Communities Clasification Whitacker Et. Al

Classification of Natural Communities
Author(s): Robert H. Whittaker

Source: Botanical Review, Vol. 28, No. 1, Classification of Natural Communities (Jan. - Mar.,
1962), pp. 1-239
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THE BOTANICAL REVIEW
VOL. 28 JANUARY-MARCH, 1962 No.1
CLASSIFICATION OF NATURAL COMMUNITIES'

ROBERT H. WHITTAKER
Department of Biology, Brooklyn College
Brooklyn 10, N. Y.
Historic Background ............................................................ 2
Introduction ........................................................... 2
Early History and the Physiognomic Tradition ......................................................... 4
The Southern Tradition ........................................................... 9
The Northern Tradition ........................................................... 23
The Russian Tradition ........................................................... 38
The British Tradition ........................................................... 42
The American Tradition ........................................................... 50
Other Approaches to Classification of Ecosystems ...................................................... 55
Animal Communities ........................................................... 62
Major Units of Classification ............... ............................................ 69
The Ecology of Ecological Traditions ............................... ............................72
The Individualistic Dissent ............. .............................................. 78
The Theory of Synecological Classification ...........................................................83
Evidence ............................................................................................................................................
83
Types of Evidence ................................................ 83
Similarity and Dissimilarity of Stands ........................... ..................... 85
Continuity and Discontinuity .... ............................................ 88
Distributional Relations of Species ................ ................................ 90
Dynamic Relations of Species ............ .................................... 96
The Basis of Classification ................................................ 102
Premises ................................................. 102
The Stand and the Landscape ............ .................................... 103
Abstract Patterns and their Meaning ...................... ...........................109
Community Units as Classes and Types .................................................114
The Process of Classification ............ ..................................... 118
Conclusion ................................................. 123
Application ........................ 125
Principles in the Classification of Natural Communities.................................... 125
Formal Classification in the School of Braun-Blanquet .................................... 128
The Method ................................................ 128
Evaluation ................................................ 134
Informal Classification by Dominance and Physiognomy ..................................140
Evaluation ................................................ 141
Application ................................................ 148
Summary ..............156
Literature Cited ............160
'A contribution from the Biology Department, Brooklyn College, the City
University of New York. Cost of publication was supported in part by a
grant from the National Science Foundation. The author is indebted to T. W.
Bocher, F. E. Egler, R. F. Fosberg, and J. Hospers for comments on the
manuscript.
2 THE BOTANICAL REVIEW
HISTORIC BACKGROUND
INTRODUCTION
No aspect of synecologicalscience has been the subject of more dis-

cussion and argument,or has had a more crucialrole in the evolution
of ecological schools, than the classificationof natural communities.
This monographis a review, and an attemptto provide a currentunder-
standing, of this area of ecological problems. The title may suggest
two things that this paper is not. It is not a manual of how to classify
communitiesby a particularviewpoint or system,and it does not present
a classificationof the natural communitiesof the world. The primary
purposeis inquiry into the theory,the general meaning and underlying
problems, of the classificationof naturalcommunities.
It has often been asserted that the associationis the fundamental
unit of ecological science, and much has been made of the analogy of
the associationas a unit for classifyingcommunitiesand the species as
a unit for classifyingorganisms.Earlierin the developmentof the field,
associationswere seen as something very like Linnaeanspecies on an-
other level of biological phenomena;and the experiencesin hierarchial
classificationof organismswere thought to offer direct indicationsof
how communitiesshould be classified.The past decadeshave seen the
most profound changes in conceptions of species, which, no longer
taken for grantedas units, are studied in terms of underlyingphenom-
ena of genetics, population dynamics,and evolution, and are under-
stood as different kinds of genetic systems and population structures
in differentgroups of organisms.To the extent that an analogy of the
species and associationis currentlyappropriate,it may suggest some-
thing very different from the view of the associationwhich originally
made the analogy so attractiveto ecologists. It suggests, especially,the
need for studying and interpretingassociationsand other community-
types in terms of those phenomenaof populationdistributionsand the
interrelationsof communitieswhich underlie the units that ecologists
recognize.
It suggests, further, the need for a broaderperspectiveon classifica-
tions as they have been applied to differentkinds of communities.In
a particularresearchproblem it is naturalto apply consistentlya single
approachto classification.An investigatormay well profit, however,
from acquaintancewith the wide range of classificationswhich others
have applied when he chooses the approachfor a given researchprob-
CLASSIFICATION OF NATURAL COMMUNITIES 3
lem. Almost any approachto classificationhe may attempthas probably

been used before, somewhere in the world. Because of the extent of
the literatureit is nxoteasy to attain this acquaintanceor to achieve
that internationalperspectivein one's field so much to be desired by
a scientist.A secondarypurposeof this monographis that of a reference
work which may make such a broaderview of the field less difficult.
A fairly thoroughaccountof vegetationclassificationup to that time
was written 40 years ago by Du Rietz (1921). Since that time the
literatureof natural communitieshas become an almost unlimited ex-
panse of uncountedthousandsof papers, on which the unwaryscholar
may drift interminablyto no destination. In the effort to cope with
this literaturewithout writing a review of quite excessive length, the
author has limited himself to very short accountsof most approaches
and has resorted often to multiple citations, in citation-clutteredpas-
sages, of papers bearing on particularpoints. Since most studies of
natural communitiesinclude classificationin some form, this account
and bibliography could be expanded without limit. It is inevitable
under the circumstancesthat many papers have been overlooked or
unavailable,that many others which merit discussionhave simply been
includedin multiple citations,that many authorsand even some schools
have been omitted from consideration, and that accounts of many
authorsand schools are brief to the point of superficiality.Within the
limitations of this monograph the author has sought to provide not
an account sufficientin itself, but a small-scalechart of classificatory
approaches,togetherwith reasonablycomprehensivecitationof primary
sourcesfrom which better understandingof a given approachmay be
obtained.
The difficulty of the ecological literature results not merely from
numbersof papers,but from the remarkablefragmentationof the study
of plant communitiesinto schools. Probablyin no other field of natural
science has there been such proliferation of local schools with dis-
tinctive viewpoints and techniques. The arbitrarinesswhich often at-
tends grouping into schools of authors whose individual viewpoints
differ is well known; but such grouping is in this field necessaryto an
intelligible account.The authorhas chosen to group schools into seven
major ecological "traditions"-five of them regional (the Southern
and Northern Traditionsof Western Europe,the Russian,British, and
American Traditions), two based on approachesto classificationand
overlappinginto all the regional traditions (the tradition of physiog-
nomic classification,based on structureor morphologyof naturalcom-

munities,and the traditionof multi-factoralor landscapeunits). Group-
ing into traditionsis also somewhatarbitrary;one may well consider,
for instance, that British and Americanecology form together a single
traditionof no greater inherent diversitythan the others. But the tra-
ditions permit observationof some major featuresof ecologicalhistory;
and the discussion which follows is organized in terms of these tra-
ditions, beginning with the physiognomic,in which the classification
of natural communitieshad its origin.
The first of the three parts of this study is an extended historical
review of the origin, differentiation,and dispersal of ideas on classi-
fying communities,the schools in which they have been applied, and
the major units of classificationthat have emerged. The second part
is the theoretical discussion, first examining evidence bearing on the
nature of associationsand other units, and then presenting an inter-
pretation of the classificatoryprocess as it is applied to natural com-
munities. The third part considerscertainapplications.Some principles
of classificationof communities are suggested, and two major ap-
proaches- Braun-Blanquet'ssystem of floristic associations,and the
British and American approachthrough physiognomyand dominance
- are examinedin more detail as illustrationsof classificatoryproblems
in practice.
EARLY HISTORY AND THE PHYSIOGNOMICTRADITION
Although vegetation units seem primarilyconceptionsof twentieth-

centuryscience, their origins may be traced back through most of the
precedingcentury(vide Hult, 1881; Moss, 1910; Clements,1916, 1928;
Riibel, 1917; Gams, 1918; Du Rietz, 1921; Diniker, 1936; Schmid,
1936b). In his works from 1805 to 1807 Humboldtexpressedboth the
idea of growth-formsof plants, on which physiognomicunits are based
(Humboldt, 1805, 1806, 1807) and the idea of associatedplants (gesel-
liger Pflanzen, Humboldt, 1807) forming communitieswhich may be
characterizedby their dominantspecies and termed associations(". ..
diese Gruppierungder Erica vulgaris, Erica tetralix . . ." Humboldt,
1807:7; in French ". . . cette associationde l'ericavulgaris . . ." Hum-
boldt 1805:17). Grisbach (1838:160; Du Rietz, 1921:42; Clements,
1928: 117-8) gave the idea of community-unitsfurther expression,
though still in incipient form. The term formationwas applied by him
to a group of plants possessing a definite physiognomiccharacter,as a

meadow, forest, etc. Grisebachalso indicated, however, that a forma-
tion could be characterizedby a single social or dominant species, a
complex of dominantspecies belonging to one family, or an aggregate
of species of various taxonomic characterbut of some common pecu-
liarity, as the alpine meadow consistsalmostentirelyof perennialherbs.
Thus, in the earliestconceptionsof vegetationunits, both physiognomy
and species composition were used for their characterization.Only
later, as the formation came to apply only to physiognomicunits, was
the associationdistinguishedfrom it as a unit based on the alternative
criterionof species composition.
During the latter part of the nineteenth centurymany authorscon-
tributed to the developmentof the physiognomicunit of vegetation,
which came to be termedthe formnation.Heer (1835) recognizedsome
communities by physiognomy and site as Lokalit?ten, loosely corre-
sponding to later formations.Post (1844, 1862) termedmajorphysiog-
nomicgroupingsvegetationsgrupper.Sendtner(1854) suggestedthe
term Vegetationsform for such physiognomictypes as meadow, forest,
and moor; and the Unterformen or subdivisions of these correspond
to formations of later authors. Lorenz (1858) recognizedvegetation-
forms and subforms in much the sense of Sendtner, together with
Typen (1858) and Facies (Lorenz, 1863) which correspondmore nearly
to the modern association.Kerner (1863) consideredformationsto be
plant groups with regularstructureand distinctivecomposition,named
by dominantspecies; these were grouped into higher units correspond-
ing to present formations.Grisebachin a later work (1872) used the
formation for major physiognomictypes, in the sense which later be-
came widely accepted.Hult (1881, 1887, 1898) used "formation"for
very narrow physiognomic units defined by stratal structure.Drude
(1888, 1890a, 1890b, 1896) earlier conceived the formation as a
smallerunit than Grisebach's,characterizedby floristiccompositionand
habitat; Drude (1913; Morton, 1915) later defined the formation as
a unit correspondingto a certainclimate and soil and characterizedby
the dominanceof certainlife-forms. Warming (1888, 1889) described
the vegetationof Greenlandthroughphysiognomic-ecologicformations;
Ostenfeld (1908) describedthat of the Faeroesthroughphysiognomic-
ecologic formations and associations defined by species dominance.
Units such as forest, meadow, and steppe, conceivedin terms of phys-
iognomy and habitat,were also designatedformationsby Schr6ter(Steb-
ler and Schroter, 1892; Schroter, 1894; Schroter and Kirchner, 1902),
and types were recognized within these by dominant species.
During the earlier part of the twentieth century, however, the forma-
tion continued to be used in a variety of ways, with the possible criteria
of physiognomy, floristics, dominance, habitat, and dynamic relations in
rivalry with one another in the usage of different authors. The chaotic
usage of f ormation was reviewed and the need for standardization
indicated by Warburg in 1900. Clements (1902, 1905) at first re-
garded habitat as the basis of formations. Adamovic (1898, 1901, 1906,
1909) and Podpera (1902) treated Balkan vegetation through forma-
tions defined primarily by physiognomy and habitat, and types dis-
tinguished by dominance. Olsson-Seffer (1905) suggested the use of
formation for physiognomic units without reference to species compo-
sition and the subdivision of formations into associations. Moss (1907,
1910) emphasized habitat, but also brought succession into the defini-
tion, regarding the series of plant associations which eventually be-
come a stable or closed association as a plant formation. Gradmann
(1909) advocated definition of formations by floristic composition.
The term formation was not used in the first survey of plant com-
munities of the world by Warming (1895, 1896); but in a later ver-
sion (1909) the formation was defined as a community of species,
all belonging to definite growth-forms, that have become associated
together by definite external (edaphic or climatic) characters of the
habitat to which they are adapted. From this welter of conflicting
definitions, the physiognomic definition advocated and applied by
Grisebach (1872), Schrbter (1894), Olsson-Seffer (1905), and Warm-
ing (1909) gradually prevailed.
The formation has become the central concept of one of the major
approaches to the classification of communities, the physiognomic tradi-
tion. Classification of vegetation into formations must be based, how-
ever, on a second concept, that of types or forms of plants, variously
termed vegetation-forms, Hauptformen, Grundformen, life-forms, and
growth-forms, which determine the physiognomy of communities. Paral-
leling the early development of the formation concept, the concept of
plant form was originated by Humboldt (1805, 1806, 1807, 1849)
and further developed by Kerner (1863), Grisebach (1872), Hult
(1881), Warming (1884, 1895), Reiter (1885), Drude (1890a,
1890b, 1896), Krause (1891), Schimper (1898, 1903), and Pound
and Clements (1900); history of the concept is reviewed by Drude
(1913), Gams (1918), Clements (1920, 1928), Bews (1925), Riibel

(1930), and Du Rietz (1931). Twentieth-century systems of plant
forms include those of Raunkiaer (1905, 1934), Warming (1908,
1909, 1923), Diels (1908), Drude (1913, 1932), Gams (1918),
Du Rietz (1921, 1931), Braun-Blanquet (1928a, 1951a), Turrill
(1929), Riibel (1930), Shreve (1942, 1951), Ellenberg (1956),
Schmid (1956a), and Troll (1958b); and the systems for special
groups of plants of Arber (1920), Gimingham and Birse (1957),
Barkman (1958), Hosokawa et al. (1954), Horikawa and Miyawaki
(1954), and Kuwabara (1960).
Following the work of Raunkiaer (1905), the conceptions of plant
forms have evolved in two directions. The system of Raunkiaer (1905,
1910, 1934, 1937), and Braun-Blanquet's (1928a, 1932a, 1951a)
adaptation of it, is in ideal a single-factor classification, with its primary
classes defined only by the position or character of those plant parts
(buds and seeds) in which meristematic tissues survive unfavorable
seasons. The percentages of these plant-form classes, termed life-forms,
in a flora or community form a "biological spectrum"; and these spectra
have been very widely studied for their expression of climates and local
environments of communities (Raunkiaer, 1934; Cain, 1950; Dan-
sereau, 1951; Braun-Blanquet, 1951a). Only in the approaches of Raun-
kiaer, Gams, and Lippmaa (see below) have life-forms in this sense
been used as a primary basis for community-classification. In the other
direction of development, plant forms are based on many aspects of
plant morphology which may be significantly related to environment;
criteria which may be used include woody vs. herbaceous growth,
stature, evergreen vs. semi-deciduous and deciduous leaves, leaf form,
size, texture and arrangement, succulence, spinosity, epiphytic position,
the climbing habit, and sometimes membership in taxonomic group-
ings. For such multi-factoral classes, there are an unlimited number of
possible classifications; and the particular manner of classification is
determined not so much by logic as by convention and usefulness in
describing the physiognomy of communities. Many authors have called
these morphological plant forms life-forms, but the term growth-form
seems preferable in English to distinguish them from the Raunkiaer
life-forms.
The plant formation is, ideally, a community-type defined by dom-
inance of a given growth-form in the uppermost stratum (or the upper-
most closed stratum) of the community, or by combinations of domi-
nant growth-forms.The growth-formsthemselves are significantlyre-

lated to environmentin that they dominate communitiesonly under
certain conditions; interpretationswhich go beyond this in drawing
conclusions about the adaptive or "epharmonic"(Vesque, 1882) re-
lations of growth-forms to environmenthave been criticized by Du
Rietz (1931). Community-typesdefined by dominant growth-forms
are consequentlyalso significantlyrelated to environment,but in no
simple manner.Formationsare, in application,defined by varied, con-
ventionally accepted combinations of growth-form dominance and
characteristicsof environment.
Vegetationof a certainphysiognomyusuallyoccursin a certainrange,
or ranges, of environmentalconditionsto which the dominantgrowth-
forms are adapted.Physiognomicallysimilar communitiesoccuron dif-
ferent continents where similar environmentalconditions occur. This
physiognomic convergenceof vegetation in widely separatedregions
is one of the major phenomenaof plant geographyand a major justi-
fication of the physiognomicapproachto vegetation. The convergence
may be imperfect becauseof the differentkinds of plants in different
floristic regions; and absenceof a major-growth-formfrom an area to
which it might be well adaptedwill sometimesresultin the occurrenceof
very differenttypes of communitiesin closely similar climates (Beadle,
1951; Whittaker, 1956). The similarityof plant formationson differ-
ent continentsmakes possible, however,their groupinginto world-wide
physiognomicunits which have been variouslytermed formations,for-
mation-types,formation-groups,and formation-classes(Drude, 1888,
1890a, 1890b; Schimper, 1898; Schimper and Faber, 1935; Schroter
and Kirchner, 1902; Diels, 1908; Warming, 1909; Warming and
Graebner,1930-33; Brockmann-Jerosch and Riubel,1912; Riibel, 1913b,
1930; Herzog, 1933; Lieth, 1956; Schmithiisen,1959; Fosberg, 1961),
Vereinsklassen(Warming, 1896), vegetation-types(Drude, 1913), iso-
coenoses and isocies (Gams, 1918, 1954; Du Rietz, 1932), Hauptfor-
mationen (Cajander, 1922), panformations(Du Rietz, 1932), homo-
logous formations (Braun-Blanquet,1932a), and panclimaxes (Clem-
ents, 1936). The most widely used English term for this unit is for-
mation-type (Nichols, 1917; Burtt Davy, 1938; Tansley, 1939; Dan-
sereau, 1951). When the biome or biotic community-typeis substituted
for the formation, the correspondingworld-wide unit becomes the
biome-type(Allee et al., 1949; Tischler, 1951).
The earlier physiognomic systems of Grisebach,Drude, Warming,
CLASSIFICATIONOF NATURAL COMMUNITIES 9
and Schimper have influenced innumerableother authors; the trans-

