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Estuarine Ecology A detailed and accessible exploration of the fundamentals and the latest advances in estuarine ecology In the newly revised third edition of Estuarine Ecology, a team of distinguished ecologists presents the current knowledge in estuarine ecology with particular emphasis on recent trends and advances. The book is accessible to undergraduate students while also providing a welcome summary of up-to-date content for a more advanced readership. This latest edition is optimized for classroom use, with a more intuitive mode of presentation that takes into account feedback from the previous edition's readers. Review questions and exercises have been added to assist in the learning and retention of complex concepts. Estuarine Ecology remains the gold standard for the discipline by taking stock of the manifold scientific breakthroughs made in the field since the last edition was written. It also offers: * Thorough introductions to estuarine geomorphology, circulation, and chemistry * In-depth treatments of estuarine primary and secondary production, including coastal marshes and mangrove wetlands * A holistic view of estuarine ecosystems, their modeling and analysis, as well as the impact of human activities and climate change * A companion website with detailed answers to exercise questions Perfect for students of estuarine ecology, environmental science, fisheries science, oceanography, and natural resource management, Estuarine Ecology will also earn a place in the libraries of professionals, government employees, and consultants working on estuary and wetlands management and conservation.

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Estuarine Ecology

Third Edition

Edited by

BYRON C. CRUMP

College of Earth, Ocean, and Atmospheric Sciences

Oregon State University

Corvallis, OR 97330, USA

JEREMY M. TESTA

Chesapeake Biological Laboratory

University of Maryland Center for Environmental Science

Solomons, MD 20688, USA

KENNETH H. DUNTON

Marine Science Institute

The University of Texas at Austin

Port Aransas, TX 78373, USA

This third edition first published 2023© 2023 John Wiley & Sons, Inc.

Edition History: John Wiley & Sons Inc (1e, 1989; 2e, 2012)

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions.

The right of Byron C. Crump, Jeremy M. Testa, and Kenneth H. Dunton to be identified as the authors of the editorial material in this work has been asserted in accordance with law.

Registered OfficeJohn Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA

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Limit of Liability/Disclaimer of WarrantyIn view of ongoing research, equipment modifications, changes in governmental regulations, and the constant flow of information relating to the use of experimental reagents, equipment, and devices, the reader is urged to review and evaluate the information provided in the package insert or instructions for each chemical, piece of equipment, reagent, or device for, among other things, any changes in the instructions or indication of usage and for added warnings and precautions. While the publisher and authors have used their best efforts in preparing this work, they make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives, written sales materials or promotional statements for this work. The fact that an organization, website, or product is referred to in this work as a citation and/or potential source of further information does not mean that the publisher and authors endorse the information or services the organization, website, or product may provide or recommendations it may make. This work is sold with the understanding that the publisher is not engaged in rendering professional services. The advice and strategies contained herein may not be suitable for your situation. You should consult with a specialist where appropriate. Further, readers should be aware that websites listed in this work may have changed or disappeared between when this work was written and when it is read. Neither the publisher nor authors shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages.

Library of Congress Cataloging‐in‐Publication DataNames: Crump, Byron C., editor. | Testa, Jeremy M., editor. | Dunton, Kenneth H., editor. | Day, John W., 1945‐ author of introduction.Title: Estuarine ecology / edited by Byron C. Crump (College of Earth, Ocean, and Atmospheric Sciences, Oregon State University), Jeremy M. Testa (Chesapeake Biological Laboratory, University of Maryland Center for Environmental Science), Kenneth H. Dunton (Marine Science Institute, the University of Texas at Austin).Description: Third edition. | Hoboken, NJ : Wiley, 2023. | Includes bibliographical references and index.Identifiers: LCCN 2022017156 (print) | LCCN 2022017157 (ebook) | ISBN 9781119534655 (cloth) | ISBN 9781119534624 (pdf) | ISBN 9781119534563 (epub)Subjects: LCSH: Estuarine ecology.Classification: LCC QH541.5.E8 E849 2023 (print) | LCC QH541.5.E8 (ebook) | DDC 577.7/86–dc23/eng/20220706LC record available at https://lccn.loc.gov/2022017156LC ebook record available at https://lccn.loc.gov/2022017157

