Principles of Dispersal in Higher Plants

225 Pages·1982·5.319 MB·English

by  Dr. Leendert van der Pijl (auth.)

L. van der Pijl
Principles of Dispersal
in Higher Plants
Third Revised and Expanded Edition
With 30 Figures
Springer-Verlag
Berlin Heidelberg New York 1982
Dr. LEENDERT VAN DER PUL
Emeritus Professor of Botany
University ofIndonesia and
University ofNijmegen
Sportlaan 236
The Hague/Netherlands
The cover motif shows water burrs with the double function of
transport (via attachment to animals) and of establishment (anchor
ing to the substrate).
ISBN-13: 978-3-642-87927-2 e-ISBN-13: 978-3-642-87925-8
DOl: 10.1007/978-3-642-87925-8
Library of Congress Cataloging in Publication Data. Pijl, L. van der (Leendert van der),
1905-. Principles of dispersal in higher plants. Bibliography: p. Includes index. l.
Seeds-Dispersal. 2. Botany-Ecology. 1. Title. QK929.P5 1982 582'.0467 82-848
AACR2.
This work is subject to copyright. All rights are reserved, whether the whole or part of
the material is concerned, specifically those of translation, reprinting, re-use of illustra
tions, broadcasting, reproduction by photocopying machine or similar means, and
storage in data banks.
Under § 54 of the German Copyright Law where copies are made for other than private
use a fee is payable to "Verwertungsgesellschaft Wort", Munich.
© by Springer- Verlag Berlin Heidelberg 1969, 1972 and 1982
Softcover reprint of the hardcover 3rd edition 1982
The use of registered names, trademarks, etc. in this publication does not imply, even in
the absence of a specific statement, that such names are exempt from the relevant pro
tective laws and regulations and therefore free for general use.
Typesetting, printing and bookbinding: Konrad Triltsch, Graphischer Betrieb,
Wtirzburg.
2131/3130-543210
Preface to the Third Edition
The second edition was a hasty one, prepared in hospital only with
the help of amendations on typed slips. The present revision has
been more thorough, so that contents and references were almost
doubled compared to the first edition.
Even more emphasis has been laid on ecology (including syne
cology), evolution and establishment after transport. The temptation
was great to present these points under a new title, which would also
have avoided being quoted with data (as being the ultimate wisdom)
from the 1968 edition sitting in the local library. The publisher,
though very liberal in satisfying needs, is, however, at the same time
very moral and conscientious in his marketing methods, which
means maintaining the original title. This though he, too, profits
from selling one's "principles".
I regret the time did not allow inclusion of the results of the
Symposium on Dispersal and Distribution, Hamburg, 10-12 June
1981.
It seems polite to react to criticisms by some reviewers:
A. On the loss of readability by the numerous cross-references:
The very compressed text of a textbook should be studied, not just
read.
An ecosystem is a web with many internal connections - and so
is an ecological book, where aspects from different chapters have to
be interconnected for coherence without repetition in each.
B. On complaints that an author's index and a glossary are lack
ing: Such an index would only be necessary in a more historically
orientated review - and a glossary for terms is superfluous as the
subject - index refers to definitions special to our field.
The Hague, Spring 1982 L. VAN DER PIJL
Preface to the First Edition
The work offered here is a companion volume to the work by
K. FAEGRl and L. VAN DER PUL, Principles of Pollination Ecology,
which deals with the preceding phase ofreproduction in plants.
In the present work too, the emphasis is on principles and ecolo
gy. It is neither an enumeration of mechanisms, nor a compilation of
cases. RIDLEY'S monumental work The Dispersal of Plants Through
out the World comprises 700 large pages of small print, and research
has proceeded since then. Though this work is more than just a
compilation and contains much insight and thoughts on principles
in addition to reviews, its completeness hinders its use as a textbook.
As a reference work, it is unsurpassed and the writer made frequent
use of it.
The writer paid special attention to functional backgrounds for
the use of taxonomists working with "characters" and to biosystem
atics at the macro level.
