The Ecology of Seeds
How many seeds should a plant produce, and how big should they be? How
often should a plant produce them? Why and how are seeds dispersed, and
what are the implications for the diversity and composition of vegetation?
These are just some of the questions tackled in this wide-ranging review of the
role of seeds in the ecology of plants. The authors bring together information
on the ecological aspects of seed biology, starting with a consideration of
reproductive strategies in seed plants and progressing through the life cycle,
covering seed maturation, dispersal, storage in the soil, dormancy,
germination, seedling establishment and regeneration in the field. The text
encompasses a wide range of concepts of general relevance to plant ecology,
reflecting the central role that the study of seed ecology has played in
elucidating many fundamental aspects of plant community function.
M i c h a e l F e n n e r is a senior lecturer in ecology in the School of Biological
Sciences at the University of Southampton, UK. He is author of Seed Ecology
(1985) and editor of Seeds: The Ecology of Regeneration in Plant Communities,
2nd edition (2000).
K e n T h o m p s o n is a research fellow and honorary senior lecturer in the
Department of Animal and Plant Sciences at the University of Sheffield, UK.
He is author of The Soil Seed Banks of North West Europe (1997) and An Ear to the
Ground: Garden Science for Ordinary Mortals (2003).
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© Cambridge University Press
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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The Ecology
of Seeds
Michael Fenner
University of Southampton, Southampton, UK
Ken Thompson
University of Sheffield, Sheffield, UK
www.cambridge.org
© Cambridge University Press
Cambridge University Press
0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
Frontmatter
More information
p u b l i s h e d b y t h e p r e s s s y n d i c a t e o f t h e u n i v e r s i t y o f c a m b r i d g e
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Ruiz de Alarc´
on 13, 28014 Madrid, Spain
Dock House, The Waterfront, Cape Town 8001, South Africa
http://www.cambridge.org
C
M. Fenner and K. Thompson 2004
This book is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without
the written permission of Cambridge University Press.
First published 2004
Printed in the United Kingdom at the University Press, Cambridge
Typeface Swift 9.5/14 pt.
System L
A
TEX 2
ε [tb]
A catalogue record for this book is available from the British Library
Library of Congress Cataloguing in Publication data
Fenner, Michael, 1949--
The ecology of seeds/by Michael Fenner & Ken Thompson.
p.
cm.
Includes bibliographical references and index.
ISBN 0 521 65311 8 (hardback) -- ISBN 0 521 65368 1 (paperback)
1. Seeds -- Ecology.
2. Plants -- Reproduction.
I. Thompson, Ken, 1954--
II. Title.
QK661.F45
2004
581.4 67 -- dc22
2004045661
ISBN 0 521 65311 8 hardback
ISBN 0 521 65368 1 paperback
The publisher has used its best endeavours to ensure that URLs for external
websites referred to in this book are correct and active at the time of going to
press. However, the publisher has no responsibility for the websites and can make
no guarantee that a site will remain live or that the content is or will remain
appropriate.
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© Cambridge University Press
Cambridge University Press
0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
Frontmatter
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Contents
List of boxes page
viii
Preface
ix
1 Life histories, reproductive strategies and allocation
1
1.1 Sexual vs. asexual reproduction in plants
1
1.2 Life histories and survival schedules
3
1.3 Variability of seed crops
12
1.4 The cost of reproduction
16
1.5 Reproductive allocation and effort
20
1.6 Seed size and number
23
1.7 Phenotypic variation in seed size
29
2 Pre-dispersal hazards
32
2.1 Fruit and seed set
32
2.2 Incomplete pollination
33
2.3 Ovule abortion
34
2.4 Resource limitation
39
2.5 Pre-dispersal seed predation
40
3 Seed dispersal
47
3.1 Wind dispersal
47
3.2 Dispersal by birds and mammals
51
3.3 Myrmecochory
60
3.4 Water and ballistic dispersal
62
3.5 Man, his livestock and machinery
63
3.6 Evolution of dispersal
67
3.7 Some final questions
72
v
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© Cambridge University Press
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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vi
Contents
4 Soil seed banks
76
4.1 Seed banks in practice
78
4.2 Dormancy and seed size
80
4.3 Predicting seed persistence; hard seeds
82
4.4 Seed-bank dynamics
86
4.5 Serotiny
89
4.6 Ecological significance of seed banks
89
5 Seed dormancy
97
5.1 Types of seed dormancy
97
5.2 The function of seed dormancy
98
5.3 Defining dormancy
99
5.4 Microbes and seed dormancy
103
5.5 Effects of parental environment on dormancy
104
6 Germination
110
6.1 Temperature and germination
110
6.2 Responses of seeds to light
116
6.3 Water availability during germination
121
6.4 The soil chemical environment
123
6.5 Effects of climate change
131
7 Post-dispersal hazards
136
7.1 Post-dispersal predation
136
7.2 Loss to pathogens
140
7.3 Fatal germination at depth
141
7.4 Loss of viability with age
143
8 Seedling establishment
145
8.1 Early growth of seedlings
145
8.2 Seedling morphology
146
8.3 Relative growth rate
148
8.4 Seedling mineral requirements
152
8.5 Factors limiting establishment
155
8.6 Mycorrhizal inoculation of seedlings
159
8.7 Facilitation
160
8.8 Plasticity
161
9 Gaps, regeneration and diversity
163
9.1 Gaps, patches and safe sites
163
9.2 ‘Gaps’ difficult to define and detect
164
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- The Ecology of Seeds
Michael Fenner and Ken Thompson
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Contents
vii
9.3 Limitations to recruitment in gaps
167
9.4 Microtopography of soil surface
172
9.5 Gaps and species diversity
176
References
179
Index
241
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© Cambridge University Press
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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Boxes