Cover Design: WileyCover Image: © Brent Durand/Getty Images (front cover);courtesy of Kenneth H. Dunton & Byron C. Crump (back cover)

We dedicate this book to the memory of our dear colleagues and friends Alejandro Yáñez‐Arancibia and W. Michael Kemp. Alex and Michael were editors and authors of the first and second edition of this book, and their spirit and energy live on in the third edition. Many of the authors of this book were influenced beyond measure by Alex and Michael in many ways, through collaboration, inspiration, and mentorship. We hope this book continues to serve as a living reflection of their innumerable contributions to the field of estuarine science.

Preface

This book is a textbook for a course in estuarine ecology designed to introduce students to the interdisciplinary study of estuarine ecosystems. The textbook is not only designed for upper‐level undergraduate students and introductory‐level graduate students, but it is also a useful reference for estuarine scientists across the fields of physical, biological, chemical, and ecological sciences. This book is the third edition of Estuarine Ecology; the first edition was published in 1989 and was the first book focused on estuarine ecology; the second edition was published in 2012 and expanded in length and scope to account for the explosion of literature and knowledge related to estuarine ecology; this third edition not only builds upon the expanded scope of the second edition but also represents a transformation to a textbook geared toward a wider range of students (upper‐level undergraduates) and a more complete set of teaching tools (study questions, exercises).

We continued the spirit of the second edition by engaging with a broad range of experts to sufficiently communicate the latest knowledge in a wide array of topics, and we also engaged a new cohort of early career scientists who carry on the tradition and knowledge of their graduate advisers and senior colleagues. The book begins with an introduction to estuaries and their ecology, with a new emphasis on how estuaries are in many ways a network of linked environments over salinity gradients, land–water interfaces, water columns, and underlying sediments, and from enclosed waters out to the sea. This introduction is followed by chapters that emphasize the hydrodynamic and chemical features of estuaries. These introductory perspectives are followed by a collection of chapters on estuarine primary producer communities, including phytoplankton, benthic algae, seagrasses, coastal marshes, and mangrove ecosystems; these chapters give students and faculty alike a detailed view of the key ecological features of these ecosystems that serve as the foundation of estuarine productivity. Next, several chapters detail the key consumer communities in estuaries, ranging from microscopic communities through zooplankton and on to benthos, nekton, and wildlife. The third edition includes a combination of the two previous microbial chapters, an expanded zooplankton chapter and a reorganization of the book into a sequence of consumers beginning with microbes and ending at megafauna. The final section of the book includes chapters that either provide synthetic treatments of key ecological topics or address more holistic aspects of estuarine ecosystem ecology. These include chapters on food webs and ecosystem metabolism, followed by tools‐focused chapters of ecosystem modeling and fisheries sciences, and culminating in a broad summary of how estuaries respond to global changes.

The third edition of Estuarine Ecology continues the challenge of providing a comprehensive treatment of a diverse array of topics relevant for the field of estuarine science. In the preface to the second edition, those authors aimed to consider the book “a work in progress” that provides a framework that can be routinely updated to keep pace with both the fast pace of global change and the scientific literature that describes it. We took on the task of this first rapid iteration and also took the opportunity to make the book more widely adaptable for undergraduate and graduate coursework. At this time of publication, we can already see the light of the fourth edition emerge.

Byron C. CrumpJeremy M. TestaKenneth H. Dunton

Acknowledgments

This book is a product of years of sustained efforts by the chapter authors, who we thank for their voluntary contributions to the community. We fondly recall our initial meeting with the authors at the Providence 2017 meeting of the Coastal and Estuarine Research Federation to launch the third edition. It is no small feat that all chapter authors contributed substantially to the process of writing the book, working with us to edit their previous chapters toward a broader audience, generating review questions for their chapters, and offering their time to review other chapters to make sure they are as widely understood as possible. We would also like to recognize the previous editors and authors of this book, John Day, Jr., Charles Hall, W. Michael Kemp, and Alejandro Yáñez‐Arancibia, whose intellectual contributions and spirit live on in the third edition. We also recognize the Coastal and Estuarine Research Federation, who has strongly supported the publication of this book.

