J. Pan & P.D. Pratolongo 2022: Marine biology.

2022·29.8 MB·English

Front cover: Oral disc and tentacles of the sea anemone Bunodosoma zamponii, from an intertidal pool. The sea
lettuce, Ulva (Enteromorpha) compressa, is seen in the upper background.
Photo credit: Jerónimo Pan
First edition published 2022
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Library of Congress Cataloging‑in‑Publication Data
Names: Pan, Jerónimo, editor. | Pratolongo, Paula D., editor.
Title: Marine biology : a functional approach to the oceans and their
organisms / Jerónimo Pan, Paula D. Pratolongo.
Other titles: Marine science series.
Description: First edition. | Boca Raton, FL : CRC Press, 2021. | Series:
Marine science | Includes bibliographical references and index.
Identifiers: LCCN 2021028238 | ISBN 9780367024987 (hardcover)
Subjects: LCSH: Marine biology.
Classification: LCC QH91 M25 2021 | DDC 578.77--dc23
LC record available at https://lccn.loc.gov/2021028238
ISBN: 978-0-367-02498-7 (hbk)
ISBN: 978-1-032-19390-8 (pbk)
ISBN: 978-0-429-39924-4 (ebk)
DOI: 10.1201/9780429399244
Typeset in Times New Roman
by Radiant Productions
Series: Marine Science
Series Editor: Andrés Arias
Volume 1
Marine Biology
A Functional Approach to the Oceans and
their Organisms
Editors
Jerónimo Pan
National Scientific and Technical Research Council (CONICET)
Instituto de Geología de Costas y del Cuaternario (IGCyC)
Instituto de Investigaciones Marinas y Costeras (IIMyC)
Mar del Plata, Argentina
Paula D. Pratolongo
Centro de Recursos Naturales Renovables de la Zona Semiárida (CERZOS-CONICET-UNS)
Dto. Biología, Bioquímica y Farmacia-Universidad Nacional del Sur (BByF-UNS)
National Scientific and Technical Research Council (CONICET)
Bahía Blanca, Argentina
p,
p,
A SCIENCE PUBLISHERS BOOK
A SCIENCE PUBLISHERS BOOK
Preface to the Series
Covering more than 70% of the Earth, the oceans regulate our environment including the climate,
water purification, carbon storage and primary production. In addition, they also provide provisioning
and cultural services, including harvestable goods (fish, shellfish and seaweed), mangroves timber,
algae, minerals, recreation, beauty and plenty of cultural benefits. At present, the human population
is reaching 8000 million while about half of the planet’s suitable land has already been cultivated for
crops, exacerbating climate change, biodiversity loss and compromising other ecosystems services. In
this scenario, our oceans, which hold 97 per cent of the Earth’s water, emerge as the immediate supply
answer, not only in the food production sector, but also as a source of energy, biomaterials and even
biochemicals/pharmaceuticals.
In these times, when global attention is focusing on oceans and their role in present and future human
development, we are humbled to deliver this Marine Book Series through Taylor & Francis, aiming to
cover all aspects of oceans and marine research. The present series will provide state-of-the-art coverage
of important topics in marine biology and biodiversity, marine ecology, marine biotechnology, marine
bioproducts, marine bioprocessing and marine chemistry. We believe this series will be of interest to
specialists and nonspecialists alike as well as to scientists and managers engaged in the study of marine
biosphere, addressing natural phenomena occurring at coasts, wetlands, estuaries, continental slopes,
continental rises, abyssal regions, mid-ocean ridges, along with neritic and pelagic waters of the ocean.
We are pleased to deliver the Marine Series Collection starting point “Marine Biology: A Functional
Approach to the Oceans and their Organisms”, edited by Dr. Jerónimo Pan and Dr. Paula Pratolongo,
which includes contributions from an impressive group of international scientists which will bring a
comprehensive analysis ranging from the ocean fundamentals to the coupling and functionalities of
different ecosystems—from primary producers to the oceanic megafauna. This first volume will provide
a well-rounded perspective of the ocean essentials at a glance, including a reader-friendly style that does
not lose any scientific accuracy.
