20 THE BRAUN-BLANQ.UET APPROACH
VICTOR WESTHOFF AND EDDY VAN DER MAAREL
Contents
20.1 Introduction 289
20.2 History 290
20.3 General Concepts . 293
20.3.1 Concrete vs. Abstract Units 293
20.3.2 Floristic-Sociological Classification Units 295
20.3.3 Diagnostic Species. 296
20.3.4 Sociological Progression 298
20.3.5 Natural Classification 299
20.4 Analytical Research Phase 301
20.4.1 Reconnaissance and Choice of Plots 301
20.4.2 Boundaries. 303
20.4.3 Minimal Area and Plot Size 305
20.4.4 Description of Structure and Strata 307
20.4.5 Floristic Analysis 308
20.4.5.1 Cover and Abundance. 308
20.4.5.2 Sociability. 310
20.4.5.3 Vitality and Fertility. 311
20.4.5.4 Periodicity. 312
20.4.6 Releve Protocols 312
20.5 Synthetical Research Phase. 313
20.5.1 Steps in Synthesis. 313
20.5.2 Primary Table 315
20.5.3 Presence, Constancy, Homotoneity 316
20.5.4 Species Weights. 319
20.5.5 Table Rearrangements. . . . . . 320
20.5.6 Mechanical Means for Rearrangement. 323
20.5.7 Determination of Fidelity. 324
20.5.8 Geographical Effects on Fidelity. 326
20.5.9 Phytocoenon Characterization 328
20.6 Syntaxonomical Research Phase. 329
287
R. H. Whittaker (ed.), Classification of Plant Communities
© Dr W. Junk bv Publishers, The Hauge 1978
20.6.1 Syntaxon Table Characterization . . . 329
20.6.2 Characterization of the Association . . 329
20.6.3 Geographic Problems with Associations. 330
20.6.4 Higher Units (Alliance, Order, Class) 332
20.6.5 Units above the Class . . . . . . . . 334
20.6.6 Horizontal and Vertical Classification . 336
20.6.7 Lower Units (Subassociation, Variant, Facies). 336
20.6.8 Nomenclature . . . . . . . . . . . 339
20.6.8.1 Construction of Names. . . . . . . . 340
20.6.8.2 Validity, Changes, and Authors' Names 342
20.6.9 Scheme of Syntaxa . . . 343
20.7 Extension of the Approach 344
20.7.1 Other Geographic Areas. 344
20.7.2 Biotic Communities. . . 348
20.7.3 Microcommunities and Synusiae 348
20.7.4 Community Complexes . 349
20.7.5 Applied Phytosociology . 351
20.7.6 Structural Considerations. 352
20.7.7 Indicator Groups. . . . 353
20.7.7.1 Indicators for Classification. 354
20.7.7.2 Indicators for Gradient Relations 355
20.8
20.8.1
20.8.2
20.8.3
20.8.4
20.8.5
20.8.6
20.8.7
20.8.8
20.9
20.9.1
20.9.2
20.10
20.11
20.12
288
Numerical Techniques.
Storage of Releves .
Species Correlation
Fidelity Tests. . .
Releve Similarity .
Measurement of Homotoneity .
Numerical Classification .
Table Rearrangement. .
Numerical Syntaxonomy.
Ordination. . . . .
Informal Ordination
Formal Ordination
Conclusion.
Summary .
Additions to the Second Edition.
355
355
357
358
359
360
360
361
363
365
365
366
371
372
374
20 THE BRAUN-BLANQ.UET APPROACH
20.1 Introduction
We shall, in this last chapter, treat the floristic-sociological
or BRAUN-BLANQUET approach to classification and interpretation
of communities. Before elaborating the details of the approach, we
may state its essence in three ideas:
(i) Plant communities are conceived as types of vegetation,
recognized by their floristic composition. The full species compositions
of communities better express their relationships to one another
and environment than any other characteristic'.
(ii) Amongst the species that make up the floristic composition
of a community, some are more sensitive expressions of a given
relationship than others. For practical classification (and indication
of environment) the approach seeks to use those species whose
ecological relationships make them most effective indicators; these
are diagnostic species (character-species, differential-species, and constant
companions).
(iii) Diagnostic species are used to organize communities into
a hierarchical classification of which the association is the basic unit.
The vast information with which phytosociologists deal must, of
necessity, be thus organized; and the hierarchy is not merely
necessary but invaluable for the understanding and communication
of community relationships that it makes possible.
The reader will see how far the elaboration of this three-part
theme has led the members of the 'school of BRAUN-BLANQUET.'
The results are (like community relationships) complex. We fear
lest the reader should lose sight of the heart of the approach - the
floristic perspective - for the abundance of technical details that
follow. The latter are only the skeleton of the approach. Furthermore,
if we do not press the physical comparison too far, its body
is the still-growing corpus of basic and applied research and the
resulting knowledge of and insight into communities. This,review
must deal mainly with the skeleton of the approach; we cannot in
the space we have discuss adequately the kinds of understanding
of communities that have come from it. We ask the reader to
recognize wherein this- account must stop short, and to seek jf he
is interested a further feeling for what the approach offers by
observing or reading studies applying it. On the other hand, we
289
shall pay some attention (especially in sections 20.8 and 20.9) to
recent developments in phytosociology that are not part of the
BRAUN-BLANQ.UET approach as such but may be of value since they
introduce concepts and methods that relate the floristic-sociological
approach to other approaches.
Textbooks and reviews on the floristic-sociological approach
have been published in most European countries, Argentine, India,
Japan and the U.S.A. (see the bibliography by MAAREL, TUXEN &
WESTHOFF, 1970, Exc.1). Recent accounts include GOUNOT (1969),
WESTHOFF & HELD (1969), and KNAPp (1971). Useful accounts
in English are BECKING (1957), WHITTAKER. (1962), KUCHLER
(1967) and SHIMWELL (1972).
20.2 History
The origin and history of the approach have been reviewed
elsewhere and can be stated only briefly here (see GAMS 1918,
BRAUN-BLANQ.UET 1921, and Du RIETZ 1921, as well as BECKING
1957, WHITTAKER 1962, and SHIMWELL 1972). The systematic description
of plant communities and the idea of community types can
be traced from great students of plant geography, HUMBOLDT
(1805), SCHOUW (1823), HEER (1835) and GRISEBACH (1838). From
their work two main approaches developed, the physiognomic and
the floristic. The physiognomic approach developed as its principal
unit the formation, a unit characterized by physiognomy or vegetation
structure (POST 1862, GRISEBACH 1872, WARMING 1895; see
article 13). Through the work ofstudents dealing with smaller-scale
units (esp. LEcoQ. 1844, 1855, THuRMANN 184-9, LORENz 1858,
KERNER 1863 and DRUDE 1890, in southern and central Europe,
POST 1862,HuLT 1881 and CAJANDER 1903 in northern Europe) there
developed the essential idea. of the floristic-sociological approach:
plant communities as units of classification based primarily on
species composition. From LORENZ (1863), MOBIUS (1877) and
DAHL (1908) a parallel approach to biotic communities can be
traced. (See BALOGH 1958 and WHITTAKER 1962 for review and
references.)
Much of the further development leading to the BRAUNBLANQ.UET
approach was centered in Zurich (STEBLER & SCHROTER
1893, SCHROTER 1894-, SCHROTER & KIRCHNER 1902, BROCKMANN-
1) Th.is and many other bibliographies to be mentioned in this contribution
have been published in Excerpta Botanica Sect. B Sociologica (R. TuxEN Ed).
They will not be listed in the references, but indicated with 'Exc'.
290
JEROSCH 1907) and Montpellier (FLAHAuLT 1893, 1898, 1901,
PAVILLARD 1901, 1912). A main outcome of this period was a full
hierarchy for vegetation classification-vegetation-type, formationgroup,
formation, subformation, stand-type down to local variants
(Nebentypen) and geographic variants (Fazies). The stand-type
was called the 'association' and considered the basic unit. FLAHAULT
& SCHROTER (1910) agreed on the following definition, presented
to the Third International Botanical Congress in Brussels, 'An association
is a plant community of definite floristic composition,
presenting a uniform physiognomy, and growing in uniform habitat
conditions. The association is the fundamental unit of synecology.'
(translation following PAVILLARD 1935b).
From this background BRAUN-(BLANQ,UET) carried out a
monographic study of alpine vegetation (1913). In this, and the
essay by BRAUN-BLANQ,UET & FURRER (1913), attention was focused
on 'Charakterpflanzen' or character species - species that possess
'fidelity' (relative restriction) to a given association. GRADMANN
(1909) had advocated the approach through floristic composition
and the use of character species ('Leitpflanzen'). The key ideas in
BRAUN-BLANQ,UET'S treatment were: (i) The study of communities
should be based on a fundamental unit, comparable to the species.
(ii) This unit should be the association, and associations should be
defined by their possession of character-species. (iii) Each association
consists (like a species) of 'individuals,' and the association
(like the species) can be described from samples of its individuals.
(iv) Each sample ('Aufnahme,' releve) should be chosen so as to
represent adequately such an individual, and it should include
analysis of the complete species assemblage. (v) Associations should
be grouped into higher units not by physiognomy, but by floristic
composition.
Additions to the approach came in further publications. BRAUNBLANQ,UET
(1915) added the 'Assoziationsgruppe' (later called
the Verband, alliance) as a unit above the association, also defined
by character-species. BRAUN-BLANQ,UET (1918) added the subassociation,
as a deviation from the typical association expressed
in a constant floristic difference, and the facies as a subordinate
unit possessing merely quantitative differences. BRAUN-BLANQ,UET
(1921) outlined essentially the full system, including the analytical
scales (20.4.5) and the 'sociological progression' (20.3.4), for arranging
communities by their levels of organization. BRAUNBLANQ,UET
(1925) pressed the claims of fidelity or 'Gesellschaftstreue'
as the key to vegetation systematics, against the disagreement
of northern schools. In the same essay he introduced the 'characteristic
species combination' (charakteristische Artenkombination
291
after SCHMID, 1923) as the ultimate community diagnosis, comprising
both character-species and constant companion species. KOCH
(1925) and BRAUN-BLANQ,UET & JENNY (1926) added the concept of
differential-species for the characterization of subordinate units,
and the order as a unit above the alliance. BRAUN-BLANQ,UET &
PAVILLARD (1922, 1925, 1928) in their Vocabulaire codified the
analytic and synthetic procedures of the approach. Finally the
science of pl~lllt sociology and its application were spelled out more
fully in the first edition of the textbook of BRAUN-BLANQ,UET (1928,
1932).1)
From these beginnings the influence of the approach spread
through western and central Europe. We cannot review that spread,
but will mention such leaders as PAVILLARD, ALLORGE and MOLINIER
in France, TUXEN and OBERDORFER in Germany, SZAFER and PAWLOWSKI
in Poland, FURRER in Switzerland, So6in Hungary, KLlKAin
Czechoslovakia, HORvAT, HORVATIC and WRABER in Yugoslavia,
BORZA in Rumania, GIACOMINI and TOMASSELLI in Italy, BOLOS
in Spain, LEBRUN in Belgium, and DE LEEuw in the Netherlands.
References for these authors are found in BRAUN-BLANQ,UET (1964,
1968). BRAUN-BLANQ,UET'S own work began at Zurich and continued
from 1927 onwards at Montpellier, where he still leads the
S.LG.M.A., Station Internationale de Gcobotanique Mcditerrancenne
et Alpine. A second centre was established 1932 at Stolzenau, West
Germany under Reinhold TUXEN, a pre-eminent leader of the approach
and former director of the Zentralstelle (later, Bundesanstalt)
fur Vegetationskartierung, who has done much to give the
approach its value in application. From TUXEN'S centre the approach
spread to various non-European countries as well, and
notably to Japan.
During its spread the approach has been known by several
names: French-Swiss (or Swiss-French) school, Zurich-Montpellier
school, Middle European-Mediterranean school, and Sigmatism
(TANSLEY 1922, Du RIETZ 1936, BRAUN-BLANQ,UET 1959, 1968,
EGLER 1954 ,BECKING 1957, WHITTAKER 1962, TUXEN 1969a). Most
of these terms lack specificity or are unclear. Moreover we feel
that the time has passed for the word 'school' with its implications of
a fixed system. We prefer in this article the most direct name, viz.
BRAUN-BLANQ,UET approach, whilst we may characterize the essential
ideas in the term, the 'floristiC-sociological approach.'
') One notes through this development of the full range of concepts the
continuing contribution of BRAUN-BLANQ.UET himself, that has given the approach
its present character (GAMS 1972 to the contrary).
292
20.3 General Concepts
20.3.1 CONCRETE vs. ABSTRACT UNITS
Much dispute has centered on the nature of the plant community.
This will be briefly reviewed as far as the BRAUN-BLANQUET approach
is concerned. (See further PAVILLARD 1935, WESTHOFF 1951,
1970, WHITTAKER 1962, BRAUN-BLANQUET 1964.)
Concepts of the plant community include: (i) the organismal
concept (CLEMENTS 1936, T ANSLEY 1920): the community as a
'superorganism.' (ii) the concept ofsocial structure (PACZOSKI 1930)
and many early Russian authors such as SUKATSCHEW (1929). (iii)
the individualistic concept (GLEASON 1926): the community as a
changeable mixture of 'individualistically' distributed plant species.
(iv) the concept of population structure (WHITTAKER 1953, 1962,
1970): the community as a system of interacting species and vegetation
as a complex population pattern.
The BRAUN-BLANQUET approach takes a practical, intermediate
position that recognizes the heterogeneity of species distributions
but emphasizes nonetheless the interactions between
plants in the community, which has a certain individuality because
of relative discontinuities between communities in the field. Definitions
of plant community or phytocoenose range from the more
superficial, 'any collection of plants growing together which has
as a whole a certain unity' (TANSLEY 1935), to the more profound,
'a plant community (+animal community = biotic community)
is a working community; the species composition of which
is in a sociologic-dynamical equilibrium in competition for space,
minerals, water and energy, in which each component affects all
others and which is characterized by harmony between environment
and production and phenomena expressing life in form,
colour and temporal course' (TtiXEN 1957).
As shown by TtixEN's definition, the community concept may
be broadened from the plant community to the biotic community
of plants and animals: producers, consumers and decomposers. In
a further broadening of perspective the biotic community (= biocoenose)
plus its environment or biotope is treated as a functional
unit, the ecosystem. The concept of ecosystem has become most
familiar from its expression in English-language ecology by TANSLEY
(1935), but the concept is largely identical with FRIEDERICH'S
(1927, 1958) 'holocoene' and the 'biogeocoenose' of SUKACHEV
(e.g. 1954, 1960) and other Russian authors. Though by far the
greatest development of the BRAUN-BLAN~UET approach has been
293
in application to plant communities, biotic communities are clearly
amenable to parallel study (section 20.7.2). TuxEN (1957, 1965b)
has emphasized this possibility and stressed the influence of
FRIEDERICHS and THIENEMAN (e.g. 1956) on his own views.
The term plant community and its German equivalent 'Pflanzengesellschaft'
have been used in both concrete and abstract senses,
which has caused dispute and confusion (see TANSLEY 1920, 1935,
Du RIETZ 1921, ALECHIN 1926, PAVILLARD 1935, WESTHOFF 1951,
1965, WHITTAKER 1956, 1962). The floristic-sociological approach
has always stressed the distinction between the concrete and the
abstract community. The classification units were, naturally, abstract
units. Concrete plant communities were often, especially in
forestry, referred to as stand (Bestand), whilst concrete representatives
ofassociations were even called association-individuals (PAVILLARD
1912, BRAUN-BLANQ.'uET & PAVILLARD 1922). This term has
been much disputed and criticized (most thoroughly by WHITTAKER
1962) and we consider it now as of only historic interest.
WESTHOFF (1951, 1965) has acknowledged the distinction between
concrete and abstract communities in separate definitions,
which were linked to a general definition of vegetation. For the
concrete plant community the term phytocoenose (GAMS 1918) was
proposed, for the abstract community the term phytocoenon
(MAAREL 1965, specifying the general term 'coenon' proposed by
BARKMAN et al. 1958, WESTHOFF et al. 1959).
The term (phyto)coenon may be proposed as a suitable international
term which may replace the terms community-type
(WHITTAKER 1956, 1962) and nodum. The latter term has been suggested
by POORE (1956, 1962), but has been used in a more specific
meaning by WILLIAMS & LAMBERT (1961) in their 'nodal
analysis' (CfIVIMEy-COOK & PROCTOR 1966).
For biotic communities parallel terms: biocoenose and biocoenon
may. be used (MAAREL 1965). For partial communities
MORZER BRUYNS' (1950) term merocoenose can be used with
merocoenon as the abstract equivalent. Layer communities could be
called, stratocoenose-stratocoenon. For specific subcommunities consisting
ofplants ofthe same stratum, life-form and seasonal relations
the couple, society-synusia may be reserved. (See further article 16.)
In conclusion we present WESTHOFF'S (1951, 1970) definitions
in a slightly adapted form. Vegetation is defined as a system of
largely spontaneously growing plant populations, growing in coherence
with their sites and forming an ecosystem with these sites and
all other forms of life occurring in these sites. (Thus are excluded all
assemblages of mobile plants and collections of plants growing in
arrangements set up by man such as flowerbeds and arboreta.)
294
A phytocoenose is defined as a part of a vegetation consisting
of interacting populations growing in a uniform environment and
showing a floristic composition and structure that is relatively
uniform and distinct from the surrounding vegetation.
A phytocoenon is defined as a type of phytocoenose derived from
the characterization of a group of phytocoenoses corresponding
with each other in all characters that are considered typologically
relevant.
20.3.2 FLORISTIC-SOCIOLOGICAL CLASSIFICATION UNITS
Phytocoena include such various kinds of vegetation units as
formations defined by physiognomy, dominance-types defined by
major species, forest site-types defined by undergrowth composition,
and noda derived by quantitative comparisons or numerical procedures.
The approach of BRAUN-BLANQUET has its own, formal
hierarchy of units, of which none of those just mentioned is a part.
The fundamental unit of the hierarchy is the association, a unit that
corresponds in function to the species as the fundamental unit of
idiotaxonomy, or the classification of individual organisms. The
word 'association' has had a long, complex, and argument-afflicted
history as different schools sought to determine its meaning to their
preference (WHITTAKER 1962, SHIMWELL 1972, see also articles 14,
17 and 18). In an earlier definition by BRAUN-BLANQUET (1921)
'the association is a plant community characterized by definite
floristic and sociological (organizational) features which shows, by
the presence of character-species (exclusive, selective, and preferential)
a certain independence.' MEIJER DREES (1951) defined
the association as ' a plant community identified by its characteristic
taxon combination, including one or more (local) character-taxa
or differentiating taxa.' A similar conception was agreed on during
a Symposium on Plant Sociological Systematics at Stolzenau in
1~64 (in TuxEN 1968b, especially OBERDORFER 1968) and a later
colloquium at Rinteln (see DIERSCHKE 1971). We shall return to
characterization of the association (20.6.2) but emphasize that, in
keeping with the floristic-sociological perspective, the association
is defined by its characteristic species combination including characterand
differential-species as well as companions (Begleiter) with high
presence values (over 60%). The Sixth Botanical Coq.gress at Amsterdam,
1935, accepted definition of the association by characteristic
or differential species in the sense of BRAUN-BLANQUET as
one of three resolutions given in article 18.3.6.
295
As basic units associations, like species, are to be grouped into
a hierarchy of higher units. Associations are classed into alliances,
alliances into orders, orders into classes, and classes into divisions
(20.6.4-5); associations are divided into lower units of the hierarchy
(20.6.7). All these units are coena as defined above; but the coena
of the formal system of BRAUN-BLANQUET may be termed .ryntaxa
(BARKMAN et al. 1958, WESTHOFF et al. 1959). Thus the parallelism
in the classification of organisms and phytocoenoses is realized:
to the species as a fundamental unit corresponds the association, to
the taxon for units on any level corresponds the syntaxon, to
(idio) taxonomy for the practice of classification corresponds
syntaxonomy, to (idio)systematics as the broader study of relationships
among organisms corresponds synsystematics.
20.3.3 DIAGNOSTIC SPECIES
Syntaxa are defined by diagnostic species (character-species,
differential species, and constant companions). Character-species are
species that are relatively restricted tb the stands (or samples) of a
given phytocoenon, and therefore characterize it and indicate its
environment (Fig. 1). In ideal, a group of character-species is used
for the characterization of a syntaxon of theBRAUN-BLANQUET classification
on any level from the association to the class. To serve as
character-species for an association, a species should have a relatively
narrow distribution even if it is not simply restricted to the association.
(Degrees of restriction to a given syntaxon, which are
termed degrees of fidelity or Treue, are discussed in 20.5.6.) Note
that the concept does not say that the species need be important in
phytocoenoses; very minor species may have diagnostic value.
The German term is 'Charakterart'; this has been variously
translated into English as characteristic species (BRAUN-BLANQUET
1932), which seems unsatisfactory, faithful species (POORE 1955a,
BARKMAN 1958a,BEEFTINK 1962, MOORE 1962,WESTHOFF 1959), and
character-species (BECKING 1957, WHITTAKER 1962). The latter,
most direct translation, is our preference. HErMANs (1939) introduced
'kensoort' in Dutch to 'avoid the germanism 'karaktersoort';
and the Dutch kensoort returned to German as Kennart (TtiXEN
1950: 99). Most phytosociologists writing in German now use that
term, which is accepted in the third edition of BRAUN-BLANQUET'S
(1964) text. Syntaxa are most often defined by the fundamental
units of idiotaxonomy - species -, but this is not always the case.
Sometimes plant subspecies, varieties, or ecotypes may contribute
to the definition of lower-level syntaxa or geographically vicariant
296
Associations
~A-. subassociations
B ---Ca
I b I c
-~
I
I3
I -=::;;.:;., 5
2
.1
4
I
I I 7
6 8
I I
9
I I
wet moist I fresh I dry very dry
Fig. 1. Diagnostic species along a moisture gradient. Species I and 2
are character-species for association B, and have their populations centered in
(or largely confined to) that association. Species 3 and 4 are character-species
for association A, and species 5 is a character-species for association C. Species
4 and 6 are differential-species for subassociation a of association B, and species
7 and 8 are differential-species for subassociation c of association B. In each case
presence of the differential-species distinguishes the moist, or dry, subassociation
from the "typical" (fresh) subassociation b. Species 9 is a more widely distributed
species that might help characterize association B as a constant companion for
that association. Species 9 might also be a character-species for a higher syntaxon,
such as an alliance uniting associations Band C with other associations.
associations. Sometimes a genus or subgenus may be used to define
a higher syntaxon, as the alliance Spartinion is defined by the
fidelity of any species of the genus Spartina (BEEFTINK 1965, 1968,
see alSOPIGNATTlI968a: 72, WESTHOFF apUdPIGNATTI 1968a: 74). It
is therefore appropriate to broaden character-species to charactertaxon
(BARKMAN et al. 1958) as we shall do here following Dutch
use of 'kentaxon' (WESTHOFF & HELD 1969). The German equivalent
is of course 'Kenntaxon'.
Character-species characterize syntaxa by their normal occurrence
in phytocoenoses of that syntaxon, contrasted with their
absence less frequent occurrence or smaller total estimate in phytocoenoses
of all other syntaxa. It is possible also to distinguish closely
related syntaxa by the presence and absence ofcertain species without
concern for the broader distributions of those species. Two subassociations
of an association may be tharacterized by the fact that
samples of one normally include species 4 and 6, whereas samples of
the other normally lack these species (Fig. 1). These species function
as differential-species that distinguish the two subassociations. Dif-
297
ferential-species, as the second class of diagnostic species, define
syntaxa on the basis of the distributional boundaries of species
(without regard to fidelity for the syntaxa in question), and are
used primarily to define lower syntaxa.
Since being introduced by KOCH (1925) and BRAUN-BLANQ.UET
(1925) as 'Differentialart', the concept has entered English as differential
species (BRA'UN-BLANQ.UET 1932, WHITTAKER 1962) or
differentiating species (BECKING 1957), whilst in German the term
Trenllart has come into general use (TuxEN 1950, 1952, BRAUNBLANQ.UET
1959, 1964a) in parallel with Kennart. The use of differential-species
has been further developed by SCHWICKERATH
(1931, 1942, 1954a) and TUXEN (1937, 1950, 1952). Differentialspecies
and Trennart are synonyms, and these are largely synonymous
also with differentiating species; but we shall here use 'differentiating
species' for the more informal use of diagnostic species
of unassigned syntaxonomic position in the synthetic phase of
research (20.5), and speak of differential-species and differentialspecies
groups for the formal characterization of syntaxa (20.6).
A further recent development is the concept of 'character-combination'
(after the Dutch kencombinatie) introduced by WESTHOFF
& HELD (1969) following BEEFTINK (1965, 'differentiating
species combination'). The essential concept is the exclusiveness to
a syntaxon of the combination of species (or taxa), whilst none of
them need be a character-taxon.
A third group of species with diagnostic value are the constant
companions (20.5.3) which occur in most releves of a syntaxon
but are not designated as character- or differential-species. Constant
companions are added to the character-taxa to form the
'characteristic species (taxon) combination' for associations and
higher taxa. Although BRAUN-BLANQ.UET (1925) introduced this
concept rather early and repeatedly (e.g. 1959, see MOORE 1962)
stressed its relevance there has been doubt amongst critics (e.g.
POORE 1955-1956). Of course, as BRAUN-BLANQ.UET (e.g. 1932: 68)
remarked for associations, these syntaxa are established best which
possess a high proportion of both characteristic and constant
specIes.
20.3.4 SOCIOLOGICAL .PROGRESSION
BRAUN-BLANQ.UET (1921) sketched an 'arrangement of the
plant communities according to their sociological progression.' The
ide!! of this arrangement was again a parallel with idiobiology,
specifically with the arrangement of taxonomic groups by evolu-
298
tionary level. Levels of structural and organizational development
were chosen as criteria for arranging phytocoenoses from moving
aero- and hydroplankton communities on the lowest level to the
tropical rain forest at the highest level. Other communities are arranged
between these according to increasing stratification, complexity,
presence of dependent communities, species richness,
diversity of growth-forms, and assumed intensity of species interactions.
