Table Of ContentPLANT SPECIES AND PLANT COMMUNITIES
PLANT SPECIES
AND
PLANT COMMUNITIES
Proceedings of the International Symposium held at Nijmegen, November 11-12, 1976
in honour of
Professor Dr. Victor Westhoff
on the occasion of his sixtieth birthday
edited by
Eddy van der Maarel & Marinus J. A. Werger
Dr. W. Junk bv Publishers The Hague - Boston - London 1978
ISBN-13: 978-90-6193-591-9 e-ISBN- 13: 978-94-009-9987-9
DOl: 10.1007/978-94-009-9987-9
© Dr. W. Junk bv Publishers 1978
No part of this book may be reproduced and/or published in any form, by print, photoprint, microfilm or any other means
without written permission from the publishers.
This Symposium was sponsored by the Faculty of Science of the University of Nijmegen.
The articles in this book have also been published in our periodical Vegetatio (see page 177).
Prof. Dr. Victor WesthofT
CONTENTS
Editorial note .............................................................................................................................. ...... IX
E. van der Maarel, Opening address ....................................................................................................... .
M. J. Adriani, Personal word to Victor Westhoff ................... .................. ... ....... ...... ....................... ........... 2
M. J. Adriani & E. van der Maarel, Plant species and plant communities: An introduction ....... ............... ...... ..... 3
J. Braun-Blanquet, Die Quellflur-Gesellschaft des Cratoneuro-Arabidetum bellidifoliae (Koch 1928) in der subalpinen
Stufe Graubiindens..... ... ............ .. ...... ....... ... ..... ........................ ............. .. .... .. ................ .. .... ............. 7
J. H. Willems, Observations on North-West European limestone grassland communities: Phytosociological and
ecological notes on chalk grasslands of Southern England ........................................................................ 11
Martin Meyer, Vergleich verschiedener Chrysopogon gryllus-reicher Trockenwiesen des insubrischen Klimabereiches
und angrenzender Gebiete ... , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Michael B. Dale, Planning an adaptive numerical classification..................................................................... 29
C. H. Gimingham, Cal/una and its associated species: Some aspects of co-existence in communities....................... 35
W. G. Beeftink, M. C. Daane, W. de Munck & J. Nieuwenhuize, Aspects of population dynamics in Halimione
portulacoides communities..... ......... .. ............ ...... ............ ..................... ....... .. .... .. ....... .. .... .. .... ......... .. .. 43
Nils Maimer, Lennart Lindgren & Stefan Persson, Vegetational succession in a South Swedish deciduous wood............ 55
Rudolf Hundt, Untersuchungen zur Entwicklung von Geholz-Aufforstungen auf Bergbaukippen in der Diibener
Heide (DDR) .............................................................................................................................. ... 69
G. Sissingh, Optimal woodland development on sandy soils in the Netherlands .. ............... .......................... ..... 81
A. O. Horvat, Die Bedeutung des Klimas fUr die Zusammensetzung der Vegetation SW-Ungarns, des Elsasz und der
Umgebung von Brianc;on, Alpes Maritimes. ... .............. ............... ............. ......... ..................... ................ 87
A. W. H. Damman, Geographical changes in the vegetation pattern of raised bogs in the Bay of Fundy region of
Maine and New Brunswick................................................................................................................. 91
Elias Landolt, The importance of closely related taxa for the delimitation of phytosociological units.................... ....... 107
Wolfgang Holzner, Weed species and weed communities.............................................................................. 119
a
Jean-Marie Gehu, Le concept de sigmassociation et son application l'ettlde du paysage vegetal des falaises atlantiques
franc;aises . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
A. H. P. Stumpel & J. T. R. Kalkhoven, A vegetation map of the Netherlands, based on the relationship between
ecotopes and types of potential natural vegetation............................................. ...................................... 137
J. Lebrun, Applications phytosociologiques a I'amenagement du territoire ...................................................... 149
Sandro Pignatti, Evolutionary trends in Mediterranean flora and vegetation......... ............... .................. ........... 157
M. 1. A. Wergner & E. van der Maarel, Plant species and plant communities: Some conclusions........................... 169
Reference to original publications in VEGETATIO .................................................................................... 177
List of unpublished contributions to the symposium.................................................................................... 178
Editorial note
This volume is not just a liber amicorum, a Festschrift. It is a report on a Symposium, a being together of friends, in this
case friends of Professor Dr. Victor Westhoff. We thought that such an event would do justice to the extraordinary personality
of Westhoff. And so we not only discussed scientific matters, on a theme that has enjoyed Westhorrs interest for a long time,
but we also had food together, made fun, made music and read poetry. The special atmosphere of these few days cannot be
described appropriately in this book.
