Table Of ContentMARINE MAMMALS
BIOLOGY AND CONSERVATION
MARINE MAMMALS
BIOLOGY AND CONSERVATION
Edited by
Peter G. H. Evans
Sea W'I/ell Foundation
Oxford, Eng/and
and
Juan Antonio Raga
Unil'ersity of Valencia
Va/ellcia, Spain
Springer Science+Business Media, LLC
Library of COl1gn:ss Cataloging-in-Publication Data
Marine mammals: biology and conservation/edited by Peter G.H. Evans and Juan
Amonio Raga.
p. em.
Includes bibliograph.ical n:fen:nccs and index.
ISBN 978-0-306-46573-4 ISBN 978-1-4615-0529-7 (eBook)
DOI 10.1007/978--1-4615-0529-7
I. Marine mammals. 2. Wildlife oonservalion. I. Evans. Peler G.
H. [I. Raga. Juan
Amonio.
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Cover phOIO: Atlantic Sponed Do[phins underwater- B. Wiirsig.
Back cover phOIOS from left top: Mating herd of Rorida manalees- Florida Fish and Wildlife
Conservalion Commission. Sea oner bn:aking open a clam-Po Chanin.
and Two adult male New Zealand fur seals lighting- I. Stirling.
Spine photos: Adu!! male walrlls-1. Slirling and Polar bear on icc- I. Stirling.
Cover design by Susanne Van Doyne (Trade Design Group).
ISBN 978-0-3Q6..46573-4
© 2001 Springer Science+Busirn:ss Media New York
Originally published by Kluwer Academic/Plenum Publishers. New York in 2001
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Preface
Interest in marine mammals has increased dramatically in the last few
decades, as evidenced by the number of books, scientific papers, and
conferences devoted to these animals. Nowadays, a conference on marine
mammals can attract between one and two thousand scientists from around
the world. This upsurge of interest has resulted in a body of knowledge
which, in many cases, has identified major conservation problems facing
particular species. At the same time, this knowledge and the associated
activities of environmental organisations have served to introduce marine
mammals to a receptive public, to the extent that they are now perceived by
many as the living icons of biodiversity conservation.
Much of the impetus for the current interest in marine mammal
conservation comes from "Save the Whale" campaigns started in the 1960s
by environmental groups around the world, in response to declining whale
populations after over-exploitation by humans. This public pressure led to
an international moratorium on whaling recommended in 1972 by the
United Nations Conference on the Human Environment in Stockholm,
Sweden, and eventually adopted by the International Whaling Commission
ten years later. This moratorium largely holds sway to this day, and further
protective measures have included the delimitation of extensive areas of the
Indian Ocean (1979) and Southern Ocean (1994) as whale sanctuaries.
The United States was one of the first countries to introduce specific
legislation for marine mammals under the Marine Mammal Protection Act
in 1972. This not only significantly advanced conservation action in that
country, but also stimulated research through the availability of more
substantial resources for scientists. Other countries such as Australia and
New Zealand have followed with their own legislative measures.
In the last three decades of the twentieth century, various global
conventions for environmental protection have been established, including
the MARPOL Agreement (1973/78) for control of marine pollution;
UNCLOS (1982) -the UN Convention on the Law of the Sea; and the Earth
Summit (1992) -Convention on Biological Diversity.
In recent years, conservation problems facing marine species have
driven several coastal nations to develop protective measures. Specific
regional agreements that have been established in Europe include OSP AR -
the Oslo Paris Convention for the Protection of the Marine Environment of
the North-East Atlantic (1992), and, specifically for cetaceans, the
"Agreement on the Conservation of Small Cetaceans of the Baltic and North
Sea" (ASCOBANS) (1992), and the "Agreement on the Conservation of
Cetaceans of the Black Sea, Mediterranean Sea and Contiguous Atlantic
v
VI Marine Mammals: Biology and Conservation
Area" (ACCOBAMS) (1996), within the Bonn Convention -the Convention
on Conservation of Migratory Species of Wild Animals (1979).
Marine mammal science has developed especially in North America.
This is reflected in the formation in 1981 of the Society of Marine
Mammalogy (MMS) in that continent, with members drawn particularly
from the United States. Most text books on marine mammals emanate from
the USA. Ninety percent of contributors to the recent Smithsonian text
"Biology of Marine Mammals" and its companion volume "Conservation
and Management of Marine Mammals" also come from the United States.
Largely due to lack of resources, the study of marine mammals in Europe
has tended to fall behind North America.