Thus BRAUN-BLANQ.UET (1921) summed up many of the
structural-successional trends much later commented on by ODUM
(1969)! The scheme was later modified in detail, and increasing
stability was added as a criterion. BRAUN-BLANQ.UET (1964) arranges
higher syntaxa (classes and orders) in the progression.
WAGNER (1968) called the sociological progression a 'purely
artificial division principle' (cf. ELLENBERG 1963). Of course, no
phylogenetic meaning should be imputed to its sequence. Yet the
sociological progression is useful, as a principle at right angle to
the levels of the hierarchy, in arranging syntaxa in surveys. Its use
is illustrated in vegetation monographs by TUXEN (1937), OBERDORFER
(1957), ELLENBERG (1963), OBERDORFER etal. (1967),
WESTHOFF & HELD (1969), and others. It may be of interest that
it has appeared also as an axis of broad-scale ordination of syntaxa
(MAAREL 1972a, see 20.9.2).
20.3.5 NATURAL CLASSIFICATION
The problem of 'natural' classification has been discussed in
another article (12.1.2). It has been accepted by BRAUN-BLANQ.UET
and others that associations are abstract units. A degree of continuity
or gradualness in transitions between phytocoena is recognized,
but considered not to preclude classification. 'We are
convinced that the plant cover of the earth in all its dimensions can
be divided into phytosociological groupings of higher and lower
rank; their delineation may be either sharp or less sharp and
gliding.' (BRAUN-BLANQ.UET and TuxEN in comment on GOODALL
1963, translated.)
American research in gradient analysis has led to interpretations
that may seem in conflict with those ofBRAUN-BLANQ.UET. The
American work of WHITTAKER (1951, 1954, 1956, 1962, 1967,
articles 2 and 3) and CURTIS (CURTIS & McINTOSH 1951, CURTIS
1959, McINTOSH 1967, article 7) developed in parallel to Russian
work of RAMENSKY (1930, article 17) toward the concept expressed
by WHITTAKER (1970) as 'the population structure of vegetation.'
In this concept the population structure of the individual phyto-
299
coenose may be analysed through dominance-diversity and speciesimportance
relations, and niche differentiation among plant species
expressed in stratification and functional differences among them.
The phytocoenose is then conceived as a 'system ofinteracting, nichedifferentiated
and partially competitive species.' The vegetation of
an area, on the other hand, is to be approached through gradient
analysis, studying the manner in which species populations are
distributed along environmental gradients and combined into
phytocoenoses. Species are differently distributed from one another,
so that undisturbed phytocoenoses intergrade continuously in most
areas. 'In relation to patterns of environmental gradients, communities
form complex and largely continuous population patterns.'
(WHITTAKER 1970, article 3.9.)
It is further recognized that, 'Because of environmental interruptions
and some relative discontinuities inherent in vegetation
itself, the pattern may also be considered a complex mixture of
continuity and relative discontinuity' (WHITTAKER 1956: 32).
Vegetational continuity by no means precludes useful classification
(WHITTAKER 1956, 1962, 1970). Gradient analysis implies a vision
of vegetation, regarded as a coherent pattern of intergrading
phytocoenoses, different from that of phytosociologists who tend
to see it in terms of a typology of phytocoena (see, however, 20.7.4
on complexes). Yet we think that the difference between the concept
of the gradient approach and BRAUN-BLANQUET'S is clearly one of
degree - ofemphasis ofcontinuity vs. discontinuity where both are
present, of species individuality vs. species groupings where both
are realistic, and of gradient analysis vs. classification where both
are possible. We judge the difference between less extreme students
of gradient analysis, and of the BRAUN-BLANQUET approach, to
be one of emphasis and perspective, not one of fact or under-
standing.
Given the individuality ofspecies distributions and some degree
of continuity between phytocoenoses, classification is not strictly
natural in the sense defined in article 12.l.2. BRAUN-BLANQUET
(1951a: 561,1959: 147) has considered argument on the naturalness
of classification pointless. TuxEN (1955), in response to a criticism
of ELLENBERG (1954b), interpreted phytosociological classification
through the concept of types as ideal concepts, recognized in an
empirical way from "correlation concentrates,' i.e. groups of correlated
characters. That which is evident and characteristic of a
type is always its nucleus, not its periphery; types are not pigeonholes
but foci in a field of variation. GLAHN (1968, see also RAUSQHERT
1969) elaborated the concept ofvegetation type, distinguishing
as its three aspects: (i) The vegetation type as identity: repeti-
300
tions of certain observations are approached via intuitive integration,
resulting in vegetation-type concepts based on recurring
combinations of species. (ii) The vegetation type as maximal correlative
concentration: The joint floristic-sociological and ecological
approach leads to types as correspondences of recurring species
combinations with recurring combinations of environmental factors.
(iii) The vegetation type as systematic category: A hierarchic
system is feasible as a result of an integrative, inductive process
grouping the initial types into higher and higher ranks on the basis
ofcommon species combinations. The similarity of this statement to
the theory of classification developed at greater length by WHITTAKER
(1962) may be noted. Relative naturalness of classification
in this perspective is to be judged by success in embodying the
maximum number of significant relationships among phytocoena
and species in the structure of the hierarchy. We assert that compared
with other classifications the BRAUN-BLANQUET approach,
with its floristic emphasis and maximum use ofspecies distributional
relationships for classification, does not fall short.
20.4 Analytical Research Phase
20.4.1 RECONNAISSANCE AND CHOICE OF PLOTS
An analysis of the vegetation is as a rule preceded by a preliminary
survey of the area, when this is little known to the investigator.
This reconnaissance (see CAIN & CASTRO 1959) includes
a study of the general vegetation pattern, and the establishment of
the apparent relations of the various vegetation types with geology,
topography and soil conditions. The next step, 'primary survey,'
including a superficial description of the major communities, is
mostly passed over in the BRAUN-BLANQUET approach.
A vegetation analysis starts with the choice of stands (phytocoenoses)
on the basis of the reconnaissance. Within the stand one
sample plot is laid down, often covering a large part of the stand.
The analysis of this plot is called the releve. The BRAUN-BLANQUET
approach has often been criticized for the subjectivity of its
sampling procedure. However, subjectivity of stand choice must be
accepted in the procedures of many empirical sciences. A selection
of releves is desired that will effectively represent the variation in
the vegetation under study, the samples being so chosen that they
will not represent different phytocoena disproportionately and will
not include mixed, incomplete, or unstable stands. For this purpose
of equitable representation of different kinds of communities with
301
most useful reieves, a subjective, 'stratified' sample selection is far
superior to sample choice by random points on a map.
Releves may be undertaken without classification as a purpose.
They may then serve for the study of vegetation dynamics on a
given area (periodicity, fluctuation, or succession) or another
ecological purpose, e.g. the study of the ecology of a certain taxon
or ecotype by describing its pattern in relation to the pattern of the
vegetation. In most cases, however, reieves are intended to be used
for some form of classification or ordination. In that situation, the
only preconception in this choice is the demand for uniformity
of the stand.
'Uniformity' may better express what is sought than 'homogeneity.'
This chapter is not the place for a treatment of the
mathematical aspects of homogeneity in vegetation; we may refer
to POORE (1955, 1956), ELLENBERG (1956), DAHL (1957, 1960),
BECKING (1957), LAMBERT & DALE (1964), BRAUN-BLANQ.UET
(1964), MAAREL (1966b) and GOUNOT (1969) for general considerations,
to GOODALL (1952, see also his bibliography, 1962 Exc.),
DAHL & HADAC (1949), DAHL (1957), GREIG-SMITH (1964) for aspects
of homogeneity in the distribution of plant individuals, to
GOODALL (1961) and GREIG-SMITH (1964), for pattern analysis, to
RAUNKIAER (e.g. 1934), GUINOCHET (1955), DAHL (1957), CAIN &
CASTRO (1959) for the relation between homogeneity and frequency
distribution and to CURTIS (1959), GODRON (1966) and GOUNOT
(1969) for infrastand similarities and information measures as approaches
to homogeneity.
The first condition is that no obvious structural boundaries or
variation in stratification are visible within the stand. The second
criterion is uniform floristic composition. It is usual to look for
joint patterns of dominant and/or abundant species and then to
delimit a stand where ,qualitative changes in patterns occur, i.e.
where one or more species drop out and others come in. In many
cases an experienced field worker is able to judge this rapidly.
In many cases however the changes in pattern are quantitative
rather than qualitative. The species composition differs little from
one site to an adjacent one, but the relative proportions of abundance
and cover do vary. It is usual to make separate releves in any
case where the abiotic habitat factors show a clear discontinuity or
at least a marked transition.
Many species cause pattern heterogeneity only by their growth
form or their aggregation or shoot clustering. In such a case one may
try at first to study this biotically heterogeneous pattern as one single
stand, i.e. with one releve. However, the situation becomes different
as soon as the crowding of one major species leads to the establish-
302
ment of one or more other species which appear to be locally bound
to it. A simple example is the establishment of a dwarf shrub patch
in a herbaceous vegetation, e.g. a clone of Salix repens in open dune
grassland. When one or more species are found particularly bound
to that dwarf shrub patch we prefer to consider the latter as a
separate phytocoenose which has thus to be analysed with a
separate releve. However, there are exceptions to this rule. E.g.
OBERDORFER (1970, Canaries) and WERGER (1973, South Africa)
report stable savanna mosaics consisting of a dwarf-shrub and
grassland and a low tree and shrub phytocoenose. Such mosaics
are considered one vegetation type, since one does not find locally
more extensive patches of either phytocoenose without adjacent
patches of the other one.
In intermediate or doubtful cases, such as a swamp community
with tall tussocks of sedges (Carex hudsonii) alternating with
wetter hollows, it will be advisable, at least when the type ofvegetation
pattern is unknown, to follow both procedures, viz. analysing
the pattern as a whole as well as releves of tussocks and hollows
separately. It will then turn out later which releve is more useful
for the classification purpose.
20.4.2 BOUNDARIES
As was said before, the recognition of distinct stands may presuppose
the occurrence of discontinuities in the field. Although according
to the opinion of BRAUN-BLANQ,UET workers such discontinuities
are mostly to be observed, it is obvious that boundaries
between stands are less sharp in some cases than in others. In such
cases the boundary may be detected by means of a belt transect
analysis or a 'line taxation.' Such transects consist of a series of
small quadrats laid down at right angles to the extension of the
boundary zone. By comparison of quantitative data on species occurrences
in the quadrats the boundary zone can be discerned from
the uniform phytocoenoses. MAAREL (e.g., MAAREL & LEERTOUWER
1967) and FRESCO (1972) refined boundary analysis by
constructing 'differential profiles,' in which can be shown degrees
of change between adjacent quadrats along a gradient. For similar
techniques see article 5.4.4.
In general we may distinguish between two types of boundary
zones which are, according to the relation theory of LEEUWEN
(1965-1970, WESTHOFF 1971a, b, WESTHOFF & LEEUWEN 1966,
SHIMWELL 1972), the limes convergens and the limes divergens. The
limes convergens zone, or convergent limit, is characterized by sharp
303
III
c:Q)
Cl
...Q)
. ~
"III
c: III
Q) Q)
Cl E...Q)
;>
c: c:
0 0
(J
~
III ;;:Q)
0
E ...Q.
c: c:
0 0.;;
..!!
(U
....
- Q)
0 Cl
... ~Q.
c:
0
.... CD(U
....Q)
Cl
~
«
Fig. 2. Vegetation profiles corresponding to the limes convergens and lime~
divergens zones ( LEEUWEN 1965).
304
vegetational boundaries on either side of which uniform phytocoenoses
occur, with few species represented, some by many individuals,
in coarse-grained patterns. The corresponding environment
is unstable, i.e. with sharp fluctuations in vertical direction
(e.g. water-table or phreatic level) or in horizontal direction (e.g.
deposition of flood marks). A typical synsystematical unit of this
type is the Agropyro-Rumicion crispi.
The limes divergens zone, or divergent limit, is characterized by
numerous small-scale boundaries in phytocoenoses merging continuously
into each other, with many species generally represented
by few individuals in fine-grained patterns. The corresponding
environment is stable and determined by a gradient. A typical
series of divergent synsystematical units is Mesobromion-Trifolion
medii-Carpinion betuli. Fig. 2 shows a vegetation
profile belonging to these types of boundary. According to MAAREL
(e.g. 1966a), LEEUWEN (1966) and WESTHOFF (1971a) the ecotone
concept of LIVINGSTONE and CLEMENTS (e.g. WEAVER & CLEMENTS
1938, CURTIS 1959) could be applied to the limes convergens zone,
whilst the ecocline concept as derived from HUXLEY'S cline concept
(WESTHOFF 1947, MEIJER DREES 1951) may be considered an
equivalent of the limes divergens zone.
The delimitation of stands within ecotones generally gives no
difficulties, except for pattern problems already discussed. Gradients
of vegetation-and-biotopes (ecoclines) can be approached by belt
transects following the direction of variation, which transects have
to be divided into sample plots (not necessarily quadrats) small
enough to be uniform. Recently JAKUCS (1972) gave examples of
this approach in 'grassland-shrub-woodland' transitions within the
range of the Quercetea pubescenti-petraeae in Hungary.
20.4.3 MINIMAL AREA AND PLOT SIZE
The size ofthe sample plot is largely dependent on the structure
of the vegetation under study, but may be affected also by the size
of.the stand. In many cases the stand is sufficiently small to be
analysed largely in one releve. In many other cases, however, this
is not possible, and then the plot should be large enough. that all
species of regular occurrence in the stand should be present in the
sample plot. In extremely uniform phytocoenoses which are very
poor in species, e.g. salt marshes, where differences in abundance
and cover are of major concern, these differences should be
considered in establish;ng the size of the sample plot.
305
These conditions refer to the minimal area concept (see also
articles 5.2.1, arid 18.3.3). Without treating this concept in detail
we may give some comments relevant to the phytocoenose description.
First the minimal area as an analytical concept (for the
phytocoenose) has to be distinguished from the synthetic minimal
area (for the phytocoenon). The analytic minimal area should be
defined for a stand under study as a representative area, e.g. as an
adequate sample of species of regular occurrence in the stand;
the size decision may, but will not necessarily, be based on the
total number of species in the stand or the usual minimal area
curve (BRAUN-BLANQ.UET 1964, CAIN & CASTRO 1959, Du RIETZ
1921,1930, GOODALL 1952,1961, GREIG-SMITH 1964-, HOPKINS 1957,
MAAREL 1966b, TuxEN 1970c and 1970 Exc., VESTAL 194-9). A
synthetic minimal area, in contrast, is subject to many current
definitions (e.g. BRAUN-BLANQ.UET 1964, CAIN & CASTRO 1959,
and WESTHOFF 1951 - who explicitly defined it as 'the minimal
surface which has as a rule to be occupied by a sample of a plant
community if the normal specific assemblage will be able to
develop'). The size decision in this case involves not merely the
stand itself but the 'normal' composition of stands representing a
phytocoenon. It is clear that before judging this in the field one
should know from synthesis the characteristic species combination.
Nevertheless many authors, including BRAUN-BLANQ.UET (1964-),
describe the normal species-area determination, applied analytically
to the stand, as the way to find the minimal area.
TUXEN (1970c) presented numerous examples of species-area
curves ending in a horizontal part, implying that at the sample
size where the horizontal part begins, full species representation or
saturation has been reached. Tiixen consequently spoke of'saturated
communities,' which he found among homogeneous communities
of various organization levels. He cited various authors who found
curves of the same type, including Du RIETZ (1921).
Most authors, e.g. BRAUN-BLANQ.UET (1964-), CAIN & CASTRO
(1959), HOPKINS (1955), and MAAREL (1966b) have found curves
that never reach an asymptote. It is true that many of their
curves refer to either relatively small or relatively large areas. It
remains unclear in which situation which type of species relation
can be expected - the ARRHENIUs-type of a log-log relation (cf.
PRESTON 1962), the ROMELL-type of a log-linear relation (cf.
HOPKINS 1955, DAHL 1957, WILLIAMS 1964-, MAAREL 1966b), or
the KYLIN-type ofsaturation relation. BALOGH (1958), and MAAREL
(1966b), quoting FREY (1928), who presented 'the three types,
interpreted them as referring to very uniform and species-poor
environments (limes convergens type!), highly variegated and
306
TABLE I
Minimal area values in square meters for various communities
Epiphytic communities
Terrestrial moss communities
Hygrophylous pioneer communities (Isoeto-Nanojuncetea)
Dune grasslands (Koelerio-Corynephoretea)
Salt marshes (Asteretea tripolii)
Pastures (Lolio-Cynosuretum)
Mobile coastal dune communities (Ammophiletea)
Hay meadows (Arrhenatheretalia)
Heathlands (Nardo-Callunetea)
Alpine meadow and dwarfshrubs (Elyno-Seslerietea)
Calcareous grasslands (Festuco-Brometea)
Chaparral, temperate sclerophyll shrubland
Weed communities (Secalietea)
Scrub communities (Rhamno-Prunetea)
Steppe communities
Temperate deciduous forest (Querco-Fagetea)
Mixed deciduous forest (North America)
Tropical secondary rainforest
Tropical swamp forest
0.1-0.4
1-4
1-4
1-10
2-10
5-10
10-20
10-25
10-50
10-50
10-50
10-100
25-100
25-100
50-100
100-500
200-800
200-1000
2000-4000
species-rich situations (limes divergens type), and intermediate
types or (BALOGH) 'normal homogeneous stands', respectively.
Table I presents some minimal area values for community
types, taken from various sources. (See TDxEN, 1970 Exc. for a bibliography).
They range from a few dm2 for certain epiphytic communities
to one hectare or more for climax tropical rain forests.
In general the plot size is taken somewhat larger than the minimal
area. It will be clear that plot size should not vary too much within
one vegetation type. The shape of the plot is not standardized and
may depend on the situation. If possible, a quadrat of rectangular
shape is to be preferred.
20.4.4 DESCRIPTION OF STRUCTURE AND STRATA
Vegetation layering is an important structural character.
Mostly only four principal layers are distinguished as the tree,
shrub, herb, and moss layers. The latter is also named field layer
or thallophyte layer (which is less adequate, because many of the
larger fungi do not belong to it). These principal layers may be
further subdivided. In the releve as many layers are distinguished
as is considered appropriate or necessary. For each layer height and
coverage degree in per cent, mostly with 5-10% intervals, are
estimated. For tree and shrub layers the age is estimated and additional
data on stem diameter, number ofdead or fallen trees, and·
occurrence of epiphytic communities are gathered.
307
Stratification can be more accurately described with the help of
diagrams in which the total or combined cover of each layer is
indicated (HULT 1881, BRAUN-BLANQ.UET 1928, cf. CAIN & CASTRO
1959, KNAPP 1971). These diagrams are related to vertical profiles,
as have frequently been presented for tropical rain forest (see
article 13.4.2) and by ZONNEVELD (1960) for the freshwater tidal
delta of the Rhine. Both however have their own special uses. The
profile can show exactly the structure of a given phytocoenose; in
tne coverage-stratification diagram a pattern typical for a phytocoenon
can be generalized (CAIN & CASTRO 1959). A more formal
method is that of DANSEREAU (1957, DANSEREAU & ARROS 1959)
for the description and recording of vegetation on a structural
basis (see article 13.3.3).
It is possible to classify different strata into separate synusial
units. This approach is treated in detail by BARKMAN (article 16).
The BRAUN-BLANQ.UET approach however considers the main strata
of a given stand as a single phytocoenose which has to be analysed
as a whole, at least as far as terrestrial communities are concerned.
The main arguments are: (i) The different strata are rooting in a
common substratum; (ii) The layers are ecologically closely interrelated;
(iii) The plants of all other layers have originally been
part of the field layer and they must pass through one or more
layers as they develop from seedlings to their mature life-form
stature. The layered community is, then, a dynamic totality (see
WEBB et a1. 1967b on the vertical integration of the rain forest).
The situation in aquatic communities of higher plants is less
clear however. Here the coherence of layers is much looser, or
absent. Du RIETZ (1930a) has proposed a stratification scheme, which
was modified by HARTOG & SEGAL (1964) (see also article 16.4.3).
Layers are distinguished according to rooting or non-rooting of
plants and the positions of leaves in relation to the water surface.
The layers are named by growth-forms after representative genera,
e.g. the isoetid layer.
An additional analysis can be made of subterranean layers.
For a treatment of root stratification see BRAUN-BLANQ.UET (1964:
59-62). Bibliographies of root studies were given by WILMANNS
(1959 Exc., 1966 Exc.) and TUXEN & WILMANNS (1973 Exc).
20.4.5 FLORISTIC ANALYSIS
20.4.5.1 Cover and Abundance
The description ofstructure is followed by an inventory oftaxa,
at least of phanerogams, pteridophytes, bryophytes and lichens.
308
TABLE II
The combined estimation (cover-abundance) scale of BRAUN-BLANQ.UET, compared
with the cover-abundance scale of DOMIN (See EVANS & DAHL 1955). The
subdivisions 2m, 2a, and 2b are proposed by BARKMAN et al. (1964). The
ordinal transformation is discussed in 20.5.4.
BRAUN-BLANQ.UET, cover-abundance
r one or few individuals
+ occasional and less than 5 %
of total plot area
abundant and with very low
cover, or less abundant but
with higher cover; in any case
less than.') % cover of total
plot area
2 very abundant and less than
5 % cover, or 5-25 % cover
of total plot area
2m very abundant
2a 5-12.5 % cover,
irrespective of number
of individuals
2b 12.5-25 % cover,
irrespective of number
of individuals
3 25-50 % cover of total plot
area, irrespective of number of
individuals
4 50-75 % cover of total plot
area, irrespective of number of
individuals
5 75-100 % cover of total plot
area, irrespective of number
of individuals
ordinal
transform
I
2
3
4
5
6
7
8
9
DOMIN
+
I
2
3
4
5
6
7
8
9
10
one individual,
reduced vigor
rare
sparse
< 4 %, frequent
5-10 %
11-25 %
26-33 %
34-50 %
51-75 %
76-90 %
91-100 %
The taxa are listed according to the layer in which they grow.
Plants which appear to be structurally transgressive, i.e. occurring
in more than one layer, have to be recorded in each of these layers
separately.
Taxa occurring only outside the sample plot (but within the
stand) are noted in parentheses. Next, the quantitative occurrence
of each taxon is estimated. In the BRAUN-BLANQ.UET approach two
criteria are considered most useful: abundance and coverage degree.
Abundance relates to the density of the individuals of a given
species in a plot. Cover degree is measured as the vertical projection
of all aerial parts of plants of a given species as a percentage of the
309
total plot area. The term 'dominance,' often used as a synonym for
coverage, is less appropriate.
Abundance and cover degree are usually estimated together
in a single 'combined estimation' or 'cover-abundance' scale. The
five-point scale in Table II, from BRAUN-BLANQUET (1928, see also
article 18.4.1) is in general use. Several authors (e.g. TUOMIKOSKI
1942, DOING 1954, EVANS & DAHL 1955, BARKMAN et al. 1964) used
more detailed scales of combined estimation; the BRAUN-BLANQUET
symbol 2 especially was refined. The more elaborate scales are
useful for special purposes, e.g. for an accurate record of change
of abundance and cover by succession on permanent sample plots
in the course of years (DOING 1954). However, they often suggest
more accuracy than can really be justified. Only values obtained
by one investigator should then be compared (cf. CAIN & CASTRO
1959: 142-143, MAAREL 1966b). We have recently introduced as
a refinement of scale interval 2 (taken from BARKMAN et al. 1964)
the scale subdivisions 2m, 2a, and 2b given in Table II. This
elaboration, which brings the total number of scale values to 9,
has proved to be useful and reliable.
20.4.5.2 Sociability
Sociability or gregariousness is an expression of horizontal
pattern of species. It is a measure of the degree of clustering
(contagion) of the plant units of a species. A plant unit (WILLIAMS
1964) may be an individual or a shoot or a sprout-forming part of
an individual. Measurement of sociability goes back (again) to
HEER (1835).
In the floristic-sociological approach sociability is estimated
with the following scale (BRAUN-BLANQUET 1928, 1932, 1951, 1964).
1. growing solitary, singly.
2. growing in small groups of a few individuals, or in small tussocks
(caespitose), e.g. Corynephorus canescens in shifting sands.
3. growing in small patches, cushions or large tussocks, e.g. Carex
hudsonii as a hummock builder in eutrophic swamps; Silene acaulis
and SaxiJraga opPositifolia in alpine swards.
4. growing in extensive patches, in carpets or broken mats, e.g.
stands of Hedera' helix, Lamium galeobdolon, Asperula odorata etc. in
deciduous temperate forest.
5. growing in great crowds or extensive mats completely covering
the whole plot area; mostly pure populations, e.g. Erica tetralix in
~rica-heath; Sphagnum rubellum or S.pulchrum in raised bogs.
A variant of scale value 5 (5, loose or open 5) was proposed
310
for populations with a cover degree of over 75% consisting, however,
of plants which are sufficiently separated as to leave space for
other species. Similar variants can be used for loose cover of
51-75% (4) and 26-50% (3) (MELTZER & WESTHOFF 1942,
BRAUN-BLANQUET 1964).
In the releve sociability values are written immediately behind
the combined estimation values, e.g. Scirpus maritimus 1.1.
During the last decade several investigators (see FU'KAREK
1964) expressed the opinion that the diagnostic value of sociability
has been overestimated and that a certain sociability degree is a
specific character of most taxa. Other authors (BRAUN-BLANQUET
1964, SCAMONI & PASSARGE 1963, WESTHOFF 1965) do not agree
with this view. Only a few species have a fixed degree of aggregation
based upon their innate manner of growth. The degree of
gregariousness of most species is much influenced by habitat
conditions and competition, and therefore is of major phytosociological
importance. Sociability may also change considerably
during the course of a succession; many examples are given by the
authors quoted above. On the other hand, the variation in sociability
will be correlated with variation in cover degree to some
extent.