lated version of Warming (1909) has had an especially strong in-
fluenceon English-languageauthors.Revisionsof the classictreatments
of Warmingand Schimperhave been presentedin Warmingand Graeb-
ner (1930-33) and Schimperand Faber (1935). A widely influential
recent system is Riubel's(1930, 1936), developed from the classifica-
tion of Brockmann-Jerosch and Riibel (1912a; Riibel, 1913b; cf. Bri-
quet, 1920; Vierhapper,1921). Riubelrecognizesthree terrestrialvege-
tation-types-Lignosa, Herbosa,and Deserta, correspondingto the for-
mation-groupsof Schimperand Faber-and, within these, formation-
groups and 27 cosmopolitanformation-classes.A physiognomicsystem
adapted for mapping has recently been presented by Kiuchler(1947,
1948, 1949, 1950). The systemsof Schimperand Faberand of Kuchler
and the relation of the Raunkiaerlife-forms to the former have been
reviewed, and a new system offered, by Dansereau (1951, 1957). In
addition to their use in plant geography,physiognomicunits have been
used by ecologists and phytosociologistsin all the regional traditions,
as will be discussed;the formationis perhapsthe most widely used of
all vegetation units.
THE SOUTHERN TRADITION
After the first suggestion of the association by Humboldt, a few

associationswere mentioned and the widely used suffix -etum first ap-
plied to them by Schouw (1822, 1823). Lecoq (1844, 1855) applied
the term associationto vegetation plots, without grouping these into
communityunits in the modern sense. Lorenz (1858) establishedthe
Typus as a unit at the level of the association,characterizedby com-
position in terms of species and species-groupsor societies, and named
by the -etum suffix. Drude (1890b) recognizedBestdndeas local units
determinedby dominantspecies within a formation.Steblerand Schro-
ter (1892) and Schroter(1894) recognizedphysiognomicformations,
which changed in composition accordingto local conditions to form
Typen (after Lorenz, 1858) and Nebentypen, designated by species
dominant in or especially characteristicof the site. These latter are
essentiallyassociations;and the modern distinctionbetween the forma-
tion as a higher-level unit defined by physiognomyand habitat, and
the type or associationas a lower-level unit defined by species compo-
sition was thus recognizedby Lorenz,and Steblerand Schrdter.Flahault
(1898) applied the term association and advocatedits acceptanceas

a fundamental unit characterizedby dominating species (Flahault,
1901a, 1901b). Schroterand Kirchner (1902) also suggested use of
association for lower-levelunits. The agreementof Flahaultand Schro-
ter led to the definitionproposed by them in 1910 to the Third Inter-
national Botanical Congress, "An associationis a plant communityof
definite floristic composition, presenting a uniform physiognomy,and
growing in uniform habitat conditions. The associationis the funda-
mental unit of synecology." (Flahault and Schroter,1910; Pavillard,
1935a). This definition has become the basis for general acceptance
of the association as a vegetation unit defined by species composition.
"Floristic composition" can be quite variously interpreted,however;
and some very differentconceptionsof plant associationshave appeared
in different schools within the limits set by this definition.
In southernEurope,furtherdevelopmentswere guided by the notable
group of students of vegetation centered in the cities of Zurich and
Montpellier, working in the floristicallyrich and varied vegetation of
the Alps and MediterraneanFrance. From the backgroundof Heer,
Sendtner, Lorenz, and Kerner, the Swiss-Frenchschool was founded
by Schroterof Zurich and Flahaultof Montpellier. A series of mono-
graphs on the Alps by Brockmann-Jerosch(1907), Riubel (1912a),
and Braun-Blanquet(1913), and essays by Pavillard (1912) and
Braun-Blanquetand Furrer (1913), laid the basis for much of the
further evolution of the SouthernTradition.
Flahault (1901a, 1901b) accepteddominanceas the criterionof as-
sociations, but Brockmann-Jerosch(1907) considered the constant
species, those appearingin at least half the samples for a stand-type
(association), the basis of its characterization.Among the constants
formation-ubiquitists(constants of wide distributionin various com-
munities) were distinguishedfrom the more restrictedand more critical
constants, the character-plants. Very similar classificatoryhierarchies
following Schroter (Schroterand Kirchner, 1902) -Vegetationstypus,
Formationsgruppe,Formation, Subformation, and Bestandstypusor
Assoziation down to local variants or Nebentypen, and geographic
variants or Fazies-were used by Brockmann-Jeroschand by Riubel
(1912a, 1913a). Riubel'streatmentwas influenced also by a distin-
guished essay by Gradmann (1909), which indicated the need for a
floristic approachto classification,the necessarilyabstractcharacterof
the units derived, and the importanceof the restrictionof some plants
CLASSIFICATIONOF NATURAL COMMUNITIES 1l
(Leitpflanzen)to a given unit for its characterization.Early works of

the school of Ziurich-Montpellier(Brockmann-Jerosch,1907; Riibel,
1912a; Braun-Blanquet,1913) establishedalso the proceduresof com-
piling individual stand samples into tables for community-types,of
ranking species in the communityby constancyor other characteristics
to clarify their diagnosticvalue for the unit, and of naming community-
types by dominantor characteristicspecies, using the genus name with
the suffix -etum followed by the species name in the genitive. (e.g.,
"Caricetumcurvulae"for a communitycharacterizedby Carexcurvula).
These had alreadyappearedin the pioneer work of Cajander(1903a),
were adopted by other Scandinavianauthors,and came to be standard
procedures of Continental "phytosociologists,"distinguishing their
work from much of that of British and American"ecologists."
The traditionof Zurich and Montpellierhad become a well-defined
school by 1912-1913, but shortly thereafter began to branch into a
grouping of schools in these cities. One majordirectionwas represented
by Brockmann-Jerosch and Riubel(Brockmann-Jerosch and Riibel, 1912;
Rilbel, 1913b, 1922, 1930, 1934, 1936), treating the vegetation of
the world in terms of formationsand the convergentformation-types
which appearon differentcontinentsin responseto similar climates.A
second directionwas taken by the school of Braun-Blanquet,in which
floristiccompositionwas the basis for systematicclassificationof com-
munities. Braun-Blanquet(1913) carriedto its logical conclusionthe
shift of emphasisfrom dominanceto distributionalrelationsof species,
as a basis for communitycharacterization, begun by Brockmann-Jerosch
(1907) and Riubel(1912a). The concept of character-species, largely
confined to a given community-typeand therefore characterizingit,
a concept traceable from Heer (1835) through Brockmann-Jerosch
(1907), Gradmann(1909), Riubel(1912a) and others (Braun-Blan-
quet, 1921; Wendelberger, 1952), was further developed by Braun-
Blanquetand made the key idea of his school.
A third directionwas representedby Liudi(1920, 1921, 1923, 1929,
1932, 1945), with somewhat greater emphasis on succession than
other authors of Ziurich-Montpellier,the association being regarded
as a more or less stable stage in a successionalseries. Habitat relations
of communitieswere also emphasizedby Liidi (1928, 1948), and a
less one-sided approachto the definition of communitiesthan Braun-
Blanquet's was sought, considering constancy, habitat, physiognomy,
and other charactersas well as the character-species. The term character-
specieswas restrictedby Liudi(1928), as by Brockmann-Jerosch (1907)

and Wangerin (1925), to those constantsor majorspecies of the com-
munity-typewhich show relative restrictionto it. The associationessay
by Liidi (1928) agrees in general with Wangerin's (1925) in its
definitionof the associationand its recognitionof the Elementarassozia-
tion (Drude, 1913, 1919, 1932:53) as a unit within the association.
These three schools within the Southern Tradition representthree
major emphases in the approachto communities-the physiognomic,
floristic, and ecological (in the sense of habitat-relations).A fourth
possibility is representedby the school of Schmid (1922, 1940, 1941,
1942, 1949, 1950, 1952) and Diniker (1936, 1939a, 1939b) with
its emphasis of chorological units (Vegetationsguirtel)and of local
Biocoenosen, instead of associations.Still anotherapproachamong the
schools of Zurich and Montpellieris Kuhnholtz-Lordat's(1952) with
its detailed analysis of environmentaldeterminationand successional
process. Work of Jovet (1949) is also concernedwith environmental
and successionalrelations of community-typesrecognized by habitats
and dominance.
Of these schools,Braun-Blanquet's(see also Application) has gained
much the widest following. Braun-Blanquet's(1913) monograph of
high-alpine vegetation followed Brockmann-Jerosch(1907) in some
ways, but based classificationnot on constantsbut on Charakterpflanzen
or character-species, species (which need not be constants) which have
narrow ecological amplitudes and are consequently almost restricted
to a single association;this condition of relative restrictionto an asso-
ciation was termed Treue or fidelity. The major features of Braun-
Blanquet'ssystem appearedin an essay the same year (Braun-Blanquet
and Furrer, 1913). The first object in the study of communitieswas
seen as the search for a unit comparableto the species as a basis for
research.This unit should be based on character-species;associations
should in fact be defined by their possession of character-species, and
communitieslacking these are either transitorygroupings or mixtures.
Each associationis composedof individuals,just as the species is com-
posed of individuals (cf. Pavillard, 1912); the association,like the
species, can thus be describedfrom a sampleof its individuals.Associa-
tions should be grouped into higher units not by physiognomybut by
floristic comparison.The whole later history of the school of Braun-
Blanquetmay be seen as a logical developmentand elaborationof the

ideas in this first essay.
The two key ideas, of the approachthrough floristiccomparisonand
the use of Leitpfianzen or Charakterpflanzeni,had been stated also in
the essay by Gradmann (1909). The essay by Braun-Blanquetand
Furrer (1913) differsin what may seem a narrowerapproach,with the
emphasis of these same ideas almost to the exclusion of others, and
in the evident domination of the taxonomic analogy. The analogy of
the associationto the species, perhapsfirst stated by Schroter(Schroter
and Kirchner,1902; Pavillard,1927, 1954), and the consequentquasi-
taxonomicapproachto communities,came to be characteristicof both
the northernand southernschools of phytosociologyand to be a second
feature distinguishingthe approachof many "phytosociologists"from
that of most "ecologists"of English-speakingcountries.
Braun-Blanquetrepeated the emphasis on the floristic approach
through character-speciesand offered the definition of the association
as a floristicallyuniform plant-community,more or less in equilibrium
with externalfactors,and manifestingecologicalindependencethrough
the presence of character-speciesspecific to the association (Braun-
Blanquet, 1915:251, 1918, 1921:323). He presented the system in al-
most fully developed form in 1921, with the various conventionsfor
descriptionof communities (frequency, abundance,dominance,socia-
bility, dynamic value; fidelity and constancy,etc.), the grouping of
associations into Assoziationsgruppen (1915) or Verbinde with their
own character-species, and the "sociologicalprogression"for arranging
communitiesaccording to levels of organization,in analogy with the
arrangementof taxonomic units according to evolutionarylevel (cf.
Schmid, 1922; Daniker, 1928). To this systemwas added the concept
of differential-species(Koch, 1925; Braun-Blanquetand Jenny, 1926)
supplementingthat of character-species and serving especially for the
characterizationof units below the association.
The school of Braun-Blanquetwas joined by Pavillard (1919, 1920,
1927, 1928b); and the Braun-Blanquetsystemwas incorporatedin out-
line form into the well-known Vocabulaire of plant sociology (Braun-
Blanquet and Pavillard, 1922). Pavillard (1912) and Braun-Blanquet
(Braun-Blanquet and Furrer, 1913; cf. Braun-Blanquet, 1951a:49), re-
garded the formation as the characteristic physiognomy of a stable
plant community, related to external factors, and not as a classificatory
unit. The term formation later largely dropped out of use in this school;
and to replace it a hierarchy of floristic higher units (Vrerband or Al-

liance, Ordnung, Klass, and Vegetationskreis) developed (Braun-Blan-
quet and Pavillard, 1925; Koch, 1925; Braun-Blanquet and Jenny,
1926; Pawlowski et al., 1928). The alliance, order, and class were in-
tended to be floristicunits recognizedand defined, like the association,
by character-species.It was also expected, however, that these units
would have more than floristic significanceand so would in general
correspond to natural higher units that might be recognized by other
means (Braun-Blanquet and Jenny, 1926; Oberdorfer, 1937).
The central concept of character-specieshas been widely criticized
by Scandinavian authors and others (Du Rietz et al., 1918; Gams,
1918, 1939, 1941; Tengwall, 1920; Du Rietz, 1921; 1923b; Du
Rietz and Gams, 1924; Nordhagen, 1923, 1928; Wangerin, 1922,
1925; Ludi, 1928; Katz, 1929; 1930b, 1933; Almquist, 1929; Paczoski,
1930b, Meusel, 1939b; Poore, 1956). Faegri (1937) observed that,
with rare exceptions, Scandinavian phytosociologists have searched in
vain for character-species upon which to found their associations (see
also Du Rietz et al., 1918; Tengwall, 1920; Nordhagen, 1923, 1928,
1937; Kalliola, 1939). Domin (1928b, 1929) found almost no species
of complete fidelity in the Czechoslovakian vegetation and that some as-
sociations possessed no character-species whatever (cf. Zlatnik, 1928b).
Similar conclusions were reached in northern forest and bog vegeta-
tion of Russia by Katz (1930b) and in Palestinian desert vegetation
by Zohary and Orshan (1949, 1956). Liudi (1928, cf. Frey, 1922)
consideredthat in the Alps themselves, fidelity was weakly developed
in some communitiesand lacking in others, and regarded fidelity as
of limited value as a criterionof community-types.Riubel(1934) has
suggested that fidelity may be well-definedin continentalclimates,but
weakly developed or of no value in maritimeclimates. The doctrine
of character-species was defended against the strong criticisms of Du
Rietz and Gams (1924) by Braun-Blanquet (1925) with an affirma-
tion of the value of character-species as a basic concept of phytosociol-
ogy, as the only objective criterionfor communityclassification,and as
an approach to indication of development (Braun-Blanquet and Jenny,
1926), disturbance,and areas of communities.
Authors advocating balanced use of various properties of communi-
ties for their characterization have criticized the one-sidedness of Braun-
Blanquet'sapproach(Liudi,1928; Meusel, 1939a, 1939b, 1940, 1943a;
Schmid, 1941, 1942, 1950; Gams, 1939, 1941; Gradmann, 1941;
Ehrendorfer, 1954). The validity or appropriateness of the analogy be-

tween the association and species, which guides the orientation of the
school, has been questioned by Nordhagen (1920, 1923), Gleason
(1926), Negri (1926, 1927), Lenoble (1926, 1928a), Lippmaa
(1931, 1933a), Hauman (1933), Meusel (1940), Gradmann (1941),
Schmid (1941, 1942), Gaussen (1954) and Ehrendorfer (1954). The
hierarchy of units has been criticized as inadequate to express the
multi-dimensional relations among communities (Gams, 1918, 1939,
1941, 1954; Meusel, 1940; Schmid, 1944; Webb, 1954); Braun-
Blanquet (1939, cf. 1955) has expressed the opinion that ecological
relations of most communities are not multi-dimensional and may quite
well be presented linearly. The view of the stand as an "association-
individual,' advanced by Pavillard (1912, 1927, 1935a) and by Braun-
Blanquet and Furrer (1913) lhas been widely criticized, either as a term
or as a concept by Alechin (1925b), Kylin (1926), Vierhapper
(1926), Nordhagen (1928:70), Liudi (1928), Sukatschew (1929),
Paczoski (1930b), Lippmaa (1931, 1933a), Schmid (1940, 1941,
1942), Ellenberg (1950a), and especially in a series of papers by Du
Rietz (192l:72, 125, 1923b, 1925a, 1928, 1932:306). The manner
of naming units has been regarded as objectionable by some ecologists
(Bartlett, 1933). These criticized featuareswere incorporated with the
rest of the system into Braun-Blanquet's book, Pflanzensoziologie (1928a,
1932a, 1951a), which has been one of the two most influential books
on classification of communities, influencing many phytosociologists as
profoundly as Warming (1909) has influenced English-speaking ecol-
ogists.
As Braun-Blanquet's ideas were applied by other authors in the 1920's,
the influence of his school spread through Europe (Allorge, 1921-2;
Beger, 1922-3; Schustler, 1923; Szafer et al., 1923; Koch, 1925; Scher-
rer, 1925; Issler, 1926; Tiixen, 1928). The Station Internationale de
Geobotanique Mediterraneenne et Alpine (SIGMA) was founded at
Montpellier in 1930, and has since served as headquarters and training
center for the school. For some years the belt of nations from the
Netherlands west through Germany into central Europe was a zone of
competition between Braun-Bianquet's ideas and those of other schools,
especially the rival school of Uppsala. In Poland, Paczoski's (1928,
1930b) study of forests was based on groups and types defined pri-
marily by dominant trees and combinations of tree species, and on in-
tensive study of successional relations. Braun-Blanquet's system, how-
ever, found a first northernoutpost in Poland in the "Cracowschool."

