Table Of ContentPlant-animal interactions in Mediterranean-type ecosystems
Tasks for vegetation science 31
SERIES EDITORS
H. Lieth, University of Osnabruck, Germany
H.A. Mooney, Stanford University, Stanford, California, U.S.A.
A. Kratochwil, University of Osnabruck, Germany
The titles published in this series are listed at the end of this volume.
Plant-animal interactions
in Mediterranean-type
ecosystems
Edited by
M. ARIANOUTSOU and R.H. GROVES
Springer Science+Business Media, B.V.
Library of Congress Cataloging-in-Publication Data
Plant-animal interactions in Mediterranean-type ecosystems I editors,
Margarita Arianoutsou, R.H. Groves.
p. cm. -- (Tasks for vegetat i on se i ence ; v. 31>
Papers outgrowth of a conference held at Maleme, Crete, Greece,
Sept. 23-27, 1991.
Inc 1u des index.
ISBN 978-94-010-4392-2 ISBN 978-94-011-0908-6 (eBook)
DOI 10.1007/978-94-011-0908-6
1. Animal-plant relationships--Congresses. 2. Mediterranean
climate--Congresses. 3. Biotic communities--Congresses.
1. Arianoutsou, Margarita. II. Groves, R. H. III. Ser ies: Tasks
for vegetation science ; 31.
QH549.5.P54 1994
574.5'247--dc20 93-6345
ISBN 978-94-010-4392-2
Printed an acid-free paper
AH Rights Reserved
© 1994 Springer Science+Business Media Dordrecht
Originally published by Kluwer Academic Publishers in 1994
Softcover reprint of the hardcover 1s t edition 1994
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
retrieval system, without written permission from the copyright owner.
Table of contents
Preface by F. di Castri VB
Introduction by M. Arianoutsou and R.H. Groves ix
Historical introduction
Chapter 1. C.A. Thanos, Aristotle and Theophrastus on plant-animal interactions 3
Community structure
Chapter 2. R.L. Specht, Species richness of vascular plants and vertebrates in relation to
canopy productivity 15
Chapter 3. J. Blondel and P.e. Dias, Summergreenness, evergreenness and life history
variation in Mediterranean Blue Tits 25
Chapter 4. A. Legakis, Community structure and species richness in the Mediterranean-type
soil fauna 37
Chapter 5. M.L. Cody, Bird diversity within and among Australian heathlands 47
Chapter 6. D.T. Bell, Plant community structure in southwestern Australia and aspects of
herbivory, seed dispersal and pollination 63
Triangular relationships
Chapter 7. H.A. Mooney and R.J. Hobbs, Resource webs in Mediterranean-type climates 73
Chapter 8. B.B. Lamont, Triangular trophic relationships in Mediterranean-climate Western
Australia 83
Herbivory
Chapter 9. N. G. Seligman and A. Perevolotsky, Has intensive grazing by domestic livestock
degraded Mediterranean Basin rangelands? 93
Chapter 10. P.W. Rundel, M.R. Sharifi and A. Gonzalez-Coloma, Resource availability and
herbivory in Larrea tridentata 105
Chapter 11. R. Ginocchio and G. Montenegro, Effects of insect herbivory on plant
architecture 115
v
vi
Pollination
Chapter 12. A. Dafni and C. O'Toole, Pollination syndromes in the Mediterranean:
generalizations and peculiarities 125
Chapter 13. S.D. Johnson and W.J. Bond, Red flowers and butterfly pollination in the
fynbos of South Africa 137
Seed dispersal
Chapter 14. E.J. Moll and B. McKenzie, Modes of dispersal of seeds in the Cape fynbos 151
Chapter 15. R.M. Cowling, S.M. Pierce, W.D. Stock and M. Cocks, Why are there so many
myrmecochorous species in the Cape fynbos? 159
Index of key words 169
Author index 170
Systematic index 176
Preface
The Sixth International Conference on Mediterranean Climate ecosystems was held at Maleme (Crete),
Greece, from September 23 to September 27, 1991. This conference had as its theme 'Plant-Animal
Interactions in Mediterranean-type Ecosystems'. Most of the papers presented to that meeting have
already been published (see Thanos, C.A. ed., 1992, Proceedings of the VI International Conference
on Mediterranean Climate Ecosystems, Athens, 389 pp.). These 57 papers were all necessarily short.