List of Contributors

Anderson, Iris, Virginia Institute of Marine Sciences, College of William and Mary, Gloucester Point, VA, USA

Aoki, Lillian, Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, NY, USA

Benfield, Mark C., Louisiana State University, College of the Coast and Environment, Baton Rouge, LA, USA

Bianchi, Thomas S., Department of Geological Sciences, University of Florida, Gainesville, FL, USA

Blum, Linda K., Laboratory of Microbial Ecology, Department of Environmental Sciences, University of Virginia, Charlottesville, VA, USA

Borum, Jens, Department of Biology, University of Copenhagen, Copenhagen, Denmark

Cable, Jaye E., Department of Earth, Marine, and Environmental Sciences, University of North Carolina, Chapel Hill, NC, USA

Cowan, James H., Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, LA, USA

Crump, Byron C., College of Earth, Ocean, and Atmospheric Sciences, Oregon State University, Corvallis, OR, USA

Day, John W., Department of Oceanography and Coastal Sciences, College of the Coast and Environment, Louisiana State University, Baton Rouge, LA, USA

Dunton, Kenneth H., University of Texas at Austin, Marine Science Institute, Port Aransas, TX, USA

Elphick, Chris S., Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT, USA

Fleeger, John W., Department of Biological Sciences, Louisiana State University, Baton Rouge, LA, USA

Fong, Peggy, Department of Ecology and Evolutionary Biology, University of California Los Angeles, Los Angeles, CA, USA

Greenberg, Russel, Smithsonian Migratory Bird Center, National Zoological Park, Washington, DC, USA

Gregory Shriver, W., Department of Entomology and Wildlife Ecology, University of Delaware, Newark, DE, USA

Gruber, Renee K., Australian Institute of Marine Science, Townsville, QLD, Australia

Gurbisz, Cassie, St. Mary’s College of Maryland, St. Marys City, MD, USA

Hagy, James D., Center for Environmental Measurement and Modeling, Atlantic Coastal Environmental Sciences Division, Narragansett, RI, USA

Hopkinson, Charles S., Department of Marine Sciences, University of Georgia and Georgia Sea Grant, Athens, Georgia, USA

Ibáñez, Carles, Department of Climate Change, EURECAT, Technological Centre of Catalonia, Amposta, Catalonia, Spain

Justić, Dubravko, Department of Oceanography & Coastal Sciences, School of the Coast & Environment, Louisiana State University, Baton Rouge, LA, USA

Kjerfve, Björn, School of the Earth, Ocean and Environment, University of South Carolina, Columbia, SC, USA

Krauss, Ken W., Wetland and Aquatic Research Center, U.S. Geological Survey, Lafayette, LA, USA

Lovejoy, Connie, Department of Biology, Laval University, Québec, Canada

Martínez‐Eixarch, Maite, Marine and Continental Waters, IRTA Institute of Agrifood Research and Technology, Sant Carles de la Ràpita, Spain

McGlathery, Karen J., Department of Environmental Sciences, University of Virginia, Charlottesville, VA, USA

Mendelssohn, Irving A., Department of Oceanography and Coastal Sciences, College of the Coast and Environment, Louisiana State University, Baton Rouge, LA, USA

Michael Kemp, W., Horn Point Laboratory, University of Maryland, Center for Environmental Science, Cambridge, MD, USA

Mills, Aaron L., Laboratory of Microbial Ecology, Department of Environmental Sciences, University of Virginia, Charlottesville, VA, USA

Morris, James T., University of South Carolina, Belle W. Baruch Institute for Marine and Coastal Sciences, Columbia, SC USA

Morrison, Elise S., Department of Environmental Engineering Sciences, University of Florida, Gainesville, FL, USA