1.1 Trade-offs
page 4
2.1 Low seed set in sparse populations: the Allee effect
36
3.1 Why do plants have poisonous fruits?
58
3.2 Parent-offspring conflicts in germination and dispersal
74
4.1 Is seed persistence in soil a plant trait?
85
6.1 Response to smoke
129
9.1 Seed traits and plant abundance
169
9.2 Role of leaf litter in regeneration
175
viii
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© Cambridge University Press
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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Preface
In 1985 one of us published a small book called Seed Ecology. It contained
42 000 words and cited 334 references. It was successful in introducing a genera-
tion of ecologists to our subject, but it is now seriously out of date and has been
out of print for some time. The book you are now holding contains 94 000 words
and cites 1117 references. Only a small part of this expansion can be attributed
to covering any part of the subject in more detail; nearly all of it reflects simply
the massive increase in interest in seed ecology in the past 20 years. One sign of
this expansion was the launch in 1991 of the journal Seed Science Research, provid-
ing a major platform for fundamental work in seed biology, including ecology;
37 of our cited references are from that journal. More recently, the International
Society for Seed Science was founded in 2000. This society sponsors meetings on
all aspects of seed science, including, for the first time in 2004, a major inter-
national meeting on seed ecology. Our cited references also reflect this recent
growth: 82% are from the past two decades, while 15% are post-1999.
Recent work in this field has transformed our understanding of many aspects
of seed ecology, especially dispersal, storage in the soil and the ecological role
of seed dormancy. There has also been increasing recognition that regeneration
from seed has fundamental impacts on the diversity and composition of plant
communities; seed ecology has never seemed more relevant to ‘mainstream’
plant ecology. Nevertheless, we have not lost sight of our debt to the pioneers
who laid the foundations for later work. Many of the most important figures in
the history of ecology, including Darwin, Harper and Salisbury, made significant
contributions to seed ecology.
In this book, we attempt to synthesize the current information available
on the ecological aspects of seed biology, starting with a consideration of
reproductive strategies in seed plants and the costs and compromises involved.
Special attention here is given to the interesting topic of seed size. The text
then follows the progress of seeds through the stages of their life cycle in a
ix
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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x
Preface
roughly chronological sequence: seed maturation, dispersal, storage in the soil,
dormancy, germination and seedling establishment. The final chapter gives an
account of the role that canopy gaps play in the regeneration of plants in the
field. Throughout the book, various specialized topics (which might otherwise
have interrupted the flow of the text) are presented in self-contained boxes.
We aimed to present a broadly representative overview of the current lit-
erature rather than a comprehensive review of it. We have tried to present a
reasonably balanced account of the field in a way that reflects current thinking.
Nevertheless, where we have strong feelings on particular topics, we have not
refrained from nailing our colours to the mast. When this happens, for exam-
ple concerning the definition of dormancy, we hope you find our arguments
convincing.
We hope that this text will be useful to students of plant ecology at all levels.
The regeneration of plants from seed involves a very wide range of ecological
concepts of current interest, from reproductive strategies to the maintenance of
species diversity. Pollination, seed dispersal and seed predation all offer inter-
esting insights into the evolution of plant--animal interactions. The numerous
trade-offs encountered (e.g. between seed size and number, early reduced repro-
duction vs. delayed increased fecundity, early high-risk germination vs. delayed
safer germination, etc.) offer scope for theoretical investigations and modelling.
We hope that the work reported here from the literature will stimulate students
into devising their own experimental investigations. Excellent undergraduate
projects on seed ecology can often be carried out in the field with a minimum
of technical resources.
Several colleagues provided substantially new figures or new versions of pub-
lished figures. We would particularly like to thank Costas Thanos, Mary Leck
and Bego˜
na Peco. Otherwise, the text is entirely our own work. We can there-
fore assert cheerfully that any errors and omissions are ours alone.
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© Cambridge University Press
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0521653681
- The Ecology of Seeds
Michael Fenner and Ken Thompson
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