We still have some things to say. First of all we were very much pleased, almost embarrassed by the tremendous response
we received from almost all Dutch friends, pupils and colleagues and from the selection of foreign friends we approached
with the idea of the symposium. This response made it necessary to ask some of the potential contributors to content them
selves with poster stands or with the presidency of one of the four scientific sessions. We are very grateful indeed to all these
colleagues and friends for their enthusiasm and readiness to contribute to the symposium in whatever way.
Secondly we enjoyed the effort of our publishers. Most contributions presented at the symposium or displayed at the poster
stands have been published in VEGETATIO in the meantime; they are collected in this volume. Only a few contributions
have not been included according to the authors' wishes; these were either summaries of previously published work, or
preliminary reports considered not yet suitable for publication.
Dr. W. Junk Publishers are gratefully acknowledged for their rapid cooperation in collating the available papers in this
volume.
Thirdly, we have to thank the Director of the Faculty of Science, Dr. C. J. M. Aarts, for his immediate agreement and
support, Prof. dr. ir. J. J. Steggerda, chairman of the Faculty's Board, who kindly addressed Prof. Westhoff at the symposium
dinner, and Prof. dr. H. F. Linskens, Director of the Botanical Laboratory, for his help with the initiation of the programme.
A very special word of thanks we would like to say to Peter Toll, LL.D., of the Faculty's Directorate who took care of the
general organisation of the symposium and the festivities involved in a both cheerful and very effective way! Through him
we also thank the many Faculty employees who contributed in various ways to the success of the event. Without their help
the symposium would not have been possible at all!
Finally we wish to express our feelings of gratitude to staff and students of our own department of Geobotany, especially
to Mrs. A. van de Zand-Barten. Without their continuous help the symposium would not have become what it still is in our
own memory: an unforgettable feast!
The editors
Eddy van der Maarel
Marinus J. A. Werger
IX
Plant Species and Plant Communities
OPENING ADDRESS
Ladies and Gentlemen,
Professor and Mrs. Westhoff: Dear Victor and Nettie: Prof. P. van der Veken, Ghent, and Prof. G. Wendelberger,
I welcome all of you at this special symposium in honour Vienna. We are happy indeed that most of them are on
of a colleague who is to celebrate his 60th birthday and who the list of speakers, demonstrators and symposium presi
occupies a very special place in our life as a plant ecologist, dents.
vegetation scientist or (Westhoffs preference!) geobo Further, I may welcome two colleagues from abroad
tanist. At the same time we have come together in a rela who have become more or less 'nijmegenised', Dr. Mike
tively quite atmosphere (what we really do too seldom) to Dale, St. Lucia/Brisbane and Prof. Wolfgang Holzner,
listen to lectures and to discuss, both publicly and privately, Vienna, both guest lecturers at the Department of Geo
a theme that is of special concern to our 'hero of the feast' botany here.
