and gentle slope provides an ideal setting for the wetland development. Also passive margins are less prone to receive large episodic events like tsunamis. Tsunamis and storm surges, in particular, are major coast modifiers, but when they act on low coasts their effects are more far reaching than they are on higher relief coasts. For a wetland to form, there is a need for a particular geomorphological setting such as an embayment or estuary providing a relatively low-energy environment favoring sediment settling, deposition and preservation. However, that is only the beginning of a large and complex life where many geological (i.e., sediment supply, geological setting and isostasy), physical (i.e., oceanographic, atmospheric, fluvial, groundwater processes and sea level changes), chemical (i.e., nutrients, pollutants), biological (i.e., intervening flora and fauna), and anthropic factors play a wide spectra of roles. Coastal wetlands are areas that have combined physical sources and biological processes to develop structure that continues to take advantage of natural energy inputs.

This book has been planned to address in an integrated way all these processes and their consequences on the characterization and evolution of coastal wetlands. It aims to provide an integrated perspective on coastal wetlands as ecosystems for the public, engineers, scientists and resources managers. It is only after acquiring this perspective that scientists can confidently propose ecohydrologic solutions for managing these environments in an ecologically sustainable way. This is but one small step toward encouraging humanity to look beyond purely technological, and often failed, solutions to complex environmental problems.

This is done by focusing on the principal components considering the full range of environments from freshwater to subtidal and from polar to tropical systems. The book has been divided into seven parts starting from a synthesis chapter that integrates the whole book. Part I covers, in three chapters, the general description of the wetlands structured according to broad climatic regions and introduces the most important physical processes that are common to all coastal wetlands including some geomorphologic and modeling principles. Part II are specific to each particular type of wetland (tidal flats, marshes and seagrasses, and mangroves). Within each part (Parts III to V), there are chapters dealing with their particular geomorphology, sedimentology, biology and biogeochemistry. Finally Parts VI and VII provide insight into the restoration and management and sustainability and landscape dynamics.

As editors, our work was greatly facilitated by the tremendous cooperation and enthusiasm from each of the authors to complete this process that began mid 2006. Each author, an authority in his or her specialty, was specifically invited to write a review chapter. Therefore, the challenges were much larger than in the case of typical contributed articles. But the reward, we think, is much more beneficial for the student, professor, or researcher employing this book for his or her particular interest. Readers will not only be able to find a specific topic but will find related information to complement and enhance the understanding of the topic.

We are in debt to the more than 50 reviewers (most of them are not authors in the book) who have agreed and provided graciously and unselfishly their valuable time. Some took on the responsibility of two chapters, and their efforts are rewarded with improvements of each contribution. In many cases, reviewers gave us interesting ideas that helped in the general structure of the book. A list of the reviewers is provided.

We also thank Elsevier Science and the various Publishing Editors who were in charge of our book along the period since we first proposed our idea to the final result that you are reading now. First of all to Kristien van Lunen who first believed that our proposal was realizable and then to the important contributions and patience of Jennifer Hele, and also to Pauline Riebeek, Linda Versteeg and lastly Sara Pratt. Stalin Viswanathan did an excellent job copyediting the whole book.

List of Contributors

Kenneth F. Abraham

Wildlife Research and Development Section

Ontario Ministry of Natural Resources

Peterborough, Ontario, Canada

Paul Adam

School of Biological, Earth and Environmental Sciences

University of New South Wales

NSW 2052, Australia

John R. L. Allen

Department of Archaeology

School of Human and Environmental Sciences

University of Reading

PO Box 227

Whiteknights, Reading RG6 6AB, UK

Daniel M. Alongi

Australian Institute of Marine Science

PMB 3

Townsville M.C.

Queensland 4810, Australia

Rebecca J. Aspden

Sediment Ecology Research Group

Gatty Marine Laboratory

St Andrews University

Fife KY16 8LB, UK

Andrew H. Baldwin

Department of Environmental Science and Technology

University of Maryland

1423 Animal Science Building

College Park, MD 20742, USA

Donald M. Baltz

Coastal Fisheries Institute

Louisiana State University

Baton Rouge, LA 70803-7503, USA

Aat Barendregt

Department of Environmental Sciences Copernicus Institute for Sustainable Development and Innovation

Utrecht University

PO Box 80115

3508 TC Utrecht, the Netherlands

Kevin S. Black

Partrac Ltd.