However, as a result of the increased interest in scientific and
conservation aspects of marine mammals in Europe, and the need to
disseminate this knowledge in the region, the European Cetacean Society
(ECS) was founded in 1987. This society has progressively consolidated as
an organisation interested in fostering cetacean (and other marine mammal)
research. The ever increasing and enthusiastic attendance of students at ECS
annual meetings raised the need to provide them with specialised up to date
information on marine mammals that is usually absent from most university
courses. For this reason, and with sponsorship from the International
University Menendez Pelayo, the First European Seminar on Marine
Mammals: Biology and Conservation was organised in 1996 in Valencia
(Spain). After this course, the possibility of producing a book for
undergraduate and graduate students was considered, and this eventually
materialised in 1998, following contributions to the Second European
Seminar.
During the 1990s there has also been growing interest in marine
mammals from other parts of the world, particularly South America, Asia,
and Australasia. In 1996, for example, a new society called Sociedad Latino
Americana de Especialistas en Mamiferos Aquaticos (SOLAMAC) was
founded, bringing together marine mammalogists from throughout that
subcontinent, and a society catering for the Pacific region is in the progress
of being formed. We have tried to take account of this in the contributions
to this volume.
Our book is aimed particularly at advanced undergraduate and graduate
students, although it should also be of value to teachers and indeed anyone
with a special interest in marine mammal science. We have tried to provide
a wide range of views and perceptions on the biology and conservation of
marine mammals, but with emphasis upon applied aspects of research given
the pressing conservation problems that marine mammals face at present.
Within the confines of a single book, it is impossible to cover
extensively every subject within marine mammal science. We have
Preface Vll
therefore been selective, opting away from a more comprehensive but
necessarily less detailed book on the subject (for which there is already an
excellent text by Annalisa Berta and James L. Sumich .entitled "Marine
Mammals: Evolutionary Biology", published by Academic Press). Our
emphasis has been on evolution, ecology, behaviour, and various aspects of
marine mammal health and conservation, not only because of the needs for
applied research in those subject areas in this modem world but also
because these have proved to be of particular interest to students. We only
briefly touch on aspects of anatomy and physiology, for example. For
people specialising in those areas, we recommend the relevant chapters in
the recent book "Biology of Marine Mammals", edited by John E. Reynolds
III and Sentiel A. Rommel, and published by the Smithsonian Institution,
USA.
Our book is structured around seventeen chapters, grouped into five
sections. Each section has a short introduction that attempts to provide an
overview of that theme, filling in some gaps, and identifying key literature
references.
The first section is on Life History and Ecology and includes chapters
under the following headings: "Life history strategies of marine mammals"
by Peter Evans and Ian Stirling; "How persistent are marine mammal
habitats in an ocean of variability? - habitat use, home range and site
fidelity in marine mammals" by Arne Bj0rge; and "Ecological aspects of
reproduction of marine mammals" by Christina Lockyer.
The second section on Sensory Systems and Behaviour consists of three
chapters dealing with "Sound and cetaceans" by Jonathan Gordon and Peter
Tyack; "Behavioural ecology of cetaceans" by James Boran, Peter Evans
and Martin Rosen; and "New perspectives on the behavioural ecology of
pinnipeds" by Humberto Luis Cappozzo.
The third section concerns Survey and Study Techniques, including also
three chapters: "The assessment of marine mammal population size and
status" by Philip Hammond; "Acoustic techniques for studying cetaceans"
by Jonathan Gordon and Peter Tyack; and "Applications of molecular data
in cetacean taxonomy and population genetics with special emphasis on
defining species boundaries" by Michel Milinkovitch, Rick LeDuc, Ralph
Tiedemann and Andrew Dizon.
Under the fourth section on Health, Parasites, and Diseases, are three
chapters: "Marine mammal health: holding the balance in an ever-changing
sea" by Joseph Geraci and Valerie Lounsbury; "Living together: the
parasites of marine mammals" by Javier Aznar, Mercedes Fernandez, Juan
Balbuena, and Juan Antonio Raga; and "Mass mortalities in marine
mammals" by Mariano Domingo, Seamus Kennedy and Marie Van
Bressem.
viii Marine Mammals: Biology and Conservation
The fifth and last section focuses upon Conservation and Management,
and features five chapters on "Marine mammals and ecosystems: ecological
and economic interactions" by Enrique Crespo and Martin Hall;
"Environmentalists, fishermen, cetaceans and fish: is there a balance and
can science help to find it?" by Martin Hall and Gregory Donovan;
"Organohalogenated contaminants in marine mammals" by Ailsa Hall;
"Cetaceans and humans: influences of noise" by Bernd Wiirsig and Peter
Evans; and "Global climate change and marine mammals" by Bernd
Wiirsig, Randall Reeves and J.G. Ortega-Ortiz.
We owe a great debt of gratitude to the panel of almost thirty scientists,
drawn from nine countries, who contributed chapters on the specific topics
within their field of interest.