Sociability is commonly considered as an expression of
vitality. However in various situations this is not so. JAKUCS (1970,
1972) remarked that character-taxa of the Trifolio-Geranietea
(thermoxerophilous woodland fringe communities or 'Saumgesellschaften')
tend to grow with sociability 1 in their optimal habitat,
whilst they form polycorms with reduced vitality in suboptimal
habitats. Facies of species are often connected with extreme or
disturbed habitats (see 20.6.7).
20.4.5.3Vitaliry and Fertiliry
Further variables in the performance of a species are its
vitality and its fertility, representing vegetative and generative
development respectively. BRAUN-BLANQU'ET (e.g. 1932, 1964)
developed a scale of relative 'thriving' (Gedeihen) with four
categories indicated by symbols (BRAUN-BLANQU'ET 1932).
e, 1 Well developed, regularly completing the life cycle (an extraordinary
vitality is indicated with 'lux', luxurious).
0, 2 With vegetative propagation but not completing the life
cycle.
0, 3 Feeble with low vegetative propagation, not completing the
life cycle.
311
00, 4 Occasionally germinating but not vegetatively propagating.
BRAUN-BLANQ.UET (1964) slightly altered the symbols and gave
them numbers from 1 to 4, as above.
BARKMAN et al. (1964), following VARESCHI (1931) and ZOLLER
(1954), stated that for many species vitality and fertility are independent
or even negatively correlated parameters. They
proposed separate scales for these.
20.4.5 .4 Periodiciry
In addition to vitality the seasonal phase in the life cycle of
each plant, its 'phenological state' is recorded. Various scales are
in use, the first one being that of GAMS (1918). BRAUN-BLANQ.UET
(1964: 67 and 510) presented two scales (which are very similar).
As was stated by GAMS (1918), ELLENBERG (1939, 1954a) and others
(see also the bibliography by BALATOVA-TuLACKOVA, 1970 Exc.)
one incidental record during the releve is really not sufficient, a
complete phenological diagram should be desired for each community.
For a releve at a given time, however, appropriate abbreviations
may be used - v. (vegetative), fl. (flowering), fro (fruiting),
etc.
20.4.6 RELEVE PROTOCOLS
The various structural and floristic data discussed so far are
written in standardized form either in field note books or on special
protocol forms. These notes are preceded by some notes on the
following items:
1. Date; running number; topographic locality (as detailed as
possible); altitude; exposure and inclination; geology of substrate.
If possible the location should be indicated on a detailed map.
2. Size and shape both of the plot and of the entire uniform stand;
character of adjacent vegetation; soil profile; phreatic level; depth
and differentiation of root system.
3. Character and'intensity of human and animal influence, e.g.
pasturing, burning, mowing, treading, manuring, irrigation.
Table III presents an example of a releve, taken from MELTZER &
WESTHOFF (1942). For each species are recorded the scale numbers
fqr cover-abundance (before the period) and sociability (after the
period), the phenological abbreviation, and the vitality symbol.
312
TABLE III
Protocol of a releve (translated from MELTZER & WESTHOFF 1942)
Nr. 39462. 1st August 1939. Terschelling, Bessenplak S of beacon near beach
mark 6. Gridnr. G5.61.43 in IVON-system (Institute for Vegetation Research
in the Netherlands). Stand very uniform, Empetrum heath on slope of 6 m tall
parabolic dune, exposition NNE, inclination 30°.
Habitat: shadowed, moist soil, by day not strongly heated and rarely desiccating.
Slight shifting of sand. Little human and animal influence.
Profile: Ao: 2 cm semi-decayed material.
AI: 5 cm dark humus containing sand.
C: bright, white dune sand.
Sample plot 100 sq m.
Herb layer cover 100 %,20-40 cm
Polypodium vulgare 2.3
Empetrum nigrum 4.4
Hieracium umbellatum 1.1-2
Festuca rubra subvar. arenaria +.1
Hypochoeris radicata +.1
Calamagrostis epigeios +.1
Jasione montana +.1
Carex arenaria 1.1
Ammophila arenaria 2.2
Salix repens +.2
Viola canina var. dunensis r
Moss layer cover 100 %, 2-5 cm
Hypnum cupressiforme var. ericetorum 3.3
Pleurozium schreberi 3.3
Dicranum scoparium 2.3
Mnium hornum 2.3
Lophocolea bidentata 2.2
Eurhynchium stokesii +.3
Plagiothecium denticulatum +.2
Polytrichum juniperinum +.3
Peltigera canina +.2
Parmelia physodes +.1
Cladonia alcicornis +.2
20.5 Synthetical Research Phase
20.5.1 STEPS IN SYNTHESIS
v.
•fro
•fl.
•fro
•fro
•fl.
•fl.
•v. 0
v. 0
fro
•v. 0
v. 0
v. 0
v. 0
v.
•fro
•v. 0
fro
•v. 0
v.
•v. 0
v. 0
The analysis of stands is only the first step in the description of
vegetation units. After releves have been collected, they must be
compared. This is the start of the synthetical phase which leads to
the distinction of coena and, if wished for, the final classification
of syntaxa. To this purpose, a number of releves are tabulated in a
matrix, which is usually named a releve table cir community table.
313
314
environmental
description
dif'ferentiated
table
comparison
with
syntaxonomical
literature
syntaxon
table
synoptic
table
presence
table
comparison
with other
synoptic tables
synoptic
syntaxon
table
Fig. 3. Scheme of steps in the synthetic procedure.
Such a table is also called a primary or raw table (Rohtabelle).
Each primary table is then rearranged into a structured table in
which one or more uniform phytocoena are distinguished and
characterized. After consultation of the relevant syntaxonomical
literature (and possible further revision), this phytocoenon table or
community table may then be presented as a syntaxon table, often
for an association.
When environmental data such as soil analyses are available
the vegetation types arrived at can be characterized synecologically.
Finally the coena, or syntaxa, are checked in the field, e.g. whilst
mapping the vegetation of the area under investigation or during
the reconnaissance of a neighbouring area. This may lead to the
collection of new releves. Thus vegetation classification in the
BRAUN-BLANQ,UET approach is essentially an iterative process, or
one of successive approximation in the sense of POORE (1956). Fig.
3 shows the various steps in synthesis. SHIMWELL (1972) also outlines
in English (cf. KNAPP 1958 and ELLENBERG 1956) the steps
of the procedure.
The checking of units is an essential part of the method; this
point seems not to be recognized by all critics of the BRAUN-BLANQ,trET
approach (e.g. POORE 1955~56). MOORE (1962) explained
that the misapprehension that the fidelity concept rests on a circular
argument (e.g. POORE 1955a) is based on the failure to recognize
the checking step: 'In present practice, associations are not
distinguished in the field at all, but only when editing the tables
of releves. The first step involves describing uniform tracts of
vegetation, not representative stands of a presumed association.
Only when sufficient releves have been accumulated and analysed
one can discern the associations. Of course, in observing continental
phytosociologists at work in their own homeland whose vegetation
they know, one may be misled as to their methodology. They will
now have reached the second stage of checking the reality of units
already distinguished. They will not necessarily make this clear to
a visiting inquirer.'
20.5.2 PRIMARY TABLE
In the primary table all taxa are listed at the left hand side of a
sheet of squared paper; to each releve a single vertical column is
assigned. Each item in any column .should contain either a dot or
dash (in the case that a taxon is absent in the corresponding releve)
or else a combined estimation value (or an actual coverage per
cent) and preferably a sociability value too. It may be convenient
315
at this preliminary stage to group the species under separate
headings according to strata and to separate phanerogams and
cryptogams. The addition of new species from the later releves
produces a characteristic tailing off to the right of the table. Each
row of the table represents a species (in a given stratum). Comparison
of the rows of the table enables us to judge the distribution
of any taxon over the releves; comparison of the columns may lead
to preliminary conclusions on relative similarities of the releves.
The primary table should be rearranged several times, in order
to establish groupings of releves by rearrangement of columns as
well as to group taxa with similar distributions in the table by rearrangement
of rows. Before we discuss this rearrangement procedure
(20.5.5) some concepts and techniques should be discussed.
20.5.3 PRESENCE, CONSTANCY, HOMOTONEITY
Presence is the occurrence of a taxon in a vegetation table. It
is usually measured as a degree by the number of releves in which
the taxon occurs (regardless of abundance and cover) expressed
as a percentage of the total number of releves compared. Presence
degree (Stetigkeit) can thus be calculated from any number of
reieves, regardless of the difference in size of the plots.
When plots of equal size are compared the corresponding per
cent occurrence for a taxon is named 'constancy' (Konstanz).
(Frequency, in contrast with these, is an analytical concept dealing
with the distribution of a taxon within subsamples from a given
stand.) Presence degree and constancy are determined from stands
of different and often widely distributed localities. Constancy
determinations on plots of equal s'ize were especially favored by the
Uppsala school (article 18), starting with Du RIETZ et al. (1920),
although the first such measurements, by BROCKMANN-jEROSCH
(1907) had used plots of various sizes. In the BRAUN-BLANQUET approach
presence degree has been termed 'Stecigkeit' and distinguished
from 'Konstanz'; but confusion has resulted from the translation
of both cO!lcepts as 'constancy' in some statements in English
(e.g. BRAUN-BLANQU'ET 1932, MORAVEC 1971).
Constancy and presence degree can be given in exact percentages,
or in percentage classes. It is usual to distinguish five
classes, noted in Roman figures:
316
Class:
I
II
III
IV
V
Percentage of plots in
which taxon occurs:
1-20
21-40
41-60
61-80
81-100
The numbers of taxa falling into these classes are often presented
in a 'constancy diagram.' Such diagrams are impor'tant characteristics
of vegetation units and provide useful tools to check the
uniformity or homogeneity of a table. This synthetic homogeneity
concept, however, should be distinguished from analytical homogeneity.
BARKMAN (1958a: 316-317) designated these concepts as
'intensive homogeneity' and 'extensive homogeneity.' However, as
DAHL (1970, apud TuxEN, 1970c: 101) pointed out, the latter term
has been designated as homotoneity (NORDHAGEN 1943, DAHL 1957,
1960). This term was long overlooked in the BRAUN-BLANQ.UET approach,
but now receives common use (e.g. TuxEN 1970c, MORAVEC
1971). Homogeneity is an analytic concept, based on comparing
different plots of the same size taken from an individual stand,
whereas homotoneity is a synthetic concept, based on comparing
similar plots from different stands of the same community-type
or phytocoenon.
Homotoneity (see also 20.8.4) has been judged with the help
of constancy diagrams; especially in Nortl}-European approaches
(see further article 18). The classical interpretation, based on
RAUNKIAER'S (e.g. 1934) 'law offrequency,' ofa constancy diagram
is that the following relation between the constancy classes exists:
SI> SII> Sill ~ SlY < Sy (the reversed J-shape). In very
homotoneous tables class V may equal class I (V-shape). When
classes III and IV include more species than class V, the table is
considered heterotoneous (cf. GUINOCHET 1955, CAIN & CASTRO
1959, MORAVEC 1971). According to MAAREL (1972, in MAAREL
& TuxEN 1972: 209), the reasoning of WILLIAMS (1964) and observations
of BARKMAN (1958a) showed that constancy class I is more
or less dependent on the total number of releves. Between the other
classes the following relation exists in homotoneous tables:
Sly+Sy/SII+SIII is slightly over 1, whilst SIIl+Sly+SV/SII is
mostly about 2.
A second feature with which the homotoneity of a table can
be easily checked is the variation in the number of taxa within the
releves of the table. In most cases well-developed stands of a
phytocoenon do not differ much in numb~r of species. A high
317
variance of that number is reason to suppose that the table is
heterotoneous, which may mean that more than one phytocoenon
is represented in it (HOFMANN & PASSARGE 1964). After the releves
have been classed into groups according to their number of taxa,
we may obtain the frequency distributions of these classes, which
will indicate this variation. A table comprising releves belonging
to two different coena may produce a curve with two or more peaks
(KNAPP 1971).
Apart from superficial inspection of homotoneity some simple
measures have been proposed on the basis of constancy figures and
species numbers (see MORAVEC 1971 for a survey). The following
symbols, all corresponding to one vegetation table, are used: M
is the number of releves: Sm is the total number of species and 8
the mean number of species per releve; S., etc. is the number of
species falling into constancy class V, etc. Cj refers to the constancy
per cents for species of the constancy class(es) indicated by the
subscript to the summation sign, or for the prevalent species P those
belonging to the group of species with the highest constancy
values, the number of which (Sp) approaches the average number
of species (8).
Amongst the measures based on constancy figures are:
1. The ratio SyjS[y (DAHL 1957).
1
2. Basic homotoneity coefficient -S 2 Cj (MORAVEC 1971).
m IV+V
1
3. 'Homogeneity value'S I Gj (RAABE e.g. 1952).
p p
ICj
4. Index of homogeneity (CURTIS 1959) p - -
IGjSm
Sm
5. Mean constancy for all species I CdSm
So
6. Corrected homotoneity coefficient (MORAVEC 1971). This is
coefficient 2 with an 'oscillation factor,' determined by the difference
in species number between the richest and the poorest
releve.
TUXEN (1970c) presented curves for various communities
ending in a horizontal line (section 20.4.3); one may derive a
'minimal releve number' from such curves. In fact TUXEN based
his saturation quotient on tables containing about this minimal
number of releves. Other observations suggest that the number of
species grows continuously with the number of releves (e.g. ETTER
1949, BARKMAN 1958a, DAHL 1957, 1960, WILLIAMS 1964). DAHL
(1961) based an 'index of uniformity' on this relation (see 20.8.4)
318
and BARKMAN adapted KLEMENT'S heterogeneity index by omitting
species with a constancy ofless than 10 %.
7. 'Saturation quotient' 100S/Sm (Ti..iXEN 1970c).
As Tiixen mentioned this measure is the reciprocal of the homogeneity
(i.e. heterogeneity) value of DAHL & HADAC (1941).
WHITTAKER (1972) suggested use of BD = (Sm/S) -1 as a measure
of beta diversity (article 3.6). PFEIFFER'S (1957) homogeneity
value was reduced to TUXEN'S measure by MORAVEC. KLEMENT
(1941) used the same measure under the same name as DAHL &
HADAC did, as was mentioned by BARKMAN (1958a), who concluded
that this index is largely dependent on the total number of releves
in the table.
In conclusion we think it would be more realistic to base a
saturation quotient explicitly on species with a minimum constancy
value, e.g. species from constancy classes II-V, and to measure the
rate of increase in total species number separately. For measurement
of homotoneity we suggest the mean similarity coefficient for
the releves (see 20.8.5).
20.5.4 SPECIES WEIGHTS
Species may be weighted in synthetical procedures such as
fidelity determination, assignment of syntaxa to higher units and
calculation of spectra. Weighting is usually with the combined
estimation value or some transformation of this. SCHWICKERATH
(1931) used 'Artmachtigkeit' - and when species groups were
concerned 'Gruppenmachtigkeit' (species and group importance
value, MAAREL 1972b) - based on BRAUN-BLANQUET figures and
arbitrary numerical values for symbols + and r. TUXEN & ELLENBERG
(1937) and BRAUN-BLANQUET (1946) used the Deckungswert
(cover value) and Gruppenwert (group value) by taking average
coverage percentages for BRAUN-BLANQUET values 2-5 and ar':'
bitrary values for 1, + and r. The latter procedure has received
more application than that of SCHWICKERATH, although TUXEN &
ELLENBERG stressed its limited applicability, which was also
criticized by WESTHOFF (1947), -MEIJER DREES (1949), SISSINGH
(1950).
"DAGNELIE (1960) suggested the use of an arc-sine transformation
as is usual with percentage scores. His rounded figures appear
to be identical with the original BRAUN-BLANQUET values. Various
other transformations have been applied, including those of ETTER
(1949), BARKMAN (1958a), BARKMAN et al. (1964), MAAREL
(1966b), SCHMID & KUHN (1970), MOORE (MOORE & O'SULLIVAN
1970) and LONDO (1971). (See MAAREL, 1972b for a review.)
319
Without discussing this problem here (see MAAREL 1972b) we are
in favour of a simple 'ordinal transformation' of the extended ninepoint
BRAUN-BLANQUET scale (Table II). This produces also
partial accordance with the DOMIN scale in its modification by
EVANs & DAHL (1955), which is much used in Scandinavia and
Britain.
20.5.5 TABLE REARRANGEMENTS
The development of phytosociological tables has been stressed
by the leaders of the floristic-sociological approach as a crucial
procedure. E.g. BRAUN-BLANQUET (195la, b) wrote (transl.): 'Appropriately
elaborated association tables are comparable with
thorough species diagnoses. By means of the tables the considerable
detailed work from the minute floristic analyses of the lower
vegetation units becomes accessible and evaluable. From the table
it also appears whether one has worked seriously and reliably;
the tables are the proper touch-stone of the concerned plant
sociologist.'
At the same time the table method is said to be difficult to
learn, at least by written instructions. TUXEN & PREISING (1951)
spoke of 'special aptitudes, a view of the interrelations in the
tabular picture, and broad experience in the sociology, synecology
and syngenetics of plant communities,' that are all needed for
mastering the ordering of tables. Although personal instruction
in the technique is important, as numerous pupils of BRAUNBLANQU'ET
and TUXEN can confirm, it may be admitted that early
BRAUN-BLANQUET sociology paid insufficient attention to explicit
statements of the table techniques.
KNAPp (1948, 1958) and ELLENBERG (1956) presented detailed
accounts of the table technique as it had been developed by TUXEN
in the 1930's at the Zentra1stelle fiir Vegetationskartierung. ELLENBERG'S
scheme (1956: 46, slightly altered) is as follows (see Fig. 3):
(1) rearrangement of the first matrix to a 'presence table';
(2) tracing of table-differentiating species, mostly with the aid of
'partial tables';
(3) rearrangement of taxa and releves according to the empirically
established occurrence of groups of differentiating species; the
result is a 'differentiated table';
(4) compilation of every relevant differentiated vegetation table
into a 'synoptic table' ('Vbersichtstabelle');
(5) determination of fidelity by comparing an adequate number of
320
synoptic tables ('zusammenfassende Dbersichtstabellen' in the
sense of ELLENBERG l.c.);
(6) rearrangement of the differentiated table into a final syntaxon
table ('charakterisierte Tabelle' in the sense of ELLENBERG l.c.).
Similar schemes were presented by MOORE (1962), SCAMONI &
PASSARGE (1963), FUKAREK (1964), KUCHLER (1967), GOUNOT
(1969), KNAPP (1971) and SHIMWELL (1972). The steps 1, 2
and 3 are the basic ones; they are essential for any local classification
of releves into phytocoena. Step 4 is preliminary to 5.
Steps 5 and 6 have to do with the next level of investigation:
the establishment of a (perhaps provisional) syntaxonomical clas-
sification.
TUXEN (1970b) recommended to start with a preliminary
procedure aiming at the exclusion of releves with deviant numbers
of species that suggest fragmentary or heterogeneous stands. Such
releves are thus excluded from successive rearrangements and
fidelity calculations and are not considered in an association
diagnosis.
The first rearrangement of a primary table aims at listing the
species after their degree of presence. In the presence table it may
become apparent, whether the releves vary irregularly, or whether
certain combinations of species are found recurrently. In the latter
case such groups may be more or less mutually exclusive and serve
as groups of differentiating species. Those are not likely to be found
in the presence classes V and I; they will mostly belong to the intermediate
classes II, III and IV. The supposed differentiating
species are underlined or boxed in the presence table. These differentiating
species, which are provisional sets of diagnostic
species (see 20.3.4), are used to rearrange the table.
The next step is to set up a partial or extracted table in which
only the species thought to have diagnostic value are given and used
to rearrange both columns and rows. During the rearrangement
of a provisional partial table, the differentiating species are written
on the left hand side of the paper and the rows arranged in their
corresponding groups, and the releves (columns) are rearranged
to show a diagonal order of species groups from the left to the right
of the table. To simplify the rearrangement of the releves dictation
or transfer strips are used.
When the partial table has been worked out, the complete
table (thus containing again the temporarily omitted species) IS
rearranged accordingly so that the differentiating groups appear
clearly. Now we have the 'differentiated table.' The species with
a high presence values indicate the degree of coherence or homo-
321
toneity of the table. The species with intermediate and lower
presence values used in differentiating groups represent the
heterotoneity ofthe table. After comparing the sizes ofthe two groups,
one may decide how many phytocoena the table includes and
whether the table should be divided, or combined with others for
further treatment.
Step 4 from ELLENBERG'S scheme comprises the replacement of
each uniform table or partial table by a column in which for each
participating species the presence degree is indicated (either as a
percentage or a class value). Such a table (which may consist of
only one column) is called Dbersichtstabelle by ELLENBERG but
is also known as Sammeltabelle, synthetische Tabelle and Stetigkeitstabelle.
Following a recent denomination by DOING (1969a)
and TUXEN (1970b) we propose the neutral name 'synoptic table.'
After comparison of the synoptic table with those from other
types of vegetation from the same region an idea can be formed
about the local diagnostic species groups in the table under study
(step 6). The differentiating groups used may represent recognized
diagnostic groups, or may represent approximate groupings that
should be revised on the basis of broader knowledge of diagnostic
groups, or may repre'lent diagnostic groups for newly recognized
phytocoena. With those species, as well as the constant taxa, a
phytocoenon may then be identified or described. Here we could
speak of a synoptic phytocoenon table, from which the corresponding
releve-table or 'phytocoenon table' (Gesellschaftstabelle)
can be rewritten. When moreover the synoptic table is compared
with the literature and with tables from similar vegetation types
from other regions, character-taxa may be found and a syntaxonomic
interpretation may follow. The synoptic table now
serves as a synoptic-syntaxon-table and the corresponding (rewritten)
releve-table is now a syntaxon-table. The latter will often
be an association table. 'The fidelity research implied in steps 5 and
6 will be discussed in 20.5.7-8.
The rearrangement method might be criticized with the objection
that the table pattern may be 'an artefact imposed by the clever
shuffling of data and not mirroring any reality in nature,' as it
has been expressed by MOORE (1962) in his refutation of this objection.
In reality, the characterized table is, first, an effective organization
of information, an ordering of species and releves in relation
to one another (and biotope properties). Second, it is considered
a working hypothesis to be tested by further observation. It should
receive confirmation: (i) from furcher floristic data, critically and
impartially collected; (ii) from mapping the units distinguished;
and (iii) from the evidence ofrelationships between the vegetational
322
units and environment. The groups of differentiating taxa may be
presumed to indicate ecological differences in one or more specific
major biotope factors or correlated sets of factors, such as phreatic
level, humus, carbonate content, amount of soil hardening; salt
content, or frequency and duration of periodical inundation by sea
water; exposure; shade; rate of pasturing or mowing; etc. The
resulting hypotheses have to be checked by continued ecological
research. If the work has been well done, the differentiating groups
should have indicator value in relation to environmental factors
(see 20.7.6), and the phytocoena as kinds of communities should
bear consistent, reliable relations to kinds of biotopes.
20.5.6 MECHANICAL MEANS FOR REARRANGEMENT
The 'classical' rearrangement technique by manual ordering
of taxa and releves, as it has been rendered above, is a timeconsuming
procedure and a potential source of errors. Attempts
to simplify and mechanize the procedure have been made by several
investigators. The most simple means is the use of cardboard strips,
usually for the releves and thus for the columns ofthe table.
Various systems of blocks (wooden, plastic, or aluminium) have
been developed (WILMANNS 1959, MARGL 1967, MULLER et al.
1972). TRENTEPOHL (1968) showed a magnetic device which
seemed to be less vulnerable to manual disordering of the blocks.
For tables with a large number of releves or a large number of
species the use of punched cards is obvious. Three types of cards
are in use - the edge-punched card, the visual punched card, and
the machine punched card. COTTAM & CURTIS (1948) initiated
use of cards with the third and most advanced type. Examples of
the use of edge-punched cards are found with EMBERGER et al.
(1957), GOUNOT (1957) and ZONNEVELD (early sixties, internal undated
reports Netherlands Soil Survey Institute).
ELLENBERG & CRISTOFOLINI (1964) intensively used visual
punch cards, mainly for sorting releves. The corresponding cards
are laid on a light-table and then compared with a standard card e.g.
one containing the characteristic species combination of a
certain syntaxon. ELLENBERG (1968) published a version of this
technique where combined estimation values of species are taken
into consideration, however at the expense of space on the card.
According to ELLENBERG, this apparatus is only efficient if the
number of releves is large, at least 200 to every.table. As MULLER et .
al. (1972) stated, this technique may be useful where the vegetation
has already largely been classified and the vegetation units de-
323
suibed, and where additional releves need only to be arranged and
included in existing tables. Further use of machine punched
cards in phytosociology has been described by DAGNELIE (1960),
BECKING (1961), GROENEWOUD (1965) and many others; whilst
recently a number of computer-based table rearrangement programmes
have been developed; this will be discussed in 20.8.1 and
20.8.6.
20.5.7 DETERMINATION OF FIDELITY
The next step in the synthetic procedure is the determination
of fidelity. By comparing a number of tables - if possible all
available tables covering the vegetation of a given area - it is
possible to discern character-species in the sense discussed in 20.3.3.
As indicated there, the criteria are the distribution of the species
in different coena; fidelity is the degree to which a species is concentrated
in one coenon, vs. dispersed with more even occurrence
in several coena. Species distributions iri the tables are observed on
the basis of measures already discussed - presence degree, combined
estimation value, vitality, and sociability. It should be
emphasized that these measures are relevant to a species' fidelity only
as they differ between different coena. Character-species can as
well be minor species as major ones; there is no necessary relation
between fidelity and dominance. In the classification of European
forests, more significance is attached to the fidelity of shrubs,
herbs, and mosses, both because tree species are often widely
distributed in different coena, and because disturbance has brought
extensive replacement of natural tree species.
Five degrees of fidelity have been distinguished (BRAUNBLANQ.UET
1921, 1928, 1932, 1951, 1964) in the relation of a species
to the given coenon.