141 St James Road

Glasgow G4 0LT, UK

Laurence A. Boorman

L.A.B. Coastal

Holywell, St. Ives

Cambridgeshire PE27 4TQ, UK

Mark M. Brinson

Biology Department

Howell Science Complex

East Carolina University

Greenville, NC 27858, USA

Stephen W. Broome

Department of Soil Science

North Carolina State University

Box 7619

Raleigh, NC 27695-7619, USA

Donald R. Cahoon

U.S. Geological Survey

Patuxent Wildlife Research Center

c/o BARC-East, Building 308

10300 Baltimore Avenue

Beltsville, MD 20708, USA

John C. Callaway

Department of Environmental Science

University of San Francisco

2130 Fulton St.

San Francisco, CA 94117, USA

Luca Carniello

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and

International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Marcelo C. L. Cohen

Laboratory of Coastal Dynamics and Centre for Geosciences

Federal University of Pará

Avda Perimetral 2651

66077-530 Belém (Pa), Brazil

Perran L. M. Cook

Water Studies Centre

Monash University

Clayton

3800 Victoria, Australia

Christopher B. Craft

School of Public and Environmental Affairs

Indiana University

1315 E. 10th Street

Bloomington, IN 47405, USA

Andrea D’Alpaos

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Luigi D’Alpaos

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Gareth Davies

School of Earth and Environmental Sciences

University of Wollongong

Wollongong, NSW 2522, Australia

Dirk de Beer

Max Planck Institute for Marine Microbiology

28359 Bremen, Germany

Andrea Defina

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Joanna C. Ellison

School of Geography and Environmental Studies

University of Tasmania

Locked Bag 1376

Launceston, Tasmania 7250, Australia

Mark Fonseca

NOAA/NOS Center for Coastal Fisheries and Habitat Research

101 Pivers Island Road

Beaufort, NC 28516-9722, USA

Shu Gao

School of Geographic and Oceanographic Sciences

Nanjing University

Nanjing 210093, China

Angus Garbutt

NERC Centre for Ecology and Hydrology Environment

Centre Wales

Deiniol Road

Bangor, LL57 2UP Wales, UK

Richard S. Hammerschlag

U.S. Geological Survey

USGS Patuxent Wildlife Research Center

Laurel, MD 20708, USA

Marianne Holmer

Institute of Biology

University of Southern Denmark

Campusvej 55

5230 Odense M

Denmark

Robert L. Jefferies

Department of Ecology and Evolutionary Biology

University of Toronto

Ontario, Canada

Samantha B. Joye

Department of Marine Sciences

University of Georgia

Room 220 Marine Sciences Bldg

Athens, GA 30602-3636, USA

Jason R. Kirby

School of Biological and Earth Science Liverpool John Moores University

Byrom Street

Liverpool L3 3AF, UK

Stefano Lanzoni

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Rubén J. Lara

Centre for Tropical Marine Ecology Fahrenheitstrasse 6

28359 Bremen, Germany

Roy R. Lewis III

Lewis Environmental Services, Inc.

P.O. Box 5430

Salt Springs, FL 32134, USA

Karen McKee

National Wetlands Research Center

U. S. Geological Survey

Lafayette, LA 70808, USA

Marco Marani

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

I. Peter Martini

Department of Land Resource Science

University of Guelph

Ontario, Canada

Yoshihiro Mazda

Department of Marine Science

School of Marine Science and Technology

Tokai University

3-20-1 Orido

Shimizu, Shizuoka 424-8610, Japan

J. Patrick Megonigal

Smithsonian Environmental Research Center

P.O. Box 28

647 Contees Wharf Road

Edgewater, MD 21037, USA

R. I. Guy Morrison

Canadian Wildlife Service

National Wildlife Research Centre

Hull, Quebec, Canada

Scott C. Neubauer

Baruch Marine Field Laboratory

University of South Carolina

PO Box 1630

Georgetown, SC 29442, USA

Julian Oxmann

Centre for Tropical Marine Ecology Fahrenheitstrasse 6

28359 Bremen, Germany

Eric I. Paling

Marine and Freshwater Research Laboratory Environmental Science

Murdoch University

Murdoch 6150, Western Australia

David M. Paterson

Sediment Ecology Research Group

Gatty Marine Laboratory

St Andrews University

Fife KY16 8LB, UK

Gerardo M. E. Perillo

CONICET – Instituto Argentino de Oceanografía and Departamento de Geología

Universidad Nacional del Sur

CC 804

B8000FWB Bahía Blanca, Argentina

María Cintia Piccolo

CONICET – Instituto Argentino de Oceanografía and Departamento de Geografía

Universidad Nacional del Sur

CC 804

B8000FWB Bahía Blanca, Argentina

Andrew Plater

Department of Geography

University of Liverpool

P.O. Box 147

Liverpool L69 7ZT, UK

Paula D. Pratolongo

CONICET – Instituto Argentino de Oceanografía and Departamento de Biología, Bioquímica y Farmacia

Universidad Nacional del Sur

CC 804

B8000FWB Bahía Blanca, Argentina

Enrique Reyes

Department of Biology

Howell Science Complex

East Carolina University

Greenville, NC 27858, USA

Andrea Rinaldo

Dipartimento di Ingegneria Idraulica, Marittima Ambientale e Geotecnica and International Centre for Hydrology Dino Tonini