Authors of this school found the Braun-Blanquetsystem, developed in
the Alps, well suited for study in the Tatra Mountains (Szafer et al.,
1923, 1927; Motyka, 1925, 1926; Pawlowski, 1926, 1935; Szafer and
Pawlowski, 1927; Szafer and Sokolowski,1927; Pawlowskiand Stecki,
1927; Pawlowski et al., 1928) and other areas (Juraszek,1928; Koz-
lowska, 1928; Kulczyn'ski,1928; Kleist, 1929; Walas, 1933; Wilzek,
1935; Pawlowski, 1947; Kornas, 1950). Some of these authorswere,
however, influencedby the northernschools as well as by Braun-Blan-
quet (Juraszek, 1928; Kleist, 1929). In Hungary, early work of Ra-
paics (1922, 1927) and Soo (1929, 1930a, 1930b, 1930c) was based
on dominanceand influencedby the school of Uppsala; later work of
Soo (1934, 1936a, 1936b, 1939, 1949, 1952, 1954a, 1954b), Zolyomi
(1934, 1936, 1950) and others has followed the school of Braun-
Blanquet.The Braun-Blanquetsystem has been applied in Bulgariaby
Horvatic (1930, 1931, 1934, 1939, 1954) and Horvat (1931, 1934,
1936, 1954; Horvat et al., 1938) and in Jugoslavia by Horvat (1930a,
1930b) and Wraber (1954). Because of its spread into Poland, Hun-
gary, Czechoslovakia,and Bulgaria,the school of Braun-Blanquetwas
at one time termedthe "French-WestSlavic" (Gams, 1939) or "middle
European-Mediterranean" (Braun-Blanquet,1939) school, in contrast
to the rival "Nordic-Alpine"school (Gams, 1939).
In Czechoslovakia,the earliestwork of Domin (1905) was based on
physiognomicformations;in later work Domin (1926, 1928a, 1928b,
1929) rejectedthe character-species concept as impracticaland agreed
with the school of Uppsala on the importanceof constancyfor the
characterizationof associations.Hilitzer (1925, 1927) followed the
school of Uppsala; Podpera (1928) recognized physiognomic units.
Studies of Bohemian bogs by Rudolph and others (Rudolph, 1928;
Rudolph et al., 1928; Rudolph and Firbas, 1927; Firbasand Sigmond,
1928) followed the school of Uppsala in the use of micro-associations
or sociations.Earlywork of Zlatnik (1926, 1928a) and Klika (1927,
1928, 1929a, 1929b) was influenced by the Northern Tradition, but
the Braun-Blanquetapproachwas advocatedby Schustler (1923) and
increasingly followed in later Czech work (Zlatnik, 1928b; Klika,
1930, 1931a, 1931b; Mikygka, 1932; Krajina, 1933; Kmonicek, 1936).
The extensive work of Klika (1931a, 1931b, 1932b, 1933, 1936a,
1936b, 1938; Klika and Vlach, 1937) used the Braun-Blanquetclassi-
fication; and work of Klika (1932a, 1937, 1939), Mikyska (1930,
1933, 1935, 1937, 1939, 1953), Svoboda (1935a, 1935b), and others
on the classificationof forests was largely within the frameworkof
the Braun-Blanquetunits. Some of these types were characterizedby
strataldominantsand constantspecies as well as by differential-species,
however;the influenceof the Northern Traditionon Czech forest typ-
ology is evident (vide Zlatnik, 1928a; Konsel, 1929; Mikygka,1930,
Hilitzer, 1934; Muller, 1938). Recent papers have presented an ap-
proach to forest classificationthrough differential-speciescombinations
and analysisof ecological and distributionalrelations of undergrowth
species (Zlatnik, 1954, 1956, 1961). Foresttypes, classifiedinto Wald-
typengruppenand other units, are relatedin a patternor grid of eleva-
tion and soil conditions (Zlatnik, 1961). The approachrepresentsa
synthesisof conceptsfrom Braun-Blanquetand Schmidin the Southern
Tradition with the biogeocenose and other viewpoints of the Russian
Tradition (cf. Klika, 1953, 1954).
In the Netherlands a stratalapproachrelated to the Northern Tra-
dition was applied by de Vries and associates(see below), but other
Dutch authors have actively applied the system of Braun-Blanquetto
the vegetation of the Low Countries (Braun-Blanquetand Leeuw,
1936; Vlieger, 1937; Vlieger and Kruseman,1937; Leeuw et al., 1938;
Diemont et al., 1940; Weevers, 1940; Meltzer and Westhoff, 1944;
Adriani, 1945; Westhoff et al., 1946; Barkman,1949, 1954; Sissingh,
1950; Hartog, 1951; Hoffmannand Westhoff, 1951; Boerboom,1957)
and Surinam (Lindeman, 1953). The system has been applied also in
Belgium (Louis and Lebrun, 1942; Lebrunet al., 1949; Duvigneaud,
1949; Noirfalise, 1952; Bodeux, 1954, 1955; Mullenders, 1954; Heine-
mann, 1956) and the Belgian Congo (Lebrun, 1947; Germain, 1952;
Leonard, 1952, 1954; Mullenders, 1953; Lebrun and Gilbert, 1954).
Duvigneaud (1946, 1949) emphasizedphysiognomyand dominance
more stronglythan most authorsof the school, and he (1946) has also
interpretedvariabiiityas an essentialcharacteristic
of the associationand
suggesteda systemof lower-levelunits. The associationis interpretedas
a more or less stable plant grouping, of definite floristic composition
and physiognomybut variablewithin limits, characterizedby a nuclear
group of species bound to one another by strong sociological affinity
which determines the typical physiognomy of the association,while
distributionalrelations of other groups of species produce variations
in the associationthat can sometimesmodify its physiognomy.
Lebrun (1947) considered the methods of Braun-Blanquet,devel-
oped in temperatesouthern Europe, entirely suitable for the study of

tropical vegetation because of their flexibility, just as Tuxen (1951)
has asserted their suitablenessfor the northern vegetation of Scandi-
navia. Lebrunand Gilbert's (1954) classificationis, like most othersof
African vegetation, physiognomicin basis; physiognomictypes recog-
nized by others are fitted into the Braun-Blanquethierarchyas alliances
and orders. Mangenot et al. (1948) also consideredthe concepts of
the school of Braun-Blanquetapplicableto the vegetation of tropical
forests and recognizedfour edaphic groups of forests, which they in-
terpretedas orders. Emberger(1954; see also Embergeret al., 1950a,
1950b) has arguedon the basis of frequencydistributionsthat tropical
forests consist of true associations,ratherthan the variableand change-
able mixtures suggested by Aubreville (1938).
Although some German authors (Hueck, 1925; Kaiser, 1926) fol-
lowed the Uppsala school, and Wangerin (1925), Drude (1926), and
Gradmann(1941) have criticizedthe Braun-Blanquetsystem,this has
had very extensive application in Germany following the leadership
of Tiuxen (1928, 1937, 1954a, 1955). In Austria, Scharfetter(1911,
1918) first approachedvegetation through formations;in later work
Scharfetter(1921, 1932, 1936) emphasizedgeomorphic-edaphicand
chorological relations. Gams (1918, 1927, 1933, 1936) developed a
stratal or synusial approachto vegetation related to the Northern Tra-
dition and later allied himself with the school of Uppsala (Du Rietz
and Gamis,1924; Gams, 1933, 1936, 1939). Vierhapper(1918, 1921,
1925, 1935) emphasizedphysiognomyand constancyin general agree-
ment with the school of Uppsala; and Zumpfe (1929) applied the
approachof the Uppsala school to bog vegetation. Recent work of the
"Vienna school" (Wagner, 1941, 1950b, Wendelberger, 1951, 1952,
1953a, 1953b, 1954a, 1954b; Wendelberger-ZelinkaandWendelberger,
1956; Kielhauser, 1954a, 1954b, 1956) in general follows Braun-
Blanquet. Wendelberger (1951), however, has consideredthe Braun-
Blanquet system in a broadercontext of possibilities of classification,
and is one of the few authors of the school to make explicit use of
the formation concept (Wendelberger, 1954a, 1955, 1959). Wag-
ner's (1950a, 1950c, 1954a) work is distinctive in its emphasis on
ecological series and suggestion of the need for relating the units of
Braun-Blanquetin a multi-dimensionalcoordinate system (Wagner
1954b, 1958).
Some of the most impressiveachievementsof the school, especially
in Germany,are in applied vegetation study and the use of vegetation

maps in relationto land management.Tiixen's Bundesanstalt(formerly
the Zentralstelle)fur Vegetationskartierunzgat Stolzenau/Weserhas be-
come a second center for the school along with the SIGMA at Mont-
pellier, and has been the source of many studies in applied phytoso-
ciology. The system of Braun-Blanquethas been adapted for detailed
studies of forests (Hartmann, 1932, 1933, 1934, 1936, 1954; Braun-
Blanquet, 1936a, Diemont, 1938; Trepp, 1947; Buchwald, 1951a,
1951b; Lohmeyer,1951; Moor, 1952; Oberdorfer,1953; Rubner,1953;
Braun-Blanquetet al., 1954; Tiuxen, 1954a; Seibert, 1954; Becking,
1956; Richard, 1961), grassland communities (Knoll, 1932; Braun-
Blanquet, 1936b; Soroceanu, 1936; Marschall, 1947; Knapp, 1949,
1954; Klapp, 1949, 1949-50, 1956; Klappet al., 1954; Wagner, 195ob;
Tiixen and Preising, 1951; Ellenberg, 1952a; Wendelberger, 1953b,
1954a; Tiixen, 1954b; Schneider,1954; Speideland van 'Senden,1954),
and weed assemblages(Krusemanand Vlieger, 1939; Sissingh, 1950;
Tiuxen, 1950b; Ellenberg, 1950a; J. Tiixen, 1958). A modificationof
terminology is introducedinto the system by Knapp (1948a, 1948b;
Wendelberger, 1954a), who regardsas the basic unit the widespread
'Hauptassoziationen,appearinglocally in the form of associations.The
Hauptassoziationand Assoziation are defined by regional and local
character-species(in the sense of Braun-Blanquet,1921), respectively.
The Hauptassoziationenmay be subdivided also into edaphic Haupt-
subassoziationen,which in turn appearlocally in the form of subasso-
ciations.
Aichinger's (1951a, 1951b, 1951c, 1952a, 1952b, 1952c, 1954;
1956-7; Wendelberger, 1951) system for applied forestry represents
a further departurefrom the doctrines of Braun-Blanquet.Although
Aichinger indicateshis adherenceto the school, his approachis funda-
mentally distinct from it; be considersthat the associationcannot serve
as the basis of forest management,and character-species appearin his
system only secondarilyand incidentally.Forests of the AustrianAlps
are grouped first into dominance-types(Obergruppen)by the major
canopyspecies;these are divided into edaphic-floristicGruppendefined
by combinationsof soil moisture,acidity, and nutrient conditions, and
differential-speciesgroups which indicate these conditions.Within the
groups the unit of primaryemphasisis the Vegetationsentwicklungstyp,
a very narrowly defined unit which brings together all those stands
which are alike in physiognomy (i.e., dominance), which correspond
both in floristic and sociologic charactersand in site relations, and

which belong to the same stage of a developmentalseries. While these
types may be named in the mannerof sociations,a given sociationmay
have differentroles in differentsuccessions(Aichinger, 195 ib). Charac-
terizationof the Vegetationzsentwicklungstyp must considerdifferential-
species indicating successionalrelations and must express the relation
of the type to other types (e.g., LaricetumcallunosumA PICEETUM
myrtillosumA Piceetumoxalidosum). As a systemadaptedto study of
the variouslychanging forests producedby human disturbanceAichin-
ger's approachis as strongly dynamicas the Americansystemof Clem-
ents, which it otherwiselittle resembles.Hartmann(1952) has devel-
oped anothersystemof forest types for the Austrianmountains,related
to the Aichinger system in its emphasisof environmentaland dynamic
relations,to the Braun-Blanquetsystem in its use of diagnosticspecies.
Forestsare classifiedby elevationzones and moisture-gradientrelations,
and within the major environmentalgroupings which result by under-
growth types and successionalrelations.
In addition to its general triumph in the area from the Low Coun-
tries to the Balkans,conceptionsof the school of Braun-Blanquethave
had increasingacceptancein Northern Europe (see below). The school
has also spread south froin France into Italy (Tomaselli, 1947, 1951,
1956; Giacomini, 1950), Spain (Braun-Blanquet,1948; Rivas Goday,
1948; Rothmaler, 1954; Bolos, 1954a), Portugal (Myre, 1945; Myre
and Pinto da Silva, 1949; Malato-Beliz, 1954; Braun-Blanquetet al.,
1956; Barreto,1958), and acrossthe Mediterraneanto Israeland Egypt
(Tadros, 1953). Although early work by Eig (1927), founder of the
Palestinianor Israeli school, was physiognomic,the later work of Eig
(1933, 1938, 1939, 1946), Boyko (1945, 1947, 1949, 1954), Zohary
(1945, 1946, 1950, 1953, 1960; Zoharyet al., 1947, 1949, 1951, 1954,
1956; Orshan and Zohary, 1955; Feinbrun and Zohary, 1955) and
Oppenheimer(1949, 1952) is generallywithin the form of the Braun-
Blanquet classification.The units are in large part determined,how-
ever, by physiognomyand dominance,as in the British ecological tra-
dition, ratherthan by character-species(vide Zoharyet al., 1949, 1956).
The Braun-Blanquetsystemhas been applied in Japanby Suzuki-Tokio
(1954a, 1954b), Usui (1955), Miyawaki (1960), and Sasaki (1957);
and the units have been used by Hosokawa (1952a, 1952b, 1954a,
1954b, 1954d) and Yamanaka (1955), although their approach is
primarilythrough dominanceinstead of character-species.In addition
to the basicunits of the Braun-Blanquetclassification,Hosokawa (195 1,

1952a, 1953, 1954b, 1954c, 1955a; Hosokawaet al., 1954) and Omura
(1953; Omura et al., 1955) have sought to develop a system of units
for epiphyte synusiae. There has been very little application of the
Braun-Blanquetsystem in orthodox form in the English-languagena-
tions. Recent work of Barker (1953), Chapman (1954b), and Moore
(1954) in New Zealandand of Bharucha(Bharuchaand Satyanaryan,
1954, 1958; Bharuchaand Leeuw, 1957; Satyanarayan,1958) in India
apply conceptsof the school. Conard (1935a, 1939, 1952, 1954), Dan-
sereau (1943, 1946) and other American authors, and Tansley and
Adamson (1926) and Poore (1955a, 1955b, 1955c, 1956) in England,
have followed the systemin part, but not as a whole. Becking (1956)
has recentlyapplied the systemto the Douglas-firforests of the north-
western United States.
When Braun-Blanquet'ssystem has been used outside its European
homeland, it has usually experienced considerabledilution with con-
cepts from other ecological traditions. In Europe and overseas it is,
however, by far the most extensivelyused single approachto intensive
study of vegetation; the school of Braun-Blanquethas produced, all
together, a literatureof ratheroverwhelmingextent (Braun-Blanquet,
1951a). The system is consideredto apply to animal communitiesas
well as to plants (Braun-Blanquet,1932b, 1953; Tiixen, 1932; Braun-
Blanquetand Tiixen, 1943; Rabeler, 1937a, 1947; Franz, 1950; Gisin,
1951; Tischler, 1951; Quezel and Verdier, 1953; Rabelerand Tiixen,
1955), to aquatic communities as well as terrestrialones (Allorge,
1925; Roll, 1938, 1942; Panknin, 1945; Margalef, 1949; Symoens,
1951; Braun-Blanquet,1951a; Molinier, 1960) and, with additional
units, to epiphytes (Barkman, 1958). In its developmentand spread,
the original ideas of Braun-Blanquethave had limited modification.
Physiognomyand dominanceare somewhatmore emphasizedin some
currentwork; and a unit based on dominance,the sociation,has been
acceptedas appropriatefor Scandinavianand other vegetation poor in
species (Braun-Blanquetand Tiixen, 1943; Braun-Blanquet,1951a;
Wendelberger, 1951, 1952). The concepts of diagnostic species have
been complicatedby various distinctions (Schwickerath,1940a, 194ob,
1942, 1954; Drees, 195la; Becking, 1957). The intensivework in cen-
tral Europe has suggested significant restrictionson the concepts of
association and character-species(Ellenberg, 1954a, 1956; Scamoni,
1954). Some recent work places more emphasis on vegetationalcon.
tinuity and relations to environmentalgradients in ecological series or

coordinatesystems,and on "ecologicalgroups" of species with similar
distributionalrelations (Duvigneaud, 1946; Ellenberg, 1948, 1950a,
1952a, 1954a, 1956; Wagner, 1950a, 1950c, 1954a, 1954b; Hauff et
al., 1950; Schonhar,1954; Klausing, 1956). But no other school has
followed so faithfully and for so long its original concepts.
Some recent works carrythe quasi-taxonomicapproachof the school
to its conclusion. Associationsare named in exact analogy to species,
with rules of priority and synonoymyand with the author and date
attachedto the name of the association (see recommendationsof Ar-
widsson, 1929; Moor, 1938; Dahl and Hadac, 1941; Braun-Blanquet,
1948-9; Drees, 1951a, 1951b, 1953; Barkman,1953). The first fascicle
of the Prodromedes Groupebzenets Vegetaux, an immenselyambitious
project to systematizethe plant communitiesof the world, was pub-
lished in 1933. In this and other works surveying the communities
of a region or higher category (Braun-Blanquet,1933; Meier and
Braun-Blanquet,1934; Braun-Blanquet,et al., 1936, 1939; Moor, 1937;
Tiixen, 1937; Westhoff et al., 1946; Knapp, 1948b; Braun-Blanquet,
1948-9; Lebrunet al., 1949; Klement, 1955), associationsare classified
into higher categories, listed with pertinent citations and synonymy,
and defined in terms of character-species, and the ecology and distri-
butions of the associationsare briefly noted and sub-unitsmentioned.
In these works the ideas of Braun-Blanquetfind their ultimate expres-
sion-treatment of natural communitiesin a form exactly paralleling
manuals of floras.
With the SouthernTraditionmay be grouped also many authorsof
diverse approachesin MediterraneanEurope. Extensive work in Italy
has been influencedby Negri (1905, 1912, 1914, 1926, 1927, 1932).
Negri's emphasison habitat and propertiesof communitiesother than
character-speciescontrastswith the system of Braun-Blanquet,and is
more nearly comparableto the approachesof Schmid and Ludi among
the schools of Zurich and Montpellier.Negri (1926, 1927, 1954) has
indicated also agreementwith the individualisticdissent of Ramensky,
Gleason, and Lenoble. Community-typesare informally characterized
by varied charactersof environment,physiognomy,and dominancein
studies by Negri and other Italian authors (e.g., Pichi-Sermolli,1948;
Zangheri, 1950). Classificationby physiognomy and dominance was
applied by Gaussen (1926) and Cuatrecasas(1929), and the concepts
of del Villar (1925, 1929a, 1929b) in Spain and Gaussen (1933,
1954) in Francehave influencedother authors(e.g., Cuatrecasas,1934;