He is indebted to Dr. P. MULLER-SCHNEIDER (Chur, Switzer
land) for the permission to translate parts of his Verbreitungsbiologie
der Bliitenpjlanzen - of which permission a modest use has been
made. Thanks are also due to the Director of the Rijksherbarium at
Leyden, and to its librarian for the use of the library.
Mr. F. J. NATAN was so kind as to take a number of photographs
at the author's request.
Prof. Dr. H. F. LINSKENS stimulated the work actively and made
completion possible with the collaboration of the members of his
staff at Nijmegen University, amongst whom Miss I. DE Roos should
be thanked especially.
Prof. Dr. B. J. D. MEEUSE (Seattle) was of great service, criticiz
ing the manuscript.
The Hague, Summer 1968 L. VAN DER PUL
Contents
I. Introduction . . . . . . . . . . . . . . .
A. The Place of Dispersal in the Chain of Life
B. Limitations and Objections . . 3
C. History and General Literature 4
II. General Terminology . . . . . . . . . . . . . . . . 6
III. The Units of Dispersal ............ 10
Vegetative Parts in Dispersal and False Vivipary 11
IV. The Relation Between Flowers, Seeds and Fruits 15
A. Seed and Fruit . . . . . . . . . . . . . 15
B. Morphological Fruit Systems . . . . . . . 16
C. Morphological Interaction Between Fruit and Flower 16
l. General. . . . . . . . . . 16
2. Position. . . . . . . . . . 18
3. Monovuly and Monospermy 19
4. Inferiority and the Calyx . . 19
D. Inadequacy of Current Fruit Terminology 20
V. Ecological Dispersal Classes, Established on the Basis
of the Dispersing Agents 22
A. General. . . . . . . . 23
B. Invertebrates . . . . . 23
C. Fishes and Ichthyochory 23
D. Reptiles and Saurochory 25
E. Birds and Ornithochory 28
1. Epizoochory by Birds 28
2. Synzoochorous Bird Diaspores 29
3. Endozoochory. . . . . . 31
Non-adapted Diaspores. . . 31
Adapted Diaspores. . . . . 33
The Syndrome of Bird Diaspores 35
Oil-containing Fruits . . 38
Remarks on Evolution . 39
Mimesis (Imitative Seeds) 40
VIII Contents
F. Mammals and Mammaliochory 45
l. General. ....... . 45
2. Dyszoochory and Rodents 45
3. Accidental Endozoochory . 46
4. Adaptive Endozoochory 47
Ungulates ...... . 48
Bats and Chiropterochory 49
Primates . . ... 53
Various Mammals . 54
G. Ants and Myrmecochory 55
H. Wind and Anemochory 60
l. General. ... 60
2. Dust Diaspores . . . 63
3. Balloons . . . . . . 64
4. Plumed (Comose) Diaspores 64
5. Winged Diaspores . . . . . 65
6. Tumbleweeds . . . . . . . 67
7. Wind-Ballists (Anemoballists) 69
J. Water and Hydrochory ... . 70
l. General. ........ . 70
2. Rain Wash (Ombrohydrochory) 71
3. Rain-Ballists. . . . . . . . . 72
4. Submerged Transport in Water 73
5. Floating Diaspores. . . . . . 74
K. Epizoochory, Transport on the Outside of Animals
in General . . . 78
1. Diverse Origins 78
2. Trample Burrs. 79
3. Water Burrs. . 81
4. Burrs and Other Adhesives Above Ground Level 81
5. Other Spiny Fruits. . . . . . . . 82
L. Autochory, Dispersal by the Plant Itself 83
1. General. . . . 83
2. Active Ballists. . . 84
3. Passive Ballists 86
4. Creeping Diaspores 87
M. Barochory, Dispersal by Weight Only 89
N. Retrospective View . . . . . . . 90
VI. Dispersal Strategy and the Biocoenosis 91
A. Atelochory . . . 91
l. General. . . . 91
2. Synaptospermy 92
3. Basicarpy . 94
4. Geocarpy. . . 94
Contents IX
B. Polychory and Attendant Phenomena 96
1. General. . . . 96
2. Heterodiaspory . . . . . . . . 98
3. Tachyspory . . . . . . . . . . 100
C. Concluding Remarks on Synecology 101
1. Deserts. . . . 10 1
2. The Rain Forest 102
3. Epiphytes. . 106
4. The Arctic . . 107
5. Island Floras . 107
6. Plant Sociology and Dispersal 112
7. Coordinated Dispersal 113
VII. Establishment 115
A. General. 115
B. Fixation . 116
C. Vivipary . 117
D. Germination 120
1. General Importance 120
2. Span of Life and Dormancy 121
3. Influence of Dispersing Agents and Other Stimuli 122
VIII. The Evolution of Dispersal Organs in General 125
