Table Of ContentSuccession in Abandoned Fields
Geobotany 15
Series Editor
M.1.A. WERGER
Succession in Abandoned Fields
Studies in Central Bohemia, Czechoslovakia
edited by
1. OSBORNOvA
M. KovAROvA
J. LEPS
K. PRACH
KLUWER ACADEMIC PUBLISHERS
DORDRECHTI BOSTON I LONDON
Library of Congress Cataloging-in-Publication Data
Succession in abandoned fields studies in central Bohemia,
Czechos I ovak i a I ed i ted by J. Osbornov,; ... [et a 1.l.
p. em. -- iGeobotany 15)
BiblIography, p.
Inc I udes Index.
1. Eco log i ca I success i on--Czechos I ovak i a--Bohe. ia. I. Osbornov,;,
J. II. Series.
QH 178 . C8S83 1990
581 .5' 26404' 094371 --dc20 89-15 407
ISBN-13: 978-94-010-7603-6 e-ISBN-13: 978-94-009-2444-4
DOl: 10.1007/978-94-009-2444-4
Published by Kluwer Academic Publishers,
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Table of contents
List of contributors vii
Foreword by Marcel Rejmanek ix
Preface xv
1. Introduction
by J. Leps and K. Praeh 1
2. General characteristics of the region
by J. Osbornova and H. Rambouskova 3
3. Abandoned fields in the region 7
3.1 General characteristics
by A. Klaudisova and 1. Osbornova 7
3.2 Intensively studied fields
by J. Osbornova 10
3.3 Principal plant species of the studied fields
by J. Osbornova 11
4. Dynamics of populations and communities 23
4.1 Changes in vegetation during succession 23
4.1.1 Plant populations
n
~Kfu~
(Vegetation dynamics in the newly abandoned field by J. LepS)
4.1.2 Life histories of principal plant populations, including their aUelopathic interferences
by L. Soukupova 32
4.1.3 Horizontal structure
byJ. Lept 38
4.1.4 Successional and seasonal changes in biomass and production
by J. Osbornova and K. Praeh 42
4.2 Heterotrophic organisms 53
4.2.1 Soil micromycetes
by O. Fassatiova 53
4.2.2 Collembola and other microarthropods
by J. Rusek 56
4.2.3 Small mammals
by J. KoneCny 58
5. Selected ecosystem processes and functions 61
5.1 Microclimate
M
~Kfu~
vi
5.2 Water balance 62
5.2.1 Soil moisture, transpiration, water saturation deficit and sublethal deficit
by H. RambouskoVii 62
5.2.2 Root water potential of dominants
by J. Slavikova 68
5.3 Mineral nutrients 69
5.3.1 General soil chemical characteristics
by M. Kovtifova 69
5.3.2 Seasonal dynamics of phosphorus
by N. Sojkova 72
5.3.3 Dynamics of potassium, calcium and sodium
by M. Krkavcova 77
5.4 Decomposition and release of nutrients
by V. Hadincova, M. Kovafova and J. Pelikanova 80
6. Reaction to perturbations 93
6.1 Mowing
by H. Koblihova-Baumova 94
6.2 Vehicle perturbation
by O. Rauch 97
6.3 Nutrient additives 101
6.3.1 Effects of phosphorus addition on interactions between the plant dominants
by M. Kovafova and E. Brabec 101
6.3.2 Phosphorus leaching experiments
by M. Kovtifova 105
6.3.3 Potassium additives
by M. Krkavcova 110
6.4 Fly ash application
by O. Rauch and V. Hadincova 112
6.5 Effect of herbicides
by 1. Michalek, V. Burianek and V. Hadincova 116
7. Synthesis 127
7.1 Vegetational dynamics
by K. Prach 127
7.2 Water and nutrient economy
by M. Kovtifova 134
7.3 Stability
byJ. Leps 143
7.4 Abandoned fields in the landscape
by H. Rambouskova 148
8. Summary
by K. Prach and 1. Leps 151
Appendix 153
References 159
Subject index 167
List of contributors
(all from Czechoslovakia)
BRABEC Eduard, Institute of Botany, Czechosl. Acad. Sci., CS-252 43 Pnihonice u Prahy
BURIANEK Vac1av, Research Institute of Forestry and Game Management, Jiloviste-Strnady,
CS-255 01 Zbraslav
FASSATIOVA Olga, Department of Botany, Fac. Sci., Charles University, Benatska 2, CS-128 01 Praha
HADINCOVA Vera, Institute of Botany, Czechosl. Acad. Sci., CS-252 43 Prilhonice u Prahy
KLAUDISOvA Alexandra, Institute for Monument Protection and Nature Conservancy, Valdstejnske
nam. 1, CS-110 00 Praha