Ciferri, 1936; Gomes and Grandvaux, 1954-5). Study of vegetation
through regional vegetation types-zones in mountains and climax
areas in the lowlands-has been of special importancein the Medi-
terraneanregion in work of Emberger (1930, 1936, 1942), Negri
(1934, 1947) De Philippis (1937), Gaussen (1938-9, 1951), Gia-
cobbe (1938, 1947-9), Mathon (1949), Negre (1950), Tomaselli
(1951), Bolos (1954b, 1954c), and Zangheri (1954). Some of these
authors (Emberger, Tomaselli, Negre, Bolos) are associated with the
school of Braun-Blanquet; others are not. Gaussen's approach to vegeta-
tation mappingis relatedto that of Schmid (Kiuchler1953); Zangheri's
(1954) definitionof climax areasis similarto those of Schmid (1949)
and Liudi (1935), and Schmid's system has been adopted in Italy by
Sappa (1951, 1952). In treating the vegetation of Africa, Trochain
(1940, 1953), Chevalier(1948), Pitot (1949), Schnell (1952), Ozen-
da (1954), Eeckhout (1954), and Gomes and Grandvaux (1954-5)
have used physiognomic units comparableto those of the American
Shantz (Shantz and Marbut, 1923) and British authors (Chipp, 1926;
Burtt Davy, 1938); Pichi-Sermolli(1955) has consideredapplications
of the Beard (1944a, 1955) physiognomicsystem.
THE NORTHERN TRADITION
While the evolution of vegetation units from Humboldt and Grise-

bach to Braun-Blanquetand others in southernEurope representsone
of the major traditions of phytosociology,the schools of northern
Europe are descendantsof a differenttradition. In the early treatment
of vegetation of middle Sweden by Post (1842, 1844, 1851, 1862),
individualplots were groupedinto vegetationslokaler,on approximately
the level of later Scandinavianassociations;and these locales were
grouped into vegetationsgruppercorrespondingto later formationsor
other higher groupings. With the work of Hult (1881, 1887, 1898)
some distinctivefeaturesof the northernschools and their principalunit
(the Scandinavianassociation, later the sociation) were already ap-
parent. Hult (1881) tracing the backgroundof vegetation-formsand
formationsfrom Humboldt through Grisebachand Kerner, outlined a
systemof narrowlydefinedvegetation-formsand vegetationunits based
on these. Emphasizingthe need for classificationbased on the vegeta-
tion itself, in opposition to the "deductive"classificationby habitatof
Norrlin (1870, 1872) and Waino (1878), Hult grouped stands by

their similarityin stratal structureand termed the resulting units for-
mations. These were, however, far smaller units than the formations
of southern authors and correspondedto the associations(sociations)
of Fries (1913) and later Scandinavianauthors.Hult's approachand
methods were further developed by Sernander (1898, 1899, 1901,
1912, 1915; Post and Sernander,1910), under whose leadershipthe
Uppsala school of plant sociology was founded, with characteristics-
the use of small quadratsfor quantitativestudies and of narrowvege-
tation units defined by stratalstructure-setting it apartfrom the south-
ern schools.
Nilsson (1902) classified Swedish communities into four major
series, primarilyby characterof the undergrowth,and subdivisionsof
these; Nilsson's classificationwas the basis of work also by Sylven
(1904), Fries (1913), Samuelsson(1917a), and Smith (1920). The
monograph of Fries (1913) applied the distinction made by Warm-
ing (1909) between the associationand formationto Swedish vegeta-
tion. The associationwas regardedas a vegetationtype uniform, on the
whole, in physiognomyand floristic composition; the formation was
characterizedthrough physiognomiccorrespondencewithout reference
to floristiccomposition.In practice,uniformitywithin associationswas
based on correspondencein stratalstructure.These small and numerous
units, definedwithout referenceto character-species, were fundamentally
different from the contemporaneousassociations of Braun-Blanquet
(1913), and were a first clear definition and systematicapplicationas
such of the northernor Scandinavian"association."Samuelsson(1917a)
followed Nilsson and Fries in his classificationand regardedthe as-
sociationas an abstraction,a grouping of similarstandswhich represent
the associationonly more or less incompletely,and which may well be
comparedwith the poorly bounded plant species of difficultgenera.
In the decade following Fries' study a series of major works ap-
peared from the Uppsala school (Du Rietz et al., 1918, 1920; Teng-
wall, 1920; Du Rietz, 1921; Osvald, 1923) and from Nordhagen
(1920, 1923) in Norway further applying and clarifyingthe northern
association.In the paperof Du Rietz, Fries, and Tengwall (1918) these
"youngerUppsala authors"of that time sought to give modernformu-
lation to the Swedish-Finniishtradition of Post, Hult and Sernander.
In a set of definitions of vegetation units (cf. Samuelsson,1916; Dui
Rietz, 1917; Cajander, 1922), the associationwas defined as a plant
communityof definite floristiccompositionand physiognomy,whereas

the "abstract"conceptions of Melin (1917) and Samuelsson (1916,
1917a, 1917b) were criticizedand the inclusion of habitatin the defi-
nition of Flahault and Schroter(1910) rejected. It was indicatedalso
that the classificationof plant communitiesis in and for itself a goal,
as is the grouping of organismsinto species in taxonomy. The asso-
ciation was later defined (Du Rietz et al., 1920; Du Rietz, 1921:144;
Osvald, 1923) as a plant communitywith definiteconstants and definite
physiognomy. The two major groupings above the association were
the complex of associationsrelated ecologicallyand sometimesdynam-
ically, as in bog vegetation (Du Rietz, 1917, 1930a, 1932; Du Rietz
et al., 1918; Osvald,1923, 1925), and the formation of physiognom-
ically equivalent associations (Fries, 1913; Samuelsson, 1916, 1917a,
Du Rietz, 1917, 1921, 1925b, 1930a, 1930b, 1932, 1949; Du Rietz
et al., 1918).
The associationwas regardedas a fundamentaland "concrete"unit
to be analyzed,especiallyin termsof constancy,throughquadratstudies.
From the extensive sampling work of these authors (Du Rietz et al.,
1920; Du Rietz, 1921), it appearedthat a set of constancy-lawscould
be established,further defining the nature of the associationand pro-
viding a secure methodologicalbasis for plant sociology. It was indi-
cated that each associationpossesses a number of species, or at least
one species, that occur in all the plots of the association,provided the
plots exceed a certain minimum area. The associationmay be defined
by these species, the constants, which occur in 90 per cent or more of
samples, either locally or throughoutthe range of the association.The
numberof constantsnormally exceeds the number of species in inter-
mediate constancyclasses;the associationis thus made up of a nucleus
of constant species, forming the dominant part of the vegetational
structureof the community,and a variablegroup of incidentaland ac-
companyingspecies. The associationwas thought also to be character-
ized by a definite minimum-areafor sampling, related to its richness
in species and the size of individual plants. When sample size was in-
creased, below the minimum-area,the number of constants increased
rapidly; but the number of constantsremainedthe same with further
increasein sample size beyond the minimum-area.Associationswere
thought to be in contactwith one anotherwith sharpboundaries,where
the group of species characterizingone abruptly replaced the group
characterizinganother;the transitionbetween associationswas normally
extremely small in relation to the area of the associationsthemselves.

Fromthese principlesa full definitionof the associationcould be formu-
lated (Du Rietz et al., 1920; Du Rietz, 1923a): An associationis a
complex of species-combinations which recurwith especialfrequencyin
nature and possess a common nucleus of species (constants) almost
never lacking and present in more or less definite quantitativerela-
tions; this complex is as a rule sharply bounded in relation to other
similar species-combinations-i.e., through the lack or relative rarity
of intermediatespecies-combinations.
The decade following the publicationsof Fries (1913) and Braun-
Blanquet (1913) thus saw the differentiationin the Swedish-Finnish
and Swiss-Frenchtraditionsof two clearly defined schools led respec-
tively by Du Rietz and Braun-Blanquet,each with its own techniques
and doctrines,but agreeing in their emphasison classificationand con-
ception of the association as the fundamental unit for the science
of naturalcommunities.The "associations"in questionwere, however,
very different units. As the means of studying and interpretingthese
also differed, a whole series of contrastsdistinguishedthe two schools
before later compromise: 1) the small associationsor "microassocia-
tions" (Riibel, 1925) of the Uppsala school vs. the larger associations
or "macroassociations"of the school of Braun-Blanquetand other
authorsof Ziirich-Montpeilier;2) characterizationof the northernas-
sociationsby constantspecies vs. characterizationof the southernasso-
ciations by character-species; 3) emphasisof stratalstructureand phys-
iognomy vs. emphasisof floristiccomposition;4) emphasisof quadrat
studies to determineconstancyvs. emphasisof estimationand the need
for experiencein judging fidelity; 5) attemptsat quantitativeresearch
into the nature of associationsvs. acceptanceof the convention of
character-species as sufficient;6) assertionof the "concrete"nature of
associationsvs. conceptionof associationsas abstractunits; 7) assump-
tion of sharp boundaries separating these units vs. acceptance of
some intergradationof associations;8) rejectionvs. acceptanceof the
term and concept of the "association-individual"; 9) grouping of as-
sociations by topographicand dynamic relations into vegetation-com-
plexes and 10) by physiognomyinto formationsvs. a floristichierarchy
of alliances, etc., above the association.The two schools might agree
in basic classificatoryperspective,but they disagreed on almost every
detail in actual applicationof classificationto vegetation. Thus, while
ecologists and phytosociologistshave on occasion offered criticismsof
one another, among the phytosociologiststhemselvesthe argumentbe-

tween the northern and southern camps was waged with the special
intensityof a civil war.
Each of the differencesabove contributedto the enlivening of phyto-
sociologicalliteratureduring the 1920's and 1930's. The large numbers
of small associationsin studies of the Uppsala school, often more than
a hundred in a limited area (see Fries, 1913; Osvald, 1923; Sterner,
1925; Waren, 1926; Kaiser, 1926; Regel, 1923a, 1923b; Nordhagen,
1928) have been a poiIntof criticism by phytosociologists (Braun-
Blanquet, 1925; Ludi, 1928) and ecologists (Tansley and Chipp,
1926:9). The concept of minimum-areahas been the subjectof much
discussion,reviewed by Goodall (1952), and little agreement.The use
of constancyas a criterion,togetherwith the indicationof a fundamental
distinctionbetween the constantsand accompanyingor incidentalspe-
cies, has been widely criticized (Arrhenius, 1921; Nordhagen, 1923,
1924, 1928; Kylin, 1923, 1926; Braun-Blanquet,1921, 1925; Pearsall,
1924; Alechin, 1925b; Riibel, 1925; Wangerin, 1925; Pavillard,1927;
Lippmaa, 1931; Katz, 1933; Ashby, 1936; Kalliola, 1939). In the
later developmentof the Uppsala school, emphasisin the definitionof
community-typesgradually shifted from constancyto dominance.Du
Rietz (1930a, 1932; see also 1936, 1949) later offereda full hierarchy
of units defined by dominancein rivalry to the floristic hierarchyof
Braun-Blanquetand Koch. The use of dominantswas, in turn, criti-
cized by supportersof the southern schools (Braun-Blanquet,1921,
1925, 1951a, 1951b; Lddi, 1928; Pavillard, 1935a, 1936; Tuomikoski,
1942). The assertionthat sharp boundariesseparatecommunities(or,
at least, the intensitywith which this was assertedby Du Rietz) found
a number of critics (Frodin, 1921; Frey, 1923; Kylin, 1923; 1926;
Kujala, 1926; Nordhagen, 1928; Liudi,1928; Kalliola, 1939). Southern
authors have considered arithmeticaloperation a weak basis for the
determinationof fundamental units (Riubel, 1925; Braun-Blanquet,
1925; Wangerin, 1925; Pavillard, 1927), and the quantitativeempha-
sis of the Uppsala school inspired Braun-Blanquet(1925) to describe
his rivals as the Herren Quadratiker.The conceptionof the association
as a "concrete"unit (Du Rietz et al., 1918; Du Rietz, 1921:15, 1923a,
1928, 1929) has been opposed by Nordhagen (1923, 1928), Regel
(1923a, 1923b), Wangerin (1925), Hueck (1925), Kylin (1926),
Waren (1926), Paczoski (1930b), Kalliola (1939), Kalela (1939),
and various southern authors (Braun-Blanquet,1921; Koch, 1925;
Schroter, 1926; Zlatnik, 1928b; Liidi, 1928; Pavillard, 1935c) re-

garding it as abstractin character.Answers to some of these criticisms
have been offered by Du Rietz (1923a, 1923b, 1925a, 1928). The
work of Du Rietz and his associatesgave the phytosociologistsfor the
first time a coherentbody of theory on the natureof the associationas
a fundamentalunit; but this apparentachievementwas not permitted
by the phytosociologiststhemselvesto stand for long.
The northern association, defined by major species of the various
strata,was applied in manystudiesin Sweden (Du Rietz, 1925a, 1925b,
1925c; Du Rietz and Nannfeldt, 1925; Sernander,1925; Vallin, 1925;
Blomgrenand Naumann, 1925; Almquist, 1929; Thunmark,1931, etc.)
and Norway. From Sweden and Norway this approachto vegetation
spread eastwardto the Baltic nations in work of Regel (1921, 1923a,
1923b, 1927a, 1927b, 1932, 1933, 1935, ;944) from Lithuania,Waren
(1926), Kalela (1939), Kalliola (1939), and Paasio (1939, 1941)
in Finland, and Vilberg (1927) in Estonia,and into Russia.From the
Scandinavianand Baltic area,the influenceof the Uppsala school spread
southward to the Netherlands (Vries, 1926, 1929, 1932), Germany
(Hueck, 1925; Kaiser, 1926; Krieger, 1937), Austria (Vierhapper,
1925; Zumpfe, 1929), Czechoslovakia (Domin, 1926; Firbas and
Sigmond, 1928; Hilitzer, 1927) and Hungary (Rapaics, 1927; Soo,
1929, 1930b). Throughoutmuch of this area ideas of the school of
Uppsalacame into directcompetitionwith those of the school of Braun-
Blanquet; and, as has been described, the latter gradually prevailed.
The approachthrough sociationshas been used also in the East Indies
(Kooper, 1927), North Americaand the British Isles (Osvald, 1933,
1949, 1954; Hanson, 1953, Gjaerevoll, 1954), and Japan (Ito, 1959;
Kitagawa, 1960; Tatewaki et al., 1960).
The intense rivalry of the two schools has in recent years largely
worn itself out, or been resolvedin compromise.Variousauthorssought
to contributeto the reconciliationof the two approaches(Riubel,1925,
1927; Vierhapper, 1926; Nordhagen, 1937, Vries, 1939). Du Rietz
(1935, 1936) came to the view that there are many, equally funda-
mental units in phytosociology,not one unit for which the term as-
sociation must be reserved (cf. criticism of Pavillard, 1936). Recog-
nition that vegetational conditions in northern and southern Europe
differ and might call for differentunits has made compromisepossible.
The term sociation was proposed by Riibel (1927) for the society of
ecologists and elemenitary-association of Drude, and was accepted by
Du Rietz (1930a, 1930b, 1932, 1935, 1936) and Gams (1933, 1936)
for the northernassociation.The agreementwas expressedin three reso-
lutions passed at the Sixth Botanical Congress in 1935 (Du Rietz,
1936; Cain, 1936a): (1) To use the term sociationfor vegetation-units
characterizedmainly by dominancein the differentlayers, in the sense
of Scandinavianplant sociologists. (2) To use the term associationfor
vegetation-unitscharacterizedmainly by characteristic-and differential-
species in the sense of Zurich-Montpellier,or at least for units of the
same order of sociological value; subassociationand facies can, where
necessary,be used for their subordinateunits. (3) To unite sociations
and associationsinto alliances in the sense of Ziwrich-Montpellier, and
the alliances into higher units. Agreement was thus possible on the
basis that the associationin the sense of Ziirich-Montpellierwas suitable
as a general unit and for the conditionsof southernEurope,while the
sociationwas an appropriateunit for the relativelypoor flora of Scan-
dinavia and other countries where community-typesmust be defined
by dominance.The sociationis acceptedas a unit in the recent edition
of Braun-Blanquet'stext (1951a), although with continued insistence
on the primacyof the associationin the sense of Braun-Blanquet.
A critical point of the compromisewas the possibility of fitting
sociationsinto a hierarchyof higher units defined in a fundamentally
differentmanner.This third recommendationof the Congresswas im-
plementedby anotherof the Scandinavianauthors,Nordhagen (1937)
in Norway. Nordhagen was one of the contributorsto establishmentof
the northernassociationor sociationas a unit (Nordhagen 1920, 1923,
the Soziotypus), and one of the most active of the Scandinaviantra-
dition in applying it (Nordhagen, 1923, 1928, 1937, 1940, 1943,
1954b, 1955; Faegri, 1934; Knaben, 1952). Nordhagen (1924, 1928)
has also criticisedthe school of Uppsala in some respects,while seek-
ing to bridge the gap between it and the school of Braun-Blanquet.In
his classificationof alpine vegetation (1937) the sociation was used
as the basic unit; but sociationswere grouped into alliancesand higher
units defined by character-species.Thus, if the sociationis regardedby
some Scandinaviansas a finer instrumentof ecological researchthan
the association (Faegri, 1937), the sociation may be fitted into a
phytosociologicalsystem of wider application.
Some recent Scandinavianwork has gone yet further toward agree-
ment with the Southern Tradition. Du Rietz (1942, 1945) adopted
the conceptsof Scheidearten, for differential-species,and Leitarten,cor-
responding in general to character-species, but limited to species almost