Each chapter has been reviewed additionally by at least two recognised
authorities in its respective field, and for their incisive and helpful
comments, we thank: John R. Baker, Juan Antonio Balbueno, Arne Bj0rge,
John Calambokidis, Philip J. Clapham, Christopher W. Clark, Justin G.
Cooke, James D. Darling, Douglas P. DeMaster, Karen L. Forney, Roger L.
Gentry, Joseph R. Geraci, David 1. Gibson, Eric P. Hoberg, A Rus Hoelzel,
Aleta A Hohn, Vincent Janik, Paul Jepson, Robert D. Kenney, Robin J.
Law, Burney J. Le Boeuf, David J. Marcogliese, Simon P. Northridge,
Gianni Pavan, William F. Perrin, Andrew J. Read, Peter J.H. Reijnders, W.
John Richardson, Randall S. Wells, Koen van Waerebeek, Graham AJ.
Worthy, and Bernd Wiirsig.
Photographs to accompany this book were kindly provided by A
Aguilar, 1. Birks, A Bj0rge, J.-M. Bompar, J. Boran, L. Cappozzo, P.
Chanin, E. Crespo, P. Evans, M. Fedak, J. Geraci, D. Glockner, J.A Gomez,
J. Gordon, B. Haase, A Hall, M. Hall, T. Henningsen, J. Heyning, B. Hicks,
S. Kraus, F. Larsen, V. Lounsbury, J. Moncrieff, P. Morris, K. Norris
Library, R. Pitman, J.A Raga, F. Ritter, W. Rossiter, M. Scheer, E. Secchi,
T. Simila, C. Smeenk, 1. Stirling, N. Tregenza, F. Trujillo, P. Tyack, T.
Walmsley, B. Wilson, J. Wang, S. Wright, B. Wiirsig, K. Young, and E.
Zuniga.
Support for the publication of this book was generously provided by the
Conselleria de Medio Ambiente of Generalitat Valenciana, Ministerio de
Medio Ambiente of Spain, and the University of Valencia.
Finally, we would like to thank Bridget Eichoff for her valuable
assistance in the formatting of chapters, and Joanna Lawrence of Kluwer
AcademiclPlenum Publishers for her continuous help and encouragement
during this book's production.
Peter G.H. Evans and Juan Antonio Raga
Contents
A) INTRODUCTION: LIFE HISTORY & ECOLOGY 1
1. Life history strategies of marine mammals 7
Peter G. H. Evans and Ian Stirling
2. How persistent are marine mammal habitats in an ocean of
variability? 63
Arne Bj¢rge
3. Ecological aspects of reproduction of marine mammals 93
Christina Lockyer
B) SENSORY SYSTEMS AND BEHAVIOUR l33
4. Sound and cetaceans l39
Jonathan Gordon and Peter Tyack
5. Behavioural ecology of cetaceans 197
James R. Boran, Peter G. H. Evans, and Martin 1. Rosen
6. New perspectives on the behavioural ecology of pinnipeds 243
Humberto Luis Cappozzo
C) SURVEY AND STUDY TECHNIQUES 265
7. Assessment of marine mammal population size and status 269
Philip S. Hammond
8. Acoustic techniques for studying cetaceans 293
Jonathan Gordon and Peter Tyack
IX
x Marine Mammals: Biology and Conservation
9. Applications of molecular data in cetacean taxonomy and
population genetics with special emphasis on defining species
boundaries in cetaceans. 325
Michel C. Milinkovitch, Rick LeDuc, Ralph Tiedemann,
and Andrew Dizon
D) HEALTH, PARASITES AND PATHOGENS 361
10. Marine mammal health: Holding the balance in an ever
changing sea 365
Joseph R. Geraci and Valerie Lounsbury
11. Living together: The parasites of marine mammals 385
F. Javier Aznar, Juan A. Balbuena, Mercedes Fem{mdez,
and J. Antonio Raga
12. Marine mammal mass mortalities 425
Mariano Domingo, Seamus Kennedy, and
Marie-Fran~oise Van Bressem
E) CONSERVATION & MANAGEMENT 457
13. Interactions between aquatic mammals and humans in the
context of ecosystem management 463
Enrique A. Crespo and Martin Hall
14. Environmentalists, fishermen, cetaceans and fish: Is there a
balance and can science help to find it? 491
Martin Hall and Gregory P. Donovan
15. Organohalogenated contaminants in marine mammals 523
Ailsa J. Hall
16. Cetaceans and humans: influences of noise 565
Bernd Wiirsig and Peter G. H. Evans
Contents xi
17. Global climate change and marine mammals 589
Bernd Wiirsig, Randall R. Reeves and J. G. Ortega-Ortiz
SYSTEMATIC LIST OF MARINE MAMMALS 609
INDEX 615
