Intertidal Ecology

366 Pages·1996·17.012 MB·English

by  David Raffaelli, Stephen Hawkins (auth.)

Intertidal Ecology
Ecology
~ntertidal
,-----
David Raffaelli
Senior Lecturer
Department of Zoology
University of Aberdeen
Aberdeenshire, UK
and
Stephen Hawkins
Professor of Environmental Biology and
Director, Centre for Environmental Sciences
University of Southampton
Southampton
Hampshire, UK
Formerly at Port Erin Marine Laboratory, University of Liverpool,
Isle of Man, UK
KLUWER ACADEMIC PUBLISHERS
DORDRECHT I BOSTON I LONDON
Library of Congress Catalog Card Number: 96-83049
ISBN-13: 978-0-412-29960-5 e-ISBN-13: 978-94-009-1489-6
DOl: 10.1007/978-94-009-1489-6
Published by Kluwer Academic Publishers,
P.O. Box 17, 3300 AA Dordrecht, The Netherlands.
Sold and distributed in North, Central and South America
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In all other countries, sold and distributed
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First edition 1 996
Second edition 1999
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All Rights Reserved
© 1999 Kluwer Academic Publishers
Softcover reprint ofthe hardcover 1st edition 1999
No part of the material protected by this copyright notice may be reproduced or
utilized in any form or by any means, electronic or mechanical,
including photocopying, recording or by any information storage and
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Contents
Preface vii
Acknowledgements
Xl
1 The shore environment: major gradients 1
1.1 Four major environmental gradients 1
1.2 Interactions between gradients 16
1.3 Modifying factors 27
1.4 Comparisons with other ecosystems 32
Overview 35
2 Patterns of distribution 36
2.1 Universal zonation schemes for rocky shores 36
2.2 Rocky shore zonation in different parts of the world 39
2.3 Distribution patterns on other shore types 50
2.4 The meiofauna 61
2.5 Macrophytes on sheltered shores 63
2.6 Zonation along the estuarine gradient 66
Overview 69
3 Causes of zonation 71
3.1 Vertical zonation on rocky shores 71
3.2 Determinants of zonation on other kinds of shores 86
3.3 Causes of zonation along other shore gradients 93
Overview 96
4 Community dynamics 98
4.1 Rocky shores 99
4.2 Sediment shores 121
vi Contents
4.3 Community succession 137
Overview 145
5 Coping with the physical and biological environment 147
5.1 Finding a place to live 147
5.2 Coping with the vertical and horizontal gradients on
rocky shores 151
5.3 Coping with life on sandy shores 159
5.4 Coping by modifying behaviour 164
5.5 Dealing with other organisms 169
5.6 Associations, mutualisms and symbioses 179
Overview 184
6 The shore as a system 185
6.1 The oceanographic context 185
6.2 The flow of material and energy in shore systems 190
6.3 Links between the shore and other systems 206
6.4 Energy flow and functional interactions on shores 211
Overview 213
7 Human impact on the shore 214
7.1 Collecting marine plants and animals for food, bait
and curios 215
7.2 Chronic pollution 220
7.3 Acute effects on shores 227
7.4 Introduction of new species 238
7.5 Coastal zone change 245
Overview 253
8 Studying shores 255
8.1 Describing the shore 256
8.2 General methods for use on both rocky and sandy
shores 265
8.3 Methods applicable to rocky shores 267
8.4 Methods applicable to sediment shores 270
8.5 Long-term studies 274
8.6 Field experiments 275
Overview 283
References 284
Species index 327
Subject index 335
Preface
The seashore has long been the subject of fascination and study - the
Ancient Greek scholar Aristotle made observations and wrote about
Mediterranean sea urchins. The considerable knowledge of what to eat
and where it could be found has been passed down since prehistoric
times by oral tradition in many societies - in Britain it is still unwise to
eat shellfish in months without an 'r' in them.
Over the last three hundred years or so we have seen the
formalization of science and this of course has touched intertidal
ecology. Linnaeus classified specimens collected from the seashore and
many common species (Patella vulgata L., Mytilus edulis L., Littorina
littorea (L.)) bear his imprint because he formally described, named and
catalogued them. Early natural historians described zonation patterns in
the first part of the 19th century (Audouin and Milne-Edwards, 1832),
and the Victorians became avid admirers and collectors of shore animals
and plants with the advent of the new fashion of seaside holidays
(Gosse, 1856; Kingsley, 1856). As science became professionalized
towards the end of the century, marine biologists took advantage of low
tides to gain easy access to marine life for taxonomic work and classical
studies of functional morphology. The first serious studies of the ecology
of the shore were made at this time (e.g. Walton, 1915) at the new
marine biological stations established throughout the world, at first
using a qualitative approach but increasingly using quantitative
methods. The shore has always been a training ground for young
scientists - both for taught courses and for the first steps in research. For
example, Russell, who in the 1930s was instrumental in introducing an
analytical approach in fisheries science, began his career working on
limpets (Russell, 1907).