Dr. Andrés H. Arias
Preface
Why a functional approach to marine biology?
Upon opening this book, you are about to embark on a trip into the world’s oceans that may lead you from
a familiar sandy beach to the far reaches of the open ocean and the deep sea, on a fascinating journey
studying marine life and how living creatures, from giant whales to tiny microbes, interact with the
environment in which they thrive. There are many possible approaches to the study of a certain subject in
the life sciences, and when it comes to a complex, interdisciplinary subject such as Marine Biology, the
approaches can be even more challenging. Biology has traditionally put an emphasis on structure over
function. Taxonomy and systematics have played a significant role in the development of Marine Biology
as a discipline, especially in the 19th and 20th century, following the Linnean tradition established amidst
the Romantic Naturalism movement.1
Are Marine Biology and Biological Oceanography the same thing? No. Marine Biology is the
study of the biological aspects of marine organisms, meaning that the focus is put in the biota itself
rather than the marine environment which they inhabit. On the other hand, Biological Oceanography
is the branch within the science of oceanography that focuses on the processes mediated by the biota,
which have implications for the oceans. Traditionally, scientists who go on cruises look into distribution
patterns of biota and relate them to environmental properties are considered biological oceanographers,
and those who explore the biological functioning of organisms, e.g., conducting laboratory experiments,
consider themselves marine biologists. Are these two disciplines too far apart from each other? No, not
necessarily. In fact, they only differ in their focuses, while they have many points of contact and overlap.
As it has been proposed, the combination of the two disciplinary approaches briefly sketched above leads
to a better understanding of the ecology of the oceans.2 In that sense, this book tries to “reconcile” (for
lack of a better term) these two seemingly different disciplines, in the sense that we believe the study of
marine organisms may be incomplete, and even limited, without considering them in the context of the
environment that they thrive in, and how they interact and modify it through their metabolic, behavioral
and reproductive processes.
Undoubtedly, the 20th century saw the advent and consolidation of Ecology as a new discipline in
the biological sciences, drawing much of its conceptual framework from experimental developments
in marine systems (e.g., island biogeography experiment by MacArthur and Wilson,3 the concept of
keystone species and bottom-up vs. top-down controls elegantly introduced by Robert Paine4 through
rocky-shore communities). Can we separate the study of marine organisms from their environment?
No. The study of Marine Biology necessarily requires an ecological approach; it is even implicit in the
very term (Marine refers to the environment, Biology refers to the life that thrives in and alters such
environment). Remember that the term ecology was coined by the German zoologist Ernst Haeckel,
after the Greek term οἶκος, meaning “house”. That is enough reason for us to have primed an ecological
approach to the treatment of each chapter in this book.
1 Larson, J.L. 1967. Goethe and Linnæus. J. Hist. Ideas 28(4): 590–596.
2 Kiørboe, T. 2008. A Mechanistic Approach to Plankton Ecology. Princeton University Press, Princeton, NJ. 209 pp.
3 MacArthur, R.H. and E.O. Wilson. 1967. The Theory of Island Biogeography. Princeton University Press, Princeton, NJ.
203 pp.
4 Payne, R.T. 1966. Food web complexity and species diversity. Am. Nat. 100: 65–75.
vi Marine Biology: A Functional Approach to the Oceans and their Organisms
So why focus on function over structure? In its early stages, the blooming science of Ecology heavily
relied on the characterization of communities from a structural point of view; the Braun-Blanquet cover-
abundance scale5 for the analysis of plant communities is perhaps the most remarkable example of such
disciplinary approach, and this approach was imprinted into many ecological studies throughout the 20th
century. More recently, the “ecosystem service” framework, introduced some 15 years ago,6 emphasized
the (economic) value that the natural functioning of ecosystems provides not only to the biosphere, but
also to mankind. It has since been successfully adopted in academic and environmental management
circles. What would be the economic costs of clearing a tidal prism of suspended particles in an enclosed
embayment if we should need to resort to engineering devices rather than relying on benthic suspension
feeders? On a closer look, the ecosystem service framework is inherently linked to ecosystem functioning
rather than structure. The Anthropocene has witnessed many changes in the way biogeochemical processes
and rates work at a planetary scale. As such, it is the functioning of marine ecosystems that is being
compromised.7 The case of an invading seagrass in the Caribbean Sea, mentioned in Chapter 8, is one
example of how ecosystem functioning may be altered, while the structure might remain quite the same.