KOBLIHOvA-BAUMOvA Helena, Institute of Botany, Czechosl. Acad. Sci., CS-252 43 Prilhonice u
Prahy
KONECNY Josef, Obrancil miru 63, CS-170 00 Praha
KovAROvA Marcela, Institute of Botany, Czechosl. Acad. Sci., CS-252 43 Prilhonice u Prahy
KRKAVCovA Miloslava, Karlovarska 5, CS-160 00 Praha
LEPS Jan, Department of Biomathematics, BioI. Res. Centre, Czechosl. Acad. Sci., Branisovska 31,
CS-370 05 Ceske Budejovice
MICHALEK Jaroslav, Regional Museum, Zamecka 1, CS-356 00 Sokolov
OSBORNOvA Jana, Department of Botany, Fac. Sci., Charles University, Benatska 2, CS-128 01 Praha
PELIKANOvA Jifina, Institute of Experimental Botany, Czechosl. Acad. Sci., Branisovska 31,
CS-370 05 Ceske Budejovice
PRACH Karel, Institute of Botany, Czechosl. Acad. Sci., Dukelska 145, CS-379 82 Tfeboii
RAMBOUSKOvA Hana, Institute of Landscape Ecology, Czechosl. Acad. Sci., Mala plynarnf 2,
CS-170 00 Praha
RAUCH Ota, Institute of Landscape Ecology, Czechosl. Acad. Sci., Na sadkach 702, CS-370 05 Ceske
Budejovice
RUSEK Josef, Institute of Soil Biology, Czechosl. Acad. Sci., Na sadkach 702, CS-370 05 Ceske
Budejovice
SLAVIKOvA Jifina, Department of Botany, Fac. Sci., Charles University, Benatska 2, CS-128 01 Praha
SOJKOVA Nadezda, Institute of Experimental Botany, Czechosl. Acad. Sci., Ke dvoru 16,
CS-166 30 Praha
SOUKupovA Lenka, Institute of Botany, Czechosl. Acad. Sci., Dukelska 145, CS-379 82 Tfeboii
Foreword
Old and New Fields of Old-Field Ecology
In ecology, succession occupies a place similar to
that of evolution in general biology.
Ram6n Margalef, 1968.
It was a great honor for me to have been asked by Marinus Werger to write an introductory note to this very
special volume. Presumably my friends and former students in Europe felt that a few words from the New
World might put the results presented in this exciting book into a somewhat broader perspective. My
perspective (or retrospective), however, is neither impersonal nor original; it is an eclectic reflection of
recent developments in ecology and in old-field ecology in particular.
The ecological generalizations and theories of Ram6n Margalef and Eugene P. Odum, as we perceived
them in Prague in the early 1970s, were for some of us so attractive and promising that we even started to
believe it would not take too long until we had a truly unifying general theory of ecological succession. All
that was needed - we thought - were data clarifying a few controversial issues. This is how our studies of
old-field succession began in 1973. We viewed old-fields as a sort of 'Drosophila' for terrestrial ecology.
What was our starting point? (1) We were enthusiastic about the succession-production-diversity-stability
oriented 'new' ecology represented by seminal publications of Margalef (1968, 1969), Whittaker (1970),
Odum (1971), and McNaughton (1974). (2) We had some fresh experience with achievements and short
comings of the International Biological Program actively carried on in Czechoslovakia - even though
succession was never really addressed in this program. (3) We were eager to apply the quantitative methods
of Greig-Smith (1964) and Kershaw (1973). (4) Trained in Braun-Blanquet phytosociology, we had an
intimate knowledge of the central European flora (note 323 species of vascular plants used for ordination in
Chap. 3.1). Unfortunately, we did not have enough understanding for conducting adequate experiments
with competing plant popUlations; only a few ecologist outside of England, Japan, and Holland had such
understanding at that time and, as a matter of fact, satisfactory methods as we understand them now were
available only in Japan (Suehiro and Ogawa 1980, Rejmanek et al. 1989). Also, we had only limited
equipment for more demanding ecophysiological studies.