At the turn of the century, ecologists began to appreciate the
tremendous opportunities the shore provided to test ideas developing in
their very young science. Shore organisms were taken into the
viii Preface
laboratory and experiments carried out (e.g. Herdman, 1890). Just a few
years later, Baker (1909) summarized her culture experiments on
intertidal brown algae in what Connell (1972) called a prophetic passage:
'On the whole it seems as though the greatest competition has been
called into play in the lowest zones, the dry and uncongenial regions of
the upper shore being left to the most tolerant forms, which, if left to
themselves, are able to grow anywhere on the shore'. This neatly
summarized the end product of another 70 years of often contradictory
endeavour!
In the 1920s one of the first British textbooks on shore ecology
appeared (Flatteley and Walton, 1926), with descriptions of zonation
patterns and hints on how to survey shores. Several books aimed at the
general reader (Wilson, 1937; Yonge, 1949), but also jolly useful for
students, were produced. Seaside field courses were well established by
this time and identification guides were beginning to be produced (for
example, in the UK, Eales, 1950; Barratt and Yonge, 1958).
The period between the two world wars saw much research interest
in shore ecology. Classic broads cale studies of rocky shores were
undertaken in this period (e.g., Stephenson, 1936; Ricketts and Calvin,
1939; Stephenson and Stephenson, 1949). In Europe these set the scene
for more detailed quantitative studies of zonation (Colman, 1933) and of
individual species including distribution, population structure,
reproductive cycles and sex change (Orton, 1929; Moore, 1934; Moore
and Kitching, 1939). Similar studies were under way in the United States
(Hewatt, 1935, 1937) and South Africa (Broekhuysen, 1941). The first
experiments and detailed long-term descriptions were made in the 1930s
by pioneering French workers (Fischer-Piette, 1932; Hatton, 1938) and
South African workers (Bokenham, 1938). Fischer-Piette also undertook
broadscale biogeographic surveys on both sides of La Manche (English
Channel) and in France, Spain, Portugal and North Africa.
After World War II the experimental approach gathered steam but
did not attract much attention Ganes, 1948; Lodge, 1948; Southward,
1956; Connell - but not published until the 1960s; Kitching and Ebling,
1967). Quantitative surveys of zonation were made (Southward and
Orton, 1954) and biogeographic studies undertaken (Southward and
Crisp, 1954; Crisp and Southward, 1958; Muus, 1967). Experimental
studies of the ecophysiology and behaviour of shore animals were
pursued (Knight-Jones, 1953; Crisp and Barnes, 1954; Barnes and Barnes,
1957) and the Stephensons inspired broadscale descriptions of zonation
throughout the world (Dakin, 1953; Morton and Miller, 1968;
Stephenson and Stephenson, 1972; also Dahl, 1952 on soft shores). Much
of this work on rocky shores is summarized in the excellent reviews of
Preface ix
Southward (1958) and Lewis (1964), and that for the soft shores in
Eltringham (1971). Lewis (1964) and Stephenson and Stephenson (1972)
mark the zenith of the broadscale qualitative descriptive approach.
Lewis in partkular also asks many questions in his book which were the
subject of study by his group over the next 30 years. Crisp, Southward
and Lewis all anticipated the current interest in 'supply-side ecology' -
but in a jargon-free way.
The 1960s saw renewed interest in ecology sparked by the work of
Connell (1961a,b) and Paine (1966, 1969). They and their kindred spirits
in North America were essentially ecologists using the shore to
experimentally test ecological ideas and theories (e.g. Paine and Vadas,
1969; Dayton, 1971, 1972; Menge, 1976; Peterson, 1977, 1979). This
approach spread rapidly to Australia (Underwood, 1975, 1976a,b, 1978),
New Zealand (Luckens, 1976), Germany (Reise, 1985) and South Africa
(Branch, 1976; Griffiths, 1981).