Università di Padova

via Loredan 20

I-35131 Padova, Italy

Victor H. Rivera-Monroy

Wetland Biogeochemistry Institute

Department of Oceanography and Coastal Science

Louisiana State University

Baton Rouge, LA 70803, USA

Kerrylee Rogers

Rivers and Wetlands Unit

Department of Environment and Conservation, NSW

PO Box A290

Sydney South NSW 1232, Australia

John M. Rybczyk

Department of Environmental Science

Western Washington University

516 High St.

Bellingham, WA 98225, USA

Neil Saintilan

Rivers and Wetlands Unit

Department of Environment and Conservation, NSW

PO Box A290

Sydney South NSW 1232, Australia

Bettina B. Schmitt

Centre for Tropical Marine Ecology, Fahrenheitstrasse 6

28359 Bremen, Germany

Pedro W.M. Souza Filho

Centre for Geosciences

Federal University of Pará

Av. Augusto Correa 1

66077-110 Belém (Pa), Brazil

Claudio F. Szlafsztein

Laboratory of Coastal Dynamics and Centre for Geosciences

Federal University of Pará

Avda Perimetral 2651

66077-530 Belém (Pa), Brazil

Craig Tobias

Department of Earth Sciences

University of North Carolina at Wilmington

601 S. College Rd.

Wilmington, NC 28403, USA

Robert R. Twilley

Wetland Biogeochemistry Institute Department of Oceanography and Coastal Science

Louisiana State University

Baton Rouge, LA 70803, USA

Marieke M. van Katwijk

Environmental Science

Radboud University

Nijmegen

Postbus 9010

6500 GL Nijmegen, the Netherlands

Mike van Keulen

School of Biological Sciences and Biotechnology

Murdoch University

Murdoch 6150, Western Australia

Jenneke M. Visser

Department of Oceanography and Coastal Sciences

School of the Coast and Environment

Louisiana State University

Baton Rouge, LA 70803, USA

Present address: Institute for Coastal Ecology and Engineering University of Louisiana at Lafayette PO Box 44650 Lafayette, LA 70504-4650, USA