Victor Westhoff: Plant species and plant communities. I would like to add those Dutch friends who are on the
Dear Victor, this symposium was planned as a surprise. list of speakers: Prof. Jan Barkman, Dr. Wim Beeftink,
However, one of the personal characteristics you excel in Dr. Geerd Sissingh and last, but not least, Dr. Marcus
is your natural curiosity. Therefore, I presume that you Adriani, our nestor and talented speaker, who will present
have been more or less aware of what is going to happen the introductory lecture. *
here! We, your colleagues and friends, wish to express Finally, I must mention some distinguished foreign
our acknowledgements for your achievement as an eminent colleagues who would have liked to come but were unable
geobotanist and a remarkable human being. We do this by to attend the symposium: Dr. D. Ranwell, Norwich, Prof.
organising this symposium for you. S. Rivas-Martinez, Madrid, Prof. H. Sukopp, Berlin and
Our feelings towards you are clearly demonstrated by Prof. R.H. Whittaker, Cornell University.
the number of participants, which is well over 150. It At the ena of my opening speech I cannot resist taking
would go too far to mention all of them by name, but you the opportunity to say some personal words to you, Victor,
will agree with my wish to make some exceptions, especially as a token of our twenty years of friendship. I do this 'short
for the colleagues from abroad. and snappy' by connecting your name with that of your
First of all, I welcome Prof. J. Lebrun, Universite de main teacher and great example: Josias Braun-Blanquet.
Louvain-La-Neuve, president of our International Society He is still active, as you know, and I am happy to announce
for Vegetation Science. Monsieur Lebrun: soyez Ie bienvenu a small paper of his for this symposium. In a recent paper
ici! in Vegetatio on the Braun-Blanquet approach in perspe.;ti\e
Then I would like to welcome Dr. E. Duffey from the I made a comparison between the history of vegetation
Institute of Terrestrial Ecology (Nature Conservancy) science and the history of music and there. I dared to
at Monks Wood, one of our outstanding colleagues in compare Braun-Blanquet's significance for phytosociology
nature conservation and vice-president of the British with that of Johann Sebastian Bach for music. Well, Victor,
Ecological Society, that old and famous organisation, I feel it appropriate now to continue along these lines and
which nominated Victor Westhoff an honorary member to connect your name with that of the musical genious you
earlier this year. admire so much and even rank above all others: Wolfgang
From the wide circle of University colleagues, professors Amadeus Mozart.
of Geobotany or its equivalents, I may welcome the With the wish that it will be both serious and cheerful,
following: Prof. J. M. Gehu, Lille, Prof. C. H. Gimingham,
Aberdeen, Prof. R. Hundt, Halle, Prof. G. J ahn, Gottingen,
* The scientific part of this lecture will be placed at the beginning
Prof. E. Landolt, Zurich, Prof. N. Maimer, Lund, Prof. of the series of contributions, whilst the personal words Adriani
J.J. Moore S.J., Dublin, Profs. S. and E. Pignatti, Trieste, added will follow after these opening words.
both harmonious and melodious, I open this symposium!
Eddy van der Maarel
Jeuillez me permettre, Mme et M. Westhoff, mesdames et
messieurs, d' ajouter encore deux mots en fran9ais. Tout
a
d' abord pour vous feIiciter, tous les deux, I'occasion de
votre anniversaire remarquable: voyez cequi se p<'sse au
jour d'hui: regardons cequi se passera demain! Nous
vous admirons, M. Westhoff, vous, I' homme de la science
geobotanique, I' homme qui a demerites. II y a une grande
reconnaissance de la part de vos amis et de vos eleves:
c' est beaucoup et c' est tres serieux. II y a presque trop:
les deuxjours du symposion ne suffisent pas. Mais quand
meme: In der Beschrankung zeigt sich der Meister!
a
Votre amitie m' est tres precieuse cause de votre
inspiration continue pour les conceptions scientifiques,
par votre inspiration pour les grandes responsabilites
des chercheurs.
Quel deIice de regarder ensemble des tableaux, des
dessins, des gravures, d' ecouter ensemble la musique
d' un Scarlatti, d'un Franz Schubert, d' un Cesar Frank,
de se dire qu' un Paul Verlaine, 'comme poete a touche
aux profondeurs de la joie et de la misere de I' homme. Et
ce tout, cette totalite, Victor, pour vous impregne par votre
sens inne d' un holisme. Mes felicitations cordiales pour
vous deux, Victor et Nettie!