Both the authors (D.R. and S.J.H.) were research students in the 1970s
when intereslt in shore ecology was re-awakened in Britain (e.g. Lewis,
1976; Hartnoll and Wright, 1977; Hughes, 1980a). This renewed interest
was because shores were amenable to studies of energetics (Wright and
Hartnoll, 1981), of ecophysiology (Foster, 1971a), of animal behaviour
and behavioural ecology (Hughes, 1980b). There was also considerable
debate about the use of shore communities for monitoring pollution in
coastal waters (Lewis, 1976) and in measuring human impacts on shores
(Southward and Southward, 1978), especially estuaries (Knox, 1986). On
soft shores much work was prompted because of their importance as
nursery grounds for fish (McIntyre, 1970; McIntyre and Murison, 1973)
and feeding grounds for birds (Baird et al., 1985). Low-shore and shallow
subtidal kelp beds were also being studied because of their considerable
contribution to nearshore productivity (e.g. Mann, 1973, Field et al.,
1980a).
The use of shores for developing and testing ecological theory has
proceeded apace in the 1980s and 1990s (Reise, 1985; Paine, 1994); their
potential as an ecological laboratory seems limitless. Emphasis remains
on understanding dynamic processes, with ecologists now addressing
interactions between physical oceanography, larval supply and
community dynamics (Gaines et ai., 1985), as well as relationships
between patterns and processes at different spatial and temporal scales
(Giller et al., 1994). These are exciting developments.
Our book is an attempt at interpreting studies on both rocky and
depositing shores, in particular their usefulness in demonstrating and
exploring general ecological principles. However, we do feel that the
coastline and nearshore waters often need to be considered as a single
x Preface
system. The low water is a convenient and arbitrary dividing line at
which we have stopped (with occasional straying) to keep things
manageable and within the experience of most students. Moreover,
most students taking courses in biology do a field course involving shore
work at some time. We hope that pressures on budgets do not further
reduce this admirable tradition.
Chapter 1 outlines the major environmental gradients on the shore
and Chapter 2 describes distribution patterns along these gradients of
tidal height, wave action, sediment and salinity, as in estuaries. Salt
marshes and mangroves are also briefly considered to widen the view.
The rest of the book focuses mainly on rocky and depositing sandy and
muddy shores. Chapter 3 discusses the causes of the distribution
patterns along these gradients, emphasizing the importance of
manipulative field experiments. Chapter 4 explores the factors involved
in structuring shore communities, again emphasizing experimental
approaches. Chapter 5 considers how shore organisms cope with the
physical environment and respond to biological interaction, and
Chapter 6 is concerned with how shores function as ecosystems.
Chapter 7 then widens the scope of the book with an account of human
impacts on the shore. Chapter 8 contains some advice on studying
shores. We hope some of our readers will be inspired to put their wellies
on, get out on the shore and do some experiments!
David Raffaelli and Stephen Hawkins
Acknowledgements
Many people have contributed to and maintained our interest in
intertidal systems over the years, but we are especially grateful to early
guidance from Roger Hughes, Richard Hartnoll, Harry Milne and
George Russell. This interest has been sustained by our own research
students and fellows, all of whom have played a vital role in expanding
our horizons and· keeping us on our toes. We thank them all. The
atmosphere and friendships at Culterty and Port Erin Marine
Laboratory (where a lot of the book was written) are much appreciated.
Many friends and colleagues read chapters and provided much
helpful criticism and literature: Dan Baird, Pat Boaden, Debora Cha,
Bruce Coull, John Crothers, Paul Dayton, Bob Elner, Teresa Fernandes,
Steve Hall, Terry Holt, Roger Hughes, Stuart Jenkins, Sara Lawrence,
Sarah Lawrie, Anton McLachlan, Alasdair McIntyre, Roger Mitchell,
Geoff Moore,. Derek Murison, Trevor Norton, Bob Paine, Keith Probert,
Sarah Proud, Karsten Reise, John Taylor, Richard Thompson, Simon
Thrush, Tony Underwood, Richard Warwick and Bev Wilson. Thanks.
During the last frantic weeks of assembly, the efforts of Sara
Lawrence, Debbie Jones, Mark Williams, Elspeth Jack and Sue Way saw
the book home. Thanks are due to their patience, tolerance and rigorous
checking. Bruce MacGregor produced most of the illustrations. Various
editors at Chapman & Hall who initiated the project, nagged us along
the way and eventually bullied a final manuscript deserve thanks,
especially Clem Earle and Bob Carling. Chuck Hollingworth copy-edited
the final manuscript, and Martin Tribe saw it through production.
Finally, we wish to thank Professor A.J. Southward for his help with
the preface, and to acknowledge the inspiration he has provided over
the last 45 years by his endeavours in both rocky and sediment shore
ecology, amongst many other research interests.