The field of Marine Biology certainly is not new; in fact, there are some excellent textbooks on the
matter that have become classics over the decades since their publication. For our book, we have strived
to seek the contributions of renowned and up-and-coming professionals in the different fields of Marine
Biology, aiming to produce a fresh text for the new audiences of students and young academics. This was
done in our hopes to deliver consolidated knowledge and contribute new perspectives to the young actors
that will make a difference in the way we study, make use and preserve our oceans in the future. This
book was conceived as an introduction to Marine Biology, for a new generation of scientists, teachers
and academics interested in the world’s oceans. Of course there are many remarkable Marine Biology
textbooks out there, which paved the way for the enlargement of this discipline, and we hope this book
measures up to the legacy of renowned authors who conceived those wonderful books.
The book is organized into three sections, namely, an introductory part separated into two chapters
and providing a general background to the marine environment; a mid-section consisting of 10 Chapters,
which delve into the functional biodiversity of marine organisms (this represents the core of the book);
and finally two closing chapters that focus on current topics of special interest such as climate change.
We hope to have produced a high-quality text that will appeal both to upper-level undergraduate and
first-year graduate students, professors teaching those courses, and members of the scientific community.
For such endeavor, we have taken special care contacting renowned specialists on different topics, which
is reflected in the diversity of institutions around the world that are represented among the authors
included in this book. With contributions from authors from North and South America, Europe and Japan,
we made a conscious effort as editors for the book to be appealing to a broad audience of readers from
around the world. We are thankful to the authors who produced high-quality material, mostly based on
primary literature sources (i.e., publications from specific scientific journals), covering examples from all
major ocean basins and coastal seas on Earth. We are also thankful to those researchers who contributed
original photographs and illustrations to enrich the text (credits are given where appropriate).
This is a peer-reviewed book, meaning that all chapters have been evaluated by experts in each
field, on the basis of their scientific quality, innovative approaches and potential contributions to each
sub-discipline within Marine Biology. Accordingly, this preface would not be complete without the
acknowledgement of the superb job carried out by the many reviewers who contributed their expertise
and insight to the improvement of each chapter and the book as a whole. They come from broad areas of
academia, scientific research, conservation charities, NGOs, and environmental business and technology.
They are:
N. Aberle-Malzahn: Department of Biology, Norwegian University of Science and Technology
M. Ahrens: Universidad de Bogotá Jorge Tadeo Lozano
5 Braun-Blanquet, J. 1932. Plant Sociology (Transl. G.D. Fuller and H.S. Conrad). McGraw-Hill, New York. 539 pp.
6 Millenium Ecosystem Assessment. 2005. Ecosystems and Human Well-Being: Biodiversity Synthesis. World Resources
Institute, Washington DC.
7 Margalef, R. 1963. On certain unifying principles in ecology. Am. Nat. 97: 357–374.
Preface vii
J.M. Bedmar: Argentinian Geological Service (SEGEMAR)
C.F. Berghoff: National Institute for Fisheries Research and Development - Argentina (INIDEP)
G.S. Blanco: Marine Organismal Biology Institute (IBIOMAR)
P.E. Bourdeau: Humboldt State University
G. Cavieres P.: Center of Applied Ecology and Sustainability, Pontificia Universidad Católica de Chile
G. Cepeda: National Institute for Fisheries Research and Development - Argentina (INIDEP) & IIMyC
R.R. Christian: East Carolina University
G. Giardino: Marine and Coastal Research Institute (IIMyC)
S. Gonçalves Leles: University of Southern California
A. Jordi: Jupiter Intelligence
R. McBride: NOAA Fisheries, Northeast Fisheries Science Center
A.C. Morandini: Center for Marine Biology (CEBIMar) & Zoology Dept., Universidade de São Paulo
V. Reyes: Whale and Dolphin Conservation UK & Fundación Cethus
A.F. Rosenthal: Asociación Naturalistas Geselinos-NGO
V. Segura: INIDEP
I. Sunesen: CONICET, Phycology Division at La Plata Museum
Some others have opted to remain anonymous. Through their contributions, they have made this
book much more sound and robust; of course they are not liable for any inaccuracies and errors that may
remain in the text.