Dennis F. Whigham

Smithsonian Environmental Research Center

P.O. Box 28

647 Contees Wharf Road

Edgewater, MD 21037, USA

Eric Wolanski, DSC (Hon. Causa), FTSE, FIE Aust

Australian Centre for Tropical Freshwater Research and Department of Marine

Biology and Aquaculture

James Cook University and Australian Institute of Marine Science

Townsville, Australia

Colin D. Woodroffe

School of Earth and Environmental Sciences

University of Wollongong

Wollongong, NSW 2522, Australia

List of Reviewers

Susan Adamowicz

Carl L. Amos

Shimon C. Anisfeld

Edward Anthony

Andrew H. Baldwin

Thomas S. Bianchi

Henry Bokuniewicz

Henricus T.S. Boschker

Mark M. Brinson

David M. Burdick

Jaye Cable

Donald R. Cahoon

John Calloway

Marcelo Cohen

Christopher Craft

Steve Crooks

Carolyn A. Currin

Stephen E. Davis, III

Donald D. DeAngelis

Monique Delafontaine

Norman C Duke

Keith Dyer

Susana Enríquez

Stuart Findlay

Jon French

Carl Friedrichs

Keita Furukawa

Carl Hershner

John Jeglum

Samantha Joye

Ruben Lara

Mary A Leck

Jorge Marcovecchio

James Morris

David Osgood

Morten Pejrup

Gerardo M. E. Perillo

James Perry

John Portnoy

Denise J. Reed

Peter Ridd

Ken Ross

Lawrence Rozas

Neil Saintilan

T.J. Smith

Charles Tarnocai

Stijn Temmerman

Robert Twilley

Reginald Uncles

Steven Victor

Melanie Vile

Robert Whitlatch

Rob Williams

Eric Wolanski

Chapter 1. Coastal Wetlands

A Synthesis

Eric Wolanski, Mark M. Brinson, Donald R. Cahoon and Gerardo M.E. Perillo

Contents

1. Introduction 1

2. A Synthesis of Coastal Wetlands Science 2

2.1. Geography 3

2.2. Geomorphology evolution under climate change 4

2.3. The influence of vegetation on the geomorphology evolution with climate change 6

2.4. The stabilizing role of vegetation 9

2.5. State change and coastal evolution 13

2.6. The role of physical disturbances 14

2.7. The role of herbivores 18

2.8. Observations across ecosystem types 21

2.9. The human impact 28

2.10. Modeling and predictions 31

2.11. Coastal wetland ecosystems as a component of estuaries 32

2.12. Coastal wetland socioeconomics 34

2.13. Coastal wetlands are essential for our quality of life 38

3. Lessons from the Chapters in This Book 43

3.1. Coastal wetlands as ecosystems 44

3.2. Physical processes 45

3.3. Tidal flats 47

3.4. Marshes and seagrasses 48

3.5. Mangroves 51

3.6. Coastal wetland restoration and management 53

3.7. Coastal wetland sustainability and landscape dynamics 56

References 57

1. Introduction

Our understanding of the functioning of coastal wetland ecosystems has grown rapidly over the past decade. We have gained insight into the roles of geomorphic processes, hydrologic dynamics, biotic feedbacks, and disturbance agents in creating and molding a variety of coastal wetland ecosystems across climatic gradients. The variety is expressed not so much in the more obvious differences in vegetation cover, but rather how physical processes both facilitate and constrain a diversity of plant and animal communities. At one level, coastal wetlands are the product of tidal forces and freshwater inputs at the margin of continents. At another level, biotic factors exert feedback controls through biofilms, bioturbation of sediments, the buffeting of currents and waves, organic enrichment of sediments, and the closing of nutrient cycles. Few ecosystems provide us with such clear examples of feedback controls. Still, much remains to be learned and understood. What we do understand about the structure and functioning of coastal wetlands should provide the theoretical underpinnings for effective management in protecting them for their many contributions to ecosystem goods and services. And what we do not understand should compel us to focus our attention and energies toward seeking optimal solutions to some of the most perplexing and urgent problems facing natural resource management.

What are coastal wetland ecosystems and what are their limits of distribution? There are several general definitions for wetlands, but the Ramsar definition is likely the most broadly encompassing (http://www.ramsar.org/), while others are more focused definitions tailored to country-specific protection and management policies (Mitsch and Gosselink, 2000). We offer a very general approach rather than a precise definition: coastal wetlands are ecosystems that are found within an elevation gradient that ranges between subtidal depths to which light penetrates to support photosynthesis of benthic plants and the landward edge where the sea passes its hydrologic influence to groundwater and atmospheric processes. At the seaward margin, biofilms, benthic algae, and seagrasses are representative biotic components. At the landward margin, vegetation boundaries range from those located on groundwater seeps or fens in humid climates to relatively barren salt flats in arid climates. This book focuses on commonly recognized ecosystems along this hydrologic gradient: seagrasses, tidal flats, tidal salt and freshwater marshes, and mangrove and tidal freshwater forests. Coral reefs are not covered at all because they are so physically and biologically distinct from the foregoing list, and in part because they have received research attention equivalent to the totality of all of the wetlands covered in this book (Duarte et al., 2008). Little direct reference is made in this book to lagoons that are intermittently connected to the sea; regardless, all of the ecosystem types (seagrass meadows, mudflats, marshes, and mangroves) can and do occur in these and other more specialized geomorphic settings. They would all comprise the array that we recognize as coastal wetlands.

This book addresses the pressing need to quantify the ecological services provided by coastal wetlands as a tool to press for better management worldwide, because coastal wetlands are disappearing worldwide at an alarming rate; in some countries, the loss is 70–80% in the last 50 years (Frayer et al., 1983 and Duarte, 2002; Hily et al., 2003Bernier et al., 2006Duke et al., 2007 and Wolanski, 2007a).

2. A Synthesis of Coastal Wetlands Science

In this section, we provide an overview of the structure and functioning of coastal wetlands with emphasis on key forces and processes that interact with their coastal geographic location. It is difficult to discuss these forces in isolation because climate change influences sea level, sea level forces changes in vegetation structure, disturbance and herbivory affect vegetation, and so forth. As such, comparison across major ecosystem types can provide insight into the differences and the relative importance of both extrinsic forces and intrinsic structure. We also discuss the role of modeling in elucidating the relative importance of key processes and in predicting the effects of alteration by humans. This last effect presents us with challenges of how to best protect and manage coastal wetlands for their attributes of life support, and importantly for the natural beauty and their contribution to cultural values. To this end, we outline key research needs that recognize the usefulness of working beyond single factor cause and effect.

2.1. Geography

Coastal wetlands include salt marshes, mangroves, tidal flats, and seagrasses. They are found in all continents and at all latitudes. Cliffs and rocky shores are probably the only coasts with minimal wetlands.