wholly restricted to a certain community-type (see also Waldheim,
1944b, 1947; Albertson, 1946, 1950; Gjaerevoll, L949, 1950; Sjogren,
1954). Unlike the practice in the South, however, the Scheidearten
are used to define associations and higher units; they are thus used
by Du Rietz (1954) to define bog formations, equivalent to the classes
of Braun-Blanquet. In some recent studies emphasis is shifted from
the sociation to the association (Sjors, 1948, 1954; Albertson, 1950;
Horn af Ranzien, 1951; Nordhagen, 1954a; Dahl, 1957) although
the latter is not necessarily, in the different vegetational conditions of
Scandinavia, the same unit as that of the school of Braun-Blanquet.
In Denmark and Iceland, development of vegetation classification
took directions diffcrent from, though related to, those in Sweden and
Norway. In Denmark, the country of Warming, the use of "formation"
for small vegetation units defined by species composition persisted
longest in the school of Raunkiaer. One of the pioneers of quantitative
approaches (1909-10, 1910), Raunkiaer shared with the Uppsala school
the emphasis on vegetational statistics and the use of small vegetation
units. Raunkiaer (1913, 1917) considered that the association and
formation were essentially the same unit, which might be characterized
floristically, physiognomically through growth-forms, and biologically
in terms of life-forms (Raunkiaer, 1918; cf. Negri, 1914). His nar-
rowly defined units, termed formations, were in general characterized
and named by their dominant species (1909-10, 1918); the formation
was later (1928) defined as comprising those stands agreeing in the
occurrence of "frequency dominants"-species present in 81-100 per
cent of the quadrats. Like other phytosociologists, Raunkiaer (1918)
compared his fundamental unit with the species of taxonomy. Forma-
tions could be grouped into series of formations by dominant life-
forms (1909-10); and a hierarchy of groups, classes, and branches
of formations was also suggested (1918).
"Formations" in the sense of Raunkiaer have been used by a number
of Danish phytosociologists (Raunkiaer, 1909-10, 1913, 1918, 1934;
Vahl, 1912, 1913, 1919; Olsen, 1914; Gr0ntved, 1927; Hansen, 1930,
1932) and the Norwegian, Resvoll-Holmsen (1912, 1914a, 1914b,
1920). The formations of these authors vary in scope and manner of
definition, but in general correspond to the sociations and associations
of other phytosociologists, and not to broadly defined physiognomic
formations. Convergence with other schools is suggested in the work
of Iversen (1936), in which the Danish formation is identified with

the sociation of Du Rietz, and sociations are grouped into Soziations-
Verbinde corresponding somewhat to the associations of Braun-Blan-
quet. Sorensen (1948) has sought to relate the various units through
quantitative comparison of samples.
In early work of Bocher (1933) in Greenland, community-types
defined by frequency-dominants were termed associations, and these
were grouped into formation-types in the sense of Riubel (1930). In
later work Bocher (1940, 1941a, 1941b, 1942, 1943, 1952a, 1952b;
Bocher et al., 1946) has used the sociation as a lower-level unit, to-
gether with types and series characterized by dominating species, phys-
iognomy, and ecology (B6cher, 1943; Bocher et al., 1946). Bocher
(1933, 1938, 1940, 1943, 1945; de Lesse, 1952) has also analyzed
geographic relations of species in communities, in work related to that
of Meusel (1939a, 1943c), but having its background in earlier work
of Vahl (1904), Ostenfeld (1926), and Hansen (1930) in Danish
plant geography. In the monograph on vegetation complexes of South-
west Greenland (B6cher, 1954), the character-species of Braun-Blan-
quet are rejected in favor of four other types of diagnostic species: (1)
Area species or area-geographical differential species-species which oc-
cur in related situations and communities in different areas, and by
their presence and absence differentiate the communities of these dif-
ferent areas. (2) Climatic species or climatic indicators-species which
indicate by their broader geographic distribution and climatic relations
something of the affinities of the community in which they occur. (3)
Habitat species or ecological differential species which, in a given area,
are associated with particular edaphic and microclimatic conditions
(and may be character-species for the communities of these). (4)
Ecogeographical guiding species, which are of special significance in
reflecting both local site or ecological, and broader geographic or cli-
matic, affinities of a community in which they occur. These ccn2epts
are used in defining higher units of a hierarchy: sociations, sociation
groups, vegetational types, vegetational complexes, and vegetational
regions.
Through the early work of Gronlund (1890), Stefansson (1895),
Jonsson (1895, 1901, 1905), and Ostenfeld (1899, 1905), Icelandic
vegetation types were distinguished by combinations of habitat charac-
ters with physiognomy. Within these types, or formations, Ostenfeld
(1905) recognized lower-level facies and associations by their major
species. The approach through formations, locally recognized within

the arcticvegetationof Icelandand in most casesgiven Icelandicnames,
was carried on by Thoroddsen (1914), Gall0e (1920), Oskarsson
(1927), Humlum (1936), Falk (1940), and Davi3sson (1946). Ideas
of Ostenfeld were combinedwith those of the school of Raunkiaerin
the study by the Danish author M0lholm Hansen (Hansen, 1930).
Quantitative samples were taken by the methods of Raunkiaerand
classified into formations recognized by Icelandic authors; these for-
mations were then arrangedinto ecological series in relation to soil
moisture and snow cover. The Icelandicapproachof Stefansson,Jons-
son, Ostenfeld, and Hansen has been developedinto a distinctiveclassi-
ficationof arcticvegetationin a seriesof studiesby Steindorsson(1935,
1936, 1937, 1942, 1945, 1946). Steindorssonrecognizesthe Icelandic
formations-Floi and Myri (sedge marsh and bog), Mosapemba
(Grimmia heath), Mo and Ja3ar (heaths, etc.), Melur (fjeldmark or
gravel flat), Flag (clay flat), Valllendi (dry lowland meadows), Brek-
kur or Graesli (grassy hillsides), Geiri (snow patch), etc. These are
grouped in the mannerof Nilsson (1902) and Fries (1913) into four
major series (Steindorsson, 1945). Within the formations relatively
numerous,narrowlydefined dominance-typesare recognizedand named
by one, two, or three major species. These associations are regardedas
equivalentto the associationsof Fries (1913), and correspondin gen-
eral to the sociations of other Scandinavianauthors;they are termed
sociations in a recent paper (Steindorsson, 1954).
In Estonia the emphasisof stratalunits becamethe centralcharacter
of the distinctive school of Lippmaa (1931, 1933a, 1933b, 1935a,
1935b, 1935c, 1939; Pastak, 1935; Sirgo, 1936; Tomson, 1937; Vaga,
1940). Gams (1918, 1927, 1932, 1933) had proposed basing vegeta-
tion study on synusiae, units made up of species of similar life-forms
and similar ecological requirements.Gams recognized three levels or
degreesof synusiaeand also distinguishedthese "ecological"units from
the "topographic"units, such as the association,formed usually from
combinationsof synusiae. The approachwas applied in the Alps by
Bolleter (1921; cf. Gams, 1936), who treated communitiesboth in
terms of ecological units or synusiae(Vereine) and in terms of spatial
or topographicunits (Gemneinde)often consisting of several of these.
The Gemeindewere thus some of the more important,among the many
possible,combinationsof Vereineinto stands,characterizedby the dom-
inant layerand componentsynusiae(Bolleter, 1921) . Approachesbased
on stratalunits, and complexesof these, were applied by Regel (1923b,

1927a) from Lithuaniaand by Vries (1926, 1932) in the Netherlands.
The use of stratal or synusial units was increasinglyacceptedby Du
Rietz (1930a, 1930b, 1932) in later works, in which he suggested a
whole hierarchyof names for stratalunits (socion, consocion,associon,
federion, formion), parallelingthe names for community-unitsdefined
by all strata.Of these the socion was later dropped,and consocion,as-
socion, and federion replacedby society or Verein, union, and federa-
tion (Du Rietz, 1936). Stratalunits have been used by many Scandi-
navian and Baltic authors (Kujala, 1929; Lindquist, 1931, 1938;
Paasio, 1941; Waldheim, 1944a; Regel, 1944, 1952; Krusenstjerna,
1945; Arnborg, 1940, 1953; Sjogren, 1954, etc.). Thunmark (1931),
Vaarama(1938), and Roll (1945) have consideredthe applicationto
aquatic communities; J0rgensen (1948) and Hayren (1956) have
treated algal communitiesas sociations.
In the Estonian school Lippmaa (1933b) considered that the ele-
mental units of vegetation were associations-unistrates or unions con-
fined to a single stratum,with one or two closely related life-forms
dominant.The union might consistof many speciesrelatedby life-form
and ecology, or of only one or two species. These single-layerassocia-
tions were thought to show an evident independenceof one another,
and to combine themselves in a quite variable fashion into forest
stands, in which environmentalrelationswere better revealedby lower
stratathan by canopy (Lippmaa, 1935a, 1935b, 1935c). Such complex
vegetation as that of a forest was regardedby Lippmaa (1935b) as
a complicatedassemblageof superimposedassociations,in which each
stratumis composed of one or more elemental associationsor unions.
Since species are differentlydistributed,floristiccompositionalone was
considered inadequatefor the definition of associations.He felt that
classificationof communitiesmust be ecological,and that the association
must be characterizedby habitat and life-form as well as floristiccom-
position (Lippmaa, 1931, 1933a, 1939). Within the union Lippmaa
(1939) recognizedgeographicfacies. For classificationabovethe union,
Lippmaa (1933b, 1939) proposed a hierarchy based on grouping
unions of a given area in which the same life-form (or two similar
life-forms) dominates and which are ecologically related into an As-
soziationsgattungor genus of unions. Hligherunits were suggested up
to the division comprisingall the unions of the earthin which the same
life-form or two closely similar life-forms dominate.
Lippmaa'sunion was considered by him a different and narrower

unit than the synusia of Gams, although the two terms (and Verein,
Bolleter, 1921; Du Rietz, 1936; Braun-Blanquet,1951a:47) have often
been used as alternativenamesfor stratalunits. The union differedalso
from the stratalcomponentsof the sociation,the socion and consocion
of Du Rietz (1930a, 1932); for the unions were broaderunits which
might be of varied floristiccomposition (cf. the associon of Du Rietz,
1932). The approachesof the Uppsala and Estonianschools are other-
wise closely related, since both treat communitiesin terms of stratal
units and also of "topographic"units built up of these. The two
approachesdiffer accordingto which of the two types of units receives
primaryand which secondaryemphasis.They differalso in the fact that,
while the necessityof basing vegetation units on the vegetation itself
was a central doctrine of the Uppsala school (Du Rietz, 1921), the
use of habitat in the definition is advocatedby Lippmaa.
The synusial approachhas been criticized by a number of authors
(Beger, 1922-3:43; Wangerin, 1925; Gleason, 1936; Cain, 1936a;
Pavillard, 1936; Krieger, 1937; Kalliola, 1939; Tuomikoski, 1942;
Schmid, 1942; Braun-Blanquet,195la: 16, 49), especiallyfor its separa-
tion of the interactingstrataof a single communityinto differentunits.
The approachas it has developed in the Alps, the Scandinavianand
Baltic area, in Russia, and elsewhere represents,however, more or less
independentrecognitionby differentauthorsof a fundamentalvegeta-
tional circumstance.Plant species which characterizeunits of different
strata, and hence these units themselves, are differently distributed;
and the stratalunits are in this sense partially"independent"(Cajander,
1909; Gams, 1918; Bolleter, 1921; Waren, 1926; Lippmaa, 1933b,
1935a, 1935b, 1935c; Vaarama, 1938; Motyka, 1947; Daubenmire,
1952, 1954; Whittaker, 1956). Recognition of stratal units and of
combinations of these into "topographic"units (sociations, etc.) is
thus one major possible approachto the study of vegetation, perhaps
most effective in some northern and mountain vegetation.
The Dutch school of de Vries and his associatesmay also be grouped
with the Northern Tradition,althoughVries (1938, 1939, 1953; Dam-
man and Vries, 1954) has sought also to relate his concepts to those
of the school of Braun-Blanquet.Earlier work of Vries and others
(Vries, 1926, 1929, 1932, 1935, 1939; Peeters, et al., 1927; Schey-
grond, 1932; Feekes, 1936) was based on stratalunits and their dom-
inants, and their combinationinto stands and complexes.The approach
CLASSIFICATION OF NATURAL CONMMUNITIES 35
was closely related to that of Du Rietz (1932, 1936), the Estonian

school of Lippmaa, and the Lithuanianof Regel. More recently ex-
tensive research into the ecology of grasslands has been carried out by
Zijlstra (1928, 1940) and Vries. Character-species
are consideredless
useful than quantitativemeasurementsof frequencyand weight (Vries,
1940, 1949a, 1949b, 1954; Vries and Ennik, 1953). Frequencies in
different communities,indicatorvalue, and agriculturalvalue are con-
sidered in selecting those species ("frequency-indicators")which are
to be used for the characterizationof community-types.The grassland
types recognizedare combinationsof frequency-indicators; among them
are distinguishedmain-typeswith one, types with two, and sub-types
with three characteristicspecies. The types are regardedas intermediate
in some respectsto the units of Braun-Blanquetbased on character-spe-
cies and those of the Northern Traditiornbased on dominants (Vries,
1948, 1949a, 1954; Vries et al., 1951).
The Finnish authorCajanderwas one of the pioneersof the Swedish-
Finnish or Scandinavianand Baltic tradition,in the broadersense, in-
cluding the work of Norrlin (1870, 1872, 1873) and Nilsson (1902)
as well as the line of descent from Post through Hult to the Uppsala
school (Cajander, 1923, 1925b). Cajander (1903a, 1903b, 1903c)
used an early form of the compiled tables of stand samples which ap-
pear in most later phytosociologicalstudies and establishedthe manner
of naming associations (1903a) and sociations (1903c). Cajander rec-
ognized associationsby dominantspeciesor combinationsof dominants;
and on the basis of characteristics of other layers he distinguished
Fazies within these, corresponding more nearly to the sociations of
Uppsala (Cajander, 1903a, 1922; Cajander and Ilvessalo, 1921). The
concept of ecological series or chains of communities along environ-
mental gradients, characteristic of much later Finnish work, appears in
his studyof the relationsof alluvialvegetationto waterlevel (Cajander,
1903a). Cajander'searly work (1902, 1903a, 1903b, 1903c) also con-
tains extensive observationson the relations of undergrowthtypes to
soil conditions, and these observationsbecame the basis of the forest
type system published in 1909.
Cajander(1909) observedthat the ground vegetationof forestswas
to a considerableextent independent of the tree species forming the
canopy, so that the same undergrowthtypes might appearunder pine,
spruce, or birch, even though some differencesmust be expected be-
cause of the differentlight conditionsand litter. Since the ground vege-
tation is much more expressive of site than canopy composition is, a

system of forest site-types relating undergrowth communitiesto site
was proposed. In this approachall forest stands are classifiedtogether,
the undergrowthsof which have largely identical floristiccomposition
and the same ecological-biologicalcharacterat felling age and normal
density, as well as all those stands the undergrowthof which differs
from that just defined only in such respectsas are consideredtransitory
or accidental-e.g., in consequenceof differentage of stands, of fell-
ing, of differencein tree species,etc. (Cajanderand Ilvessalo, 1921:17;
Cajander,1949:31; Ilvessalo, 1949). Although namedfor majorunder-
growth species, the site-typeis characterizedthrough the whole compo-
sition of its lower strataand through severalLeitpflanzenarten appear-
ing always or very frequently,though with changing abundance.Thus
the plant for which a type is named may not infrequentlybe lacking
from a stand of the type (Cajanderand Ilvessalo, 1921; Linkola, 1924;
Cajander,1925a), as is possiblealso in the associationsof Zurich-Mont-
pellier but not in the sociation.Other authors,however, (e.g., Kalela,
1954) regard constancyand dominanceof species as more important
than diagnostic species for characterizingtypes.
The site-type approachhas been used extensivelyfor forest research
and practicalforestryin Finland (Cajander,1909, 1925a, 1949; Cajan-
der and Ilvessalo, 1921; Ilvessalo, 1922; Palmgren, 1922, 1928; Riuhl,
1936; Kujala, 1938; Keltikangas, 1945). A system of bog types was
also set up by Cajander (1913) and further developed by Lukkala
(1929), Aario (1932) and Paasio (1933, 1936, 1940, 1941); the
latter work (Paasio, 1941) investigates the relation of unions and
sociationsto bog types. In a study of Canadianforests, Kujala (1945)
based comparisonof forest patternsin differentareason parallel type-
series (Kujala, 1938). Three classes of site-typesin relationto moisture
conditions were recognized by Cajander;and Kujala consideredthat,
with subdivision of these, approximatelysix steps or types could be
recognizedalong the moisture gradient in a given area as a basis for
comparingseries and determiningthe equivalenceof types in different
areas. The middle membersof the series are normallybest developed
and most extensive in an area, as affected by its climate; these inter-
mediate types or Leittypen are thus equivalent to the climatic climax
in the Americansystem of Clements (Kujala, 1945). Kalela (1954)
also distinguishesregional or climaticfrom local or edaphic communi-
ties, but considersthat each forest site-type,regional or local, comprises
a climax community-typeand the various successionalcommunitiesde-

veloping toward it.
Forest type approachesbased more or less on Cajander'shave been
applied in many other countries: Sweden (Malmstrom, 1926, 1936),
Denmark (Bornebusch,1923-5, 1929), Estonia (Linkola, 1929, 1930),
Latvia (Kirstein, 1929; Mallner, 1944), Germany (Bjorkenheim, 1919;
Wiedemann, 1929; Kotz, 1929; Gaisberg and Schmid, 1933), Austria
(Hufnagel, 1954), Switzerland (Linkola, 1924), Czechoslovakia,
(Konsel, 1929), Scotland (Anderson, 1926), South Africa (Phillips,
1928), the United States and Canada (Ilvessalo, 1929; Heimburger,
1934, 1941; Sisam, 1938; Ray, 1941; Kujala, 1945; Spilsbury and
Smith, 1947; Crandall, 1958), and India (Puri, 1954). Studies of
German forest-types by Meusel (1935, 1943b, 1951a; Meusel and
Hartmann, 1943) follow the Northern Tradition in recognition of
undergrowthtypes by constants and life-forms, and arrangementof
these in a two-dimensionalpattern of ecological series in relation to
soil moisture and chemical properties. Meusel (1951-2, 1954a) has
also approachedforest classificationthrough the tree stratumand diag-
nostic species in the undergrowth.
In Denmark a system of forest types following Cajander'swas de-
veloped by Bornebusch (1923-5); and in Sweden a system of types
and main-typesbased primarilyon undergrowthwas applied by Malm-
strom (1926, 1936; Lundblad, 1927). A somewhat different system
based on combinations of stratal unions, i.e. sociations, was de-
veloped in Sweden by Eneroth (1931, 1934, 1936) and Arnborg
(1940, 1953). In this approach,the forest site-typecomprisesa whole
series of plant communitiesof which only one stage-present under
certain conditions, particularlyin normal density of the tree layer-
is normative for the type. A number of different undergrowthcom-
munities are grouped together in relation to a kind of site and a
"normative"communitywhich occurs in such sites under certain con-
ditions (Arnborg, 1940:130-131). The site-type approach is thus
carried to its logical conclusion; it is a classificationnot simply of
communities,but of sites as characterizedby groups of related com-
munities (cf. Kalela, 1954). From studies of the Swedishbeech forests
Lindquist (1931) has indicatedthe limitationsof Bornebusch'ssystem,
and applied the synusialsystemsof Du Rietz (Lindquist, 1931, 1938)
and Enerothand Arnborg (Lindquist, 1954).
The schools of Du Rietz, Lippmaa,Raunkiaer,Cajander,and others
representdifferent approachesto the study of vegetation, based on at

least three differentmajor units; but they have in common greaterem-
phasis of stratalstructureand life-forms than in southernphytosociol-
ogy. The tendencytowardgreateremphasisof undergrowththan canopy
of forests is in contrastalso to the practiceof many English-speaking
ecologists. Although no one of the northern approacheshas been so
much used as Braun-Blanquet's,the combinedinfluenceof these north-
ern schools on the developmentof ecology and phytosociologythrough-
out the world may be even greaterthan that of the southern schools.
The ecology of the English-speakingcountries has been much more
influencedby the northernschools than by Ziirich-Montpellier-in the
great influenceof Warming and the use of dominance-typesand forma-
tions, the use of Raunkiaer'slife-forms and of quantitativemethods
from the schools of Raunkiaerand of Uppsala, and the applicationby
individual authorsof site-types,unions, and sociations.And, while the
sociationhas had very little use among ecologists, it becamethe major
vegetation unit of Russian phytocenology.The magnitudeof the con-
tribution of the smaller nations of the Scandinavianand Baltic arei
(and of Switzerland) is one of the striking features of the history of
ecology as a whole.
THE RUSSIAN TRADITION