Marcus J. Adriani
2
PLANT SPECIES AND PLANT COMMUNITIES: AN INTRODUCTION·
M. J. ADRIANI! & E. VAN DER MAAREU
! Van Itersonlaan 50, Oostvoorne, The Netherlands
2Division ofGeobotany, Toernooiveld, Nijmegen, The Netherlands
This symposium is dealing with various relations between (ii) Amongst the species that make up the floristic compo
plant species as basic systematic units and plant communi sition of a community, some are more sensitive expressions
ties as basic vegetation units. It will thereby touch upon of a given relationship than others. For practical classifi
relations between the respective ecological disciplines cation (and indication of environment) the approach seeks
as well, disciplines we used to call autecology and syneco to use those species whose ecological relationships make
logy. This twofold theme has been a favourite one for a them most effective indicators; these are diagnostic species
long time to the colleague and friend whose 60th birthday (Character-species, differential-species, and constant com
we are celebrating: Victor Westhoff. When we take his panions).
thesis in 1947, on the vegetation of dunes and salt marshes (iii) Diagnostic species are used to organize communities
on the Dutch islands of Terschelling, Vlieland and Texel, into a hierarchical classification of which the association
as a starting point, Westhoffs ecological endeavour spans is the basic unit. The vast information with which phyto
a period of 30 years. sociologists deal must, of necessity, be thus organized;
Throughout this period he has approached plants and and the hierarchy is not merely necessary but invaluable
plant communities in his very personal way: both causal for the understanding and communication of community
and final. His 'Leitmotiv' can be. expressed by Goethe's relationships that it makes possible.
words 'Wie fass ich Dich, unendliche Natur", which we Although it is believed here and there that this threefold
find as a motto for a section of the book 'Inleiding tot de essence may lead to a sterile system of abstract community
Plantensociologie', 'Introduction to Plant Sociology' by types, Braun-Blanquet (e.g. 1959, 1964, 1968) has made
Meltzer & Westhoff (1942). With this early work (but how it clear all over again that this approach is leading to the
mature it was already!) we may link Westhorrs oeuvre diagnosis of a real integrated plant community. This
with that of Josias Braun-Blanquet, his great example and integration is based on the structure of the plant com
teacher: the 1942 book was in fact a thorough introduction munity, that is its organisation in space. With Mueller
to the Dutch reader of Braun-Blanquet's (1928) 'Pflanzen Dombois & Ellenberg (1974) we may consider the biomass
soziologie' . structure including the stratification of vegetation layers,
As Westhoff & van der Maarel (1973) pointed out once as a most important aspect. Besides we may distinguish a
again in the Handbook of Vegetation Science, the essence horizontal, a pattern structure.
of the Braun-Blanquet approach can be stated in three A second essential element of structure is the set of
ideas: functional relations between plants. To Braun-Blanquet
(i) Plant communities are conceived as types of vegetation, this element was even of primary importance: he treated
recognized by their floristic composition. The full species 'the social life of plants' in the very first chapter of his
compositions of communities better express their relation book. Westhoff has given many a new impulse to the study
ships to one another and environment than any other of what we may call the integration, identity and stability
characteristic. in the plant community (Langford & Buell 1969). We may
mention here his Dutch contribution on biotic factors
* Nomenclature of plant species follows Heukels-van Oost
stroom. 1975. Flora van Nederland. 18e druk. Noordhoff, (Westhoff 1958), and particularly the papers on the use
Groningen. of structural characters in the classification of vegetation
3
(Westhoff 1967, 1968). His ideas have led to an attempt to personal reason that he has little of an 'apparatnik' West
arrange syntaxonomical units in larger structural-physiog hoff holds a number of more general considerations. One of
nomic units, i.e. formations. And so in the book 'Planten them is the typically Braun-Blanquetian awareness of the
gemeenschappen in Nederland,' 'Plant communities in historical factor. This temporal dimension of a species'
the Netherlands', the syntaxa are described in 13 forma environment cannot be understood through a snapshot
tions (Westhoff & den Held 1969). of the present physico-chemical complex.