Mangroves cover about 230,000 km ² worldwide (Diop, 2003 and Duke et al., 2007), and are restricted to a mangrove belt at low and mid-latitudes (Figure 1) as controlled by the combination of continental (no frosts or only very rarely, typically less than one frost every 10 years; Lugo and Patterson Zucca, 1977) and oceanic climates (warm waters; Duke et al., 2007). Salt marshes cover a larger area worldwide; in North America alone this area is 300,000 km ² mainly in Canada and Alaska (Mitsch and Gosselink, 2000), and possibly a similar large area of salt marshes exists in northern Russia. Salt marshes are also found scattered in the mangrove belt, usually in the upper intertidal areas landward of mangroves. Seagrasses cover at least 165,000 km ² worldwide, possibly up to 500,000 km ² (Green and Short, 2003). The worldwide area of tidal flats and freshwater coastal wetlands seems unknown though order-of-magnitude estimates suggest that they may reach 300,000 km ².

2.2. Geomorphology evolution under climate change

Wetlands continuously evolved in time and space. The story of the present-day coastal wetlands starts 120,000 years ago, which is an interglacial period that lasted about 15,000 years and when the mean sea level was at a level comparable to the present one (Figure 1). Where natural conditions allowed, coastal wetlands probably flourished with a somewhat similar flora and fauna as today, though the macrofauna was very different. The human population was tiny at that time and had a negligible impact on the evolution of the coastal wetlands.

Then came a 100,000-year-long ice age that buried much of the present temperate coasts and continental shelves under hundreds of meters, sometimes several kilometers, of ice. Water was taken away from the ocean and stored on the continents. As a result, the mean sea level decreased and was at its lowest, about 120 m below the present mean sea level, about 20,000 years ago (Figure 2). At that time, coastal wetlands would only have existed along those coasts that were not buried by permanent ice, mainly along the then tropical belt. Even there, the coastal wetlands could only have existed on the upper continental slope, which is where at that time the sea met the land (Figure 3). Because the inclination of the continental slope is much steeper than the continental shelf where the coastal wetlands are now located, the space for accommodating coastal wetlands would have been very limited. At that time, the area occupied by coastal wetlands would have been much smaller and in many cases restricted to estuaries that developed along the present-day submarine canyons (Perillo, 1995). Taking all of this into account, coastal wetlands would have occupied perhaps as little as 5% of the area covered just prior to human alterations.

Then, 20,000 years ago came a period of high disturbance for coastal wetlands. The mean sea level rose rapidly, much faster than it fell in the previous 100,000 years (Figure 2). The mean sea level reached its present, world-averaged value about 6,000 years ago. At that time, the bulk of the continental ice outside of polar and near-polar regions had melted, which released a huge weight from those continental shelves that were burdened by ice. In those areas, the continental shelf rose by isostatic rebound of the land, with the largest rebound occurring where the ice burden was the greatest. In those areas (zone A in Figure 1), the land rose faster than the sea, and the relative mean sea level decreased over the last 10,000 years, the coast prograded, and new land emerged (Figure 4). In other areas, the relative sea level rose about 20 m during the last 10,000 years. In some areas (zone B in Figure 1), this increase was asymptotic, being largest 10,000 years ago and minimal at present (Figure 4); in such areas, the coast retreated from an advancing sea. In the other areas (zone C in Figure 1), the relative sea level reached a maximum about 5,000 years ago, typically 3 m higher and up to 7 m higher than at present, and it decreased smoothly to its present elevation until about 2,000 years ago. In those areas, the coast initially receded from an advancing sea and then prograded slightly during the last 5,000 years. Within each of these zones, there were local exceptions, especially in delta regions where sediment supply was sufficient to compensate for coastal retreat.

Worldwide, the estuaries responded to these changing conditions of sea level, river and sediment discharge, changing sea ice, water currents, storms, and waves brought about by a new climate. The elevation of the mean sea level relative to land level determined at any time the location of the estuary. The net sediment budget, the balance between the sediment inflow (from the river and an eventual import from the sea) and the sediment outflow from the estuary to the sea, determined the evolution of the estuary. Where the relative sea fell rapidly (zone A), new land emerged constantly and the estuary migrated seaward, continually reinventing itself; the old estuary rose above the sea level and became part of the landscape; this evolution is still proceeding in zone A (Figure 5 a). Where sea level rose, old estuaries were drowned and new estuaries formed landward where the land met the sea (Figure 5b).

In zone A, no steady-state has yet been reached. The estuary is still steadily moving seaward as new land emerges. In zone B, a quasi-steady state is being reached only recently as the relative mean sea level stabilized only in the last 1,000 years or so. In zone C, the estuary is still evolving, albeit much slower than in the past because the changes in the relative mean sea level slowed down considerably during the last 6,000 years; estuaries are, however, still not at a steady state and coastal wetlands continue to migrate.