Nesslage, Geneviève, Chesapeake Biological Laboratory, University of Maryland Center for Environmental Science, Solomons, MD, USA

Nolte, Stefanie, School of Environmental Sciences, University of East Anglia, Norwich, UK; Centre for Environment, Fisheries and Aquaculture Science, Lowestoft, UK

Paerl, Hans W., University of North Carolina at Chapel Hill, Institute of Marine Sciences, Morehead City, NC, USA

Paerl, Ryan W., North Carolina State University, Department of Marine, Earth, and Atmospheric Sciences, Raleigh, NC, USA

Pauly, Daniel, Institute for the Oceans and Fisheries & Department of Zoology, The University of British Columbia, Vancouver, BC, Canada

Pierson, James J., University of Maryland Center for Environmental Science, Horn Point Laboratory, Cambridge, MD, USA

Reyes, Enrique, Department of Biology, East Carolina University, Greenville, NC, USA

Rose, Kenneth A., University of Maryland Center for Environmental Science, Horn Point Laboratory, Cambridge, MD, USA

Rovai, Andre, Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, LA, USA

Rybczyk, John M., Department of Environmental Science, Western Washington University, Bellingham, WA, USA

Snedden, Gregg A., Wetland and Aquatic Research Center, U.S. Geological Survey, Baton Rouge, LA, USA

Stæhr, Peter A., Department of Ecoscience, Marine Diversity and Experimental Ecology, Roskilde, Denmark

Sundbäck, Kristina, Department of Marine Ecology, University of Gothenburg, Göteborg, Sweden

Testa, Jeremy M., Chesapeake Biological Laboratory, University of Maryland Center for Environmental Science, Solomons, Maryland, USA

Twilley, Robert R., Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, LA, USA

Weston, Nathaniel, Department of Geography and the Environment, Villanova University, Villanova, Pennsylvania, USA

Wilson, James G., Zoology Department, Trinity College, Dublin, Ireland

Woodland, Ryan J., Center for Environmental Science, Chesapeake Biological Laboratory, University of Maryland, Solomons, MD, USA

Yáñez‐Arancibia, Alejandro, Red Ambiente y Sustentabilidad, Instituto de Ecologia, A.C. (CPI‐CONACYT), Xalapa, Veracruz, México

About the Companion Website

This book is accompanied by a companion website.

www.wiley.com/go/crump/estuarine3

This website include:

Figures PPTs

MCQs

Short Questions and Answers

CHAPTER 1Introduction to Estuarine Ecology

Kenneth H. Dunton1, Byron C. Crump2, Jeremy M. Testa3, and John W. Day4

1 University of Texas at Austin, Marine Science Institute, Port Aransas, TX, USA

2 College of Earth, Ocean, and Atmospheric Sciences, Oregon State University, Corvallis, OR, USA

3 Chesapeake Biological Laboratory, University of Maryland Center for Environmental Science, Solomons, MD, USA

4 Department of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, LA, USA

Whooping Cranes (Grus americana) along the marsh edge within the Aransas National Wildlife Refuge on the coast of Texas, USA. These endangered native North American birds over‐winter in South Texas and migrate annually to their summer breeding and nesting grounds at Wood Buffalo National Park in northern Canada.

Photo credit: K.H. Dunton.

1.1 Background, Theory, and Issues

We begin this description of estuaries and their functions by defining estuaries very broadly as that portion of the earth’s coastal zone where there is interaction of ocean water, fresh water, land, and atmosphere. Large estuarine zones are most common in low‐relief coastal regions such as the expansive coastal plains of Europe and the east coast of North America. On glaciated coastlines at higher latitudes and on uplifted coastlines such as the Pacific coasts of Asia and the Americas, we refer to these estuarine systems as fjords. We begin our assessment as widely as possible to include all portions of the earth that interact at the edge of the sea and have produced a wide diversity of estuarine types, from coastal plain salt marshes to fjords (Figure 1.1).