A. Aims. . . . . . . . . . . . . . . . . 125
B. Isosporous Pteridophytes . . . . . . . . 125
C. Heterosporous Pteridophytes with Free Megaspores 125
D. Pteridosperms. . . . . . . . . . 128
E. Gymnosperms (or Pre-Angiosperms) 130
F. Angiosperms . . . . . . . . . . 132
1. The Seed . . . . . . . . . . . 132
2. The Seed Escaped from Angiospermy 133
3. The Sarcotesta Maintained in Conventional Fruits 135
4. Arilloids . . . . 138
5. Pulpa. . . . . . . 143
6. The Pericarp Fruit . 145
Shift of Function 145
Autonomous Cycles 147
Further Evolutionary Influences and Processes 147
IX. Ecological Developments in Leguminous Fruits 150
X. Dispersal and the Evolution of Grasses 160
A. Comparison with Cyperaceae . . 162
B. Return to Gramineae (i.e. Oryzeae) 163
C. Bam busoid Grasses . . . . . . . 166
x
Contents
D. Bamboos ................ . 168
E. Open Plains and Higher Grasses . . . . . . . 169
F. Some More Remarks on Awns and on Establishment 176
G. Retrospective Views . . . . . . . . . 180
XI. Man and His Plants in Relation to Dispersal 183
References. . . . . . . . . 188
Subject Index . . . . . . . 201
Index of Scientific Plant Names 206
Index of Scientific Animal Names 215
I. Introduction
A. The Place of Dispersal in the Chain of Life
Microbiologists' concepts are seldom concerned with dispersal and areas of dis
tribution. One of their rules, sometimes indicated as "Beijerinck's law", states:
everything is everywhere, but the milieu (environment) selects. In microbiologi
cal terms, this means that a special substrate can demonstrate the presence, and
promote the development, of certain microbes specialized for that substrate.
Such lower organisms are evidently so easily spread that they are in principle not
limited by the dispersal factor. Some fungi and mosses requiring special sub
strates (e.g. dung) have advanced beyond this point, developing devices for
directed transport of spores to preferred substrates; and in the higher plants, in
which spores have lost their general function of dispersal, the same purpose is
served by the microspores which are brought to the stigma. This directed trans
port of microspores is dealt with in pollination ecology. In higher plants, coloniz
ing dispersal is a factor limiting distribution more severely.
In this book, we have to study the ways and means which the higher plants
employ to reach, with their newly developed dispersal organs, sites where a new
generation can be established. More specifically, this involves the methods used
to keep their descendants separated in space and to provide each with its own
site, where it can compete with other plants; it also concerns the methods em
ployed to defend the future of the species by exploring new territories, par
ticularly following climatic fluctuations, or conversely to maintain a foothold on
a favourable site.
Another feature of spores, the power to withstand and survive unfavourable
conditions over a long timespan, is also evident in the reproductive organs of
higher plants. The products of these organs, such as seeds, may combine the ad
vantages of dormancy with those of genetic variation, in contrast to vegetative
resting organs, which merely continue the life of the individual without hope for
the future of the species in a changing environment.
In the present study, we shall have the opportunity to emphasize two dif
ferent aspects, viz. actual dispersal as studied in the field, and the structural basis
needed to attain this dispersal; both will be considered in an ecological context.
All too often, the second aspect has predominated so strongly in works on dis
persal that they remain examples of herbarium ecology, or worse, writing-desk
ecology. Nevertheless, the structural aspect cannot be dismissed as the starting
point, the more so since a mere enumeration of findings in certain sites (as by
Heintze, 1932) produces an unmanageable chaos.