2.3. The influence of vegetation on the geomorphology evolution with climate change

Estuaries trap some portion of the riverine sediment input (Perillo, 1995 and Wolanski, 2007a). Estuaries in zone A are thus silting as soon as they form in a new position; small estuaries not flushed by the spring snow melt of large rivers are rapidly turning into mud flats that are stabilized by biofilms and they themselves can rapidly be colonized by salt marsh vegetation, which creates new high latitude salt marshes (Martini et al., 2009). The evolution of estuaries in zones B and C was controlled by the geomorphology when the rising seas flooded new land and by the tides, river runoff, and oceanographic conditions at the coast. Because these conditions varied from site to site around the world, a large number of estuarine evolution pathways occurred (Ellison, 2009 and Woodroffe and Davies, 2009). At some locations (Figure 5c), a wide, shallow estuary formed; this enabled a vast coastal wetland to form that fringed the main channel as the estuary filled with sediment from both the river and the sea (through processes such as tidal pumping). Because of vegetation and adequate sediment supplies, this wetland type accelerated siltation and enabled the estuarine system to keep up with sea level rise. Later on this estuarine, saline sediment was capped by freshwater sediment; the estuary had by then reached old age because it had used all the accommodating space on land and it can then only evolve further by moving offshore to form a delta; or it can occasionally be rejuvenated by opening a new channel or by isostatic or structural movements. In other cases (Figure 5d), the estuary moved upland with a rising sea level until about 6,000 years ago when the sea level stabilized; as the estuary migrated, so did the coastal wetlands, leaving behind their signature in the form of a well-preserved organic-rich mud layer. This mud was capped by inorganic sediment, which provided a substrate on which the wetland prograded, and in the process forming a near-surface, organic-rich mud layer. Still in other cases (Figure 5e), principally in microtidal areas and in areas where the geomorphology formed semienclosed coastal waters as the sea level rose, riverine sediment filled a lagoon-type environment, and this area was colonized by wetlands, leaving behind an organic-rich mud layer. The relative sea-level evolution over the last few thousand years has created in sediment-starved estuaries of zone C a varied succession of wetland habitats, including (Figure 5f) the die-off of mangroves and their capping by calcareous sediment later colonized by seagrass, and (Figure 5g) the die-off of mangroves, their capping by calcareous sediment with an even later capping by a new mangrove swamp with accreting, organic-rich mud.

Thus, coastal wetlands have survived, adapted, and greatly expanded their distribution area during the last 10,000 years.

2.4. The stabilizing role of vegetation

Several authors in this book (D’Alpaos et al., 2009Ellison, 2009Gao, 2009Lara et al., 2009 and Woodroffe and Davies, 2009) argue that these coastal wetlands are not just opportunistic colonizers of intertidal areas formed by the accumulation of mud in sheltered estuaries and coastal waters. As long as the mud flat elevation remains below the mean sea level, generally the mud flat remains fairly flat and uneventful and no tidal creeks may form (Figure 6a). Once the mud flat rises above mean sea level, the tidal hydrodynamics change profoundly. A large volume of water floods and drains the mud flat in a short time compared to the entire tidal period. This flow becomes funneled in a natural depression; this generates large currents, which in turn quickly erode the banks to form a tidal course (Figure 6b). The tidal course grows by the tidal asymmetry in it, with larger ebb than flood tidal currents, a situation characteristic of shallow water hydrodynamics (D’Alpaos et al., 2009 and Mazda and Wolanski, 2009). In addition, intense erosion at the head of the tidal course is driven by small waterfalls that form there at ebb tide although other forms of erosion also help in the growth of the tidal creek over unvegetated mud flats (Perillo, 2009). At ebb tide, frictional forces retard the flow over the mud flat more than the ebb flow is retarded in the tidal channel (Figure 6c). A tidal creek forming in an unvegetated mud flat develops a dendritic pattern with several erosional hot spots, each one located at the tip of a creek (Figure 6d).

Before a mud flat becomes vegetated, it can be stabilized by biofilms or destabilized by bioturbation. Once the mud flat is vegetated by pioneer species of mangroves or salt marsh vegetation, a quasi-steady state develops characterized by a sharp discontinuity separating the vegetated wetland from the unvegetated tidal channel (Figure 7; Perillo, 2009); the exception is Arctic coastal wetlands because scouring by ice reshapes the bathymetry yearly (Figure 7d). The patterns of erosion and siltation change profoundly once the vegetation is established for three main reasons. First, the roots of the trees or the grass lock the soil together and this slows or even stops bank erosion by tidal currents and waves to a depth of several meters in mangroves in well-aerated soils near the banks and several tens of centimeters deeper in the forest (Figure 6e), typically a few tens of centimeters in salt marshes, and 15 cm in seagrass (Holmer, 2009). This process is apparent in Figure 6c that shows the tidal creek in the muddy boundary where it meets the vegetation. Second, the wave erosion of the vegetated substrate is decreased because hydraulic energy is dissipated by vegetation (Figure 6 f–h for the case of mangroves; salt marsh vegetation is less tall, but in the lower 30 cm, the vegetation density can be higher than that of mangroves) and effectively absorbed by wave energy (Figure 8 a–c for mangroves, seagrass, and salt marshes, respectively). Thirdly, the tidal currents are slowed down by the vegetation to values equal to 5% than in the tidal creek; also the currents around the vegetation generate eddies and stagnation zones where the suspended sediment brought in at rising tide will settle; the tidal currents are too small at falling tide to resuspend all that sediment; thus the vegetated tidal flat silts (D’Alpaos et al., 2009; Lara, 2009). From the wetland edge to the interior, waves are attenuated, thus reducing their capability to erode and transport sediments.