Last but not the least, we would like to express our gratitude to our families and support systems,
without whose help the road leading up to the completion of this project would have been even longer and
more winding. JP would especially like to thank his twins, Lena and Braulio, for being understanding,
encouraging and patient whenever their Daddy had to take time off to work on this book, and also for
being the best help on field trips. PP wants to mention the inspiring questions of her son Ulises, a 10-year-
old eager explorer of the beautiful oceans.
The final stages of writing and editing for this book took place during the COVID-19 pandemic,
a time when most of humankind halted their daily lives and had to take a step aside. During this time,
a deep collective conscience on the impacts we have created on our planet started to arise, and many
people, perhaps millions who have never given a thought to such matters, noticed their magnitude and
importance for the first time. Ours is a pivotal time for the care and preservation of the Earth’s oceans,
and we hope this is collectively embraced among other new paradigms; as such we hope to have made a
small contribution with this book.
February 2021 Jerónimo Pan
Paula D. Pratolongo
Mar del Plata & Bahía Blanca

Contents
Preface to the Series iii
Preface v
Section I: Introduction to the Marine Environment
1. Introduction to the Marine Environment from a Geological Perspective 3
Paula D. Pratolongo, Jerónimo Pan and Diana G. Cuadrado
2. Introduction to the Marine Environment from Physical and Chemical Perspectives 21
Paula D. Pratolongo and Jerónimo Pan
Section II: Functional Biodiversity of Marine Organisms
3. Marine Phytoplankton Diversity and Ecology 43
Jerónimo Pan
4. Planktonic Metabolism and Primary Production in the Oceans 72
Aurore Regaudie‑de‑Gioux and Aurea M. Ciotti
5. Microzooplankton: Functionally Diverse Protists in the Pelagic Realm 88
María Celeste López‑Abbate
6. Zooplankton: The Ocean Drifters 113
Danilo Luis Calliari Cuadro, Anabela Anahí Berasategui and María Clara Menéndez
7. Gelatinous Zooplankton 150
Miodeli Nogueira Júnior, Everton Giachini Tosetto, Letícia Cazarin Baldoni, Sofia Dutto,
Mitsuko Hidaka, Dhugal J. Lindsay and Renato Mitsuo Nagata
8. Soft-bottom Marine Benthos 180
Jerónimo Pan and Paula D. Pratolongo
9. Salt Marshes and Mangroves: Tidal Saline Wetlands Dominated by Vascular Plants 211
Paula D. Pratolongo
10. Marine Hard Substrate Communities 232
Jorge L. Gutiérrez, Sally D. Hacker, Martin A. Coombes, Christian Wild,
Guilherme H. Pereira‑Filho and M. Gabriela Palomo
x Marine Biology: A Functional Approach to the Oceans and their Organisms
11. Fish Ecology 274
Hannes Baumann
12. Marine Megafauna: Sea Turtles, Seabirds and Marine Mammals 297
Rocío Mariano Jelicich, Paula Berón, Sofía Copello, Natalia A. Dellabianca, Germán García,
Vanessa Labrada-Martagón, M. Natalia Paso Viola, Jesica A. Paz, Luciana Riccialdelli,
Analía San Martin, Juan Pablo Seco Pon, Mónica A. Torres and Marco Favero
Section III: Topics of Current Interest
13. Thermal Plasticity in Marine Organisms 327
Santiago Salinas and Nishad Jayasundara
14. Climate Variability and Change, their Impacts on Marine Life and Ecosystems 349
Jerónimo Pan and Reinaldo A. Maenza
Index 375
About the Editors 383