The timing of our intensive field research and subsequent data analyses coincided with a substantial shift
from the 'new' ecology of the 1960s and 1970s to the 'new-new' and 'pluralistic' ecology of the 1980s
(McIntosh 1987). Specific answers to specific questions have replaced, to a large extend, the hope for a
'universal' ecological theory. Odum (1989) and Margalef's (1989) recent generalizations about succession are
rather cautious. Several specific theories of succession have emerged in the meantime (Grime 1977, Connell
& Slatyer 1977, Gorham et al. 1979, Shugart 1984, Tilman 1985, Walker & Chapin 1987) and experimental
microcosms proved to be a useful tool for testing of specific hypotheses (Van Voris et al. 1980, Grime et al.
1987). The following pages mirror this transition.
The present volume is a result of many inevitable compromises between our intentions and what was
practically possible. Some deductions about temporal trends from studies of different aged fields ('space-for-
J. Osbomova et aI. (eds), Succession in Abandoned Fields. ix-xiii.
© 1990 Kluwer Academic Publishers, Dordrecht-
x
time substitution', Pickett 1989) were not without problems (and we were not ready to wait 50 years!). A
promising study of nitrogen cycling was interrupted by the tragic death of our student Marcela Kavkova in a
car accident. Some of our experimental plots were destroyed several times by tractors carrying garbage to a
neighbouring illegal dump (a partial compensation for this damage was a 'vehicle perturbation experiment'
as described in Chap. 6.2). After a few years, there simply was not space for more experimental plots in some
of our old-fields. Only a few phenomena studied in the old-fields were simultaneously studied in an arable
field. I am sure that everybody who has ever tried to organize and conduct similar research will understand.
Although the first detailed description of old-field succession was published from the Annaberg area in
Austria (Stossner 1859, see Clements 1928), an impressive number of studies have been conducted in the
United States during this century and readers of Ecology, Oecologia, or Journal of Ecology may easily get an
incorrect impression that old-field succession is an exclusive domain of American ecology. However, many
interesting studies have been conducted also in other countries including Japan (e.g., Hayashi 1977,
Matumura et al. 1988), German Federal Republic (Bornkamm 1984, Schmidt 1986), France (Lepart &
Escarre 1983), Italy (Ubaldi 1976), Sweden (Olson 1984), Finland (references in this volume), Poland
(references in this volume), Mexico (Purata 1986), Venezuela (Saldarriaga 1987), the Soviet Union (Tuga
naev & Pestereva 1976, Cibanova 1982), Malaysia (Kochummen & Ng 1977), and Czechoslovakia (refer
ences in this volume). A bibliography of publications from all countries (Rejmanek 1989a) was used for
completion of the 'time period/phenomena studied' summary presented in Table 1, which illustrates major
research trends in old-field ecology.
Description of changes in species composition represents a backbone of successional studies. An increased
number of publications from the present decade seems to be due to an increasing amount of data available
from permanent plots and the application of several new multivariate methods.
Many studies have also been dedicated to the understanding of relationships between species diversity and
productivity. As results presented in this volume show, successional changes of these two characteristics may
be rather complicated and dependent on the position on a moisture gradient.
Research in plant population dynamics in a broad sense is mainly a matter of the last two decades. Specific
and exciting hypotheses in this area have been tested in old-fields in recent years by influential research
schools of Fakhri A. Bazzaz, Reinhard Bornkamm, Wolfgang Schmidt, David Tilman, and Patricia A.
Werner (e.g., Bazzaz 1987; Parrish & Bazzaz 1982; Bornkamm 1984; Bornkamm & Schrade 1981; Schmidt
1981, 1983, 1986; Inouye & Tilman 1988; Tilman 1988; Gross & Werner 1982; Goldberg & Gross 1988).
Plant ecophysiology, biogeochemistry, and all types of experimental perturbation studies in old-fields
have also been flourishing in the last two decades (e.g., Bazzaz 1979, Vitousek 1983, Bekelaar & Odum 1978,
Wakefield & Barrett 1979, Goldberg 1987, Carson & Pickett 1989). The strength of the present volume lies
especially in studies in successional changes of phosphorus biogeochemistry and in evaluations of several
perturbation experiments.