A. Character-taxa.
Fidelity degree 5: Exclusive taxa (treue): taxa completely or
almost completely confined to one phytocoenon (vegetation
unit) ;
Fidelity degree 4: Selective taxa (feste): taxa occurring with clear
preference for one phytocoenon but also, though with a considerably
lower presence degree, in other coena;
Fidelity degree 3: Preferential taxa (holde) : taxa present in several
coena, perhaps with about equal presence degree, but with a
higher combined estimation value and (or) with a higher
vitality degree in one particular coenon;
324
B. Companions.
Fidelity degree 2: Indifferent taxa (vage): taxa without pronounced
preference for any coenon.
C. Accidentals.
Fidelity degree 1: Strange taxa (fremde): taxa having a definite
presence degree optimum and mostly also a cover-abundance
optimum outside the considered coenon. These are often accidental
intruders from neighbouring coena or relics from a
coenon that preceded in succession.
TABLE IV
Determination of fidelity according to SZAFER and PAWLOWSKI
(BRAUN-BLANQ.UET 1932)
F = fidelity degree;
A = cover-abundance combined estimation;
C = presence or constancy class;
V = vitality.
F in phytocoenon under consideration in comparable phytocoena
C A e A
5 I-II +-2(1)IV-V 3-5
4
3
2
I
IV-V +-2
I-III +-5
IV-V 3-5
IV-V +-2
III-IV +-2
I-III +-2
I-V 3-5
C, A various V normal
C, A, V· various
I +-1
V reduced
- outskirts and disturbed
parts of the stand(s) I
+-2
absent or very
rare
II-III (rV) +-2(1)
(relic or pioneer)
II-III +-1 (2)
I-II (III) +-1 (2)
I (rare) +
I-V +-2
C, A lower V reduced
similar
higher
As early as 1927, SZAFER & PAWl.OWSKI gave a quantitative
outline for the determination of the fidelity class. It has been taken
over, with slight modifications, by BRAl1N-BLANQ.l1ET (1928, 1932,
1951, 1964) and it is presented here with slight alterations (Table
IV). In this table the comparison is between a phytocoenon that
is being studied and comparable' phytocoena, It seems logically
incorrect to specify in such a table 'the given association' versus
'other associations;' as BRAl1N-BLANQ.l1ET (l.c.) has done in all
editions of his manual.
325
BECKING (1957) has modified the scheme of SZAFER & PAWLOWSKI
(1927) considerably, especially in omitting the possibility
of obtaining fidelity degree 5 with a low presence degree. However,
it has been crucial in the BRAl1N-BLANQUET approach that faithful
species need not be constants, and apart from diagnostic implications
(see 20.6.1) one cannot simply change this principle.
20.5.8 GEOGRAPHICAL EFFECTS ON FIDELITY
Since the beginning ofthe BRAUN-BLANQUET approach (BRAUNBLANQUET
1921) it has been recognized that the diagnostic value of
character-taxa is geographically limited. Naturally enough this
recognition has become more and more general with the growing
geographical extension ofphytosociological knowledge. The vegetation
of a region with a uniform climate and a uniform geological
history usually shows great uniformity in the ecological amplitudes
and therefore in the sociological positions of its species. Most
species however occur in larger, climatically and geologically
heterogeneous areas and present different ecological amplitudes
or habitat relations in different parts of their areas. In many cases
these differences in local habitat response within the species are
correlated with genetic differences: weare dealing then with
ecotypic variation. In other cases genetic differences, if present, are
not evident. Geographic variation in habitat response may involve
such shifts in topographic or other local distribution, as tend to
compensate for difference in climate (law of relative habitat constancy,
cf. WALTER & STRAKA 1970, article 3.2).
A number of species which behave as xerophytes in Central
Europe, e.g. Bromus ereetus, Koeleria gracilis, Carex humilis, Peucedanum
cervaria, are bound to humid soils of cool northern exposures in the
Mediterranean. Species which are real woodland plants in a
continental climate may present much wider amplitudes in oceanic
climates: e.g. Osmunda regalis, Listera ovata, D~yopteris jilix-mas.
On the contrary a number of thermoxerophilous species are indifferent
to soil conditions in Central Europe, whereas in the cooler
and more humid climate of NW-Europe they are strictly confined
to calcareous soils. Iris pseudacorus, a species ofwet eutrophic swamps
in Central and W~st~rn Europe, grows in moderately damp grassland
in the perhumid Western Irish climate. Schoenus nigricans, in
Central Europe faithful to calcareous marshes or fens, occurs in
Ireland both in fens and in ombrotrophic blanket bogs. Even within
a much more limited geographical range the differences in ecological
amplitude may be considerable. Within The Netherlands,
326
Artemisietum mari timae typicum
Ini tial phase with Artemisia maritima
,) 5)
Number of releve 53136 53134 53133 52390 52371
Locality l) Ka Ki Ka Ka Ka
Habitat 2) cr cr cr dike dike
Salinity flood-water3) p p p
Date 2-9-53 2-9-53 2-9-53 21-8-53 20-8-52
Surface in m2
Coverage in "
Olaracter-taxon of the association
Artemisia maritima
2xI2 2xl2 3xl0 2!x20 4xI2
98 95 9> 95 100
,-- --- - - - - - - ---- - - - - --,: 5.4 4-5.4 5.4 4.5.5 4.3-4:
L __________________ • ____I
Differential-taxa of the subassociations
Armeria maritima
Agrostis stolonifera var. compacta
subvar. salina
Atriplex hastata
Character- taxa Armerion mari timae
Festuca rubra f. li toralis
Glaux maritima
Juncus gerardii
Parapholis strigosa
Character-taxa Puccinellion maritimae
j-------- -,
: 2-3.2 3.3 t
L ___ • ______ • .:
-.2 +.2 1.2-3 +.2
+.2 +.2 +.1-2
Typical form
52338 54081 53022 55!
He Ko Ba
cr cr ab
5-8-52 13-8-54 5-8-53 5-8-
2x25 3x20 3xI5 3)
100 100 100
2.1 2-3.1-2 2.1-2 2.:
5.5 5.5
_ .1
5.5
+.1-2
Puccinellia maritima
Halimione portulacoides
Bostrychia scorpioides
:2.2-3 - 2-3.3 - 2~i:-3: -.1-2 1.1-2 +_1-2 1.1-2l _ _ _ _ _ -.oJ +.2 1· ~
+.2
Olaracter-taxa Glauco- Puccinell ietal ia
Spergularia marginata -.1 1.1 1.2 +.1
Limonium vulgare ssp. vulgare +.2 +.1-2 +.1-2 1.1-2 -.2 1.1
Character-taxa Asteretea triEolii
Aster'tripolium +.1 +.1-2 +.1 +.1 -.1 1.1-2 2.1-2 1- ~
Triglochin mari tima -.2
Plantago maritima -.2 +.2 -.2 +.1 +.1-2 +.2
Character-taxa Thero-Salicornionl Thera-Suaedion
and SEartinion
Salicorni a europaea
Suaeda mari tim. +.1 +.1 +.1-2 ·.1
Spartina townsendii _.2
Other taxa
Elytrigia punaens +.1-2 1.1-2 1-2.1
Atriples li ttoralis -.1
Lolium perenne
1) Localities Ka + Kaloot near Borssele (South Beveland);
He • "Slikken van de Heene" (St. Philipsland);
Ko • Marshes near Kort,ene (North Bevlland);
I. • Mar,hes near Baarland (South Beveland) •
Sp • Spiering5chor near Kallperland (North Beveland);
Kats • Marsh near Kah (North Beveland);
Wa • Marshes east of Waarde (South Beveland);
lath • Marshes west of Bath (South Beve1and).
Os • Marshes vest of Ossendrecht and
Li • ItGalleschoor" north of Lilloo (Belliua).
IS
l.
or
;5
12
10
·2
2
2
:z
2
2
Artel'lisietUlll lIariti.ae Arte.isietu. uritilUe .,rostidetosWl
8l'1HrietosUlll Typical fo.... Frapent
52343 54092 54089 52051 53043 53054 52207 53071 54130 54133 54139 54152
He Sp Sp Kats Wa W. Bath lath Os O. O. Li
ac cr cr cr ab ab cr cr cr ac or a.
p m II III III
• • •
5-8-52 14-8-54 14-8-54 4-9-52 6-8-53 6-8-53 3-7-52 6-8.53 2-9-54 2-9-54 2-9-54 2-9-54
100 3x25 4.20 2!xl2 5xl0 6.8 l!xl5 2xl0 5x50 IOx20 2xl5 4x20
95 100 100 90 100 100 100 100 100 100 100 100
r---------- ... ---- ----,
1-2.1 2.1-2 2.1-2 2.1 3.1-3 3.1-2 2.1-2 3.1-2 ' ,, -.:.._ ... ____ ... _________ ..J
11- 2. 2 1.2 1.2 +.21
4.5 5.5 2.2 2.2-3 3.2-3 3.5 1.2 3.5
1.1 1.1 1.1 +.1 1.1 2.1.2 1.1.2
5.5 5.5 5.5 5.5 3.5 2.5 5.5 5.5 4.5 4.5 5.5 4.5
1.1 +.1 1.1 2.1-2 1.1-2 +.1
+.2 +-1.1
1.1-2 +.1-2
+.2 1.2
+.1-2 1.1-2 +.1-2 1. 2.1
.-- - - - - - - - - -- - --,
1.1 1.1 +.1 +.1-2 +.1-2 +.1 I
- I
I
1.1-2 2-3.2 2.2 +.1-2 +.2 2.1-2 +.2 I
- II ,___________
------..1
2.1-2 +.1-2 +.1-2 +.1-2 +.1 1.1-2 +.1 +-1.1 +.1 2.1-2
+.2 +.2 1.2 +.1-2 +.2 +.2
2.2 +.2 +.2 1.2 2.2 1-2.2 +.2 +.1-2 +.2 +.1·2
+.10 +.10
1.1
+.1
+.1-2 +.1 +.2 +.1-2 +.1-2 +.2 +.2 +.2 2.1·2
+.2
2) Habitats cr "on elevated creek banks i
dike. at the foot of the dike, whe.. tidal drift 11. washed ..hore;
ab • on elev_ted parts with an abrasion .d,.;
ac III on hi.hly accreted parts in the ..rsh.
3) Salinity flood water : e • euhalinei
p • polyhaUne and
• ••sohaline.
It) Varian.t with Mali.ione portulacoid••
S) Variant with Atrlplex hastata
Orchis morio is faithful to the Meso bromion of dry chalk in Southern
Limburg, whereas it is bound to moorland (CallunoGenistetum
molinietosum) in the Pleistocene part of the
country, to mesohalinic peat soil (community of Orchis morio and
Ophioglossum vulgatum) in the brackish Holocene parts of Holland
(sensu stricto) and to a narrow ecotone at the margin of wet
heather dune slacks on the West Frisian islands (cf. WESTHOFF &
HELD 1969). Hymenophyllum pellatum is in the major part of Ireland
bound to damp oakwood, but on the extremely oceanic cliff of
Slievemore on Achill Island (Co. Mayo) it thrives freely in the open
air in between Calluna vulgaris and Empetrum nigrum (PRAEGER 1934).
Consequently, one and the same species can be a charactertaxon
for association A in one area and for association B in another
one. BRAUN-BLANQ.UET (1964: 100) concluded that the validity and
diagnostic value of character-taxa for a given association are
mostly restricted to a climatically uniform area. It follows that a
typology of character-taxa with respect to their geographical extension
of their validity may be useful.
BRAUN-BLANQ.UET (1928, 1932) gave a first distinction be[ween
general and regional character-taxa. Later (1951a) he mentioned
absolute, territorial and local character-taxa (cf. ELLENBERG 1956).
The distinction between territorial and local was unclear, however;
apparently it refers to the size ofthe range ofthe association in which
the taxon shows fidelity. As BARKMAN (1958a) pointed out the
replacement of general by absolute suggests a change of criterion,
i.e. from range of fidelity to degree of fidelity, which was in fact
realized in BRAUN-BLANQ.UET'S definition of 'Absolute Treue.' With
BARKMAN we think that these two criteria should be used independently.
A third criterion is the relation between the range of
the character-taxon and the range of the corresponding syntaxon.
MEIJER DREES (1951) elaborated this criterion and came to
four possibilities: 1) ranges of taxon and syntaxon coincide, 2)
taxon range forms part of syntaxon range, 3) syntaxon range forms
part oftaxon range and 4) ranges differ but overlap. MEIJER DREES
maintained BRAUN-BLANQ.UET & MOOR'S term 'regional' which he
applied to all cases in which the taxon is a character-taxon throughout
the common range of taxon and syntaxon, contrasted with the
term 'local' for cases in which the taxon is a character-taxon only
in part of the common area.
The corresponding scheme was taken over by BECKING (1957,
1961) who incorporated it moreover in a complex scheme in which
also the degrees of fidelity were considered. BARKMAN (1958), who
wished to maintain the more obvious meaning of the terms local and
regional, adopted the eight types of MEIJER DREES (1951), but
327
largely renamed them. Finally KNAPP (1971), who only mentioned
MEIJER DREES and BECKING, refined the division into four types
of geographical character-taxa in the taxon-bound sense of
MELTZER & WESTHOFF (1942): viz. 'local', 'territorial or regional,'
'region-' and 'absolute.' The following scheme may be given as a
summary:
i) Local character-species, with fidelity restricted to an area
that forms only part of the total area of the syntaxon.
ii) Regional character-species, with fidelity for the total area
of the syntaxon, but with the species area exceeding that of the
syntaxon. Subcategories could be distinguished in relation to BRAUNBLANQ,UET'S
phytogeographical typology including provinces and
sectors.
iii) General character-species, with fidelity for the total area
of the syntaxon, and the species area coinciding with the syntaxon
area.
20.5.9 PHYTOCOENON CHARACTERIZATION
Mter a primary table has been rearranged into a phytocoenon
table, presence values have been calculated, and the fidelity and
differentiating value of the taxa towards neighbouring communities
have been established, the corresponding phytocoenon can be
characterized. First the general structure can be described. Mostly
the presence class of each species is indicated at the right hand side
of the table, in some cases the presence per cent. To the right of
the presence figure the range of the combined cover-abundance
estimation values may be given.
Then, according to procedures described above the faithful,
differentiating and constant taxa may be indicated. The diagnostic
value may be only provisional, when the releves come from a
limited area or when insufficient other material could be compared.
We are dealing here with the first of three phases of systematic
phytosociology, viz. the local phase. The taxa that are oflowest
presence degree and lack diagnostic value (mostly of presence
class I) are often presented in an addendum to the table. In many
cases the description of phytocoena does not immediately lead to a
formal classification. Without this, the phytocoenon tables may
summarize much of what phytosociology seeks to understand of
relations amongst species, kinds of communities, and environments.
When data are thus organized into phytocoenon tables, however,
th~ stage is set for the syntaxonomic treatment next to be discussed.
328
20.6 Syntaxonomical Research Phase
20.6.1 SYNTAXON TABLE CHARACTERIZATION
The syntaxonomical research phase starts when a phytocoenon
is to be fitted into the hierarchic system of syntaxa. The phytocoenon
table is then interpreted by consulting the relevant syn..
taxonomic literature, especially synoptic syntaxon tables. Questions
in this are: (i) Which already described association can be recognized
in the characteristic taxon combination of the phytocoenon;
(ii) which lower units could be recognized on the basis of
the established differentiating taxa; (iii) which taxa can be
recognized as character- or differential-taxa from higher units already
distinguished?
Careful treatment ofthese questions may lead to the conclusion
that the phytocoenon under study is one already recognized, or
that it must be described as a new syntaxon, or that part of the
existing hierarchy must be redefined. Here, as ELLENBERG (1956:
57) said 'begins the range of the 'tact' of which BRAUN-BLANQ.UET,
1951: 18 spoke.' The expression 'sociological view' ('soziologischer
Blick') is relevant. Though some of the decisions on phytocoenon
similarity are objective, the essential role of judgement and
experience in a typological discipline should be recognized.
The results of these considerations are expressed in the structuring
and labeling of the phytocoenon table, which now becomes
a syntaxon table. Usually the highest unit treated in a given
syntaxon table is the association. In such an association table the
subordinate units distinguished are presented as well. Table V
presents an example of an association table ofan already recognized
association, viz. the Artemisietum maritimae, as it occurs in
a region not systematically described before, the southwestern
Netherlands.
When a number of related associations have been described,
their regional descriptions might be presented together in a synoptic
syntaxon table. An example ofsuch a table is presented in Table VI.
Both tables, composed by W. G. BEEFTINK (cf. BEEFTINK 1962,
1965, 1968), whose cooperation is gratefully acknowledged, will
be discussed in the subsequent paragraphs.
20.6.2 CHARACTERIZATION OF THE ASSOCIATION
As was discussed in 20.3.2 the association concept of the
BRAUN-BLANQ.UET approach was always based on the presence of a
329
characteristic taxon combination, including character-species.
However, as the number of described associations grew, the role
of character-taxa gradually diminished whilst the diagnostic importance
of differential-taxa increased. In some association descriptions
character-taxa are not mentioned separately but are given
together with the differential-taxa ('Kenn- und Trennarten'). This
led to the character-combination concept which in fact recognized
the possibility that none of the participating taxa is a charactertaxon
in a strict sense, but rather the correlated occurrence of
taxa is the essential 'character' of the phytocoenon.
BRAUN-BLANQ.UET (1964: 122) seemed to confirm this development
when he stated that 'the essential association features'
should be found in a stand in order to assign it to that association
and then defined this as follows: 'primarily the normal characteristic
species combination should be present, that is a minimum
number of character- and differential-species and some of the more
important companions.' However, during recent colloquia and
symposia of the International Society for Plant Geography and
Ecology the presence of at least one character-taxon was demanded
by others, especially by OBERDORFER (1968); see also the following
discussions, during the 1964 symposium at Stolzenau (TuxEN
1968: 132-141) and DIERSCHKE (1971). DIERSCHKE (1971) even
stated that 'the basic unit for the Prodromus is the association, the
rank of which is determined by the constant occurrence of at least
one character-species and which is defined by its characteristic
species combination.' This definition may be considered the
standard from which, however, vegetational circumstance may force
some departures.
20.6.3 GEOGRAPHIC PROBLEMS WITH ASSOCIATIONS
Many of the problems with character-taxa result from complexities
in the geographic behaviour ofspecies in relation to coena,
as discussed above (20.5.8). The difficulties led ELLENBERG
(1954b) to speak of 'the crisis of the character species doctrine' and
to propose a much more local delimitation of associations without
general character-species. As ELLENBERG (1956) added, the use
of geographicaHy restricted 'local character-species' for associations
is then appropriate. The local establishment of associations
emphasized by ELLENBERG (1954b) had, in fac.., been common
practice by then for some 20 years, as TUXEN (1955) stated in a
reply to ELLENBERG, whilst referring to BRAUN-BLANQ.UET & MOOR
(1938) and TUXEN (1937). In fact OBERDORFER (1957, but also
330
Table VI
Clusification of salt6 11arsh communities in the SW Netherlands
accordin, to the Braun-Blanquet method by W.G. Beeftink
Classes Thero-Sal icornietea Spartinetea
Orders Thera-Sal icornietal ia Spartinetalia
Alliances Thera-Sal icornion Sparti nion Puccinellion maritimae
Associations Sal icornietum Spart inetum Spartinetum .Puccinellietum Halimionetum
strictae mari t imae townscndii Ilaritimae portulacoidis
CalUlln
Number of releves 14 24 30 124 40
Character-taxa of the associations
Salicorni. eUTopaea coll. l ) 1100 (1-3) 1 14 (>-2) 78(+-2) 40(+-1)
Spartina maritima 14(+) 6(+)
Fuclls vesiculosus vaT .lutarius 7(2-3) 4(+-2)
Spartina townsendii agg. 86(+-1) 79(+-2) 1100 (3-5) 1 50(+-2) 35 «-2)
Puccinelli. lIaritima 2) 25(+-1) 33( +- I) 30(+-1) 1100 (3-5) 1 92(+-2)
HaliJIIlOne portulacoides 8(+) 33(r-l) 83(+-2) 1100 (3-5) 1
Arteaisia lIaritiJlla 5(+-1)
Armeria .arit111. 6(+-1)
Carex extensa
Puccinellia distans
Puccine) lia fasciculata
Puccinell ia retroflexa
Scirpus lIaritillus var. compactus 3) 27(+-2)
Faithful taxa Puccine11ion maritimae
Bostrychia scorpioides 17(+-4) 14(+-2) 21 (+-4) 37(+-4) I
Character-taxa Arllerion mari timae
Juncus eerardi i 2(+)
Festuca rubra f Iitoralis 14(+-2) 62(+-1)
Glaux .ariti.a 10(r-l) 50(+-2) 15(.-1)
Parapholis strieosa
Agrostis stolonifera var. compacta
subvar. salina
Character-taxon Puccinel1 io-S~erlu1arion salinae
Spergularia salina
Character-taxa Glauco-Puccinel) ietalia
Sperlularia .arlinata 7(r-+) 73(+-2) 42(+-2)
LiltOniwn vulgare ssp vulgare 8(+) 8(+) 30(r-+) 73(+-2) 60(.-2)
Character-taxa Asteretea tri20lii
Aster tripoliu.
Trie10chin uriti.a
Plantalo .aritilla
Other taxa
Suaeda uriti.a
A.triplex hastata
Elytrieia puneens
Lolh. perenne
Plantaao coronopus
'hrae-ite! c~nis
50(+-2) 33(+-2) 77(r-2) 98(+-2) 97(+-2)
8(+) 27(r-2) 86(+-4) 62(+-2)
20(r-l) 65(+-4) 65(+-2)
50(+-1) 12(+-2) 37(+-1) 64(+-2) 60(+-2)
77(+-2) 25(+-2) 5(+)
7(r-+) 1(+) 22(+)
~: Collmln I: Zostera noltH 29(+-2) j Column 7: CentauriUJII pulchellum 17(+.2). Carex
CentauriUII pulche11um 57(r·2). Carex distans 29(r-2), Juncus maritimu$ 29(r-l),
rua 14{r), Sonchu5 arvensh 14(r). Trifoliwn repens 14(r). 'centaurium littorale
22(+-1), Pountilla anserina 8(+-1), Plantago lIajor (24+-2), Leontodon autumnalil
caria ;,nodora 11(+)°, Bromus mollis 8(+), Ranunculus sceleratus 16(+_1)°, Poa am
CirsiUII arvense 5(+-1), Poa trivialis 3(3), HordeUII secalinWl 3(1), TaraxacUII sp
3(+), Senecio vulgaris 3(+), Anagallis arvensis 3(+), Leontodon nudicaulis 3(+);
tricaria inadora 10(r)0. Brollus lIollis 10(+), Sagina lIaritima 40(+.2), Juncus buJ
ColUlln 11: Bromus mollis S(r); ColUlln 12: Ranuncu1us sceleratus 5(+), Cochlearia
1. In the alliances Thero..Salicornion and Spartinion represented by S. stricta Dua.
2. Preferential character·taxon of the association; also selective character taxon of the
the order Glauco-Puccine11ietalia.
3. A.lso character-taxon of the alliance HJllo~Scirpion.
Asteretea tripolii
Glauco-Puc.cinell ietalia
on maritimae Armerion maritimae Puccinell io~Spergularion salinae Halo-Scirpion
etum Halilllionetum Artemisietum Juncetum Junco-Caricetum Puccinellietum PuccinellietUIII PuccinellietWi Halo-Scirpetwa.
portulacoidis maritimae gerardii extensae distantis fasciculatae retroflexae
10 11
40 61 64 37 10 12
40(.-1) 30('_1)° 42('-1)° 14(')° 35('_2)° 70(r-2)0 100(1-2)°
35(.-2) 13(') 20(.-2) 8(.)° 17(r)
92('-2) 43('-2) 31 ('-2) 14(') 62('-1) 90(r-3) 25(r-l)
1100 (3-5) I 98('-3) 62('-1) 5(.)
~ 59('-1) 8('-1)
2(·) 1 90('-3) 1 14(r)
- 1100(1-4))
1100 (1-5) I 20(r-2)0 17(r-')
17(r)1 100 (2-4) 1
1100(l-4) I
32 ('-2) ° 10(.)°
37(.-4) I 5(·-2)
86(.-5) 100(2-4) 14(.-2) 30(.)
62(.-1) 100(3-5) 97(.-5) 86(.-3) 35(.-3) 10(r)
15(.-1) 33('-1) 98('-3) 100(2-3) 35('-3) 30(r-2)
5(.-1) 64(.-3) 43('-2) 19('-2) 10(.)
11 (.-1) 100('-3) 65(.-2) 30(.-2)
197 ('-3) 80(·-2)
42('-2) 72(.-2) 47('-1) 14(') 19('-1) 20(1-2)
60(.-2) 66('-2) 91 (.-3) 57(r-2) 3(+)0
97(.-2) 100('-2) 72(.-2) 71 (r-I) 65(.-3) 100(r-3)
62(.-2) 29('-1) 77(.-2) 29(.) 5(.-1) SO(r-2)
65(.-2) 72(·-3) 98(.-3) 100(.-2) 11 (.)
60('-2) 31(.. 1) 20(+_1)° 38(+_1)° 30(r-I)0
5(.) 11 (.) 5(')° 62(._2)° 40(r)0
22(.) 31 ('-2) 28('-2) 30(.-2)
2(.) 30('-3) 10(r)
2(.) 29(') 11('-1) 10(1)
2(.) 43(r-+)0 30(.-3) 10(r)0
11um 17(+-2), Carex distans Set), Sagina maritima 2(+), Solanum dulcamara 2(+); ColumnS:
naritirnus 29(r-I), Lotus tenuis 29(2), Hippophae rhamnoides 29(r-+), Trifolium fraeiferltaurium
littorale 14(+)0; Column 9: Polygonum aviculare 41(h2), Elytrigia repens
l.eontodon autUlinalis 5(+), Trifolium repens 11(+)°, Coronopus squamatus 8(+-2), Matri,Js
16(+_1)°, Poa annua 11(.-2), Cochlearia officinalis 11(+), Festuca arundinacea 8(+),
3(1), TaraxacUII sp. 5(+), Sonchus arvensis 3(+), Poa pratensis 3(+). Solanum nigrum
on nudicaulis 3(+); Column 10: Centaurium pulchellum 10(+). Plantago major 20(r)o. Ma-
40(+-2). Juncus bufonius 40(r-2), Hordeum marinum IO(r). Samolus valerandi IO(r);
JS 5(+), Cochlearia officinalis 5(+). Atriple~ littoralis 5(+).
stricta DUll.
racter taxon of the alliance Puccinellion lIaritimae and exclusive character-taxon of
8(r)0
8(.)