The early history of the RussianTradition is less easily traced than
those of Western nations (vide Sukatschew, 1932; Alechin, 1932a,
1946; Carpenter, 1939a; Roussin, 1948). Krassnow (1888) and
Korjinsky(1888) used the termformation;these and other authorswere
much influenced by Dokutschaeffand the early development of soil
science in Russia. The Polish pioneer Paczoski (1891, 1896, 1928,
1930a, 193Gb) applied the term formation to plant communitiesand
introducedthe study of vegetationaldynamicsinto the Russiantradition
(1891, 1930a). Paczoski (1896) also proposed the term phytosociol-
ogy, later generally accepted for the study of plant communitiesin
western Europe but replaced in Russia by phytocenologyand biocen-
ology.
In an early study in the region of the semi-desert,Keller (Dimo and
Keller, 1907) introduced two ideas which were most influential in
the later developmentof the RussianTradition-the vegetationmosaic
(see also Keller, 1936) and the ecologicalseries (see also Keller, 1925-
6; Cajander,1903a). The idea of mosaicsor complexeswas also exten-
sively developed by Osvald and others in the school of Uppsala; but

the approachthrough ecologicalseries is most characteristically Russian,
a major contributionand distinguishingfeature of the Russian Tradi-
tion. Keller (1925-6) regardedthe community-typesalong an ecologi-
cal series as associations,but felt that the associationsshould be viewed
only as arbitraryand artificialabstractionsfrom an uninterruptedchain
of naturalphenomena:It is the ecological series, ratherthan the asso-
ciations,that are fundamental.Stratalor synusialunits were applied to
arid and semi-aridzone vegetation by Keller (1923, 1932, Genossen-
schaften, Convictiones) and by Grossheim (1930; Grossheimand Pri-
lipko, 1929) at Baku(Aggregation, Agglomneration, and Semiassozia-
tion and Assoziation for synusiae of the first, second, and third de-
grees; cf. Petrovskii, 1960). The approach of the Kasan-Voronezh
school of desert and steppe ecology, led by Keller and Gordjagin,was
carried further in the study of grassland vegetation by the school of
Ramenskyat Voronezh. Ramensky (1924, 1930, 1932) rejected the
associationas a unit; his object was not classificationbut the arrange-
ment of samplesin relationto environmentgradientsthroughecological
series. The approachwas applied to bog vegetation by Gerassimow
(1928).
Alechin, leader of the Moscow school in the study of steppe vegeta-
tion, also grouped communitiesinto association-complexesand associa-
tion-series along envirionmentaland successionalgradients (Alechin,
1925a, 1926). In an earlierpaper on the association(1925b; criticism
by Nordhagen, 1928:470) he asserted, like Du Rietz, that the asso-
ciation was a concreteunit or reality,ratherthan an abstraction;but he
also rejectedthe Uppsala definitionof the associationby constantsand
physiognomy, regarding floristic composition as the most important
characteristic.In later practice,however, Alechin (1932b) named as-
sociationsby the one or two dominantspecies of each stratumin agree-
ment with Katz and the Uppsala school.
The methods of the Uppsala school were applied to bog vegetation
by Katz (Katz and Katz, 1926), with general agreementon constancy
results and the necessity of narrow associationsdefined by strata.As-
sociations were grouped also into association-complexes,like those of
Osvald, and Sammelassoziationen (Katz, 1926; Dokturowsky, 1927)
and bog types (Katz, 1930c). In treatingforest vegetationKatz applied
the conception of twin-associations (cf. Hult, 1881; Du Rietz, et al.,
1918; Almquist, 1929; criticism of Cain, 1935), differing from one
anotherin a single stratum;and he describedsome associationsas sup-

plemental to another association- i.e., possessing a supplementary
stratum in addition to those in the other association (Katz, 1929,
1930a, 1930b). The twin-associationswere arrangedby the degree of
their similarity into series of twin-associations,and these twin-series
were regardedas the next systematicunit above the association(Katz,
1929). From constancystudies Katz (1930a, 1930b) concluded that
the various twin-associationsof northern and eastern Europe had a
very high, probablypreponderantnumber of species whose constancy
was regularlylinked with the dominantspecies of strata.Of the various
possibilities for defining associations, Katz (1930b, 1933) rejected
habitat, total species composition, constants, and character-species;he
consideredthat only the dominatingspecies of stratacould serve as an
unobjectionablebasis for the establishmentand characterization of asso-
ciations. This Russian "association"is derived from the Uppsala so-
ciation and is in form identicalwith it; but the mannerof using these
units and relating them to one another differs from that of most
Scandinavianauthors.
An earlysystemof forest types was developedin Russiaby Morosow
(1907, fide Sukatschew, 1932), independently of Cajander and in
close relationto Russiansoil science.Morosow (1928) conceivedforest
types holistically, as landscape-typesin which the biological, biosocial,
biogeographic, and physical-geographicfactors merge and interrelate
into the whole. While the undergrowthis a very sensitive measureof
site factors,undergrowthmay be changedby the conditionof the forest
itself. Primaryemphasiswas consequentlyplaced on edaphicproperties
of the site; and Russianvernacularexpressionswere used for the charac-
terization of stand types-e.g. "ramenj" for spruce stands of well-
drained, podsolized sandy loams. A system of broadly-definedforest
types, also based primarilyon soil properties,was developed by Krue-
dener (1926a) and criticizedby Sukatschew(1932) in relationto his
own, more widely used, system. Kruedener (1926b) also conceived
forest types as landscape-types.
Sukatschew(1929) definedthe associationthroughits "determinant"
species-the communitydominantsplus the stratal and seasonal "sub-
dominants"of Nichols (1923). The plant associationcomprisesthose
communities which correspond in phytosocial relations or, in other
words, are alike in their compositionof determinantsand in the com-
plex of direct site factors (Dingelstedt, 1928; Sukatschew,1929:310,
1935). These associationsmay, through their ecological and spatial re-

lations, be groupedinto association-seriesand successionalseries (Sukat-
schew, 1928, 1932; Socava,1927, 1930; Sambuk,1930a, 1930b; Andre-
jew, 1932; Kortschagin, 1932, 1935; Sokolowa, 1935, 1936; Markov,
1938; Leskov, 1938) and association-complexes or association-mosaics
(Sukatschew,1929; Lavrenko,1938). Associationsare also united into
association-groupswhich may be characterizedby undergrowthsimilarity
(e.g., Pinetacallunosa,Pineta cladinosa,and Pineta hylocomiosaGesam-
tassoziationen,Sambuk,1930b; cf. Solonievicz, 1933; Sokolowa, 1935,
1936; Ljubimova, 1935; Leontjev, 1935; Schennikov, 1938) and into
formations. A full hierarchyof vegetation units (association, associa-
tion-group, association-order,formation, formation-group,formation-
order, vegetation-type) was advanced by Sukatschew (1935; Kort-
schagin, 1946); a hierarchyrepresentingassumedcommunityphylogeny
has been suggestedby Socava(1944, 1945; Haudricourt,1948). Sukat-
schew (1935) has emphasizedthe importanceof defining"associations"
by stratalstructure.His associationsare consequently,like those of the
Moscow school, sociationsin the sense of Western authors;but Sukat-
schew (1935) has rejectedthe term sociation for this unit.
A most distinctivefeature of Sukatschew'swork, as of much of the
Russian Tradition, is the practice of relating associationsto environ-
mental gradients in patterns of ecological series (Sukatschew, 1928,
1932). Since the associationsrepresentpositions along environmental
gradients,they are also site-typesfor forestry.Sukatschew(1932) con-
siders that his types converge to some extent with those of Cajander,
but the Russian associationis a differentunit from the site-type. Suk-
atschew (1932) also rejects the Finnish practiceof grouping together
stands of differentcanopies;for the differencebetween canopy species
is more important in forestry than difference in undergrowth.More
recent writings of Sukatschew(1944, 1954, 1960) develop a philos-
ophy emphasizing dynamic and ecosystemicinterpretationsof natural
communities.The forest cover of a land is regarded as in perpetual
movementand change;forest-typeschangeinto one anothernow slowly,
now more rapidly, through various interrelated and variously com-
bined biotic, climatic, and physiographicprocesses. Each stand is to
be conceived as a biogeocenose, a complex whole of organisms and
environmentand their interrelations,as suggestedby Morosow (1928).
The forest-type is a grouping of stands, or individual forest biogeo-
cenoses, which are similar in composition of trees, other plants, and
animals, in the whole complex of environmentalconditions and inter-

relationsof plants and environment,and in processesof succession,and
which therefore require uniform managementmeasures (Sukatschew,
1954).
Generalizationabout the RussianTradition is difficult,both because
of the magnitudeand diversityof the contributionsup until the middle
1930's and becauseof the relative inaccessibilityof the literature,par-
ticularlyafter that time. Some of the Russian schools are little known
to Westerners and not mentioned here. The works cited above may,
however, provide a sufficient sampling to characterizethe principal
Russianapproachto associations:(1) the use of narrowlydefined asso-
ciations (i.e., sociations) named by stratal dominants and (2) treat-
ment of these associationsnot primarilyas self-sufficientunits, but as
parts of complexes or mosaics and, especially, ecological series. The
basic features of this approachappear in both the major schools of
Sukatschewat Leningrad and of Alechin and Katz at Moscow, and
may be recognized in many Russian studies other than those already
cited. The close relationof this approachto the Northern Traditionis
evident. While the ecological series and complex are concepts indi-
genous to the Russiantradition,the former is sharedwith the Finnish
school and the latter, the stratalapproach,and the associationconcept
with the school of Uppsala.So far as the approachto associationsis con-
cerned, the Russian Tradition is in some respectssuggestive of a de-
velopmentof the Uppsalaschool without later compromisewith Zurich-
Montpellier.
THE BRITISH TRADITION
At the turn of the century,the study of plant associationswas intro-

duced into Britain by Robert Smith (1898, 1899) and regarded by
Tansley (1904) as the main subjectmatterof ecology. Smith had been
a student of Flahault who, before the development of the character-
species doctrine among later authors of Ziirich-Montpellier,defined
associationsby dominant species. The approach to lower-level com-
munity-typesthrough dominancewas thus introducedinto the British
Tradition,and applied in the earlywork of RobertSmith (1898, 1900a,
1900b), W. G. Smith (1904-5; Smith and Moss, 1903; Smith and
Rankin, 1903), and Lewis (1904). The directionof the British Tradi-
tion was strongly influenced also (Tansley, 1947) by the work of
Warming (1895) and its English translation(1909). The major fea-
CLASSIFICATION OF NAI'URAL COMMUNITIES 43
tures of Warming's approachto classification-through physiognomy

at higher levels and for treatmentof extensiveareas,throughdominance
for more intensive studies and lower levels-have characterizedmuch
British ecological work from its beginnings to the present.
Distinctive approachesemphasizing vegetational dynamics also ap-
peared in England with Crampton's(1911, 1912) study of stable and
migratoryformations and in the work of Moss (1907, 1910, 1913).
In Moss' formation concept, physiognomywas set aside in favor of
habitat and vegetational dynamics.A formation comprised a number
of progressive,successionalor subordinateassociationsculminating in
one or more stable or chief associationswhich maintainthemselvesin
a habitat (Moss, 1910; Tansley, 1911); the formation was the whole
of the vegetation occurringin a definite habitat or on a certain type
of soil (Moss, 1913; Tansley, 1911). Three edaphically determined
main series of communities were recognizedby Moss et al. (1910),
and these were divided into associations characterizedby one or two
dominatingtree species. The use of physiognomyas a criterionof for-
mations was criticizedby Tansley (1913) as subjectto the artificiality
that necessarilyattachesto all single-charactersystems.Tansley consid-
ered that habitat,however determined,must be recognizedas the basis
of any natural classificationof vegetation, because habitat is the basis
of all vegetational resemblancesand differences.The ideas of Moss,
and the edaphic and dynamic emphasis shared by him and Tansley,
were carriedinto the joint study of Types of British Vegetation (Tans-
ley, 1911) as characteristicsof what might be describedat that time
as a British school.
Tansley (1920) later indicated that the formation, like the associa-
tion, must be recognized by the actual vegetation; but he considered
growth-form unsatisfactoryas a criterion because it brings together
communitiesnot closely relatedand separatesothersthat are. Indicating
also the difficultyof using two principlesof division (such as physiog-
nomy and habitat), since these will not everywhererun parallel,Tans-
ley (1920; cf. Pearsall, 1918) regardedthe formationas the sum total
of the developmentallyrelatedcommunitiesgroupedaroundthe mature
association,a conception derived from Moss and related to Clements'
climax formation. Unlike the Clementsianformation, however, Tans-
ley's formations include mature, major communities other than the
climatic climax; thus salt marsheswith their successionalcommunities
may be regardedas a formation determinedby salt water as a major
factor (Tansley, 1941). Tansley's major work of 1939 describedthe

plant formationas a unit of vegetationformed by habitatand expressed
by distinctivegrowth-forms.The formationis composedof associations,
each with different dominants and at least some different subordinate
species. In the use of association, consociation, society, and other units
Tansley (1920, 1939, 1940) largely follows Clements except that, in
accordancewith the polyclimax theory, many self-maintaining com-
munities are termed associations which Clementswould regardas suc-
cessional and term associes.
Comparedwith the literatureof phytosociology,that of British ecol-
ogy contains a remarkablysmall amount of debate on technicalitiesof
communityclassification.While the works of Continentalauthors are
often studies of plant communitiesfor the purpose of classification,
those of British ecologists tend to be, simply, studies of communities.
Few British authors have used complex hierarchiesand elaborateter-
minologies of communitiesin the model of either Braun-Blanquetor
Clements.Many have describedthe objectsof their study only as "com-
munities," or have termed them "associations,"with very little regard
for formal classification.McLean (1935) suggestedthe termsdominion
and condominion as preferable to association for dominance-types
characterizedby one, or by two or more, dominant species. In current
work community-typesare usually recognizedand named by dominat-
ing species, or in some cases by habitat;the approachthrough charac-
ter-speciesis no more in evidence than in Americanecology.
A certain informality in the approachto communitiesthus distin-
guishes British ecology from both phytosociologyand the Clementsian
school in Americanecology. In spite of the strong influence of Tans-
ley, it is not possible to define later British ecology in terms of a
school with a definite set of beliefs and practices as one may the
schools of Uppsala and Braun-Blanquet.In the approachto classifica-
tion, however, some general tendenciesmay be recognizedas sharedby
Tansley and many other authors: (1) Definition of units primarilyby
dominant species, not character-speciesor stratal structure. (2) In-
formalityin the naming of these units, for which the terms association
and consociation and the suffix -etum may or may not be used. (3)
Grouping by habitat,physiognomy,or both, into formations,but with-
out the developmentof complex hierarchies.(4) Recognitionof larger
numbersof "associations"than among Clementsianecologists who re-
strict this term to "climaticclimax" communities.
Poore (1955a, 1955b, 1955c, 1956) has recently consideredappli-