To quantify the evolution of an estuary, models have been developed to compute the sediment dynamics over vegetated and unvegetated mud flats. The aim of these models is to simulate the evolving interaction between currents and sediment transport, since this interaction determines the evolution of the bathymetry (D’Alpaos et al., 2009 and Lara et al., 2009). Although most modelers rely on engineering formulae to calculate sediment transport from water currents and waves, in practice these formulae neglect the biology. Consequently, this approach is of little predictive value in vegetated coastal wetlands where the sediment dynamics are highly influenced by the inadequately quantified influences of biota and biotic processes. Even in the simplest case of unvegetated mud flats, a variety of biological mechanisms can result in either destabilizing the mud (e.g., by burrowing fauna) or in stabilizing the mud (e.g., by an algal mat on the substrate; Le Hir et al., 2007 and Wolanski, 2007a). Further, there are no models to predict which flora and fauna will ultimately dominate (but see Reyes (2009) for habitat switching models). Thus, for vegetated wetlands (salt marshes and mangroves), the ability of models to predict the future evolution of the linked sedimentology–ecology remains an active and challenging area of research.

2.5. State change and coastal evolution

A fluctuating sea level, and the geomorphologic evolution that it induces, generates changes of the ecological state of coastal wetlands. These changes modulate the transgression upslope from upland forest to subtidal ecosystems (Figure 9 a) as a surface expression of the scenario shown in Figure 5c. During this coming century, the historical evolution of coastal wetlands described above may be accelerated, or reactivated for systems that have reached steady state at present, by the predicted increased rate of sea level rise due to global warming. Ecological succession of vegetation occurs within each of the states as a result of such disturbances as long as hydrology and sediments do not change substantially. For example, freshwater wetland forest landward of high marsh may die back after a fire, but unless salinity has increased, ecological succession will return the vegetation to forest. Alternatively, an increase in salinity will prevent forest re-establishment and subsequently change the state to emergent salt marsh. Often, the vegetation is out of phase with physical conditions so that disturbance such as fire or wrack deposition becomes the trigger to initiate change (Brinson et al., 1995).

Over larger timescales and more divergent conditions, ecosystem types may change from one to another (Figure 9a). These coastal ecosystems have different flooding and aeration regimes that control plant dynamics (Figure 9b). Submerged aquatic vegetation (SAV, such as seagrass) has persistent flooding, mud/sand flats are intermittently exposed, and wetlands have intermittent aeration of the root zone. In contrast, uplands are not under the influence of sea level and rarely experience soil saturation. SAVs differ from flats by the presence of roots in the sediment and a requirement for transparency in the water column. State change among ecosystem types normally occurs over much longer timescales than ecological succession and may be better considered under the rubric of coastal evolution rather than disturbance.

2.6. The role of physical disturbances

The role of physical disturbances in coastal wetland evolution is recognized but seldom quantified as yet. At high latitudes, salt marshes are yearly scoured by sea ice, creating a network of patches of rich vegetation, degraded vegetation, and unvegetated mud flats (Figure 7d; Martini et al., 2009). Further, ice erodes by floating imbedded sediments and transports them elsewhere for deposition. At temperate latitudes, crabs in salt marshes can excavate local depressions that can develop into tidal creeks or become unvegetated muddy patches (Perillo, 2009). In temperate and tropical climates, however, the most common natural disturbances are storms. In mangrove areas, hurricanes can destroy the vegetation facing the ocean by the force of the wind (Figure 10 a).