From the vantage point of an orbiting satellite, several of the most basic attributes of estuaries are observable. Plumes of sediment‐laden water float seaward on the ocean surface from the largest rivers, such as the Amazon, the Ganges, and the Mississippi. Color differences among various water masses, representing waters of different histories and different biotic richness, are often apparent. Coastal waters in areas with significant riverine input and broad shelf areas generally appear more greenish‐brown than the deep blue waters adjacent to many other coastlines (Figure 1.1a). There are also atmospheric features of importance to estuaries obvious from space. Clouds commonly form directly over the edges of continents as one manifestation of the atmospheric “thermal engine” that maintains the freshwater cycle on which estuaries depend (Figure 1.1d). At the altitude of a satellite, the dense human populations that proliferate in coastal zones are outlined at night by their lights.

The most recent geological epoch, the Holocene, which started approximately 11,650 years before present, could be called the age of the estuary, for estuaries are abundant today even though they may be ephemeral on geologic timescales. It is interesting to note that all of the estuaries discussed in this book did not exist 10,000–15,000 years ago and that they will cease to exist in the near geological future. Many present‐day estuaries are less than about 5000 years old, representing the time since sea level reached near its present level following the last ice age. Since that time, they have progressively filled with sediments and that process will continue. Consequently, our present‐day estuaries will either fill with sediment or will change dramatically as sea level continues to rise.

FIGURE 1.1 Examples of common estuary types across the globe: (a) the Mission‐Aransas coastal plain salt marsh estuary includes seagrass, marsh and mangrove wetlands behind San Jose Island, a barrier island in the Gulf of Mexico, (b) the classic bar‐built estuary of Cedar Bayou (Texas coast) connects Matagorda Bay with Gulf of Mexico waters, (c) the Geiranger Fjord in western Norway, (d) an astronaut view of drowned river valleys Delaware Bay (left) and Chesapeake Bay (center), as well as coastal lagoon Pamlico Sound (top right).

Source: Earth Science and Remote Sensing Unit, NASA Johnson Space Center

(e) barrier island lagoons along the eastern Alaskan Beaufort Sea coast at breakup in June, (f) a deltaic estuary as exemplified by the vast expanses of wetlands of the Mississippi River Delta.

Source: All photos except (d) by K.H. Dunton.

Many estuaries are drowned river valleys (Figure 1.1d). Their formation began as rivers carved their way to the ocean when sea levels were considerably lower. As sea levels rose, the valleys flooded. At high latitudes, river valleys were further eroded by glaciers, resulting in profoundly deep fjords (Figure 1.1c) that became linked to the ocean through glacial melt, and the formation of a shallow entrance sill.

Human populations flourished during this same time period, in no small measure owing to exploitation of the rich estuarine resources of the coastal margin. Most “cradles of civilization” arose in deltaic and lower floodplain areas where natural biota was abundant and where flooding cycles produced the rich bottomland soils and readily available fresh water supplies on which agriculture flourished (Kennett and Kennett 2006; Day et al. 2007). Early centers of civilization that developed in estuarine or deltaic environments include the Tabascan lowlands of Mexico; the valley of the Nile, the Tigris‐Euphrates, Huang He (Yellow), and Indus River deltas, and along the Andean coast of western South America where upwelling systems bordered estuarine systems.

As human populations expanded, so did human pressure on natural environments. Today, we have entered the Anthropocene, a new epoch, defined as the period in which humans have significantly altered Earth’s environments and climate (Syvitski et al. 2020). These changes include excessive nutrient loading of our oceans (eutrophication), global climate change, rapid sea level rise, habitat loss, animal extinctions, and changes in the chemical composition of atmosphere, oceans, and soil.

Let us now continue our aerial survey of estuaries, but this time at a much lower altitude, about 1000 m, in a light airplane following the course of a coastal plain river in the temperate zone from its headwaters to the ocean. The headwater river is narrow with rapids and falls, but changes near the coast to a larger meandering form with broad marshy areas where the actual edge of the river is not always clearly evident. The color of the water changes from clear blue to yellowish‐brown as the river picks up silt. As the river water nears the coast, tidal currents become apparent and, moving seaward, the influence of tidal currents becomes greater and greater.