cation of techniques of the phytosociologists.The system of Braun-
Blanquet is criticized for lack of appreciationof the importanceof
dominantspecies, over-emphasisof fidelity, and the belief that associa-
tions can be naturally classified into a hierarchy.Poore conceives of
vegetationas forming a multi-dimensionalpatternor networkof varia-
tion; in this patternabstractpoints of referencecan be establishedwhich
coincide with frequently occurringand easily recognizableplant com-
munities. An abstractunit on any level may be termed a nodum (i.e.,
a community-type);the term nodumis thus analogousto the term taxon
in systematics.Recurringpoints in the vegetation pattern and abstract
units based on them can be defined most exactly and most readily by
the constant and dominantspecies together. Neither the character-spe-
cies nor the dominants alone are suitable; and the loose naming of
units by the dominants,prevalentin British literature,is to be depre-
cated. The approach has been applied also by Poore and McVean
(1957) and Gimingham (1961).
The work of ecologists in Commonwealthcountriesoverseasis even
less to be characterizedas a school. General characteristicsof the Eng-
lish-languagetradition--emphasisof vegetationaldynamicsand recog-
nition of community-typesby physiognomyand dominance-appear in
most work of Commonwealthauthors. Some authors have sought to
apply the Clementsiansystem,othershave used a more characteristically
British approach,or have tried the Clementsiansystem and abandoned
it in favor of an approachthrough polyclimaxesand dominance-types.
In Africa, Bews (1912, 1913, 1917) followed Warming in the
definition of formations and associationsand later (1920) used the
Clementsian system. Chipp (1927) applied the terms common to
Britishand Americanauthorsto the Gold Coastforest, but used a poly-
climax interpretation. Phillips (1930, 1931a, 1931b, 1934-5, 1954),
Scott (1934), Gilliland (1938, 1952), and West (1951) have applied
the Clementsiansystem;other authorshave indicatedtheir dissent from
the monoclimax theory (Michelmore, 1934; Gillman, 1936; Milne,
1937). Milne (1947), Morison et al. (1948), Gillman (1949), and
Thomas (1945-6) have approachedvegetation interpretationthrough
catenas,ecological series of communitiescorrespondingto gradientsof
soils in relationto topography.Burtt (1942) in East Africa and various
South African authors (Galpin, 1926; Muir, 1929; Pole Evans, 1936;
Dyer, 1937; Adamson, 1938; Louw, 1951; Story, 1952; Acocks, 1953;
Archibald, 1955) have recognized community-typesby varying com-

binations of physiognomy,dominance,and habitatcharacteristicswith-
out formal classification.Recent work of Kassas (1952, 1953, 1957;
Kassas and Iman, 1954) in Egypt is based on habitat-typesand dom-
inance-types.
Stamp and Lord (1923; Stamp, 1925) followed Kurz (1877) in
classifyingthe vegetationof Burmaand sought also to apply the system
of Clements.In India, Bor (1938), Chatterjee(1958), and Misra and
Puri (1958) have followed Clements,but with allowancefor the possi-
bility of edaphicclimaxes;other authors(Misra, 1944, 1946; Misraand
Joshi, 1952; Biswasand Rao, 1953; Joshi, 1956) and Chaudhri(1961)
in Pakistanhave based their work on dominancetypes and their suc-
cessionaland environmentalrelations.Rosayro(1950, 1958) has adopt-
ed the Clementsiansystem in Ceylon (cf. Holmes, 1951, 1958). In
Canada, E. H. Moss (1932, 1944; Moss and Campbell, 1947) and
others (Lewis et al., 1928; Dowding, 1929; Breitung, 1954; Ritchie,
1956, 1959) have recognized community-typesthrough dominance
and habitat without strict Clementsian treatment; Coupland (1950,
1961; Coupland and Brayshaw, 1953) has used the Clementsianas-
sociation.Recentwork of Moss (1952, 1953a, 1953b, 1955) treats
some communitiesas edaphicclimaxes and recognizescommunity-types
more narrowlydefined than those of Clements-associations and con-
sociations defined by dominant species and fasciations defined by
combinationsof canopydominantsand undergrowth.Hills (1960) has
developed a landscapeapproachto Ontario forest ecosystems,consid-
ering all aspects of the forest communityand its environmentand re-
lating forest types to climaticzones of the province,and locally to soils.
Many Commonwealthauthorshave found a physiognomicapproach
best suited for initial study and treatmenton a broad geographicscale.
Definition of formationsby physiognomyin combinationwith environ-
ment has prevailed, rather than the dynamic conceptions of Moss,
Tansley, and Clements.In New Zealand, Cockayne(1921) and Allan
(1926, 1927) in general followed Warming (1909); Dick (1953)
has applied the system of Kiuchler(1949), and Chapman (1954a),
that of Riubel (1930). Wardle (1960), Wraight (1960), Gillham
(1960), and Hamilton and Atkinson (1961) have defined New Zea-
land community-typesby physiognomyand dominance.Chipp (1926)
recommendeduse of a physiognomicapproachfor initial work in the
influentialbook on aims and methodsin the study of vegetation (Tans-
ley and Chipp, 1926). Haman and Wood (1928) recognizedprimarily

physiognomicformations,and types based on physiognomyand dom-
inance, in British Guiana. A physiognomicsystem was developed for
India and Burmaby Champion (1936) and has been applied to Burma
by Edwards (1950). Burtt Davy (1938; Dundas, 1938; Dale, 1939;
Fairbairn, 1939; 1943; Logan, 1946) developed a classificationof
tropicalwoody vegetation-typesinto physiognomicformationsand sug-
gested that edaphically determined, stable communitiesbe treated as
subformations.Richards (1936, 1939) recognized physiognomic-ecol-
ogic forest types; Richardset al. (1939, 1940) recommendeduse of
formations and formation-types,and distinction among climatic, eda-
phic, and biotic formations.
In the New World tropics, physiognomic classificationswere de-
veloped by Barbour (1942), Stehle (1937, 1945-6, 1954), Beard
(1944a, 1944b, 1946, 1949a, 1949b, 1953, 1955), and Allen (1956).
Beard's classificationhas been followed by Fanshawe (1952, 1954),
Asprey and Loveless (1958, Asprey and Robbins, 1953; Loveless and
Asprey, 1957; Loveless, 1960), Taylor (1959), and others. Beard
recognizesrelativelynumerous,narrowlydefined formations,including
many determinedby edaphic, as well as climatic,environment;and he
relatesformationsto one anotherand environmentin "formation-series"
-ecological series in which the formations themselves are artificially
delimited stages. Beardalso defines associationsbroadly;the association
"is a floristicgrouping, being the largestpossible group with consistent
dominantseither of the same or closely allied species" (Beard, 1955).
Boughey (1957) has presented a system of formations developed by
Aubreville for Africa, Fosberg (1961) a general classificationof trop-
ical formation-classesand formations.
The work of Australian ecologists is of special interest, for they
have tested Americanand British ideas against the vegetation of their
continent and developed from their experiencestheir own systems of
classification(Crockerand Wood, 1947; Beadle and Costin, 1952). In
South Australia, Osborn (1915, 1922, 1925; Adamson and Osborn,
1922) recognized community-typesinformally by combinations of
habitat and vegetational characters. Adamson and Osborn (1924)
sought to follow Clements but were led to recognition of stabilized
edaphic subclimaxesby some of the soil effects that have so impressed
Australianecologists.Earlierwork of Wood (1929, 1930) emphasized
climatic effects and sought to treat the vegetation of South Australia
in a basicallyClementsianpattern (Wood, 1937). Wood (1939) later

rejectedthe Clementsiansystem and emphasizedthe preponderantrole
of soils in the determinationof Australian plant communities.The
series of associationsin one climate and on immaturesoils related to
a maturesoil type, and relatedfloristically,form a closely-knitcomplex
which may be called an edaphic complex. Edaphic complexes are the
natural unit of complexity above the association;the complexes may
be grouped into formations characterizedby growth-form and struc-
ture as well as floristics.
The conceptsdeveloped by Wood were applied by Crocker(Crocker
and Skewes, 1941; Baldwin and Crocker,1941; Crocker,1944, 1946a,
1946b), Jessup (1946, 1948, 1951), and others (Murray,1931; Eard-
ley, 1943, Boomsma,1946, 1949; Spechtand Perry,1948; Specht, 1951,
1958). Crocker (Baldwin and Crocker,1941; Crocker,1944, 1946a)
suggested as an additional concept the climatic complex, to be used
where soil type remains relativelyuniform and vegetation changes are
an expressionof climatic factors;Jessup (1951) suggested sociation-
complex for a group of two or more associationswhich occur on unre-
lated soils and show mosaic distributionwithin an area. Many of the
conclusions of the South Australian school were summarizedin the
paper of Crockerand Wood (1947). The association is defined as a
constant association (i.e., grouping together) of dominant species re-
curring in similar habitats; in this the term "dominantspecies" may
refer not only to the canopy but to undergrowthspecies. In practice
these associationsare more broadly defined than this wording might
seem to imply and may be subdividedinto units correspondingto the
fasciationsand consociationsof some other authors.Two bases of sub-
division are suggested by Crockerand Wood: A type is a local change
in the dominantsof the upper stratumof an associationwhich is ac-
companiedby little or no change in the other dominants;a society is
a local change in the dominantsof the lower stratum.The type is re-
garded as a more fundamentalunit than the association;the association
is a collection of types. Within definite climatic limits, associationson
relatedsoil types and with relatedfloristiccompositionare grouped to-
gether as an edaphic complex.
Systemsof formationswhich have influencedother Australianauthors
were developed by Diels (1906) in Western Australia and Prescott
(1929) in South Australia;Prescott'swere later (1931, "major asso-
ciations") applied to the rest of the continent. Patton (1932, 1933,
CLASSIFICATION
OF NATURALCOMMUNITIES 49
1934, 1935, 1942, 1953) in Victoria recognized associationsby both

physiognomyand habitat and (1937) observedthe occurrenceof very
different communitieson different rocks in the same climate. Vegeta-
tion of Western Australiawas describedby Gardner (1944, 1959) in
terms of formations. In a local study Williams (1933) found the
distributionof eucalypt associationsand consociations,as dominance-
types, was determinedby soil and (1948) applied Wood's concept of
edaphic complexes; McArthur (1957) has recognized coastal com-
munity-typesby dominance.In Queensland,Blake (1938; cf. Domin,
1911) classifiedvegetationiinto formationsand broadly defined asso-
ciations characterizedby dominanceand habitat.
Among the authorsworking in New South Wales, Petrie and others
(Petrie, 1925; Brough et al., 1924; McLuckieand Petrie, 1926, 1927a,
1927b; Petrie et al., 1929; Jarrett and Petrie, 1929) followed Clements'
system in general, but with the addition of Tansley's (1920) concep-
tions of climax communities. They recognizedbroadly defined associa-
tions comprising all communities resembling one another in floristic
composition, and divided into consociations according to the distribu-
tions of the dominants(Petrie 1925). Fraserand Vickery (1937, 1938,
1939) recognizedphysiognomic-ecologicformationsand observedthat
in eucalypt forests well defined associationsas dominance-typesap-
peared, but that gradationof undergrowthwith elevation scarcelycor-
respondedwith these. Pidgeon (1937, 1938, 1940, 1941) was led by
difficultiesin applying the successionalapproachof Clements to con-
sider that both climate and soils were responsiblefor the differentiation
of eucalyptassociationsand to prefer the system of Tansley. Pidgeon's
associationswere, however, broadlydefinedin the mannerof Clements.
More narrowly defined associationsas dominance-typeswere used by
Davis (1936, 1941a, 1941b), whose studies of edaphic effects on
vegetationalso led him to reject the Clementsiansystem. Other studies
from New South Wales have been based on informal recognitionof
dominance-types(Collins, 1923, 1924; Davis et al., 1938; Osborn and
Robertson, 1939; Pope, 1943).
The work in New South Wales has led to the systemsof classifica-
tion presentedby Beadle (1948) and Beadle and Costin (1952). The
system has been applied by Moore (1953a, 1953b), by Costin (1957,
1959) to alpine vegetation,and Baur (1957) to rain forests (cf. Webb,
1959). These authors, like Crocker and Wood (1947), reject the
Clementsian system. In Beadle's (1948) classificationof the vegeta-
tion of western New South Wales, formationsand subformations,cor-

responding in general to those of Prescott (1929, 1931) and other
Australian authors, are recognized. The use of type and association
correspondsto that of Crockerand Wood (1947); the type is the basic
unit, defined by dominantspecies, and the associationis an aggregation
of similar types. In the system of classificationand glossaryoffered by
Beadle and Costin (1952), however, the smallerunit (type) is termed
the associationand defined as a climax communityin which the dom-
inant stratum has a qualitatively uniform floristic composition, and
which exhibits a uniform structureas a whole. The associationis named
by the one or more dominants of the tallest stratum, and the term
consociationfor communitieswith a single dominantspecies is conse-
quently unneeded. Associationsare divided into subassociationsdeter-
mined by a variationin the most importantsubordinatestratumof the
association, without significant qualitative change in the dominant
stratum.Associationsare grouped into alliances, groups of floristically
related associationsof similar structure,correspondingto the associa-
tions of Crocker and Wood (1947) and Beadle (1948), and sub-
alliances. Alliances may be grouped into formationsand subformations
as physiognomictypes which reflect similar ecological conditions, but
not climatealone as in the systemof Clements.Associes and other seral
units are recognized,but these terms are not applied to edaphicallyde-
termined communitieswhich are stabilized.
THE AMERICAN TRADITION
Some of the earliest ecological studies in North Americawere based

on zones-the large-scale,transcontinentallife-zones of Merriam(1890,
1892, 1894, 1898, 1899) and the small-scalezones surroundingwater
bodies observed by MacMillan (1896, 1897) and later given succes-
sional interpretation.In the early development of the American dy-
namic approachby Cowles (1899, 1901) the plant society was used
as a unit, derived from Warming's (1895, 1896) Plantesamfundand
defined as a group of plants living together in a common habitat and
subjected to similar life conditions (Cowles, 1899:111). Cowles and
others (Whitford, 1901; Harshberger,1901; Livingston, 1902, 1903;
Brown, 1905; Transeau,1905-6) used the term society quite flexibly,
with little concern for formal classification.During the same period,
however, other American authors (Kearney, 1900, 1901; Ganong,
1903a, 1903b; Harshberger, 1903; Hart and Gleason, 1907) were

among the first to distinguish the formation and the association (or
dominance-type)in their modern senses. These two conceptionswere
adopted, together with the dynamic approachof Cowles and an in-
terpretationof the natural communityas an organism, in the formal
system of classificationdeveloped by Clements (1905, 1916).
In Clements'earliest work (Pound and Clements, 1898, 1900) the
term fornation was used for varied communitiesand without definition
of its rank. In proposing a formal system of nomenclature,Clements
(1902) advocatedas a first principle that the division of formations
must be based on the concept of habitats and offered a considerable
numberof names for formations.Clementsstated as a second principle
that Greek and Latin alone can be used in a scientificsystemand indi-
cated the necessityof rejectingall mixed derivations;there are thus no
derivational hybrids in this group of terms, whatever else a critic
might wish to call some of Clements' words. Clements (1905:292,
1928:119) consideredthat, "As effect and cause, it is inevitable that
the unit of the vegetative covering, the formation, should correspond
to the unit of the earth's surface, the habitat." However, "The final
test of a habitat is an efficientdifferencein one or more of the direct
factors, water content, humidity, and light, by virtue of which the
plant coveringdiffersin structureand species from the areascontiguous
to it." The vegetationunit is causedby the habitatunit, but the habitat
unit is recognized by the vegetation unit. The cause-and-effectphil-
osophy has been criticized as a major fallacy of Clementsianecology
(Egler, 1951; cf. Du Rietz, 1921; Whittaker, 1954b, 1957).
In Clements'monographon plant succession (1916, 1928) his sys-
tem approachedits own climax. The cause-and-effectdoctrine is re-
tained (1928:124); but vegetational development is advanced as a
point of view including and harmonizingthe other three definitions
of the formation (by physiognomy,flora, and habitat), each of which
is inadequateby itself. The formation is a complex organism which
arises, grows, matures, and dies; the climax formation is the adult
organismof which the seral communitiesare but stages of development
(1905, 1916, 1920, 1928:125-6). The formation is thus not an ab-
straction,but "is necessarilyan organic entity, covering a definite area
markedby a climaticclimax" (1928:128). Formationswere subdivided
into regional associationsdefined by their dominantspecies or genera;
associations were divided into consociations characterizedby single
dominantsand these into societies characterizedby subordinatespecies.