The involvement of structural characters in the descrip Another consideration which is concerned with the ex
tion of plant communities is not only found in Westhoffs perimental approach is the one put forward repeatedly by
own work, but also in that of many of his collaborators Heinrich Walter, notably in the preface of his book 'Die
and pupils (e.g. Doing 1963, 1966, van der Maarel 1966, Vegetation der Erde' (Walter 1973, 1973a): by far the
Segal 1970, Londo 1971, Werger 1973). largest experimentator in nature is Nature itself. Ifwe only
Despite the voluminous oeuvre on vegetation and vege carefully describe the changes in plant communities caused
tation science Westhoff produced, he has never considered by changes in the environment and describe the latter
himself a real phytosociologist, but rather an autecologist. changes as well we have results as obtained from experi
Indeed Victor Westhoff has a passion for the plant, the ments available.
'sensitive plant' as Shelley titled one of his great poems and Along similar lines is a remark by Walter in his recent
Westhoff himself described in a section of poems in his book 'Die okologischen Systeme der Kontinente' (Walter
first and only published collection of poems called' Levend 1976). Continuous and careful observations together with
barnsteen', 'Living amber'. This parallel interest in plant relatively few but effective measurements often allow a
species and plant <;ommunities led to a sort of cross relatively rapid (and cheap!) insight in the functioning of
fertilisation in the late nineteenfifties when Westhoff ecosystems. After having expressed some doubts on the
published his first accounts on what he called the phyto rapidly increasing investments in ecophysiological equip
sociological position of plant species. The elegance of this ment and automisation of registration Walter ends with
approach was demonstrated with Carex buxbaumii, Listera words which could almost be Westhoffs. Its translation
cordata and Scirpus planifolius as examples in the interna reads: The ecologist must live together with his plants
tional literature (Segal & Westhoff 1959, Westhoff 1959, at the site as continuously as possible and while observing
Westhoff & van Leeuwen 1966) and many other species in them experience all taht happens. We are dealing with plant
Dutch journals (e.g. Westhoff & Ketner 1967, Westhoff behaviour research. Only in this way the essential can be
& van Leeuwen 1962, Westhoff & Passchier 1958) and understood' .
numerous reports from his students at Nijmegen. We would do unjustice to Westhoff, however, when we
The approach consists of a careful phytosociological would not directly add how interested he is in more
description of the vegetation in which a particular species experimental ecophysiological work as soon and as long as
occurs optimally or just marginally. Through tabular it has an immediate bearing on the real field situation.
arrangement of the releves and a detailed phytosociological (e.g. Steubing & Westhoff 1966, cf. Adriani 1945). This
interpretation, and with the help of a complete spectrum appears also from present research by his pupils (e.g. Blom
of s.yntaxonomical species groups, the phytosociological ;976).
position of that species can be understood. The environ Another link with Walter's work is Westhoffs interest
ment was not so much described directly through an in the socalled Gesetz der relativen Standortskonstanz,
analysis of soil and microclimate but rather interpreted the law of relative habitat constancy. This idea, which
by means of the powerful diagnostic value of the entire comes rather close to Vinogradov~s idea of ecological
species combination of the vegetation to which the species compensation is concerned with the changes in the con
under investigation belongs. In this respect Westhoffs stellation of habitat factors governing the occurrence of a
work comes very close to the approach developed by species towards the boundary of its distribution area (see
Ellenberg, culminating in the latter's book 'Zeigerwerte Werger & van der Maarel, this volume, for references).
der Gefasspflanzen Mitteleuropas', 'Indicator values of We now come to another important aspect ofWesthoffs
the vascular plants of Central Europe' (Ellenberg 1974). work on the relation between plant species and plant com
Unlike Ellenberg and other outstanding ecologists munities, that is the human impact on nature as it is
Westhoff has not paid much attention to soil analysis, reflected in the occurrence as well as in the disappearance.
neither to experiments with plants. Apart from the more of particular species and the accompanying changes in
4