Surprisingly few data are available on successional changes of microclimate and horizontal structure in
old-fields. Another surprise is an almost complete lack of mathematical models of old-field succession. The
present volume certainly contributes to the filling of these gaps (Table 1). The role of herbivory, pathogens,
and mycorrhizae in old-field succession (Penfound 1964, Gibson et al. 1987, Brown et al. 1988, Kirkpatrick
and Bazzaz 1979, Crowell & Boerner 1988) is much less understood than in several other types of succession.
In spite of a growing interest in the resistance of different successional stages to invasions of alien species
(Hanks 1972, Inouye et al. 1987, Rejmanek 1989b), species introduction experiments in old-fields (Squiers &
WistendahI1977, Gross & Werner 1982, Oldfather 1984) are much less common than species removal
experiments. Unfortunately, these topics, so sparsely represented in the world literature, were not touched
in the course of our studies either (Table 1). The effect of the old-field area on the rate and pattern of
colonization was investigated only once (Davis & Cantlon 1969) and is now studied systematically in Kansas
Xl
(Simmons 1989). There is also only rudimentary information about changes in genetic structure of some
old-field populations (Hancock 1977, Hancock & Wilson 1976).
Only rigorously designed comparisons of simultaneous old-field successions in many areas and diverse
environments can bring more understanding of which trends are general and which are site specific.
Comparative analysis of old-field successions in England and America (Brown et al. 1987, Hendrix et al.
1988) represents a first step in this direction. However, based on our experience, the major confounding
factor in such studies - beside differences in site histories (use of herbicides, fertilizers, etc.) - will be the
quality of and distance from a source of propagules of species belonging to successionaly more advanced
ecosystems (colonization potential). This distance is absolutely crucial and determines a continuum of
situations which undoubtedly exists between Whittaker's (1975) 'direct' and 'replacement' successions. I can
only hope that the present volume will encourage research in old-field succession on much larger scales than
those we were able to handle.
It is certainly very useful to have substantial results from this type of team research published in one
volume. What is included here could be easily divided into at least 20 papers spread over more than five
journals. Such fragmentation would have resulted in redundancy, lack of focus, and lost opportunity for
integration and synthesis. The present volume, as well as the recently published volume by Falinski (1986) in
the same series, clearly show how important long-term, stable (even if small) funding is for studies on
ecological succession.
Table 1. The number of publications presenting results from studies in old-field ecology over the period 1910-1988. Altogether 518
publications are included, some of them falling under two or more categories. The last column indicates the strength of emphasis of
studies in this volume.
Phenomena studied and approaches used Period This volume
1910 '31 '41 '51 '61 '71 '81
1930 '40 '50 '60 '70 'SO '88
Changes in species composition 8 11 17 24 23 31 57 +++
Soil properties 3 3 5 4 12 14 ++
Plant demography/population dynamics/dispersal 2 3 7 16 37 38 ++
Seed banks 2 3 3 3 ++
Biomass/production, energy flow 3 7 12 12 +++
Plant ecophysiology 4 5 17 14 +
Allelopathy 2 8 14 4 +
Microclimate 2 3 ++
Population structure of communities (diversity, dominance, etc.) 3 18 16 +++
Horizontal structure (pattern and interspecific associations) 2 3 +++
Vertical structure 2 2
Nutrient movementibiogeochemical cycles 11 13 +++
Effects of experimental fertilization 8 7 +
Species removal experiments 4 6 +
Species introduction experiments 1 3
Other types of experimental perturbation (herbicides, radiation, 4 6 9 +++
mowing, trampling, etc.)
Influences of natural perturbations (drought, fire, etc.) 4 4 5 +
Direct studies of competition 3 4 10
Phenology and pollination 3 4
Role of herbivory 2 8
Role of pathogens 1
Role of mycorrhizae 2
Mathematical modeling of succession 2 ++
Description:Old and New Fields of Old-Field Ecology In ecology, succession occupies a place similar to that of evolution in general biology. Ram6n Margalef, 1968. It was a great honor for me to have been asked by Marinus Werger to write an introductory note to this very special volume. Presumably my friends and