Along the banks of the estuary, fresh and brackish water marsh plants grow at the edges of embayments. These marshes are often flanked by rows of houses with backyards that border the bank of the estuary, often with narrow piers that extend from the bank to provide access to deeper water. Among these marshes, a variety of wading birds may be observed stalking their prey at the water’s edge. Where the water is shallow and relatively clear, dark‐colored patches indicate the presence of submersed seagrasses.

As we travel seaward, tides become more important, and the intertidal zone becomes more extensive. Larger piers and bulkheads interrupt the banks of the estuary, and brown mud flats come into view, as well as greenish‐gray oyster reefs fringing the banks or dotting the mud flats. Various birds such as oystercatchers feed on the reefs, along with an occasional raccoon. The mud flats are peppered with mud snails, and just beneath the surface are teeming communities of small worms and crustaceans. Common shore birds, such as oystercatchers and skimmers, are feeding at the water’s edge. Skimmers fly along in quiet areas, each plowing a furrow in the water with their lower bill as they fish for silversides and other small fish. The darker colored path of a deep shipping channel maintained by dredging is evident toward the middle of the estuary and contrasts with the lighter colored shallows.

The mouth of the estuary takes the form of a broad sound that opens up behind a barrier island (Figure 1.1b). The sound is shallow, and we can see porpoises herding schools of juvenile menhaden, followed by gulls trying to get in on the action. Crab pot buoys and fishing boats are much in evidence. On either side of the barrier island are narrow passes with visible eddies and strange wave patterns, indicating rapid and complex currents. In high‐latitude estuaries, river flow begins with an enormous flush at ice‐breakup, dispersing ice that formed over the previous 9 months in the estuarine lagoons protected by barrier islands (Figure 1.1e)

Along the ocean beach, several shrimp boats raise long spiraling muddy plumes of sediment as they drag their trawls along the bottom. A kilometer or so offshore of the tidal passes the water changes color from dark brownish green to a lighter, less turbid green. Further offshore it is a darker and bluer color.

On the landward side of most such barrier islands, there are flat intertidal and shallow subtidal areas colonized by salt marsh plants that are bisected with meandering tidal creeks that have developed over centuries (Figure 1.1a). In low‐latitude estuaries, these same areas are colonized with mangrove trees. Moving inland from the marsh, the highest part of the island includes larger trees. The seaward side of the island may include a series of dunes, the farthest from the ocean covered with vegetation, the nearer dunes less and less vegetated. The beach has much less vegetation because wave energy from storms makes it difficult for plants to survive. In parts of the beach–barrier system, vacation houses have replaced dunes and straight navigation channels have replaced twisting tidal channels.

Behind barrier islands, river deltas often form that support enormous wetland areas of salt marsh grasses or mangrove trees. Sometimes these deltaic estuaries have been hydrologically altered by dredging to allow access for commercial activities (Figure 1.1f). In addition to their ecological importance, marsh and mangrove vegetation often form critical natural barriers to storm surges.

In summary, estuaries are complex, dynamic, and biotically rich environments dominated by physical forces and impacted by human activity. Their study requires a consideration and knowledge of geology, hydrology, chemistry, physics, and biology. Ideally, we can integrate knowledge gained through these specific disciplines using what we call systems science, a fundamentally interdisciplinary venture. This book is an introduction to the specifics of estuarine science and its integration into a coherent view of estuaries as ecosystems. We will show how estuaries are different from one another and how they are similar, and why we need to preserve them while enhancing their value to society.

We will begin by describing a very generalized estuary, to provide the reader with an introduction to the geology, physics, chemistry, and biology of estuaries. This is done with a certain danger because, as the rest of the book will show, estuaries are characterized as much by differences as by similarities. Nevertheless, in this chapter, we attempt to describe a generalized estuary. But before we proceed further, we will define an estuary.

1.2 Definitions, Terms, and Objectives

1.2.1 Definitions of Estuary and Ecology, and Difficulties in Applying These Definitions to Estuaries

The term estuary comes from the Latin aestus