But the theme of plant-animal interactions was considered by the Organizing Committee to be so
important to a fundamental understanding of the ecology of Mediterranean-climate ecosystems and to
an enhanced management ·of those systems that various international research scientists were invited to
prepare longer contributions on major aspects of the overall theme. The Book that follows represents
the result of those invitations. All five regions of Mediterranean climate are represented - Chile,
California, southern Australia and the Cape Province of South Africa, as well as the Mediterranean
Basin itself.
Previous International Conferences on Mediterranean-type Ecosystems have had as their themes
Convergence of Ecosystems and Biota (Chile, 1971), Fire and Fuel Management (California, 1977),
Soil and Nutrients (South Africa, 1980), Resilience (Western Australia, 1984) and Time Scales and
Water Stress (France, 1987). Already, planning for the next such Conference, to be held in Chile in
1994, is under way. The International Society of Mediterranean Ecologists (ISOMED) is an active one;
these Conferences are organized under its aegis. It has been my privilege to be president of ISOMED
for the period between the Fifth and the Sixth conferences. I congratulate the Organizing Committee
for ensuring the undoubted success of the Sixth Conference at Crete, and especially Dr Margarita
Arianoutsou for her role as Chairperson of that Committee. Further, I anticipate with pleasure the
return of the next Conference to South America and to Chile, a country in which I have lived and
worked.
Montpellier, February 1993 Francesco di Castri
VI!
Introduction
The Volume that follows comprises invited contributions on major aspects of Plant-Animal Interactions
in Mediterranean Type Ecosystems, which was the subject of the Sixth International Conference on
Mediterranean Climate Ecosystems, held in Crete (Greece), from September 23 to 27,1991. The subject
of plant-animal interactions was considered fundamental to the process of understanding the structure
and the function of the Mediterranean type ecosystems and to their rational management. An attempt
to present major new contributions on the issues of the overall theme of the Conference, beyond those
already published in the Book of Proceedings (Thanos C.A. ed., 1992), seems warranted; that attempt
follows.
All five regions of the world with a Mediterranean climate are represented: the Mediterranean Basin
sensu lato, California, Chile, South Africa, Australia. Not all of the chapters have adopted the format
of a review paper, although this was the original idea. Some contributors present recent research data,
which they try to incorporate into general patterns, while others perform "challenging" interpretations
to long established perspectives, calling for criticism.
The book is divided into six parts. In the first section, Historical introduction, the author tackles
plant-animal interactions as described in the voluminous works of the founding parents of modern
Biology, Aristotle and Theophrastus, the great philosophers of Greek Classical Antiquity. Besides the
obvious tribute paid to the Conference location, this chapter reveals the important scientific accomplish
ments made by the two men during the era of 384-286 BC. Community structure includes chapters on
the patterns of species richness and diversity of plants as they influence animal communities. As it is
evident from almost all the chapters in this second section, plants that live in characteristic Mediterranean
climates and that have evolved morphological and physiological adaptations, create the habitats to which
the animals respond. The responses of animals to the high levels of temporal and spatial heterogeneity
of these habitats are usually shown by a multiplicity of adaptations of the different groups of animals.
The range of plant-animal interactions is enlarged by the innumerable soil microhabitats, caused either
by the soil per se and/or by its invisible inhabitants. This is the subject of the third section presented
under the title Triangular relationships. Plant morphology, habitat structure and seasonal fluctuations
of available resources define species richness and structure in animal communities. The latter, in
consequence, shape the form and sometimes the function and the vigour of the former. This is mostly
accomplished through their feeding processes, Herbivory, which comprises the fourth part of the book.
It is evident that no one book dealing with plant-animal interactions would be complete unless some
information is presented on the important functions of the plants which are mostly animal-mediated,
namely Pollination and Seed dispersal. These two subjects, forming the fifth and the last sections of the
book respectively, reveal some of the peculiarities of the Mediterranean environment through the
various pollination syndromes observed in plants of the Mediterranean Basin and those of the Cape
fynbos as well as the specific mode of plant dispersal, myrmecochory, observed in the Cape flora.
These six sections on plant-animal interactions reflect the major trends in the direction of research.