100(r-2) I
8(')
100('-4)
8(r)
8(')
8(1)°
2S(')0
67(r-2)0
maritimi
12
19
37('-2)
26(1-3)
1100(3-S)1
5(.)
11(')
42(.-4)
74('-2)
16('-1)
63(1-3)
32(.-2)
11('-2)
1968 and OBERDORFER et al. 1967) also worked with local character
species, though always in close connexion with alliance character-
speCles.
This local association approach as such has been criticized
from another point-of-view by SCHWICKERATH (1942, 1954a,
1963). This author emphasized that geographically differentiating
taxa should be used in the delimitation of syntaxa, but whenever
a series of geographically different syntaxa should have one or
more character-taxa in common they should be grouped together
into one association. SCHWICKERATH especially criticized the splitting
up of the Xerobrometum by BRAUN-BLANQUET & MOOR
(1938) .
To solve the problem use of different terms for more localized,
vs. more widely distributed, phytocoena has been suggested.
KNAPP (1942, 1948, 1958) introduced the concept of chief-association
(Hauptassoziation) for the type of association SCHWICKERATH
was referring to. MEIJER DREES (1951) defined it as follows:
the smallest unit possessing absolute or regional character-species.
Absolute = general or regional is meant here in the sense ofBRAUNBLANQUET
& MOOR (1938) and MEIJER DREES, i.e..not occurring
outside the association's range. KNAPP (1971) later mentioned the
concept rather incidentally as referring to character-species that
are valid 'at least within an entire flora-region.' The more local
units comprised in a Hauptassoziation were called Gebietsassoziationen,
or 'regional associations'. PASSARGE (1968) and PASSARGE
& HOFMANN (1968) elaborated the suggestion by KNAPP
towards a separate system of historic-geographical units besides the
usual edaphic-ecological units. 'Regional' (territorial or provincial)
associations could then be distinguished as parallel but subordinated
units to the 'elementary association,' with which the chief-association
is identical. PASSARGE & HOFMANN (1968) subjected the
regional association to a trinary nomenclature as is applied in
zoological systematics for geographical races.
This brings us back to OBERDORFER (1957, 1968) who sharply
distinguished between 'association,' 'regional-association' and 'geographical
race.' (See also MULLER 1968). A regional association
('Gebietsassoziation') has a limited distribution-area; it is characterized
by acornbination of regional (or local) character-taxa, corresponding
with SCHWICKERATH'S geographically differentiating
taxa, and general alliance character-taxa. A geographical race is a
regional expression of a larger, reg~onal or general association, without
such a typical combination of territorial and alliance charactertaxa.
As OBERDORFER (1968) remarked, it will often be very difficult
to distinguish between the two levels. Geographical races have
331
not yet found a place in the formal hierarchy but have nonetheless
frequently beendescribed, particularlyin Germany (e.g. OBERDORFER
1957), Czechoslovakia (e.g. NEUHAUSL & NEUHAUSLOVA.-NoVOTNA.
1972) and Poland (e.g. MATUSZKIEWICZ 1962).
We have thus a surfeit of terms and concepts. To resolve this
matter we refer to the remarks on character-taxa (20.5.8) and adopt
the distinctions made by OBERDORFER and others under three
terms for units of expanding geographic extent: the local association,
regional association, and (general) association. Clearly these
intergrade, but in principle they should possess local, regional and
general character-taxa, respectively. Groups of related regional associations,
which are in fact vicariant associations (cf. PASSARGE
1968) could be called 'vicariant association groups'; this term seems
more appropriate than either chief or elementary association.
Another acceptable name for the vicariant association-group is collective
association (cf. MEIJER DREES 1951).
We shall now briefly discuss the Artemisietum maritimae
association as presented in Table V. It is characterized by one
character-taxon, Artemisia maritima which has, according to Table
VI, a high fidelity degree (at least in the investigated area) and
which is, according to the synchorological data presented by
BEEFTINK (1965) as well as chorological data on the species, a territorial
character-taxon. It follows from Table VI that most of
the salt marsh associations in the southwestern Netherlands are
typified by few but generally 'good' character-taxa (fidelity degrees
4 or 5). The normal characteristic taxon combination of the
Artemisietum maritimae in the southwestern Netherlands
consists of Artemisia maritima, Festuca rubra f. litoralis, Limonium
vulgare ssp. vulgare, Aster tripolium and Plantago maritima. BEEFTINK
has listed most of the taxa under a specific head so that the companions
are few and simply called 'other taxa'. Note the four
releves at the right end of the table, that lack the character-taxon
and are considered a fragmentary syntaxon.
20.6.4 HIGHER UNITS (ALLIANCE, ORDER, CLASS)
When a new association is recognized, it must be placed in the
system of higher units. The assignment of an association to an alliance
(and other higher units) is primarily based on comparison of
floristic relationships. As in plant taxonomy (cf. DAVIS & HEYWOOD
1963) the terms relation(ship) and affinity refer only to the possession
of common characters. They do not refer to syngenetical
(successional) links; we may compare with SOKAL & SNEATH'S
332
(1963) emphasis of phenetic relationships. Group importance
values of the character- (and differential-) taxa of the relevant alliances
are calculated, either on a presence-absence or on a quantitative
basis, and compared (cf. WESTHOFF 1947, RAABE 1957).
It may occur that the affinity towards two alliances (or orders or
classes) is almost equal. An example of close relationship towards
two alliances is presented by the Artemisietum maritimae.
Generally the Armerion mari timae alliance is the obvious higher
unit, as follows from tables V and VI, but the subassociation
Artemisietum maritimae typicum is also clearly related to
the Puccinellion maritimae. In fact the association was once
assigned to the Puccinellio-Salicornion alliance (BRAUNBLANQUET
& LEEUW 1936) from which the Puccinellion maritimae
was split off, but later (e.g. TUXEN 1937, BEEFTINK 1965,
WESTHOFF & HELD 1969) it was considered part of the Armerion
maritimae. In cases of doubt equal use is made of structural,
physiognomical and synecological considerations (cf. OBERDORFER
1957, WESTHOFF & HELD 1969).
It may also be the case that in a little-known area anumber of
new associations are described; and some of these may bear so
little relation to known alliances (etc.) that they should be classified
into new higher units. As was introduced in 20.3.2, various
higher units above the association level, viz. the alliance, the order,
and the class are characterized in the same way as the association.
Species equally occurring in a group of related associations but
faithful to that group, become character-taxa for higher units towhich
they are faithful. Contrary to most associations the higher units generally
do possess a number ofregional or general character-taxa ofhigh
fidelity. When an alliance character-taxon shows a preference for
one association within an alliance, it may be used both as alliance
character-taxon and as differential-taxon for an association within
the alliance. The species is then termed a 'transgressive charactertaxon.'
Species may of course bear similar character- and differential-
relations to orders vs. alliances, and classes vs. orders.
Interpolated units like the suballiance and suborder are characterized
by character-taxa and/or differential-taxa. Their use is to
be avoided; but the suballiance has had some use.
. Formal definitions for higher syntaxa and instructions for their
ranking have not been agreed upon (cf. PIGNATTI 1968b, DIERSCHKE
1971). In practice the number of related associations tends to
determine the number of the higher units classifying these. Growth
in number of associations expresses its influence upward, in growth
in the number of alliances, etc. For example, TuxEN (1937) described
94 associations and 41 alliances for northwest Germariy;
333
in TtiXEN (1955) the numbers were 189 and 76. In the Netherlands
the number of described alliances grew from 39 (WESTHOFF et al.
1946) to 85 (WESTHOFF & HELD 1969). PIGNATTI (1968b) called
the attention to the 'inflationary' character of this phenomenon. His
suggestive title stimulated much discussion (TtiXEN 1968b: 89-97),
which produced few solutions apart from plans for cooperation and
standardization now being realized in the European Prodromus
of plant communities.
20.6.5 UNITS ABOVE THE CLASS
The growing number of classes, as well as the extension of the
BRAUN-BLANQ.UET approach to various, and floristically different
parts of the world, lead to the introduction of units above the class.
The grouping above the class originally proposed by BRAUN-BLANQ.UET,
the 'vegetation circle' was a phytogeographical rather than a
sociological unit. BRAUN-BLANQUET (1959) and TUXEN (1970a) on
a suggestion of SCHMITHtiSEN sought to establish a new unit, the
class group (d. TtiXEN 1963: 213-218).
The class group (TtiXEN 1970a) is a set of territorially defined,
largely vicariant classes analogous with the 'vicariant association
group' (20.6.3). The characterizing taxa are in this case genera with
vicariant species in different floristic provinces or regions; for example
the classes Querco-Fagetea silvaticae (Europe), Querco-Fagetea
grandifoliae (North America), and QuercoFagetea
crenatae (East Asia) form a class group: QuercoFagea,
with the suffix -ea probably taken fromJAKucs (e.g 1972).
Curiously, BRAUN-BLANQ.UET (1964) presented the class group under
a different name 'community kingdom' (Gesellschaftsreich) and
apparently as a somewhat different phytogeographical concept.
Although he first (p. 140) stated that this unit 'takes together classes
floristically related by vicariant species and identical higher
categories', he concluded that community kingdoms should coincide
with SCHMITHUSEN'S (1961) vegetation kingdoms established
by means of vicariant classes.
TUXEN'S class group should not be confused with the geographically
determined class-group of SCAMONI et al. (1965). However,
according to the elaboration given by PASSARGE (1968) and
PASSARGE & HOFMANN' (1968), these class groups correspond actually
to the usual European classes. It is questionable whether PASSARGE
& HOFMANN'S (1968) complicated system of two hierarchies
can be integrated in the BRAUN-BLANQ.UET system. TUXEN (1971)
strongly rejected such an integration. .
Three other treatments of higher units deserve mention. (i)
334
CHAPMAN (1952, 1959) used terms like Coeno-Salicornietalia
for groups of higher units dominated by species of one widely distributed
genus..For such a unit the name Coeno-Salicornietum
should be preferred. (ii) The concept of 'vegetation type' was
developed by HADAC (e.g. 1962, 1967) for groups of more or less
similar classes. The vegetation type is based on the occurrence of
common species (indicating or 'bezeichnende' species). However,
the floristic basis of these units seems to be rather loose, and as
HADAC admits the vegetation types are 'practically formations'.
(iii) JAKUCS (1961) also used the class group concept, which he
later (e.g. 1972) named 'division' and for which he introduced the
suffix -ea. HADAC (1967) concluded that the division is largely
identical with his vegetation type, which term should then have
priority, The identity seems to be real, but for it the term 'vegetation
type' is least descriptive and most confusing; division should be
preferred. BOLOS (1968) presented a system of divisions for classes
occurring in Spain. As term and concept the division is consonant
with the hierarchy.
In conclusion, a division may be defined as a syntaxon above
the class level that unites related classes within a floristic region (or
province) on the basis of common division character-taxa. The
character-taxa may be species, or genera, or both. Let us call this
a 'vertical' unit in the sense that it unites lower units sympatric in a
given region. The class-group concept, in contrast, is comparable
with the vicariant association group in that it joins units on the
same level from different floristic territories. It is in this sense a.
'horizontal' unit. Its character-taxa will more usually be genera
than species. Therefore the name 'vicariant class group' may be
proposed, defined as a group of classes allopatric in geographic occurrence,
but linked by the occurrence ofvicariant species in one or
more genera. Since two units would be distinguished in this way, it
may be questioned whether the suffix -e a should be used for both.
It seems logical to reserve this suffix for the division, for which it
was proposed.
The definition of the division is not physiognomic. We suggest
that, the difference in definition notwithstanding, divisions may
converge in practice with the formations of Anglo-American
ecology (article 13), as broad physiognomic units limited to a given
region or continent. Vicariant class groups, in contrast, would link
floristically related phytocoena of different continents. Thus the
Querco-Fagetea-group would comprise forest phytocoena with
genera in common in Europe, eastern North America, and western
Asia, whilst a Spartinetea-group would comprise a set of more
narrowly defined phytocoena (Spartina associations) in the salt
marshes of the world.
335
20.6.6 HORIZONTAL AND VERTICAL CLASSIFICATION
Thus we follow the suggestion of KNAPP (1948), to classify independently
by two principles, the one on an edaphic-ecological
('vertical') and the other on a historic-geographical ('horizontal')
basis. This suggestion has been amply discussed (WESTHOFF 1950,
MEIJER DREES 1951, BARKMAN 1958a, BEEFTINK 1965, SCAMONI
et al. 1965, PASSARGE & HOFMANN 1968, PASSARGE 1968). All
authors agree with the incorporation of both viewpoints in syntaxonomy.
MEIJER DREES emphasized the vertical approach, because
syntaxonomy should build upward from locally classified associations.
BEEFTINK suggested that the choice of the criterion would
depend on the peculiarities of the vegetation concerned: for some
phytocoena with disjunct distributions the stronger floristic relations
and preferred grouping would be horizontal whereas for other
phytocoena the more natural grouping would be vertical. PASSARGE
& HOFMANN use both principles on each level, with the geographical
units subordinated to the corresponding edaphic units,
which makes the system very complicated (cf. TUXEN 1971, in his
critical review of PASSARGE & HOFMANN 1968).
We would modify BEEFTINK'S suggestion in recommending
that the main-axis of the classification should be 'vertical' on all
levels ofthe hierarchy. Given the main direction ofthe classification
as vertical, secondary to this, 'horizontal' vicariant groupings may
be recognized on any syntaxonomic level. We have mentioned
such groupings for associations (20.6.3) and higher syntaxa (20.6.5);
we may now trace the problem and the principle into the lower
syntaxa.
20.6.7 LOWER UNITS .(SUBASSOCIATION, VARIANT, FACIES)
Associations are on the one hand joined into higher units, and
on the other divided into subordinate units. As in idiotaxonomy
the former procedure is compulsory, the latter facultative. The
lower syntaxa are mostly characterized by differential-taxa. Usually,
also, the syntaxa below the association level are described as
deviations from average or typical situations. Thus to complete the
syntaxonomy on a certain level a 'typical' subassociation (variant,
etc.) and one or more 'deviant' syntaxa are described and characterized
in parallel.
Variations within an association have been approached in
three directions; to the edaphic-ecological ('vertical') and historicgeographical
('horizontal') may be added a syndynamical (suc-
336
cessional) point-of-view. It has been proposed repeatedly that different
syntaxa should correspond to these approaches each ofwhich
could form a hierarchy of subordinate units. Thus the subassociation
might be based on local edaphic or micrometeorological differences
within the association, the variant on geographical or
climatological differences (with the terms vicariant and geographical
race as synonyms), and the phase on differences in successional
status. This distinction is in accordance with the original approach
of BRAUN-BLANQ.UET (1928, 1932, BRAUN-BLANQ.UET & PAVILLARD
1928). SCHWICKERATH (1942) and DUVIGNEAUD (1946, 1949) distinguished
five types of differential-taxa on the basis of differences in
nutrient status, moisture conditions, successional status, geographical
area, and human influences.
The system was further extended by the incorporation of the
sociation, a unit defined by the dominant species of its strata (see
article 18). This suggestion of NORDHAGEN (e.g. 1937) was acknowledged
by the 1935 Botanical Congress (see Du RIETZ, 1936).
In the Netherlands it has become common practice to describe
species-poor communities without character-species but with one
or more dominants as sociations or consociations (WESTHOFF 1947,
1949, BEEFTINK 1965, WESTHOFF & HELD 1969). Contrary to Du
RIETZ (1930b, 1936, 1965) the sociation and consociation are considered
of association-rank (BRAUN-BLANQ.UET 1955, MAAREL et al.
1964). They may be assigned to an alliance and higher syntaxa
(WESTHOFF & HELD 1969).
From the beginning many communities have been described
without receiving a definite syntaxonomic position. They are often
called 'community of ......' ('Gesellschaft von ......') (cf. ELLENBERG
1956, NEUHAUSL 1963, FUKAREK et al. 1964). These units can
be either provisional local coena, which might later be ranked in
the system when more evidence is available, or coena without char~
acter-species or dominants. BARKMAN et al. (1958) and WESTHOFF
et al. (1959), proposed the term 'consortium' for the latter units.
An additional term for subordinate units, 'form,' is used in different
ways for syntaxa of lower rank. It may mean a unit of no
specified rank (as seems to be the case with SCHUBERT 1960, for
example). It is also used in relation to altitude (OBERDORFER 1957,
1968, BRAUN-BLANQ.UET 1964) or moisture conditions (DUVIGNEAUD
1946). The 'Ausbildungsform' is considered the lowest unit
in the hierarchy apart from the facies by TUXEN (1970a, cf. SCHUBERT
1960, PASSARGE 1968).
The distinction ofloca1-edaphic and geographic units has been
maintained by BRAUN-BLANQ.UET (1951, 1964), MEIJER DREES
(1951), OBERDORFER (1957, 1968), BARKMAN (1958b) and others.
337
Under the influence of TUXEN (1937) the variant became a subunit
under the subassociation (BRAUN-BLANQ.UET 1951a, 1964) rather
than a differently defined (geographic) unit. In line with this practice
we suggest as the downward extension of the primary hierarchy,
for units based on local variations: association, subassociation,
variant, subvariant, and facies. The last two may be often unneeded,
but facies may be ecologically significant and useful as
indicators in some cases (KRAUSE 1954). Geographic races of lower
syntaxa would be separately indicated by the territories in which
they occur (OBERDORFER 1957, MtTLLER 1968).
The subassociation is indicated with the suffix -e t 0 sum. Of the
various ecological possibilities and corresponding symbols of DuVIGNEAUD
(1946, 1949) d and <5 may be used for subassociations differentiated
by nutrient and moisture status, respectively. SCHWICKERATH
(e.g. 1942), DUVIGNEAl1D (1946) and MEIJER DREES (1951)
showed how subassociations of an association may in fact indicate
the transitions of that association towards related associations. The
subassociation 'typicum' may then be considered as the 'nucleus'
of the association (cf. Fig. 1); in fact this term is commonly used for
the subassociation that lacks differential-species. WESTHOFF (1965,
WESTHOFF & HELD 1969) observed that many subassociations ofthe
latter type were relatively poor in species, and particularly poor in
character-species of the association. Such units were indicated as
'inops' (= having shortage of); the 'typical' Querco-Betuletum
is an example of such an 'inops'-subassociation.
The next lower rank is called the variant. According to BRAUNBLANQ.UET
(1964: 124) a variant would not be characterized by
differential-species, but 'either by a strong prominence of certain
species, which cannot be considered as differential-taxa, or by a
slightly deviating species assemblage.' This criterion is, however,
diagnostically too vague. Most phytosociologists share the opinion
that a variant also must be characterized by differential-species.
The same holds for the next lower rank, the subvariant. The variant-differentiating
taxon (taxa) may be differentiating only because
of its 'strong prominence,' i.e. its higher combined estimation
values in one variant compared with others and not because it is
clearly present resp. absent. In most cases the variant is distinguished
within the subassociation; but it is also possible to divide an association
directly into variants, if the mutual differences are considered
too small to justify the rank of subassociation. Below the variant
the subvariant may be used if needed.
The lowest unit is thefacies. It is not even characterized by differential-taxa;
it thus does not employ diagnostic-species and is not
part ofthe formal hierarchy. A facies is usually characterized by the
338
dominance, in a high cover degree (scale 4 or 5), of one of the species
belonging to the normal floristic assemblage of an association.
It is not usual to consider a stand as a facies if this dominance is a
normal feature of that association. Since, for instance, dominance
of Fagus sylvatica is normal for any association assigned to the alliance
Fagion sylvaticae, it is not appropriate to construct such
a coenon as 'a Fagus facies of the Fagetum.' A facies, therefore, is
a deviation phenomenon. It may be brought about by special, and
sometimes by extreme conditions of abiotic factors, but in many
cases it is the result of human disturbance (e.g. MELTZER & WESTHOFF
1942, KNAPp 1971). Examples are a facies of Rubus sect. Sylvatici
orRubus sect. Heteracanthi in a stand ofQu er co-C a r pin etum,
or a facies of Acorus calamus in a stand of Scirpo-Phragmitetum.
It is often possible to deduce an effect of human disturbance from
the appearance of a certain facies.
20.6.8 NOMENCLATURE
Formal nomenclature of syntaxa started with the naming of
associations by BRAUN-BLANQ.UET (1913), following precedents of
CAJANDER (1903), BROCKMANN-jEROSCH (1907) and RUBEL (1912).
The first general proposals were discussed at the Brussels Botanical
Congress (FLAHAULT & SCHROTER 1910); the nomenclature of
associations and other syntaxa developed gradually thereafter.
With the growth of the number of described syntaxa the need for
standardization and rules increased. A start toward systematic
nomenclature was made in 1933 (BRAUN-BLANQ.UET 1933b), and
DAHL & HADAC (1941) proposed a coherent set of rules based on
the code of botanical nomenclature. During the Stockholm Botanical
Congress BARKMAN (1953a) proposed a general set of rules
and MEIJER DREES (1951) a complete system of nomenclature.
These were discussed in a number of papers in Vegetatio vol. 4. Contributions
by BACH et al. (1962) and RAUSCHERT (1963) appeared
later,· and BRAUN-BLANQ.UET (1964) reported on the discussion and
presented 7 rules. The next phase started during the Symposium
on Plant Sociological Systematics at Stolzenau, 1964, where MORAVEC
(1968) introduced a set of 26 articles and urged a general
acceptance of these amongst active plant sociologists. A special
Working Group is now. preparing a code for the new Prodromus
project (see TUXEN 1968b: 152, MORAVEC 1968, 1969, 1971, NEUHAUSL
1968).
339
Some instructions for the construction of names and some proposed
general rules for nomenclature of syntaxa follow.
According to MORAVEC (1971) five necessary principles are:
1. Each syntaxon (with definite rank, position and delimitation)
has only one correct name.
2. Each name can be correctly used for one syntaxon only.
3. The correct name is established according to rules based on the
priority principle.
4. The association is the fundamental nomenclatural unit (syn-
taxon).
5. The validity of nomenclatural rules is retroactive.
20.6.8.1 Construction of Names
For the standardization of syntaxon names, the following procedure
is in general use.
To the generic part of the names of one or two (not more)
characteristic (not necessarily faithful) taxa of a syntaxon a suffix
is added. These suffixes are specific for the different syntaxonomic
ranks, see Table VII. The suffix'-etum' for the association goes
back to classical Latin and has been in use since HUMBOLDT (1805).
The suffix '-ion' for the alliance was proposed by Moss (1910) and
taken over by BRAUN-BLANQUET (1921: 347). The suffix '-etalia'
for the order was presented in BRAUN-BLANQUET (1928). For the
class, up to 1932 (BRAUN-BLANQUET, FULLER & CONARD) no specific
suffix was used, since the classes were designated by circumscriptions
such as 'communities of maritime dunes' (later the class
Ammophiletea). In 1934 MEIER & BRAUN-BLANQUET proposed
the suffix '-etales' for the class, but (BRAUN-BLANQUET et al. 1939)
this suffix was later changed into '-etea'. The suffix '-etosum' was
introduced by BEGER (1922).
If we are dealing with an association or alliance that is sufficiently
characterized by one taxon (e.g. by a faithful dominant character-taxon),
the genus name of the taxon is used with the appropriate
suffix (-etum, -ion), followed by the species name in the
genitive. For example the association Ericetum tetralicis is
named after Erica tetralix, and the alliance Alnion glutinosae
after Alnus glutinosa.
In some cases the specific epithet has been replaced by a geographical
adjective indicating the area characteristic of the syntaxon,
e.g. Agropyretum boreoatlanticum for the association
of Agrop)lron junceum (syn. Elytrigia juncea) of the North Atlantic
Coast. In this case this has been done to distinguish it from a dif-
340
ferent aSSOcIatIOn of Agropyron Junceum occurring in the Mediterranean.
In recent decades, however, there is a tendency to avoid
such geographical names as far as possible and to replace them by
bigeneric names. In the case of orders and classes and, though less
frequently, alliances the specific epithet is omitted when the context
leaves no doubt - for example, 'Fagion' instead of 'Fagion
sylvaticae' in a European context.
When a syntaxon was to be named after two characteristic
(not necessarily faithful) taxa, the name of the second taxon was
provided with the appropriate suffix; whilst the name of the first
taxon was joined with the second one by the suffix -e to. Examples
are the association Querceto-Betuletum, the alliance AlnetoUlmion,
the order Glauceto-Puccinellietalia, and the class
Querceto-Fagetea. However in many cases this suffix was abbreviated
to a connecting vowel after the stem of the first name,
e.g. Alneto-Padion became Alno-Padion. BACH et al. (1962)
proposed to do this systematically, mainly on the argument that,
for example, 'Querceto-Betuletum' could mean a mixture of, or
transition between, a Quercus stand and a Betula stand, which is of
course something quite different. This proposal has been widely accepted.
However, the correct choice of the connecting vowel, especially
in the case of the third Latin declension and with names
derived from Greek words, gives difficulties. BACH et al. (1962) and
RAUSCHERT (1963) gave directions for a large number of cases.
There are two reasons for choosing a bigeneric name. The
first and more obvious is that a combination of two names should
..haracterize the syntaxon better. The first name may be a dominant,
generally the second name represents the character-taxon considered
diagnostically most important. The other reason was already
mentioned, viz. monogeneric names with a geographical adjective
are replaced by bigeneric names, in which the first name
comes from a species characteristic for the geographical region
originally indicated. The example given above, the Agropyretum
boreo-atlanticum, has been renamed in fact the MinuartioAgropyretum
(TuxEN 1955).