People living along the edge of the coastal wetlands are particularly at threat from storm surges (Figure 11) because storm surge flood waves occur at long timescales (hours to days) and are therefore little attenuated by vegetation. Tsunamis are rare but generate a major disturbance over much shorter periods of time. The presence of extensive areas of shallow waters, such as intertidal flats, is important because it presents a wide area for dissipation of energy before the wave hits the vegetation. In mangroves, for small (<6 m) tsunamis waves the damaged area may be limited (Figure 10b) to a 5- to 10-m wide strip along the tidal creek and another, much wider, strip along the coast (Mazda and Wolanski, 2009). This occurs because the vegetation is an obstacle to swift flows and thus steers the tsunami wave along the tidal creek. Thus, people (fishermen) living along the tidal creek – with no vegetation between them and the creek – are not protected against a small tsunami; however, people living behind the mangrove belt receive some protection (Wolanski, 2007b and Alongi, 2008). Provided that the tsunami wave is less than 6 m and the trees are fully developed, the vegetation can survive and the wave at 500 m inland is transformed from rising in 10 seconds to rising in 3 minutes; this gives a chance for people to take shelter or flee the area, and the attenuation considerably reduces damage to property and infrastructure (Wolanski, 2007b). A large tsunami (wave >6 m) flattens and uproots the vegetation. In some cases, the energy extracted from the wave and used for this destruction reduces the wave energy sufficiently to provide some protection to people living in the coastal zone inland from the mangroves (Figure 12). However, for a large tsunami wave, mangroves and coastal forests are known to significantly contribute to human mortality from the physical damage of water-borne debris (Cochard et al., 2008).

In climates with distinct wet and dry seasons, the evolution of the bathymetry of a tidal creek may be determined by seasonal bank slumping principally at the end of the wet season, which is a large-scale process that vegetation cannot stop (Figure 13 a) and one which all estuarine geomorphology evolution models ignore so far. In other cases, the tidal creek develops from headward erosion at the tip of the creek during floods; during the subsequent dry season, saline water floods the eroded area, kills the freshwater vegetation, and creates bare soils that are unprotected by vegetation and thus readily erode; this allows the creek to grow in the tidal freshwater flood plains (Figure 13b). This process is also still largely ignored by estuarine geomorphology evolution models.

A rapid sediment deposit of more than 5 cm (during a storm) can kill mangroves (Ellison, 1998); by contrast, salt marsh plants seem able to cope well and plant growth is even accelerated following such events (Boorman, 2009 and Garbutt and Boorman, 2009). Both salt marshes and mangroves suffer mortality through hydrologic regime change, particularly from sustained flooding or loss of structural support through sediment erosion. The below-ground biomass of mangroves, seagrass meadows, and salt marshes offers enormous protection against erosion, thus providing a more stable medium for plant growth. The soft sediments of unvegetated intertidal flats are highly mobile and generally inhospitable to emergent plant growth. The establishment of emergent vegetation is rapid once the right hydraulic conditions occur, and from that time onward, the vegetation modifies the water and sediment circulation and promotes further biomass accumulation and the creation of a new ecosystem. What parameters control the threshold conditions for the establishment of pioneer vegetation over a soft sediment system still remain unclear. Once initial colonization has been achieved, vegetation may develop rapidly, being possibly a seagrass meadow, a mangrove swamp, or a salt marsh. This is commonly achieved by the arrival of species of fauna and flora with strong dispersal characteristics, followed by corresponding changes in food web dynamics, sedimentology, and hydrology. Humanity has accelerated this process by inadvertently or purposefully encouraging the spread of exotic invasive species and by the nutritive effects of eutrophication.

2.7. The role of herbivores

In mangroves, crabs dig burrows often at high densities (Figure 14 a). These burrows circulate tidal waters and permit salt to be flushed from sediments, thus preventing hypersalinity (Alongi, 2009 and Mazda and Wolanski, 2009) that would otherwise occur from mangroves that selectively avoid salt uptake and increase pore water salinity of the sediments. When crabs are destroyed, the groundwater becomes hypersaline and the vegetation is either destroyed or it becomes stunted. Similarly, farming on sulfur-rich soils acidifies the soil through the oxidation of sulfides; the vegetation is destroyed and cannot be recovered (Figure 14b) without massive additions of lime. Coastal wetlands with an aerial canopy of vegetation generate a microclimate at the sediment surface, which in turn affects the fauna and flora. For instance at mid-latitudes, the daily temperature variation in stagnant pools of water in wetlands exceeds 5°C if the pools are shaded by mangroves and 15°C if the pools are not shaded; this in turn affects the population dynamics of fish and mosquito in these pools (Knight, 2008).

The fate of dead organic matter is important in determining the evolution of a coastal wetland. There are large differences between the fate of above-ground plant biomass and the below-ground plant biomass. Live above-ground biomass can be consumed by crabs, snails, shrimps, and cone shells (Figure 15 a–b); for instance, shrimps can remove up to 50% of seagrass leaf production (Vonk et al., 2008) and crabs and snails climb trees to harvest mangrove leaves (Vannini and Ruwa, 1994 and Vannini et al., 2008). Another fraction can be exported as particulate organic matter outwelling that in turn enhances detrital food webs in the adjoining estuarine and coastal waters; the remaining fraction may be accumulated on the substrate for eventual burial by sediment deposition. Below-ground organic biomass is generally not exported in particulate form but rather is lost through decomposition by microbes as carbon dioxide and methane, and dissolved nutrients. Decomposition takes several months to years for labile organic matter, while refractory organic matter may take hundreds of years to