It is clear, however, that they do not cover the potential range of such kinds of relationships. Community
structure of soil fauna, rodents and other mammals as affected by plant community structure, fire (and
its direct or indirect effects on this bilateral relation), nutrient cycling and land use, are other issues
that attract the attention of ISOMED scientists. These considerations become increasingly important in
the light of increasing pressure on Mediterranean ecosystems, arising from recent economic develop
ment.
It is our wish that this volume will stimulate new and interesting research on the ecology of Mediter
ranean climate ecosystems. It is also our hope that the results of the research, which is either currently
evolving or is scheduled for the future, will reinforce the movement for the greater conservation of
these ecosystems, which have been one of the places for the origin of culture and science as we now
know them.
Margarita Arianoutsou and Richard Groves, Athens and Montpellier June 1993
ix
PART ONE
Historical introduction
CHAPTER 1
Aristotle and Theophrastus on plant-animal interactions
COSTAS A. THANOS
Institute of General Botany, University of Athens, Athens 15784, Greece
Key words: Aristotle, Theophrastus, zoology, botany, plant-animal interactions, biology
Abstract. Aristotle and Theophrastus, the last great philosophers and scientists of Greek Classical Antiquity, are the founding
fathers of Zoology and Botany, respectively; they should also be honoured as the co-founders of Biology. They were close friends
and life-long collaborators who evidently decided to pursue an organized study of the living world, probably in Lesbos at 344 BC
(the landmark for the creation of the Science of Biology). The product of their division of labour, the voluminous zoological and
botanical works of Aristotle and Theophrastus, respectively, were actually used contemporaneously as university textbooks by
the students of the Lyceum.
Besides numerous comparisons and analogies, mostly on general issues, between animals and plants, both Aristotle and
Theophrastus deal with various cases of plant-animal interactions, covering virtually all aspects of the field. Their scientific
approach is notable, although the barriers to knowledge imposed by their era did not permit a significant contribution on issues
like plant sex and pollination. Their important accomplishments on plant-animal interactions include herbivory and poisonous
plants, plant pests and use of manure, insect-repellence and gall formation, fig caprification and apiculture, seed dispersal and
seed infestation.
Introduction physician at the court of Amyntas C', father of
Philip B'. At the age of 17, Aristotle moved to
Aristotle (384-322 Be) and Theophrastus (371- Athens where he studied and subsequently taught
286 Be) should be considered as the last great in Plato's Academy. Although by far the most
philosophers of Greek Classical Antiquity. They brilliant among Plato's pupils, Aristotle was not
represent, in particular, the culmination of the appointed as the new director of the Academy
natural philosophy of the Ionian scientific tra after Plato's death (347 Be). Apparently as a
dition which was inaugurated on the Aegean result of his non-designation and with a team of
coast of Asia Minor, several centuries earlier, and colleagues and followers, he travelled to Assos of
reached its climax on the opposite coast, at the Troad (Asia Minor) and founded a new school
brightest cradle of Ionian civilization, Athens. It there. Unfortunately, the venture came to an ab
should be borne in mind, however, that both Ari rupt end three years later with the assassination
stotle and Theophrastus were not Ionians but Ma of the patron of the school, Hermeias, the hege
cedonian (Dorian) and Aeolian, respectively; ac mon of Assos.
cordingly, they never became full citizens of It is at that difficult moment in Aristotle's life
Athens. During their lifetime they experienced that Theophrastus seems to have played a promi
the decline of the City-State system of Classical nent role; probably at his suggestion (Morton,
Greece and its eventual replacement by a more 1981), the two men moved to the nearby island
or less unified Greek State dominated by the of Lesbos, in the North Aegean Sea.
northern Greeks (the Macedonians), under the Theophrastus (his original name was Tyrtamos)
leadership of Alexander the Great. was born at the Lesbian town of Eresus and his
Aristotle was born at Stagira of Chalcidice father, Melas was a local fuller. It is probable
(Macedonia) and his father Nicomachus was a that at an early age (ca. 355 Be) he went to
3
M. Arianoutsou and R.H. Groves, Plant-Animal Interactions in Mediterranean-Type Ecosystems, 3-11, 1994.
© 1994 Kluwer Academic Publishers. Printed in the Netherlands.