In some cases the nomenclature is at variance with the standard
procedure. The following deviations should be mentioned:
(1) Contractions. It has become general use to follow the proposal
by KOCH (1925) to replace clumsy terms like 'Potamogetonion'
(and, later on, also 'Potamoget.onetalia' and '~otamogetonetea')
by 'Potamion', 'Potametalia' et 'Potametea'.
However, OBERDORFER et al. (1967) proposed to return to the
complete and cumbersome names.
(2) In special cases descriptive adjectives or substantives are m-
341
cluded in the name. 'Magnocaricion' indicates an alliance characterized
by tall sedges; 'Macrophorbio-Alnetum' is meant to
characterize a moist woodland (carr) association in which 'macropho~biae'
(tall forbs, Hochstauden) form a group of differentiating
speCIes.
(3) Finally, sometimes specific epithets have been used instead of
generic ones. This was common practice in the early days of alpine
phytosociology; BROCKMANN-jEROSCH (1907) and RUBEL (1912),
for example, used the names 'Curvuletum' and 'Firmetum' instead
of 'Caricetum curvulae' and 'Caricetum firmae.'
Later this practice has been applied only very rarely; a rather recent
example is 'Alno-Padion' (KNAPP 1942, MATUSZKIEWICZ &
BOROWIK 1957), to designate an alliance characterized by Alnus
glutinosa and Prunus padus.
20.6.8.2Validiry, Changes and Authors' Names
Various rules have been proposed for the settlement of the
validity of syntaxon names. A name should have been published
after 1900 in printed form, available at least in generally accessible
libraries. Publication of a name should be combined with an
adequate diagnosis.
For associations and lower-rank units this would include the
presentation of a table with the complete floristic composition of at
least three releves and the assignment of a type-releve. For higher
units the listing of character- and differential-taxa and the assignment
of a type-unit of the next lower level would be required. Such
types are called nomenclatural types.
When two or more correctly published names appear to Iefer
to one and the same syntaxon, the oldest name has priority. When
a syntaxon is divided into two or more units of the same rank, the
original name goes to that new unit that contains the nomenclatural
type.
When the valid name of a taxon used in a syntaxon name
changes, corresponding change in the syntaxon name should be
considered. In any case a syntaxon name should be changed when
one of the name-giving taxa appears to be a younger homonym of
a different legitimate name. For example, the Isoetetum setacei
should become the Iso etetum del ilei, since Isoetes setacea (Box
ex) DELILE appeared to be a homonym of Isoetes setacea LAM. =
I. echinospora DUM. and was then renamed 1. delilei ROTHMALER.
No agreement has been yet reached on the - still very freo
quent _ simple nomenclatural changes. E.g. Carex nigra (L.) REICH-
342
ARD appears to be the correct name for C. Jusca ALL. Should the
order Caricetalia fuscae now be renamed the Caricetalia
nigrae? MORAVEC (1968) would say not; WESTHOFF would say
yes and in fact (WESTHOFF & HELD 1969) changed this name.
Rules for the citation of authors' names have also been proposed,
more or less parallel to those of idiobotanical nomenclature.
The name of the author(s) who published a valid name is added to
that name together with the year of publication. When this name
rders to a previously published but invalid name, the earlier author's
name is added in the form, Thero-Airion TUXEN ex OBERDORFER
1957. When a name is changed, e.g. when the syntaxonomic
level is changed, the author of the changed name is added between
brackets. When a syntaxon is given a new contents or delimitation,
both old and new authors are mentioned connected with the abbreviation
em. (= emendavit).
These proposals were advanced by MORAVEC (1968). PIGNATTI
(1968 b: 87) recommended dropping authors' names to avoid the
description of new syntaxa by authors who primarily want to have
their names immortalized. In the discussion following, p. 89-97,
there was agreement on omitting author's names when no confusion
could occur, especially with higher units and when a prodromus
can be referred to.
20.6.9 SCHEME OF SYNTAXA
All in all phytosociological nomenclature may be considered a
necessary evil. Contrary to the opinion of DOING (1962, 1966, 1970)
that a rigid system of nomenclature should be abandoned for practical
reasons, we would emphasize that the value of the system for
ordering information and communicating among phytosociologists
outweighs its difficulties. We would further state that descriptions
offormal phytosociological syntaxa should either be published
in strict accordance with nomenclatural rules, or they should not
be published as such.
Table VII presents the current syntaxa and their nomenclature,
as discussed so far. Examples together with their denominating
taxa are added. The variant (and the facies) are given here without
suffixes. The geographically d~termined syntaxa, (20.6.3, 20.6.5),
which have not yet been agreed upon, are not included.
343
TABLE VII
Scheme of Syntaxa
Levels and units of the formal hierarchy from highest (Division) to lowest subvariant,
with suffixes and exampIes ofthe construction ofnames after denominating
taxa.
Syntaxon
Division
Class
Order
Alliance
Suballiance
Association
Subassociation
Variant
Subvariant
Suffix
-ea
-etea
-etalia
-IOn
-IOn
(-esion)
-etum
-etosum
Examples
Querco-Fagea
Phragmitetea
Querco-Fagetea
silvaticae
Littorelletalia
Festuco-Sedetalia
Agropyro-Rumicion
crispi
Alnion glutinosae
Ulmion carpinifoliae
(Ulmesion)
Ericetum tetralicis
Elymo-
Ammophiletum
Arrhenatheretum
elatioris brizetosum
ibid., Salvia variant
ibid., Bromus sub-
variant
20.7 Extension of the Approach
20.7.1 OTHER GEOGRAPHIC AREAS
Denominating taxa
genus Fagus
Phragmites australis
Quercus robur
Fagus sylvatica
Littorella uniflora
Festuca and Sedum
L. div. spp.
Agropyron repens,
(syn.: Elytrigia
repens) and
Rumex crispus
Alnus glutinosa
Ulmus carpinifolia
Erica tetralix
Elymus arenarius,
Ammophila arenaria
Briza media
Salvia pratensis
Bromus erectus
From its 'center of origin in the Alps and the western Mediterranean
the BRAUN-BLANQUET approach spread into many European
countries and Japan. Larger parts of the Mediterranean and EuroSiberian,
and smaller parts ofthe Sino-Japanese, floral regions have
been synsystematically described. Still, a considerable part of the
original working area is insufficiently known, particularly the eastern
Mediterranean, Russia and Scandinavia (see the map presented
by DIERSCHKE 1971). Expansion of the approach's influence continues,
and we shall give a short survey of more recent literature
applying the approach outside its European homeland. The survey
cannot be exhaustive; only regions will be mentioned in which substantial
work has been done. For further references the reader is
344
referred to Excerpta Botanica Sectio Sociologia (bibliographic contributions
to which are indicated below as 'Exc.'). This series can also
serve as a survey of phytosociological work in the three regions
mentioned above, with bibliographies for almost all European
countries.
Japan may deserve special attention.The phytosociological study
ofJapan was initiated by SUZUKI/ToKIO and developed especially
by MIYAWAKI and his pupils (see MIYAWAKI 1960 and 1966a, 1971a
Exc.). Of special interest is the manner in which the Japanese communities
are compared with Mediterranean and Indo-Malaysian
communities. Other comparative studies in Japan include OHBA
(1972), in which various vicariant salt marsh associations were described,
partly belonging to the 'European' class Asteretea tripolii;
TUXEN, MIYAWAKI & FUJIWARA (1972) on the class Oxycocco-Sphagnetea
in which Europe and North America (work
of R. KNAPP and A. DAMMAN) are included; and TUXEN (1966a)
on holarctic Honckenyo-Elymetea communities. These studies
were all based on extensive reIeve tables and close cooperation between
European and Japanese phytosociologists. From these studies
the concept of vicariant class group developed (see TUXEN 1970a
and 20.6.5). Vegetation mapping on a phytosociological basis has
reached a high level in Japan (see MIYAWAKI 1966b, 1971b, Exc.
for a survey of maps).
The arctic-subarctic region has only occasionally been described.
Iceland is becoming relatively well-known; a general survey is in
preparation by R. TUXEN based on phytosociological excursions by
HADAC, BOTTCHER, the International Society for Plant Geography
and Ecology, and others. Newfoundland was partly described by
DAMMAN (1964). The research on Oxycocco-Sphagnetea by
DAMMAN in Canada was already mentioned under Japan as was
the study of TUX-EN (1966a). See also STEINDORSSON (1966, Exc.)
for Iceland, BOCHER (1961, Exc.) for Greenland, and HANSON
(1959, Exc.) for Alaska.
The west and central Asiatic (Irano-Turanian) region has been
described in a fragmentary way. An extensive survey of xerophytic
and summer-dry hygro- and mesophilous communities of Afghanistan
was presented by GILLI (1969, 1971), who could reasonably
establish associations, though not as sharply delimited as in Europe
because of the large number of species in most communities. HARTMANN
(1968, 1972) published accounts on mountain grassland and
scrub communities.
For the Macaronesian region the Canaries are reiatively wellknown
through work of RIVAS-GODAY & ESTEVE CHUECA (1965),
OBERDORFER (1965), LEMS (1968), and SUNDING (1969), as well as
345
LOHMEYER & TRAUTMANN (1970) who reviewed this literature (see
also references in SUNDING 1969-70, Exc.). Studies on the Azores
and the Cape Verdes were listed by PINTO DA SILVA & TELES (1962,
Exc.). OBERDORFER (1970) reported on some plant communities on
the Canaries and described complex, stable, mosaics of succulent
scrubs and open sderophyllous woods, which are amongst the most
complicated communities of the world and should be studied as
complexes (cf 20.7.4 and the South-African Kingdom below).
Rather numerous, though still scattered phytosociological
studies have been carried out in the North-American and Canadian
regions, starting with the description of Long Island by CONARD
(1935) including 71 associations. CONARD (1952) also published
on the vegetation of Iowa and DANSEREAU (1957) on the middle
St. Lawrence Valley. TOMASELLI (1958, quoted in KUCHLER 1967)
described plant communities in eastern Kansas. Monographs are
concentrated on forest communities (except for the studies of LooMAN
e.g. 1969 on grasslands). They include those of MEDWECKAKORNAS
(1961) in the Montreal area, GRANDTNER (1966) in
Quebec, KORNAS (1965) in North Carolina, CRISTOFOLINI (1967)
in Tennessee, JANSSEN (1967) in Minnesota and KRAJINA (1969)
in British Columbia. KNAPP (1957, see also 1965) presented a
preliminary survey of higher phytosociological units, viz. 72 classes,
some of which were divided into orders and later (KNAPP 1964)
extended it to the whole holarctic Kingdom.
The Indo-Malqysian and the Pofynesian subkingdoms are practically
phytosociological terra incognita. Some studies have been
made in India (BHARUCHA & MEHER-HoMJI 1963, Exc., GUPTA
1966, 1967, Exc., MEHER-HoMJI 1969, Exc., MEHER-HoMJI &
GUPTA 1972a,b, Exc.).
The African territory has been studied rather intensively, here
and there. French studies on the North African desert region culminate
in the monograph of QUEZEL (1965), which presents a
coherent survey of over 100 desert communities largely fitted
into a syntaxonomic system (see also ROUSSINE & SAUVAGE 1961,
Exc.). Relatively intensive studies have been performed in the
West African rain forest region by LEBRUN (e.g. 1960), LEBRUN
& GILBERT (1954) and SCHNELL (1952). The associations and higher
units described, are all based on releves and range from xerophytic
and hydrophytic pioneer communities to climax forests.
LEBRUN concludes that the BRAUN-BLANQUET approach is very well
applicable. Savanna studies in this area include those of DUVIGNEAUD
(1949), ADJANOHOUN (1962), and DEVRED (1956). In
Mo<;ambique a first phytosociological inventory was carried out
by GOMES PEDRO & GRANDVAUX BARBOSA (1955), see also PINTO DA
346
SILVA & TELEs (1962, Exc.). KNAPp (1965, 1966a) presented
preliminary surveys of higher units for West, Central and East
Africa and reviewed the literature for some countries (KNAPP 1969-
70, 1971a,b, Exc.).
Phytosociological research in South Africa started recently and is
now expanding rapidly (EDWARDS 1967, TAYLOR 1969, ZINDEREN
BAKKER 1971, WERGER1973, see WERGER for references). WERGER
et al. (1972) presented the first account of plant communities in the
Cape Kingdom, including vegetation types of the famous 'Fijnbos'
formation. Their experiences with the BRAUN-BLANQUET approach
in this extremely varied mosaic of species-rich communities are
comparable with that in European xerotherm woodland-scrubborderline
communities (cf. JAKUCS 1961, 1972).
The Caribbean region has also been treated in general phytosociological
studies, e.g. KNAPp (1965). Detailed descriptions of
savanna vegetation in Surinam have been published by DONsELAAR
(1965, 1969), who found the BRAUN-BLANQUET approach
applicable. The North Surinam open savannas could be grouped
into one class. A special problem was formed by the floristic similarities
between community types ofdeviating structure. DONSELAAR
considered the floristic relations as decisive, as did WERGER et al.
(1972) for similar situations in Fijnbos communities of the Cape.
Earlier Surinam work is mentioned in BOERBOOM (1970, Exc.).
Brazilian rain forest descriptions are very limited (VELOSO 1962 in
BRAUN-BLANQUET 1964, DONsELAAR 1965). The Andean region has
received more attention; OBERDORFER (1960), SCHMITHUSEN (1960),
ESKUCHE (1968, 1969) described various scrub and forest communities,
and KOHLER (1968, 1970) desert and coastal communities.
See also ESKUCHE (1967, Exc.) for this, and adjacent regions in
South America. KNAPp(1966b) presented a survey of higher vegetation
units in Patagonia.
The BRAUN-BLANQUET approach has thus been applied in
nearly all floristic regions, and to almost all types of vegetation.
The approach should in principle be uniyersally applicable;
and in fact no difficulties have been met in its expansion out of
Europe more fundamental than those encountered in Europe. The
principal directions of difficulty are: (i) In species-poor vegetation
in the North and the Antarctic, character-taxa may be difficult to
find and use, and species dominance (and the sociation as a unit)
may be more emphasized (see article 18) (ii) Rich tropical vegetation
confronts the approach with such a wealth of species as to
make application more difficult; physiognomy has been more emphasized
in the tropics as a means of recognizing phytocoena. (iii)
Delimitation of uniform stands becomes difficult in mosaic struc-
347
tures of open xerothermic scrubs and woodlands, such as found in
the Canaries and South Africa (cf. 20.4.1).
20.7.2 BIOTIC COMMUNITIES
Approaches to the classification of animal commumtIes are
diverse (WHITTAKER 1962); applications of the Braun-Blanquet
approach have mostly been concentrated on the analysis of one or
more animal groups in plant communities already described.
General bibliographies were presented by RABELER (1957b, 1964,
Exc.). More or less complete descriptions of biotic communities are
restricted to marine benthic environments, in which animals and
plants were used together to construct a hierarchy according to
Braun-Blanquet principles (RoGER MOLINIER 1960).
The following conclusions may be drawn from work on biotic
communities (cf. BRAUN-BLANQUET 1951, 1953, 1964, RABELER
1937, 1960, 1962, 1965, WESTHOFF & WESTHOFF-]ONCHEERE 1942,
MORZER BRUYNS 1950, WHITTAKER 1962, MAAREL 1965, T-OxEN
1965a and BARKMAN article 16 in this volume) :
1. Species from most animal groups form characteristic combinations,
often indicated as 'communities.' Mostly they are merely
taxocoenoses (cf. MAAREL 1965).
2. The local distribution pattern of such groupings may coincide
with that of the plant communities in which the groupings have
been established, but mostly it is of a larger, sometimes of a smaller
scale.
3. Most animal groupings are primarily bound to specific abiotic
factors like soil surface moisture or to vegetation structural features
(or, rather, to the micrometeorological features determined by
them).
4. The plant community can be characterized by animal species
groups, although the number of faithful animal species is low. As
long as the synusial structure of animal communities is not sufficiently
known, the phytocoenoses are the most realistic framework
for biocoenotic studies. This implies that attempts like that of
QUEZEL & VERDIER (1953) to create a separate hierarchy of 'communities'
of member-s of one - or a few - animal groups is unrealistic
when these communities are in fact taxocoenoses.
20.7.3 MICROCOMMUNITIES AND SYNUSIAE
Since these community-types are treated by BARKMAN (article
348
16), we refer only to some cllrrent literature. The BRAUN-BLANQ.UET
approach has been applied to various kinds ofmicrocommunities
and synusiae. Bibliographies have been composed for: epiphytic
communities (BARKMAN 1962, Exc., 1966, Exc.), marine algal
communities (HARTOG 1967 Exc.), epigeic moss and lichen communities
(HUBSCHMAN & TuxEN 1964, Exc., TUXEN 1964, Exc.,
1968/1969, Exc.) and soil inhabiting communities (APINIS 1969,
Exc.) Classification systems have been deVeloped by HARTOG (1959)
and BOUDOURESQ.UE (1971) for marine algal communities; HARTOG
distinguished associations which were grouped into formations,
BOUDOURESQ.UE presented a complete hierarchic system. Broad
classification systems of moss and lichen communities have been
presented by KLEMENT (1955), KOPPE (1955), BARKMAN (1958),
WILMANNS (1962). From these studies one may conclude that the
BRAUN-BLANQ.UET system is generally applicable to microcommunities
and synusiae, and that it is advantageous to approach the latter
through a classification separate from that for phytocoenoses (see
also article 16.5 for references).
20.7.4 COMMUNITY COMPLEXES
A 'complex' is a set of contiguous or continuous communities
forming a mosaic or pattern. Complexes can thus be studied on
various levels from the microtopography of bogs to whole landscapes.
Small-scale topographic complexes may often be studied
without direct concern for successional relationships, even though
the communities may have developmental relations to one another
(as in bogs) or to the potential natural vegetation (SEIBERT 1968).
In situations where man has drastically changed the vegetation
pattern as well as the substrate or in geologically young systems
such as coastal dunes it may be more realistic to approach complexes
directly in their local mosaic or zonation structure. A difficult
type of complex is the 'superposition complex' COberlagerungskomplex)
within which rapidly changing community patterns occur,
e.g. with aquatic or with ephemeral communities (cf. MULLER
1970).
. For communities related by successional process the concept
of community ring (Gesellschaftsring) was mentioned by KRAUSE
(1952) and developed by SCHWICKERATH (1954b) and SCHMITHUSEN
(1961). The ring comprises the series of communities that are syndynamically
related to a terminal community. SEIBERT (1968)
translated the concept as 'circle of communities', a term that unfortunately
invites confusion with BRAUN-BLANQ.UET'S (1932,1964)
349
'circle of vegetation' or 'Vegetationskreis'. The English-language
terms 'climax-complex' and 'successional complex' (see article
14.4.2) may well be preferred.
.Study of broader topographic and edaphic complexes in relation
to climax theory was initiated by TUXEN. (For an English
account of TUXEN'S views, see KUCHLER 1967). TUXEN & DIEMONT
(1937, see also TUXEN, 1933) developed the parallel concepts of
'climax swarm' and 'climax group' for spatially contiguous stable
communities (Dauergesellschaften) that occur in the same climate,
but differ in consequence of direction of exposure in mountains
(swarm), or soil parent material in lowland areas (group). TUXEN
(1933) proposed also the term 'paraclimax' for a widespread stable
rommunity the characteristics of which are determined by soil development,
and not climate alone. TUXEN furthermore recognized
a series of climax-regions for Europe, each characterized by a
number of character-associations bound to it in their distributions,
together with companion-associations (Begleitassoziationen) of
wider distribution in more than one climax-region. The latter idea
is a development beyond the climax-complex of BRAUN-BLANQ:UET
(1928, 1964) and others; it characterizes regions not by a single,
somewhat hypothetical climatic climax (Schlussgesellschaft), but
by associations used as diagnostic-communities.
TUXEN (1937) also recognized the coherence between the
various substitute communities (Ersatzgesellschaften), both the
completely cultural and the semi-natural ones, which may occur
in the space that could be (and may actually partly be) occupied
by a terminal community. Such locally coherent communities
were consequently called 'contact communities.' The recognition
of the coherence of substitute communities in their potential
development towards one terminal community led TUXEN (1956)
to the concept of potential natural vegetation. This concept is defined
in relation to a given habitat, as the vegetation that would
finally develop (terminal community) if all human influences on
the site and its immediate surroundings would stop at once, and if
the terminal stage would be reached at once. With the latter restriction
abiotical (e.g. macroclimatological) changes during this development
are meant to be excluded. Although in many cases the
potential natural vegetation may be identical with the original vegetation
and interpretable as such, the distinction of this concept is
useful for the following reasons: (i) There may have been no 'original'vegetation
at all, since man has influenced the ecosystem under
consideration at least as long as the present climate (and thus the
present climax complex) lasts; (ii) Man may have induced irreversible
changes in the ecosystem, so that the presumed 'original' veg-
350
etation can never establish again; (iii) reconstruction of the original
vegetation may theoretically be possible in some cases, but precarious
in practice.
Further discussion of climax interpretation is beyond our purposes
here; we refer to WHITTAKER (1953), DANSEREAU (1957),
SCHMITHUSEN (1961) and BRAUN-BLANQ.UET (1964). We mention
as significant, however, the phytosociological study of community
complexes over wide geographic areas by KRAUSE (1952). KRAUSE
observes the consistency of community composition over extensive
areas together with the occurrence of vicariant communities, particularly
between eastern and western Europe. The communitycomplex
of a given area is shaped both by its macroclimate, and by
local factors producing mosaic-like community relationships. Regions
are characterized by different complexes, and different prevailing
or dominant communities. Complexes give way to other
complexes through geographic distance in three ways: (i) compression
of communities into smaller areas and exclaves, (ii) increasing
floristic impoverishment until a given community can be traced
only in infrequent fragments, and (iii) the replacement ofdominant
communities by vicariants suited to other climates. For further
discussion of vegetation areas and regions see SCHMITHUSEN (1961),
BRAUN-BLANQ.U'ET (1964), and SEIBERT (1968).
20.7.5 ApPLIED PHYTOSOCIOLOGY
The floristic-sociological approach has been applied in many
neighbouring disciplines. Early indications of this application were
presented by BRAUN-BLANQ.UET (1930) and by TUXEN (in BRAtJNBLANQ.UET
& TUXEN 1932) who observed several non-botanical
sciences with which phytosociology might have mutual relationships:
geology, hydrology, geomorphology, climatology, soil science,
animal sociology (see 20.7.2), geography, archaeology, palaeobotany
(particularly palynology), nature conservation and land
use planning. Applications of phytosociology were ·often based on
vegetation maps. Above all, under TUXEN the Zentralstelle (later
Bundesanstalt) fur Vegetationskartierung (Federal Institute for
Vegetation Mapping) at StolzenaujWeser, West-Germany, developed
vegetation mapping and its application in land interpretation
and management and thereby contributed considerably to
the acceptance of phytosociology as a significant applied, as well
as basic science. Many studies at the Institute, models of such research,
can be found in the series Angew,andte P.fianzensoziolog~e
(StolzenaujWeser) and Mitteilungen der Floristisch-Soziologische Arbeits-
351
gemeinschaji (Neue Folge, Stolzenau). See also TuxEN (1956) for a
survey of maps, BRAUN-BLANQUET (1969) for a survey of TuxEN's
work, and KUCHLER (1967) for an English account of floristicsociological
vegetation mapping.
Since applied phytosociology will be the subject of other volumes
of the Handbook, we present here only a survey of main applied
floristic-sociological literature and bibliographies. General
publications on agriculture and forestry include textbooks by
KNAPP (1949) and ELLENBERG (1950, 1952, 1954a) as well as the
series Angewandte Pjlanzensoziologie (Stolzenau) and the symposia
on Vegetation Mapping, Anthropogenic Vegetation, Plant Sociology
and Palynology, Landscape Ecology and Experimental Phytosociology
of the International Society for Plant Geography and
Ecology (TuxEN 1963, 1966b, 1967c, 1968a, 1969b). Further
general references are found in Excerpta Botanica, Sectio Sociologica.
An important general conclusion may be drawn from applied
research: detailed floristic-sociological description of vegetation
combined with adequate analysis of its environment provides a
basis for applied phytosociology in many fields of research which
can be of great social importance, at least in the cultural landscapes
of Europe.
20.7.6 STRUCTURAL CONSIDERATIONS
BRAUN-BLANQUET established the association on a floristic-sociological
basis, but since the floristic-sociological characters of an
association are supposed to n~flect all other characters a floristicsociologically
uniform association might be expected to be structurally
uniform as well. The original association concept thus implied the
physiognomic uniformity of the association. As WESTHOFF (1967)
showed, it is not always true that floristically consistent units are also
structurally uniform. Many examples are known of considerable
structural differences - even on the formation level - within associations
or higher syntaxa (e.g. BARKMAN, 1958b). In a number of
cases the discrepancy between floristic and structural uniformity
has been solved by a refined syntaxonomic treatment.
An example is the Salicornieto-Spartinetum BRAUNBLANQUET
& DE LEEuw (1936) of NW-Atlantic mud flats. This association
was considered a mosaic of patches of two different life
forms, hence of two different structural types. New releves taken
separately from the two kinds of patches revealed them to be floristically
different, which led to separated associations (cf. BEEFTINK
1962, 1965). These associations were in fact assigned to separate
352
classes, viz. Spartinetea and Thero-Salicornetea (e.g. BEEFTINK,
1962). A second example from a very different situation is the
classification of scrub and mantle communities separately from the
woodland classes to which they used to be assigned. TUXEN (1952)
created for these the order Prunetalia spinosae which he later
(1962) placed in a separate class Rhamno-Prunetea (after
RIVAS GODAY & BORJA CARBONELL).
Besides attempts to use structural criteria in floristic-sociological
classification various suggestions for an integration of the
latter system with physiognomic systems have been put forward.
A connexion between the syntaxonomical class and the physiognomic
formation has been sought by DOING (1962) and PASSARGE
(1966, 1968, SCAMONI et al. 1965). PASSARGE (1966) adopted a
formation definition that was based on the original concept of
Grisebach (See BEARD, article 13), as well as a growth-form definition
from SCHMITHUSEN (1961): a phytosociological formation is a
sociological vegetation unit dominated by related growth-forms and
hence showing a uniform physiognomy. Informal arrangements of
classes within formations have been presented by ELLENBERG (1963)
and WESTHOFF (1967), WESTHOFF & HELD (1969). One could also
think of connexions between formations and either of the floristicsociological
units above the class, viz. the division and the vicariant
class group.
20.7.7 INDICATOR GROUPS
One of the essential bases of applied phytosociology is the use
of indicator groups of species. It is consistent with - indeed it expresses
- the perspective of the BRAUN-BLANQ.UET approach that
species are used, on the basis of their distributions, as indicators of
biotope characteristics and other factors, and that when possible
groups of species are thus used. Groups of species may often give
much more effective indication of environmental factors than dominant
species, or other single species alone, or abundance relations
that may be much affected by disturbance. Discussions of
indicator use of species have been given by, among others, DuVIGNEAUD
(1946), ELLENBERG (1950), WHITTAKER (1954a) and
TUXEN (1970b). We may consider indicator species in four contexts ....:
in connexion with synsystematic, biotope or ecological, successional,
and geographic indication. In each of these, but particularly in the
biotope-ecological function, it is possible to· use indicator species
in a perspective of either classification or gradient analysis.
353
20.7.7.llndicators for Classification
One direction of development - the synsystematic use of groups
of species as indicators for syntaxa - is implicit in the BRAUNBLANQ.UET
approach. The concept was developed further, however,
with the distinction of differentiating taxon groups (SCHWICKERATH
1942, 1954a, DUVIGNEAUD 1946), and the idea of sociological
groups (SCAMONI & PASSARGE 1959, 1963, DOING 1962, 1969b).
The latter are established as groups of species that are distributionally
related, and that consequently occur in and characterize,
particular syntaxa; the idea is related to that of 'commodal' species
(WHITTAKER 1956). SCAMONI et al. (1965) constructed a sort of
'sociological profile' by arranging releves for phytocoena and determining
the distribution of sociological groups over the units compared.
Treatment in terms of syntaxonomical groups has been
developed by SEGAL & WESTHOFF (1959) and MAAREL (1969 et seq.).
Clearly, groups of species should indicate characteristics of
biotopes or habitats, as well as the particular phytocoena that occur
in those biotopes. The concept of ecological groups - sets of
species that, because of their similar distributional response to environmental
factors, tend to occur together and to indicate properties
of biotopes - may be traced from DOVIGNEAUD (1946,
1949). Ecological groups have had most extensive application by
ELLENBERG (1950, 1952, 1956), who relates them both to environmental
factors and to the syntaxa of the BRAUN-BLANQ.UET approach.
By describing species responses to various important factorssuch
as pH, lime, moisture, nitrogen - one may arrive at integrated
ecological characterizations of species (ELLENBERG 1956,
compare article 5.4.1.) Ecological groups have been used also by
investigators of the Centre d'Etudes Phytosociologiques et Ecologiques
at Montpellier. (GOUNOT 1969, GUILLERM 1971, DAGET et
al. 1972).
A third application is recognition of geographic indicator
groups that express geographic-historic influences and may characterize
vicariant phytocoena. SCHWICKERATH (1942) and OBERDORFER
(1957) have used geographic groups in dealing with vicariant
associations. Given classification of species into geographically
defined groups, or areal types, it is possible to characterize
communities by geographic spectra, based on representation of
these groups. The geographic perspective has been developed by
MEUSEL et al. and by M0LHOLM HANSEN and BaCHER in the Danish
school (see article 18.5.3-4), and in the United States WHITTAKER
(e.g. .l954b, 1960) has compared areal-type spectra of communities.
354
20.7.7.2 Indicators for Gradient Relations
Community spectra may intergrade, perhaps continuously,
along major axes of biotope characteristics, geography and climate,
and succession. Spectra (per cent representation of different species
groups) may consequently be used to indicate position along these
axes. There seems to have been little application ofindicator groups
to the study of succession, though the recognition of 'decreaser,
increaser, and invader' groups in response to grazing disturbance
(article 4.2.1) should be noted. Study of gradients of geographic
spectra also has been limited, though ELLENBERG (1950) deals with
climatic indicator groups, and article 3 (Fig. 8) illustrates a gradient
in geographic spectra. A third possibility, use of representation
of ecological groups to indicate position along environmental
gradients, has had such extensive development as to be beyond review
here. Use of ecological groups in direct ordination is discussed
in articles 2-5 and 8; use in indirect ordination in article 9. The
most extensive application in phytosociology is that of ELLENBERG
(1950, 1952, 1956). ELLENBERG classified European species (weeds
in agricultural fields, and plants of meadows and pastures) into
ecological groups on the basis of their responses to various environmental
factors. Weighted averages (see 2.2.3.1) for samples
expressed their positions in relation to these factors. At the same
time the relation of the species to syntaxa was known, and it was
consequently possible to interpret the relations of the syntaxa to
one another and environment. The technique thus created a multidimensional
direct ordination, and belongs also to section 20.9.1.
20.8 NUDlerical Techniques
There remain to be discussed two (closely related) recent developments
in phytosociology - numerical techniques and ordination.
For review of similarity measurements and approaches to
numerical classification we refer the reader to GOODALL'S articles
(6 and 19) in this volume. We discuss briefly, liowever, work in
this direction employing the BRADN-Bu.NQUET approach.
20.8.1 STORAGE OF RELEVES
Although the numbers of releves treated in syntaxonomical
studies may be very high - often several thousands - the number of
storage systems is low. Most individual phytosocio10gists and t~eir
355
institutes work with an archive of releve note books and tables.
Such is the case at syntaxonomic centres like the Station Internationale
de Geobotanique Mediterraneene et Alpine (S.I.G.M.A.)
at Montpellier and the former Bundesanstalt fur Vegetationskartierung
at Stolzenau (data from the latter are now at TDxEN'S
Arbeitsstelle fur theoretische und angewandte Pflanzensoziologie at
Todenmann uber Rinteln, West Germany). Some archives have
been set up with visual punched cards. The only such archive of
some size seems to be that of the Geobotanical Institute Rubel at
Zurich including about 5500 forest releves from Switzerland (ELLENBERG
1968).
Besides incidental use of machine punched cards in the application
of computer techniques, two archive systems based on computer
hardware are known to us: the system of the C.E.P.E.,
Centre d'Etudes Phytosociologiques et Ecologiques at Montpellier
and the system of the Working Group for Data-Processing of the
International Society for Plant Geography and Ecology. The
C.E.P.E. system (see EMBERGER 1968) has the following characteristics:
(i) The basic unit is the rcleve, which is defined as the
whole of ecological and phytosociological observations at a particular
site. (ii) General vegetation structure, situation of the site,
general climate and soil are described by means of measurements
or estimations of numerous factors. (iii) For each factor a code is
devised with, in most cases, 10 classes. (iv) All coded determinations
of general features are punched in three 'parent' releve cards,
'Cartes-Maitresse-Releve' (CMR) of which columns 1-71 (except
for CMR 3) are used. (v) Each taxon from the releve is treated separately
for each vegetation layer in which it has been observed.
Coded determinations include vegetation layer, abundance, and
dominance (in separate 0-9 codes), pattern within the site, phenological
state, vitality and life-form. The corresponding punched
cards are called 'detailed cards', Cartes Detail (CD) and used only
from column 72 onwards. (vi) Species are enumerated from 0001-
4779 according to the Quatre flores de France by FOURNIER (1961).
Cryptogams are provisionally enumerated from 5000 onwards.
(vii) Data per species per layer are combined with CMR data,
which results in three combined cards.
In this way a relatively large archive arises, which has the
advantage of being rapidly accessible to all kinds of spectrum and
correlation calculations.
The 'Working Group' was established in 1969 during the
International Symposium at Rinteln after a proposal by S. PIGNATTI,
G. CRISTOFOLINI and D. LAUSI (Trieste). One major aim
(see MAAREL 1971) was the treatment ofreleves in such a way that
356
the data could be readily stored and retrieved. An outline of the
system has been presented by CRISTOFOLINI et al. (1969); its main
characteristics are: (i) The basic unit is a full reIeve table, either
published or in manuscript form. (ii) Rows of releve tables are
punched - from column 21 onwards. Cover-abundance values per
species are coded either as direct BRAUN-BLANQUET symbols or in
a 0-9 transformation .code (Table II). (iii) In the first 20 columns
coded data on publication, ta ble number in publication, and taxon
number are punched. (iv) Taxa are coded with 7 digits, 4 for the
genus, 3 for the species - as yet no space is left for infraspecific taxa.
The genus enumeration is according to the world survey Genera
Siphonogamarum by DALLA TORRE & HARMS (1900-1907, reprint
1963). Species enumeration is as yet restricted to European species
(Russia not included), according to Flora Europaea as far as possible,
otherwise according to a provisional enumeration based on six
large standard floras of Europe. Additional provisional codes for
cryptogams are being developed. (v) A separate punched card is
used for bibliographical data on the paper (or set of unpublished
tables). (vi) A separate set of punched cards is used for general data
on the site and its situation, for each releve in each table. This part
of the system is still in development. The Montpellier system, which
is already elaborated in this respect, could be taken as a starting-
point.
The Working Group decided to concentrate activities on saIt
marsh communities. Some 3400 releves (including 576 Spartinetum
releves) have been stored in punched card decks and on
magnetic tape.
20.8.2 SPECIES CORRELATION
The study of species correlation was introduced by VRIES (e.g.
VRIES et al. 1954, DAMMAN & VRIES 1954). The plexus of grassland
species he presented (article 7, Fig. 3, p. 169) was taken over in the
textbooks of ELLENBERG (1956) and BRAUN-BLANQ.UET (1964), who
underlined VRIES' conclusion 'that the resulting species groups
broadly coincide with the associations of the Ziirich-Montpellier
school.'
Measurement of species distributional similarities may be
based on either binary or quantitative data. Distributional similarity
based on binary (presence and absence) data may be termed
'species association,' that based on quantitative representation in
releves, may be termed 'species correlation' (see article 6). In the
BRAUN-BLANQ.UET approach the quantitative weightings may be
357
either the abundance-cover scale (20.4.5.1) in individual releves (or
a transformation of that scale), or presence degrees (20.5.3) in
different phytocoena. Species association seems to be more appropriate
for the detection of species groups in releves representing
an extensive range of community difference, whilst correlation
studies are more appropriate for intensive study within the range
of broadly overlapping species (cf. GREIG-SMITH 1964). The use of
species correlation in the establishment of sociological groups is
obvious, but phytosociologists using such groups have hardly applied
this technique. The studies of HEGG (1965; see article 7, Fig.
7, p. 174) and STOCKINGER & HOLZNER (1972) may, however, be
mentioned as examples (see also article 7).
The detection of ecological groups has been particularly developed
by phytosociologists using factor analysis and principal component
analysis. DAGNELIE (1960, see also article 9) described the
use of factor analysis for the establishment of sociological groups,
and the use of joint species-environmental factor analysis for the
establishment of ecological groups. GODRON (1966) used species
correlations for the checking of 'imbricating' ecological groups;
GUILLERM (1971) and DAGET et al. (1972) constructed ecological
profiles of species against environmental factors. LACOSTE & Roux
(1971) and ROMANE (1972) used 'factor analysis ofcorrespondences'
in which floristic and environmental variables were analysed in
combination to arrive at ecological-sociological groups.
Another such method was developed by FRESCO (1971) as
part of his 'compound analysis.' He constructed overlapping species
groups based on similarities between species with respect to
their loadings on eigenvalues extracted in subsequent factor analyses.
This method could be applied to very large data sets, e.g. the
first selection of salt marsh data (1296 releves, 245 species). From
this salt marsh treatment (with an arbitrary similarity level) 65
such groups were derived which could be interpreted roulshly as
alliance character-species groups.
20.8.3 FIDELITY TESTS
Although it seems obvious to use tests on the significance of
the exclusiveness or differentiating value of species (or species
groups) for a given vegetation type, phytosociologists have hardly
done so. MEIJER DREES (1949) suggested the use of the t-test for
the significance of the difference between the group amounts of
. species groups in different releves of a set. In his example the releves
were assigned to a particular association and the two species-groups
358
examined were syntaxonomic groups for two orders; the problem
was the assignment of the association to one or the other order.
For such situations GOODALL (1953b) used FISHER'S discriminant
function, which is more powerful but also more laborious.
GOODALL (1953b) also proposed an index of fidelity for a species
based on the ratio of constancy values for that species in two
communities, which could be tested with the X2 test or by exact
calculation of P. In addition he suggested the use of this index as
an 'indicator value' for the comparison of the constancy of a species
in one community with that in various other communities - GOODALL
spoke of 'all other communities in the area.' In usual phytosociological
terms GOODALL'S indicator value is the real fidelity
index and the application of this measure to the two-community
comparison would be an 'index of differentiation.' Such tests may
have limited value, particularly because of the non-random selection
of releves from which they are derived (GOODALL 1953b).
Still, their use may be experimented with for syntaxon diagnoses
from tabular summaries.
20.8.4 RELEVE SIMILARITY
Measurements of similarity between releves have long been
accepted as one basis of grouping these into phytocoena (BRADNBLANQUET,
1928, 1932). Various coefficients of similarity and dissimilarity
are in use in phytosociology, including the formulas of
JACCARD and S0RENSEN (article 6, formulas 39 and 40). Recently
an anonymous 'similarity ratio,' which was introduced by WISHART
(1969) in CLtJSTAN, a set of classification programmes, was used by
KORTEKAAS & MAAREL (1972). This formula reads
l>iyJ~>~+LiY~- LiXiYi,
and is thus a generalisation of JACCARD'S formula. Comparatively
little use has been made of direct dissimilarity measures (GROENEWOUD
1965, MAAREL 1966b). Similar formulas, most of them derived
from S0RENSEN'S formula, are in use for the comparison of sets
of releves or the calculation of average similarity within a set (cf.
S0RENSEN 1948, RAABE 1952, CESKA 1966, 1968, FRYDMAN &
WHITTAKER, 1968, MAAREL, 1969). Weighting of species may use
presence values or a combination of presence with a~ importance
value. ELLENBERG (1956) as well as PIGNATTI & MENGARDA, see
PIGNATTI 1964, suggested the use of a similarity coefficient in the
assignment of a releve to a type by comparing it with a 'standard'
or average rcleve, e.g. one consisting only ofthe normal characteristic
359
species combination. HOFMANN & PASSARGE (1964) and SCAMONI
et al. (1965) determined similarity between releves and sets of
releves with reference to the sociological groups ('group affinity').
20.8.5 MEASUREMENT OF HOMOTONEITY
Homotoneity we have defined as the homogeneity of a community
table, the relative consistency of the releves the table comprises
(20.5.3). In addition to the simple measures mentioned in 20.5.3,
the mean similarity coefficient for the releves, as calculated in a
direct way by CESKA (1966), is a convenient measure, particularly
in classifications based on releve similarity. Either the average of
all interreleve similarity values within a table, or the average releve
similarity with the average for the table (i.e. the centroid of
the releve cluster) may be used. See also article 6 and 20.8.4 and
20.8.8 on similarity measures.
Alternative techniques based on species relationships include
the use of positive interspecific correlation by GOODALL (1953a),
association analysis (WILLIAMS & LAMBERT 1959), and information
analysis (WILLIAMS et al. 1966) (see article 19.5.1.5 and 19.5.2.5).
MAAREL (1966b) derived an information measure that is related
to that of WILLIAMS et al. and appeared to be a useful heterogeneity
coefficient in a study of a local grassland community complex.
GODRON (1966) developed an information measure, based on
BRILLOUIN'S information formula, for determination of the heterogeneity
within a series of contiguous quadrats in the field. The frequencies
of the species occurring in the set determine the value of
this measure. It could also be used for heterotoneity in sets of re-
1eves and in that case is rather similar to the formula of WILLIAMS
et al. (1966).
DAHL (1957, 1960) derived an index of uniformity from the
FISHER model (logarithmic series) for species-individual and species-area
relations (S = a loge (l+N/a), S is number of species, N
number ofindividuals, and a a diversity index, see WILLIAMS 1964).
DAHL considered the species-releve relation to be similar and defined
the diversity parameter for a table as (Sm-S)/alogM (see
symbols and formulas in 20.5.3). The index of uniformity is accordingly
defined as Sla. A clear relationship with CURTIS' (1959) index
was shown by DAHL (1957).
20.8.6 NUMERICAL CLASSIFICATION
Of the various numerical classification techniques which are
360
described in article 19, only a few are in use in quantitative phytosociology.
Direct use of similarity matrices was developed by Polish
phytosociologists (MATUSZKIEWICZ 1948, FALI~SKI 1960) and by
GUINOCHET (1955, GUINOCHET & CASAL 1957), who spoke of the
'differential analysis of CZEKANOWSKI' (after the Polish originator
of the approach). See further article 7.3.1.
Agglomerative clustering techniques (19.5.2) based on similarity
coefficients like the JACCARD and S0RENSEN indices were used
by S0RENSEN (1948), LOOMAN & CAMPBELL (1960), MAAREL
(1966b), MOORE & O'SULLIVAN (1970), MOORE et al. (1970) and
KORTEKAAS & MAAREL (1972). Agglomerative techniques can be
profitably combined with re-allocation techniques (cf. WISHART,
1969). The releve-groups arrived at in these classifications could
generally be typified with characteristic species combinations and
connected with syntaxonomical units.
I VIMEy-COOK & PROCTOR (1966) applied nodal analysis (association
analysis followed by an inverse species clustering) to salt
marsh, fen and woodland data. They obtained tables with clear
blocks ofreleves and species and concluded that the results confirmed
the arrangement of the data arrived at by traditional phytosociological
methods. The technique was considered particularly useful
for the detection of differentiating species. Moreover they found
minor lines of division which had not been obvious in previous
studies.
20.8.7 TABLE REARRANGEMENT
A successful nodal analysis should produce an ordered table.
A number of special techniques for table rearrangement have been
developed, from the early work of BENNINGHOFF & SOUTHWORTH
(1964) to the more recent of MOORE (1972, MOORE et al. 1970,
MOORE & O'SULLIVAN 1970). The following computer programmes
are known to us:
i) BENNINGHOFF & SOUTHWORTH (1964), G. W. MOORE et al.
(1967), LIETH & MOORE (1971). This programme finds species
clusters by application of LIN'S algorithm for the 'travelling salesman
problem.' It is applied to species with intermediate presence
in the table. Additionally, releves are sorted. Final table ordering
is by hand.
ii) J.J. MOORE (1971, last version). This programme'PHYTO'
rearranges both species and releves. It finds pairs of species with
intermediate presence values and high co-occurrence values. Opposing
pairs are used as differentiators. Further arrangements are
361
dictated by the user, either through suspected mutually exclusive
species or through preferred order of species and releves.
iii) SPATZ (1969, 1972 last version). This programme forms
groups of releves on a similarity basis. Differentiating species are
defined as species with presence of > 50 % in one group and < 10%
in all others. Output includes a synoptic table with synthetic figures.
iv) SCHMID & KUHN (1970). This programme calculates D2
values and forms groups on lowest D2 basis. Differentiated tables
can be dictated.
v) CESKA & ROEMER (1971). This programme finds speciesreleve
groups through iteration. Diagnostic species are defined as
in programme 3 (values vary from 50 vs. 10 to 66 vs. 33). A releve
is considered a member of a group if it contains at least 50 % (or
66 %) of the diagnostic species of that group. Groups are arranged
in order of size and within-group similarity. Final arrangement is
usually by hand.
vi) JANSSEN & MAAREL (1972, JANSSEN 1972). This programme
is based on programmes HIERAR and RELOC of the CLUSTAN
set (WISHART 1969) with a choice of 10 similarity coefficients. It
starts with an allocation of releves to groups, either at random or
as determined by previous classifications and phytosociological experience.
Group arrangement is based on group properties, particularly
the relation between reieve number and species number, or
alternatively on position along the first component of a principal
components analysis of groups. Species are arranged so as to form
a diagonal structure in the table. Significance of positive differentiating
value of species in one or more groups is determined by
a X2 test. Final arrangement is usually through MOORE'S PHYTO-
programme.
vii) STOCKINGER & HOLZNER (1972). This programme finds
species groupings through similarity analysis. Releves are sorted
according to occurrence of species groups.
It follows from this description that all programmes require
the personal finishing touch of the user; this is felt to be not an
inconsistency but rather a matter of efficiency. Most designers
claim a considerable gain in accuracy, since re-writing tables by
hand is no longer necessary. The speed of the programmes will
depend on type of computer and size of the table. Programmes i,
ii, v, and vii are most adapted to tables with comparatively many
releves and few species, programmes iii and iv to tables of the reverse
form. Programme vi has a larger capacity, but requires preliminary
experience to be fully efficient.
362
20.8.8 NUMERICAL SYNTAXONOMY
Numerical syntaxonomy has been mainly restricted to the
characterizing of various syntaxon levels by average similarity.
Various coefficients of similarity and dissimilarity are in use in
phytosociology. 'Coefficients of community' comparing samples by
the per cents oftheir species shared have been used most, with about
equal interest in the formulas ofJACCARD and S0RENSEN (article 6,
formulas 39 and 40). Formulas using importance values for species
(cover-abundance in releves, especially) include the 'percentage
similarities,' notably the CZEKANOWSKI index (article 6, formula
50). Similar formulas are in use for comparing sets of releves (representing
different phytocoena), or calculating average similarity
within sets (see S0RENSEN 1948, RAABE 1952, CES KA 1966, 1968,
FRYDMAN & WHITTAKER 1968, MAAREL 1969, 1972a). Comparison
of sets of releves may use presence per cents, or mean cover-abundance
values, for the species in these sets. Numerical classification is
further discussed in article 19.
S0RENSEN (1948) concluded that grassland groupings with
similarity levels of 0.40 roughly corresponded with syntaxa on the
alliance level. ELLENBERG (1956) stated that the average similarity
of releves within one association is between 0.25 and 0.50,
whereas subunits may be distinguished by levels above 0.50 (JACCARD
values, which are 10 % lower than S0RENSEN values on the
average). LOOMAN & CAMPBELL (1960) calculated S0RENSEN values
of > 0.70 within subunits belonging to one grassland association,
whereas values between subunits were all < 0.50. HOFMANN &
PASSARGE (1964) presented group affinity values between various
woodland associations and subassociations. Within associations
values were mostly > 0.60, between typical subassociations of
related associations values were mostly between 0.30 and 0.50.
RAABE (1952) obtained affinity values (KULCZINSKI coefficient)
between associations and alliances of weed, salt marsh and alpine
communities. Within-alliance values were between 0.40 and
0.50, between-alliance values were 0.20 to 0.40. Within salt
marsh associations values varied more widely, from 0.30 to 0.80,
and here the influence of geographical distance between local
representatives of associations was evident.
NEUHAUSL & NEUHAUSLOVA-NovOTNA (1972) studied withinand
between- group similarities fS0RENSEN values) of many woodland
associations of the alliance Carpinion. Within"group similarities
oflocallyestablished associations and lower units were generally
over 0.55. When larger areas were involved, these values were
lower but still over 0.45. Similarities between associations and be-
363
tween lower units were always lower, mostly O.IO·to 0.20. These
calculations made possible some improvement in the delimitation
of Carpinion associations (which, as the authOls stated, must be
checked by renewed phytosociological table studies). KORTEKAAS
& MAAREL (1972) carried out a numerical analysis of European
Spartina communities involving an agglomerative classification
with relocation of 576 releves. They arrived at four separate dendrograms
each comprising releves dominated by one of the SPartina
species. On various levels of the hierarchy groups were inspected
for character-species and compared with the existing syntaxonomy
(BEEFTINK & GEHU, in prep). For these communities values of the
similarity indices between 0.40 and 0.60 could be connected with
the association level, 0.61-0.70 with the subassociation level and
0.71-0.80 with the variant level. Figure 4 presents the dendrogram
for the Spartina maritima releves. Each 'subassociation' had
exactly one 'good' differential-taxon. The examples suggest that in
related communities a reasonable parallel between syntaxonomical
level and similarity level can be established. Furthermore, the
0,00
similarity
level 0,10
0,20
0,30
0,40
0,50
0,60
0,70
0,80
0,90
1 , 00
"Spartinetum
maritimae"
perennis"
"Aster/
Itripolium"
"tYf:I~~cornia europaea"
"Li oni m
V~lgare" "Suaeda maritima"
"typicum" "tYPiCU~"
'Fig. 4. Dendrogram of Spartinetum maritimae releves. (KORTEKAAS &
MAAREL, 1972).
364
similarity values for a given syntaxonomic level in different vegetation
types are similar, suggesting that various authors have made
comparable intuitive judgments of relative similarity for various
kinds of communities. Similarity measurements should thus be
thought a useful aid to synsystematics.
20.9 Ordination
'Ordination' refers to the arrangement of entities (generally
samples, or species) in a uni- or a multidimensional order (RAMENSKY
1930, GOODALL 1954, articles 1 and 2.) Ordination is often considered
to stem from the concept of vegetation as a continuum,
whereas the floristic-sociological classification may seem to assume
discontinuity. It may consequently be argued whether ordination
is appropriate in phytosociology. Even if we assume for the moment
that associations are generally discontinuous with one another,
there is no reason ordination cannot be used as an aid to understanding
the relations of releves and lower syntaxa to one another
and environment, within the association. On a different level, ordination
and the perspective ofgradient analysis can well be applied
to the relations of associations and higher syntaxa as wholes, to
one another and environment. We feel consequently that ordination
is a fully acceptable supplementary approach in phytosociol-
ogy.
There has been some dispute, mainly between BRAUN-BLANQ.OET
(1939, 1951b, 1955) and GAMS (1918, 1941, 1954) on the
question 'linear or multidimensional system in plant sociology.'
GAMS emphasized the multidimensional relationships of plant
communities, without indicating the way of constructing an appropriate
system, as BRAUN-BLANQ.UET remarked. WAGNER (1954,
1968) observed that many syntaxa show multidimensional relationships
to various syntaxa of the next higher level. The 'flexibility'
ELLENBERG (1954b) demanded at lower syntaxonomic levels could
be thought of in this perspective. Surely it is true both that linear
arrangements into hierarchies are possible and valuable, and that
study of multidirectional relationships may be rewarding.
20.9.1 INFORMAL ORDINATION
A number of approaches that represent ordination but do not
employ similarity measurements have been a,pplied in phytosociology.
In many ordered phytocoenon tables a suspected underlying
365
environmental gradient is chosen as a basis for ordering that may
be completed by inspection of ecological groups (MOORE et al.
1970). Often a moisture gradient is chosen and a 'dry' a 'typical'
and a 'moist' subassociation are arranged within an association
table. The unidimensional approach known' as compositional ordination
(article 2) has a phytosociological precursor in the approach
OfSCHWICKERATH (1931 et seq.), in which releves of one association
were arranged on the basis of representation of diagnostic
species groups. The development of weighted-average ordinations
by ELLENBERG (1950, 1952) has been referred to (20.7.7.2).
More abstract two- or many-dimensional schemes including
Russian examples (discussed in article 5) were recommended by
GAMS (1941, 1961). The approach of DUVIGNEAUD (1946) was an
early floristic-sociological ordination without numerical basis. A
number of phytosociologists have presented patterns ('mosaic
charts', 2.3.3) of phytocoena in relation to environmental gradients
(see DUVIGNEAUD 1946, WAGNER 1954, 1968, ELLENBERG 1952,
Fig. 5, 1963, ZONNEVELD 1960, HEGG 1965, and article 18, Fig. 1,
p. 555).
20.9.2 FORMAL ORDINATION
The first ordination of releves on the basis of their similarities
is found in the Polish approach of MATUSZKIEWICZ & TRACZYK
(1958), and FALINSKI (1960). They constructed 'dendrites' as arrangements
that can be ecologically interpreted (cf. GAMS 1961,
1967, and article 7).
Ordination in the sense of axis construction was first applied in
phytosociology by DAGNELIE (1960) on beechwoods (see article 9).
A comparable factor analysis on heathlands was presented by
FRESCO (1969). GROENEWOUD (1965), MOORE et al. (1970) and
MAAREL (1972a) applied principal component analysis to similarity
matrices. The Wisconsin polar ordination ofBRAY & CURTIS (1957),
or slightly mod:fi~d forms, was applied in Dutch work (unpublished
MSc theses, Universities of Utrecht and Nijmegen, MAAREL 1966b,
1969, MAAREL & LEERTOUWER 1967, LONDO 1971) and further by
ROGERS (1970). LACOSTE & Roux (1971) applied factor analysis
of correspondences (see 20.8.2) to releves assigned to various subalpine
associations and subassociations.
Ordination can be applied also to phytocoena (or syntaxa),
with each phytocoenon treated as a composite sample with its
species composition summarized as presence per cents or mean importance
values. One may then conceive of environmental gradients
366
r
+1.00
.,H~ir
, (1'"
~l.O"""'O--'--~--'-""'~ '" • "'ul- ,--.... ,·laur 13 1,5,.'thf.
1.3 .........'" .'U'IIOII
\ "I' .... ~ ,~ ,-,.u" ·~ca . •'J -- ~--___-" :-. -- --;1\
AE8OfOIIIIII • \ A·A ptt "I
$(1£</011 FlOVI'nlls \ I
" I
.lG ..'
-tOIl
-1..'tlep.....
A,,,I..
A~,'tII"'"
I.• G·p
+111O~
Fig. 5. Ordination ofnitrophilous edge communities described by TiixEN (1967a).
from MAAREL (1969).
367
relating these phytocoena as a multidimensional habitat space
(GOODALL 1963, WHITTAKER 1967) in which the centroids of the
phytocoena (as clusters of re1eves) are ordinated. ORLOCI (1966)
carried out such an ordination by principal component analysis;
GITTINS (1965), CRAWFORD & WISHART (1966, 1967), FRYDMAN
& WHITTAKER (1968), MAAREL (1969), and WHITTAKER (1972)
applied polar ordination to community-types. Mostly the similarities
between types were measured by presence or constancy values.
All the resulting ordinations were effective and easily interpretable.
An effective approach to interpretation is the plotting of contour
lines for biotope measurements (e.g. FRYDMAN & WHITTAKER
1968), to determine the relationship between axes and environmental
factors, and the plotting of community-types and representation
of diagnostic groups (FRYDMAN & WHITTAKER 1968, MAAREL
1969, WHITTAKER 1972) to show the relation of these to environment
and one another. Fig. 5 illustrates the latter technique (see
further articles 2, 8, and 9).
When no appropriate environmental data are available it is
possible to interpret the ordination by plotting the distribution of
syntaxonomical groups. Figure 5 presents an example. Nitrophilous
edge communities, bordering woodlands and scrubs, as described
by TUXEN (1967a) are ordinated and the ordination space is characterized
by various alliances. The associations are reasonably
separated in the ordination space, except for the Agropyro repentis-Aegopodietum.
The subassociation with Chaerop~yllum
hirsutllm, in particular, is distinct and has a considerable representation
of A1no-Padion species. Patterns of three alliances are presented.
Lines with equal percentage amounts of one alliance are
called 'isocenes' and outlined. The greatest difference in the set of
types is between the relatively dry, open nitrophilous Agropyro
repentis-Aegopodietum ca1ystegietosum sepium from
anthropogenic edges at low altitudes, and the relatively moist and
shaded Geranio phaei-Petasitetum along streamlets in the
montane region. WHITTAKER (1972) and FRYDMAN & WHITTAKER
(1968) presented a similar approach with similar results. Besides
species populations, species groups, and environmental measures,
various spectra can be plotted in an ordination field. RAUNKIAER
life-forms, for example, showed clear distribution patterns when
thus plotted by FRYDMAN & WHITTAKER (1968).
Other uses of ordination include applications to succession
(MAAREL 1969, LONDO 1971) and ordinations based not on species
composition but on structural characters, life-forms or sociological
(syntaxonomic) groups (KNIGHT 1965, GOFF & COTTAM
1967, MAAREL 1969, WHITTAKER 1972). MAAREL (1972a) used
368
Amenfiflorae
5"10 >
<
+ 1.00
5"1. 5"10
< "'. > I
: > r<
5"10
> ~:' + O.SO
5" . ,
.~ i: CHA AUN > .: < legumrllosae
C~R!; ,' . G!!I '" FEll cm ....
•roll ..,'" I.~ (So ~ MOll " <
CHAo.'H ll :~MM~•.~II~ ..,. 0" .:: _ _ _ _ 12"1. Centrospermae
ME ·l lo : "' IINI' i ~ "BR~ FIU CA- IE .IGU·;"'----- >
EI!C .iiri··.... QU-IU ' .••••• •• •••••••••'" _ _ __"":: 5" Rhoeadinae
I l:O : • QII-IO INOR 050 ----- > •
l U-flo CAll ' BEIi ~ + . UCI
--~----------'~0~5~0='/~'~---' ~ ~ . I
- 100 CI -OI - . " lUi NIIO 1m " m~UC < +1.00
1I~~/ \·~:!:L._~23"1.
VlCl IINO "'~tC '" Centrospermae
> ..........~S~HA • "
liliiflorae 5'I.~- -- - 'AHU \ BIDE
... "...
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"
- 0.50
lE MN
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.
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12 "to
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---.;;5".
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23 '10
<
Fig. 6a. Ordination of 51 European associations and lower syntaxa on the basis
of their plant order spectra, dimensions 1 and 2. (MAAREL, 1972a).
higher idiotaxonomical units as attributes to compare and ordinate
vegetation types. Figure 6 presents some results of an ordination
of 51 associations and subassociations belonging to 49 alliances, and
thus covering almost the entire variation in central and western
Europe. The corresponding similarity matrix was based on representations
of plant orders in synoptic tables, mainly from OBERDORFER
(1957). The first line of variation shows the sociological
progression from pioneer comminities, with alliances such as the
Bidention (BIDE) and N anocyperion (NANO) on the right,
to mature communities like the Fagion (-FA) and Carplnion
(CARP) on the left. Most grasslands, marshes, and heathlands are
intermediate. This variation is illustrated by isolines of various
idiotaxonomic orders, .e.g. Centrospermae vs. Liliiflorae. The
second axis involves a moisture gradient from wet (Eu-Potamion,
369
-1.00
8rya/es
12 %
+ 1.00
tlicornes
5% 120;.
<~>\<vm:
• I
I
I
I
I
+ 0.50 •
QU-IO
12%
8/cornes
12 %
I
mClMO . POll
+1.00•
« -
LITO
•
5%
Umbelliflorae
5%
5%
Helobrae
- 1.00
Fig. 6b. Ordination of 51 European associations and lower syntaxa on the basis
of their plant order spectra, dimensions 3 and 4. (MAAREL, 1972a)
POTA, below) to dry, (Mesobromion, MESO, upper middle).
Between axes III and IV (Fig. 6b) two more or less oblique lines
of variation can be discerned. The third axis may be tentatively
interpreted as a shading gradient from Polygonion avicularis
(POLY) and Phragmition (PHRA) below, to CardaminoMontion
(MONT) and Vaccio-Piceion (VACC) above, from
prevalence of Helobiae to that of Bryales. The fourth axis runs
from raw humus soils with a relatively high CjN ratio to soils with
a low CjN ratio and high microbiological activity (Ericion tetralicis,
ERIC, with Bicornes, upper right, to Caucalion,
CAVC, with Vmbelliflorae, lower left). We regard Fig. 6 as representing
only a first venture in broad-scale ordination of phytocoena;
its significance may be in its demonstration of possibility.
370
20.10 Conclusion
In perspective, these numerical techniques and ordinations
should be seen as ancillaries to the BRAUN-BLANQUET approach.
We re-emphasize the essential ideas with which we began: the
floristic approach to understanding, the functional value of diagnostic
species for classification and environmental indication, and
the utility of the hierarchical classification for ordering knowledge
and expressing understanding. Given this as the approach, then
the numerical techniques serve to reduce labour, and ordinations
to enhance interpretation. Ordination without prior classification
has accomplished significant research in American ecology, but
ordinations in different areas are not easily coordinated with one
another. It is the European experience that the BRAUN-BLANQ.UET
approach provides such coordination of the work of different investigators
through its classification, whilst the classification provides
a context that increases effectiveness of ordination. On the
one hand prior classification makes possible the ordination of phytocoena,
rather than an often unmanageable number of releves; on
the other hand the known relationships ofspecies groups and phytocoena
to one another and environment give basis for interpreting
an ordination. The ordination should, in turn, express and further
clarify those relationships.
We return to another theme - the complexity of the BRAUNBLANQ.UET
approach as a technical system. The complexity has
causes; these include the numerousness of species and phytocoena,
the multifarious distributional relations of species to one another
and phytocoena, and the many directions of environmental, geographic,
and developmental relationships amongst species and
phytocoena. The complexity of the BRAUN-BLANQ.UET system is
thus a response of science to the complexity of vegetation. It is no
real statement of preference that the approaches through physiognomy
and species dominance are 'simpler'; it is not necessarily
true that they are, and to the extent that they are, they are only
because they do not treat and coordinate so wide a range of information.
We thus express our judgement (or our bias) that in the
study of vegetation one approach has most fully faced the demands
for a .method that is both detailed and generalizing, both locally
intensive and integrative of the results from different local areas and
that that one approach is BRAUN-BLANQ.UET'S.
This judgement we must balance with another. The BRAUNBLANQ.UET
approach is very demanding ofresearch effort by many investigators,
ifthe vegetation of extensive areas is to be known through
it. The effort has been ·possible in Europe and some other areas·
371
but is not yet possible everywhere - there are not phytosociologists
enough. The approach may be particularly difficult to initiate
in an area lacking prior studies to offer suggestions toward classification
and diagnostic species groupings, unless the initiator is
experienced in the approach. This difficulty is one reason for the
importance of approaches through physiognomy and dominance
in many areas; by these approaches knowledge of vegetation may
be gained that might not otherwise be obtained at all. It is true
also that the physiognomic and other approaches may reveal vegetational
relationships that are important, but are outside the main
concerns of the BRAUN-BLANQUET approach. The statement that
BRAUN-BLANQUET'S is the most fully developed approach to vegetation
is thus no rejection of the contributions of others amongst the
many ways for the ordination and classification of communities.
20.l1 SUMMARY
(20.1) Amongst the central ideas of the approach are that: (i)
classification and interpretation of communities should be based
on their full floristic composition, (ii) with emphasis on diagnostic
species, whose relative restriction to samples characterizes communities
and indicates their environments and (iii) which may be
used to organize the communities into a formal, hierarchical classification.
(20.2) This floristic-sociological approach to vegetation
had its origin in southern European phytosociology centered in the
cities of Zurich and Montpellier, but especially in the ideas of
BRAUN-BLANQUET.
(20.3) It is important to distinguish particular, concrete plant
communities or phytocoenoses, and abstract classes of plant communities;
the term phytocoenon is recommended for the latter. The
phytocoena of the formal BRAUN-BLANQUET hierarchy are termed
syntaxa, in analogy with the taxa into which individual organisms
are classified. Syntaxa are characterized by diagnostic species of
three types: character-species are centered in or relatively restricted
to a particular syntaxon compared with all others, and therefore
characterize it and indicate its environment; differential-species
distinguish two closely related syntaxa by presence in most samples
of the one and absence in most samples of the other; constant companions
are not restricted to a given syntaxon but help to characterize
it and indicate its relationships to higher units. In addition to
the hierarchy, syntaxa may be arranged along a sociological progression
from simple and poorly organized to complex and highly
differentiated communities. The BRAUN-BLANQUET classification is
372
considered natural in the sense that its syntaxa (though their limits
may be arbitrary) are consistent with a large number of relationships
between environment, species populations, and biocoenoses.
(20.4) The research procedures include three phases - analytical,
synthetical, and syntaxonomical. In the analytical phase
samples or releves representing kinds of phytocoena to be studied
are taken in the field. The releves should be chosen to represent the
phytocoena to be studied, and each should sample a uniform area
of sufficient size to represent that phytocoenose adequately. A
releve includes information on environment and location and a
list of all plant species in the sample area, with species recorded by
strata and rating scales (for combined cover and abundance, and
if possible sociability, vitality, and periodicity).
(20.5) In the synthetical phase the releves are compared with
one another to derive a preliminary classification. A set of releves
are first listed in their full species composition in a primary table.
Species of intermediate presence values are emphasized in seeking
groups of differentiating species that characterize some of the
releves but not others. Partial tables are prepared using these
species to rearrange the releves into groups characterized by the
groups of differentiating species. When the rearrangement is satisfactory,
the data from all species are recopied into a phytocoenon
table in which boundaries may be drawn to mark off differentialspecies
groups and phytocoena. A synoptic table may now be
prepared in which each column summarizes data for species
occurrence in the releves of a phytocoenon. By use of synoptic
tables the phytocoena under study may be compared with others
from the same area in a search for character-species. Problems
in the recognition of character-species, including geographic differences
in fidelity of species to phytocoena, are discussed.
(20.6) In the syntaxonomical phase the studied phytocoena
may enter the formal hierarchy. The phytocoenon table is revised
into a formal syntaxonomic table indicating the character-species
and differential-species groups represented. If the phytocoenon
represents a new association, this is named and placed in the hiera,rchy
in which associations are grouped into alliances, alliances
into orders, orders into classes. Syntaxa on all these levels may be
defined by character-species; in some cases character-genera are
used for higher units. Apart from the hierarchy, associations (and
other syntaxa) from different geographic areas may be grouped by
floristic affinities, especially as marked by vicariant 'species. Associations
are divided into subordinate units: subassociations and
vario.nts characterized by differential-species, and facies as the
lowest-level units, characterized by quantitative differences in
373
species representation. Associations are named on the basis of one
or two characteristic species and the suffix -e tum. Further rules of
nomenclature for associations and other syntaxa are discussed, and
an outline of the syntaxa is given.
(20.7) Various extensions of the approach are described. Applications
to vegetation outside the European homeland are summarized.
The approach is applied also to biotic and animal communities,
and to stratal and life-form subcommunities to produce
a separate classification of these into synusiae. Complexes (mosaics
or patterns of contiguous or continuous communities) can be studied
on various levels from the microrelief of bogs to whole landscapes.
Mapping of phytocoena is valuable in applied phytosociology and
land management. For applied work, diagnostic species groups
can be used to indicate environment. Species can also be classed
into ecological groups by their distributional responses to environmental
gradients, and representation of ecological groups can be
used to indicate position of a phytocoenose along an environmental
gradient.
(20.8) Numerical techniques are being developed to store releves,
to measure homogeneity of phytocoenon tables, and to aid
in sample rearrangement in tables and the earlier stages of classification.
(20.9) Ordinations are possible on either a less formal
basis, using releves or phytocoena along known environmental
gradients, or on a more formal basis using quantitative comparisons
amongst samples to arrange them along abstract axes. Ordinations
can be applied tei phytocoena (each summarizing data from several
releves) as well as to individual releves. Results to date indicate
that this is a promising means of clarifying relationships amongst
species and syntaxa.
(20.10) Numerical techniques and ordination are aids to the
essential floristic-sociological procedures of the BRAUN-BLANQUET
approach. Despite the complexity of the procedures, the approach
is felt to be flexible to different kinds of vegetation and research
purposes, productive of understanding of the relations of species
and samples to one another and environment, useful for applied
purposes, and valuable for its coordination of the results from different
areas into a single classification. It is judged the most fully
developed and most widely useful approach to the classification
and interpretation of vegetation.
20.12 Additions to the Second Edition
No essential changes affect the BRAUN-BLANQUET approach as
treated in the first edition. Our additions are mainly references to a
374
(selected) number of publications that came to our notIce after the
manuscript for the first edition was completed. They will be given
by sections.
(20.2-3) History and General Concepts
The history of the BRAUN-BLANQ.UET approach was studied in
further detail by MAAREL (1975) on the occasion of BRAUN-BLANQ.UET'S
90th birthday. VAN DER MAAREL emphasizes the crucial
position of BRAUN'S work in the development of phytosociology,
with the conclusion that it is a synthesis of ideas and approaches
from the entire 19th century in a framework that has incorporated
many developments of the 20th century.
(20.4)lAnalyticai Research Phase
The book by MUELLER-DoMBOIS & ELLENBERG (1974) presents
full information on the releve method and compares various
methods of measuring species quantities, without giving any preference
for one particular method. An important new French contribution
was published by GUINOCHET (1973), who paid special attention
to the choice and delimitation of sample plots.
(20.4.3) MINIMAL AREA AND PLOT SIZE
WERGER (1972) discussed minimal area and emphasized that
no analytical minimal area can be found through a species-area
curve. He suggested (cf. MAAREL, 1966b*) an optimal releve-size
representing a given fraction of the total information in the stand.
MORAVEC (1973a) tried to establish a minimal area through analysis
of similarity between multiple plots of increasing size (as was done
earlier by MAAREL, 1966b* following GOUNOT & CALLEJA, 1962).
(20.5) Synthetical research phase
Again the book by MUELLER-DoMBOIS & ELLENBERG (1974)
may be mentioned as a general reference. For homotoneity MORAVEC
(1973b) proposed an empirical correction for small sets of
releves as well as a simplified calculation for large sets.
(20.6) Syntaxonomical Research Phase
Further evidence is presented by MUELLER-DoMBOIS & ELLEN-
375
BERG (1974) on the geographical variation in habitat preference of
many species and the consequently limited significance ofcharacterspecies.
The authors regret the increasing splitting of associations
and emphasise alliances as general units, recurring in similar forms
in many areas, which largely fulfil the requirements once set up for
the association.
An important approach to plant communities with an insufficient
number ofcharacter-species was developed by KOPECKY &
HEJl'''Y (1974, see also 1973 and KOPECKY 1974). They distinguish
'basal' and 'derivate' communities in addition to the cenologically
saturated communities that can be fully characterised as syntaxa.
Basal communities are composed of species with relatively broad
habitat amplitudes. They may arise either after disturbance of
saturated communities, or during a succession on newly formed
anthropogenic sites. They lack character and differential species
on the association level, but may show them on the alliance, the
order or even only the class level. KOPECKY & HEJNY propose a
special nomenclature for those communities. E.g. a basal community
(BC) characterised by dominance of Urtica dioica and
Aegopodium podagraria in which only class character-species occur is
indicated BC U rtica-Aegopodi um (Galio- U rticetea).
A derivate community is characterised by a dominating species
showing, at least regionally, a narrow amplitude and a rapid
spread from diaspores. It may develop as a special form of a basal
community during primary succession, or, again, by disturbance
of a saturated community, or during the disintegration of a basal
community due to disturbance. Derivate communities (DC) are
indicated as are basal communities; for example, within the range
of the BC Urtica-Aegopodium (Galio-Urticetea) may
occur a DC Chaerophyllum aromaticum (Galio-Urticetea).
This approach may be applied to all kinds of anthropogenic
plant communities (cf. BRAAKHEKKE & BRAAKHEKKE-ILSINK
1976) as well as to communities in naturally disturbed or otherwise
extreme environments.
Nomenclature of syntaxa has reached a milestone through
the publication of the Code of Phytosociological Nomenclature in
Vegetatio (BARKMAN et al. 1976).
(20.7) Extension of the approach
The number of applications of the Braun-Blanquet approach
in non-European areas has further increased. The reader is referred
to numerous contributions in Vegetatio, the new periodical Phytocoenologia,
and Symposium volumes (notably GEHU 1975 on coastal
376
dunes). .
Many references are found in new issues of Excerpta Botamca
Sociologica and also in Fortschritte der Botanik (Progress in Botany)
volumes 36 and 37.
With respect to community complexes a new developme~t ~an
be mentioned: the distinction of 'sigmassociations' or assoClatIOn
complexes, coherent series of associations characterizing ce~tain
landscapes (TUXEN 1973, GEHU 1974). Also syntaxa on hIgher
levels can be described in such a framework, with the general term
sigmasyntaxon. The approach is similar to the descr~ption of~eg~tation
complexes in recent Dutch landscape ecologIcal descnptIOns
(cf. MAAREL & STUMPEL 1975).
As to indicator groups, a recent synthesis of ecological knowledge
on European plant species by ELLENBERG (1974) can be
mentioned.
(20.8) Numerical techniques
The Working Group for Data-Processing in Phytosociology
ended its research. The results are now to be published in Vegetatio,
starting with a general survey of activities and perspectives
(MAAREL et al. 1976, see also MAAREL 1974).
The use of "analyse factorielle des correspondences' (reciprocal
averaging or correspondence analysis, article 11, HILL 1973, 1974)
has shown further successes in the delimitation of plant communities
and the establishment of characterising species groups, particularly
through the work of LACOSTE (1975, 1976). The Montpellier group
proceeded in establishing ecological groups by applying correspondence
analysis to floristic and environmental data (BOTTLIKOVA
et al. 1975) (see also GUINOCHET 1973).
With respect to numerical classification COETZEE & WERGER
(1975) considered the BRAUN-BLANQ.OET approach in its essence
superior to association analysis, whilst STANEK (1973) found it as
effective as a sum-of-squares agglomeration technique. This is a
divisive polythetic classification and an approach in that direction
through the method of indicator species analysis as suggested by
HILL et al. (1975) seems very promising.
Table rearrangement procedures have been treated at length
by MUELLER-DoMBOIS & ELLENBERG (1974) mainly on ELLENBERG'S
( (1956 *) original test data, with applications of numerical methods
divised by SPATZ (1972*, SPATZ & SiEGMUND 1973). STOCKINGER &
HOLZNER (1973) proceeded with their approach through the erection
ofcorrelated species groups. DALE & QUADRACCIA (1973) used
display on monitors in an interactive table sorting. DALE & WEBB
377
(1975) applied a two-parameter analysis, i.e. an analysis which is
symmetric in its use ofspecies and releves, in such a: way that releve
groups are established during the divisive procedure at a point
where discontinuities between releve groups change to continuities
within them.
The table rearrangement program TABORD (JANSSEN 1972*)
has been further extended and implemented in various centres;
JANSSEN (1975) devised a simple clustering procedure for the division
of very large data-sets (up to 6000 releves) into smaller groups
manageable with TABORD.
The attempts towards a numerical syntaxonomy of Spartinetum
communities by KORTEKAAS & MAAREL (1973) proceeded. A comparison
with the classical syntaxonomy became possible through
the publication of the Spartinetea issue of the Prodrome of European
plant communities (BEEFTINK & GEHU 1973), the first issue of this
series, a new milestone in traditional European phytosociology. The
comparison showed substantial agreement whilst some new lower
rank syntaxa were suggested by the numerical output.
(9.9) Ordination
For the rapidly growing literature onphytosociological ordination
we may refer to ORLOCI (1975), various contributions in Vegetatio
and, of course, the companion volume of this book. Like
LACOSTE (1974, 1976) with correspondence analysis, many phytosociologists
have used principal component analysis to obtain
effective boundaries between community types (e.g. BOUXIN 1975,
FEOLI 1973, GILS et al. 1975, and PIGNATTI & PIGNATTI 1975). DALE
& CLIFFORD (1976) further explored the possibilities ofusing higher
taxonomic ranks in the classification of vegetation (cf. MAAREL
1972*).
(9.10) Conclusion
The general effectiveness and efficiency of the BRAUN-BLANQ.UET
approach was emphasised once more, by WERGER (1974).
PIGNATTI (1975), contemplating the future of phytosociology,
concluded that neither a one-sided if sophisticated mathematical
phytosociology,' nor a sterile perfecting of the hierarchic system
of syntaxa should be desired. The approach should instead seek
first summarization and fundamental understanding of relations
of plant communities to one another and environment, and
~econd increasing application of this knowledge to the conservation
and management of our natural heritage.
*References to publications already listed in the first edition are marked with *; new
publications are in a separate list at the end of the References.
378
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of the same author referred to.
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AICHINGER, E., - 1951 - Soziationen, Assoziationen und \Valdentwicklungstypen.
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AICHINGER, E., - 1954 - Statische und dynamische Betrachtung in der pflanzensoziologischen
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BARKMAN, j. j., H. DOING & S. SEGAL, - 1964 - Kritische Bemerkungen und
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BEEFTINK, W. G., - 1962 - Conspectus of the phanerogamic salt plant communities
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