These terms applied only to communitiesregardedas climax in Clem-
ents' (monoclimax) sense, and parallel units (associes, consocies, so-
cies) were recognized for developmentalcommunities.The fasciation,
lociation, and a number of other units appeared in the system later
(Clements, 1936, 1949; Weaver and Clements, 1938; Clements and
Shelford, 1939). The full formulation of Clements' viewpoint was
presentedin 1936 in an essayon the natureand structureof the climax.
The Clementsiansystemwas never universallyadopted by American
ecologists, but in the middle period of the Americantradition it was
widely accepted and applied. Three features of the system brought
widespread criticism. The organismic analogy was accepted by some
other influential ecologists (Tansley, 1920, 1935; Phillips, 1934-5;
Carpenter,1939b; Shelford, 1931), but a largernumberof criticshave
rejected it as inappropriateto the interpretationof natural communi-
ties (Gleason, 1917, 1926, 1939; Gams, 1918:457-9; Kylin, 1926;
Negri, 1926, 1927; Friederichs,1930:232, 1958; Meusel, 1940, 1951b;
Vaga, 1940; Schmid, 1940, 1941, 1942, 1955; Ellenberg, 1950b,
1954a, 1954b; Muller, 1958). The unlikely characterof some of the
successionalrelations requiredby the monoclimaxtheory and assumed
by Clements led to criticismby those preferringa polyclimaxinterpre-
tation (Gams, 1918; Du Rietz, 1919, 1921:97; Tansley, 1920; Romell,
1920; Nichols, 1917, 1923; Domin, 1923, etc.; see also Whittaker,
1953); and many have criticizedalso the erection of a formal system
of classificationon the basis of dynamicrelations,many of which are
purely hypothetical (Braun-Blanquet, 1921, 1951a:556; Allorge,
1921-2; Lildi, 1923; Du Rietz, 1924, 1929; Wangerin, 1925; Conard,
1935b; Vaga, 1940; Crocker and Wood, 1947; Beadle and Costin,
1952).
While Clements' ideas influence much currentwork, some features
of his system that were most distinctive and most objectionableto
critics have been generally abandoned.These tendencies are apparent
in the work of Braun (1935a, 1935b, 1938, 1940, 1941, 1942, 1947,
1950, 1956) on the forest vegetation of the eastern United States.
Braun treats climatic relations of the eastern forests through climatic
or regional climaxes and uses the Clementsianassociationand other
units, but with recognitionof the stability of communitiesother than
the climaticclimax and the complexityof species distributionand vege-
tation pattern (Braun, 1950, 1956). Braun indicates that, because no
two dominants are exactly alike in behavior, in competition, in reac-

tion, the componentsof an associationsegregatein all mannerof ways,
producing more or less distinct communities, the association-segregates
(Braun, 1935a, 1950:11). The concept of the association,the associa-
tion-abstract, is built up from the many diverse concrete pieces of
vegetation which form the association-concrete (cf. Nichols, 1923).
Each associationis a more or less artificialunit (Braun, 1950:524; cf.
Gleason, 1926; Cain, 1947; Braun, 1947). Braun thus uses some of
the forms of Clementsianecology with a quite different perspective
on their meaning.
Importantearly dissents from the Clementsiansystem were offered
by Gleason (1917, 1926) and Nichols (1917, 1923). Nichols, whose
views were relatedto those of Moss and other British authors,rejected
the monoclimaxtheoryand approachedthe definitionof units primarily
in terms of habitat.Thus uniformityof habitatwas consideredthe cri-
terion of the association (Nichols, 1917), and formationswere inter-
preted as association-complexes relatedto habitatunits (Nichols, 1917,
1923; Watermann, 1922). Associations resembling one another in
physiognomyand ecological structure,regardlessof their floristiccom-
position, were referred to a common association-type (Nichols, 1917,
1923), correspondingto the formationsof some other authors.Gleason
(1917) considered that phenomena of vegetation depend completely
upon the phenomena of individual plants; the association represents
merelythe coincidenceof certainplant individualsand is not an organic
entity in itself. It is probable that no two habitatshave identical en-
vironments, and that no two species make identical environmental
demands.Furtherdeveloped and clarified,these ideas becamethe basis
of Gleason's (1926, 1927, 1929, 1939) controversial"individualistic
concept of the plant association."
An early study by Shantz (1906) was based on the concepts of
Clements;but the later classificationand map of Americanvegetation
(Shantzand Zon, 1924) and studyof communitiesas indicators(Shantz
and Piemeisel, 1924) were based on non-Clementsianuse of major
physiognomic community-typesand dominance-typesas associations.
Shreve's (1914, 1915, 1927, 1942, 1951) studies of mountain and
desert vegetation were little influenced by the Clementsian system.
Mountain vegetation was variously classifiedaccordingto topographic
situations (Shreve, 1914), physiognomic types as elevation zones
(1915), and community-typesdefinedby physiognomyand dominance
(1927); studies of desert vegetation concernedphysiognomyand en-

vironmentalrelations,floristicand geographicrelations,but not formal
classification (Shreve 1925, 1934, 1942, 1951). In early studies
Shreve (1915) observed the individualistic "dissociation"of species
distributions and (1914) rejected a monoclimax interpretationof
mountain vegetation. The concepts of successionand climax were re-
garded as inappropriateto desertvegetation (Shreve, 1942, 1951); and
American vegetation was mapped by areally predominantor prevaii-
ing, ratherthan theoreticallyclimax, community-types(Shreve, 1917).
Cooper (1913, 1916, 1922, 1926), along with Clementsand Cowles,
made a major contribution to American dynamic ecology. Cooper,
however, had little concern for formal classificationand never ac-
cepted the systemof Clementswith its weaknesses.The classificationhe
applied to the broad-sclerophyllvegetation (Cooper, 1922) used
formation and association in senses unlike those of Clements and re-
lated to the conceptsof Nichols (1917). In later work Cooper (1942)
recognized community-typesinformally by dominance and accepted
the stabilizationof local communitiesas well as of the regional cli-
maxes. His approachthus resemblesthat of the British tradition, and
also of many Americanauthorswho worked without following Clem-
ents' or any other formal system. Oosting (1942) has applied the
Clementsiansystem, but with more careful quantitativetreatmentand
more exacting analysis of successionin relation to site than in many
Clementsianworks. Oosting and others (Oosting and Billings, 1943,
1951; Oosting and Reed, 1952) have studied the Americanspruce-fir
forests as dominance-typesand analyzedthe fir forests of SierraNevada
through undergrowthunions (Oosting and Billings, 1943). Dauben-
mire (1942, 1952, 1953, 1954; cf. Billings, 1949) has used zones
defined by climax associations,and associations(i.e., sociations embrac-
ing all unions superimposedon the same area)-the latter following
Russian authors (Daubenmire, 1952).
Cain (1932, 1934, 1935, 1936b) and Conard (1935a, 1939, 1952,
1954) have led in the introductionof ideas from Europeanphyto-
sociology into the Americantradition (Cain et al., 1937, 1956, 1959;
Cain and Penfound, 1938; Penfound and Watkins, 1937; Penfound
and Howard, 1940; Brown, 1941). While this work is influencedby
Braun-Blanquet,it is not based on Braun-Blanquet'scentral concepts
of diagnostic species. Dansereau (1943, 1945, 1946; Dansereau and
Segadas-Vianna,1952) has followed Braun-Blanquetmore closely, but
has approachedcommunitiesthrough physiognomyinstead of floristic

hierarchiesat levels above the association (Dansereau, 1951, 1957).
Kittredge (1938), without using the Braun-Blanquetsystem itself,
has studied environmentalrelationsof forests through groups of diag-
nostic species. Synusial approacheshave been studied by Cain (1934,
1936a; Cain and Sharp, 1938), Billings (Billings and Drew, 1938;
Oosting and Billings, 1943; Billings, 1945), and Daubenmire (1942,
1952). Following work in North America by Finnish authors (Il-
vessalo, 1929; Kujala, 1945), forest site-types have been studied by
Heimburger (1934), Spilsbury and Smith, (1947) and others. The
Scandinavianapproachthrough the sociation has been applied to arc-
tic vegetationin Alaska by Hanson (1953). Recentwork of Whittaker
(1951, 1954a, 1954b, 1956, 1960) and the Wisconsin school (Curtis,
1955, 1959; Curtis and McIntosh, 1951; Brown and Curtis, 1952; Gil-
bert and Curtis, 1953; Hale, 1955; Culberson,1955; Bray, 1956, 1960,
1961; Bray and Curtis, 1957; Orpurt and Curtis, 1957; Bond, 1957;
Clausen, 1957a; Swindale and Curtis, 1957; McIntosh, 1958; Ward,
1958; Christensenet al., 1959; Habeck, 1959; Beals, 1960; Beals and
Cottam, 1960; Maycockand Curtis, 1960; Dix and Butler, 1960; cf.
Rice and Penfound, 1959; Lindsey et al., 1961) is more concerned
with vegetationalcontinuitiesand quantitativerelationsof communities
to environmentalgradients than with classification.
The Clementsiansystem has been immenselyinfluential in the de-
velopment of British and Americanecology. Much discussion of ter-
restrialnaturalcommunities,especiallyin the two decades, 1916-1935,
concernedthat system and its applicationor the suggestion of limita-
tions and criticismsof it; and some of its major featuresappearin cur-
rent textbooks (Oosting, 1948, 1956; McDougall, 1949). Since the
middle 1930's, however, acceptanceof the Clementsiansystem steadily
declined; the later period of the AmericanTradition has been one of
active experimentationwith other means of classificationand with
quantitativeapproachesto the analysis of natural communities.
OTHER APPROACHES TO CLASSIFICATIONOF ECOSYSTEMS
The ecosystem is the functional system comprising a community

of interactingorganisms-plants, animals, and saprobes-and the en-
vironmentthat affectsthem and is affectedby them. Although the lar-
gest share of classificationsof ecosystemsby ecologists have concerned
plant communities,other aspects of ecosystemscan be classified, not-

ably climate, soils or substrates,and animal communities.Eachof these
has been extensively studied, and some authors have sought through
classificationto interrelatetwo or more of the aspects of ecosystems.
Studies of climate in relation to natural communitiesare too ex-
tensive to be considered here. Major possibilities include the classi-
ficationof climatesto study their relationto vegetation (Koppen, 1900,
1923; Thornthwaite, 1931, 1948, etc.) and the converse study of
formations,as primarilyvegetation units, in relation to climate. Many
authors have used substrate, topographic position, or other charac-
teristicsof habitat to classify communities;and many have termed the
resultingunits "associations."Classificationby communityenvironments
or habitats has perhaps been carried out with most consistency by
Pearse (1926), Villar (1929a; Dansereau, 1952, 1957:130), and El-
ton and Miller (1954).
Classificationof soils is also much too extensive a topic in its own
right to be reviewed here. Classificationof soils into regional types
related both to climate and to vegetational physiognomy, following
pioneer Russian work, is familiar. Some classificationsof vegetation
use soil characteristicsas criteriaof community-typesand their relation-
ships. Within a region, soil moistureand soil parent-material,especially,
may be bases for edaphic classification.The recognition of "postcli-
maxes" in wetter and "preclimaxes"in drier environmentsthan those
of the "climatic climax" (Clements, 1936; Braun, 1950) represents
such a quasi-dynamicclassificationrelated to soil moisture. Other ap-
plications are the recognitionof site-typesand classes of site-typesby
relations of undergrowth to soil moisture (Cajander, 1949; Kujala,
1945), vegetation catenas correspondingto edaphic catenas (Milne,
1947; Morison et al., 1948), and major groupings of temperateand
tropical forests in relation to the moisturegradient that may be inter-
preted as orders in the sense of Braun-Blanquet(Mangenot et al.,
1948).
Relations between soil parent-materialand vegetation have led to
recognitionof edaphic climaxes or subclimaxes,or paraclimaxes(sensu
Tiixen, 1933, not Villar, 1929a), as a quasi-dynamicclassification.Pro-
nounced effects of soils on vegetation in Australia led Wood (1939;
Crockerand Wood, 1947) to suggest grouping of vegetationtypes into
edaphiccomplexes.Hughes (1949) has classifiedsoils within elevation
zones into suites, each related to a specificparent rock. Each soil suite
has a specific and characteristicrange of plant communities,and the

whole can be looked upon as a soil suite-plant communitycomplex.
Responses of vegetation to serpentine soils are especially evident;
Soc'ava(1927), Pichi-Sermolli (1948), and Whittaker (1960) have
treated communities as groupings or ecological series of serpentine,
intermediate,and typical soils. Such grouping by soil materialstends to
unite some closely related communities,but also to cut across other
classificationsbased on vegetation itself. Thus the communitiesof the
EuropeanSeslerion coeruleaeare widespreadboth on serpentinesand
in other edaphicallydistinctivesituations (Zlatnik, 1928b), and Rune
(1953) observesthat serpentinecommunitiesof the north of Sweden
do not fit naturallyinto a single alliance. Consequently,it is possible
also to subdividea vegetationunit on the basisof soil material,as in the
climax-swarmof Tiuxenand Diemont (1937; Diemont, 1938; Tiixen,
1954a).
Recognition of the interrelatednessof communityand environment
in the ecosystemhas led many authorsto seek classificationsbased on
multi-factoralor landscape units. Passarge (1921, 1927) considered
that "areas"(Riume) of the earth'ssurfaceare definedby the distribu-
tion of certainobjectsand phenomena-climatic provinces,meteorologi-
cal systems,water bodies and land areas,geological substrates,political
units, etc. The study of landscapesis the study of the arrangementand
interpenetrationof these areas and their fusion into the unified object
of the landscape.Since the boundariesof individual areas do not fall
together, sharp bounding of landscapes cannot be assumed; but the
most sharplyboundedareasmay be chosen for definitionof boundaries.
Often a single outstanding area characterizesand serves to name the
landscape. Units representing subdivisions and groupings of land-
scapes were recognized; and in the treatment of plant communities
(Passarge, 1929) regional communities and local communitieswere
distinguished, paralleling the distinction by ecologists of prevailing
communities (or "climaticclimaxes') and local or edaphic communi-
ties. The study of landscapeshas been of extensive interest to both
geographersand ecologists (Regel, 1949; Troll, 1950).
Markus (1925a, 1925b, 1926, 1929, 1930) designated with the
word Naturkomplexthe whole of the phenomenaand things that are
localizedin a certainpart of the earth'ssurfaceand causallyinterrelated
with one another.The aspectsof the nature-complexwere recognizedas
elements, and parts of these as members.He recognized"species"and
"genera" of complexes, the latter correspondingto the alliance or

Verband of phytosociologists;and a group of nature-complexesthat
occurredtogether he called a landscape-complex,correspondingto the
association-complexesof phytosociologists(Markus, 1930). In the dy-
namics of complexes he consideredthat membersof nature-complexes
do not changewith equal rapidity,and the displacementof one complex
by another may be recognized in the presence in the former of an
extraneousor foreign member from the latter (Markus, 1926, 1929).
Schmid (1922, 1935, 1936a, 1940, 1941, 1942, 1949, 1950, 1952,
1955, 1956b, 1961) developed an approach of unique conceptual
complexity. Wide-ranging chorological units, based on species distri-
butions, are designatedVegetationsgiir/el;within these Hauptcoenosen
may be defined by vicariousand localized species. Definition of these
major units is, in ideal, floristic;but they correspondin general to re-
gional vegetation types that may be recognized by physiognomyand
dominance.From the flora of a regional unit, its Artengarnitur,species
are distributedinto varied, intergrading,local Biocoenosen,not classi-
fiable in any real sense, but among which abstracttypes or models may
be recognized(Schmid, 1942, 1950, 1952; Ehrendorfer,1954). Among
the species themselves, abstracttypes (Repr.isentationstypen, Korrela-
tionstypen, etc.), may be recognized by similarityin the many kinds of
ecological relations through which each species may be characterized
(Schmid, 1950, 1952; Heuer, 1949; Schwarz, 1955); these many-sided
ecological forms are preferred to the traditionallife-forms. The spec-
trum of representation-types contributesto the characterizationof the
biocenose-type or -model and permits its comparisonwith other bioce-
noses (Schmid, 1950, 1952). Schmid'ssystemhas been applied both to
intensive analysisof forest communities(Schmid, 1936b, Heuer, 1949;
Schwarz, 1955; Saxer, 1955) and to broad geographic interpretation
of Europeanvegetationin terms of Vegetationsguirtel and their historic
relations (Schmid, 1961).
In Americanecology, Dice (1938, 1943, 1952; Dice and Blossom,
1937; Goldman and Moore, 1946) has developed a system of biotic
provinces as geographic areas characterizedby climate, physiography,
soil, and natural communities.Unlike many other units of classifica-
tion, biotic provinces on land are geographicallycontinuous.Their oc-
currenceand distributionis based ultimatelyon climate;associatedwith
general uniformityof climate,however, is general uniformityof physi-
ographyand soil types. Within a region, characteristictypes of ecologi-
cal communitiesdevelop, and the biotic province is thus characterized

by the occurrenceof one or more importantecological associationsthat
differ, at least in relative area covered, from those of adjacentprov-
inces. Within each area, accordingto its peculiarecological conditions,
plants and animals develop certain adaptationsand differentiateinto
local species and subspecies;the biotic provinces are also centers of
specificdifferentiation.It is as faunal or bio-geographicunits, especially
for mammaliandistributionsand subspecificpatterns,that biotic prov-
inces have been most used. Some biotic provinces can be subdivided
into biotic districts,and in mountainousregions can be verticallysub-
divided into life-belts or elevation zones.
In the RussianTradition the landscapeconceptionhas been applied
to forestsby Morosow (1928), Paczoski(1930b), Kruedener(1926b),
and Sukatschew(1944, 1954). The forest site-typesof Fennoscandian
authors and the forest classificationsof Zlatnik (1961) and Hills
(1960b) are also multi-factoralunits in their joint considerationof
properties of environmentsand communities.Many phytosociologists
have developed interpretationsthat depart from the single-factorclas-
sificationsof the schools of Uppsala and Braun-Blanquettowardmulti-
factoralclassification;e.g., the many-sidedinterpretationsof Gradmann
(1909, 1941) and Negri (1914, 1927), the topographicapproachof
Gams (1918, 1927) and Bolleter (1921) and the geomorphic-edaphic
approach of Scharfetter(1921, 1932), the use of habitats and life-
forms, as well as species composition, for classificationby Raunkiaer
(1918, 1934), Wangerin (1925), and Lippmaa(1931, 1933a, 1935b),
the comparative-typological approachof Meusel (1939a, 1939b, 1940,
1943a, 1951b, 1954b, 1954c), the interpretationof vegetationin rela-
tion to landscapesby Regel (1939, 1949, 1959), the intensive analysis
of environmental-community relationsby Liudi(1948) and Kuhnholtz-
Lordat (1952), the biochore approachby Etter (1954), and the eco-
systemicclassificationsof S0rensen (1937) and Sjbrs (1955).
Implicit in the work of all these authorsis some recognitionof the
functional whole formed by communityand environment.They have
often workedindependentlyof one anotherand the conceptof holocene
(Friederichs 1927, 1930) or ecosystem (Tansley 1935); but all have
sought interpretationand classificationbased on both communityand
environment,and not on a single propertyof vegetation.The ideal of
this multi-factoralor landscape approach to classificationis an inte-
grative conceptionthat takes into accountall propertiesof ecosystems,
and evaluatesthem in relationto one anotherto recognizethese group-

ings of ecosystemsbased on most criticalfactors and with the broadest
possible significance. The multi-factoralapproach to classificationis
one major possibility that has been applied, like the physiognomic
approach,by authors of otherwise varied viewpoints in the different
traditions.
One other approachtaken in varied forms by authors of different
traditions is that through zones. The concept of elevation zones was
applied to Europeanmountainsin the nineteenth centuryby Wahlen-
berg (1813), Schouw (1823), Boissier (1839-45), Sendtner (1854),
Flahault (1893), and others (Braun-Blanquet1951a:525). Since then
manyphytosociologistshave recognizedclimaticand distributionalzones
in Europeand altitudinalzones in Europeanand North African moun-
tains. Zones of tropical mountains have been considered by Steenis
(1935), Troll (1953, 1958a, 1961), Hedberg (1951), Salt (1954),
Keay (1955), Jackson(1956), Wace and Holdgate (1958) and others.
In North America, however, the use of zones as the primary basis
of the study of natural communitieshas had its most extensive ap-
plication (Merriam, 1890, 1892, 1894, 1898, 1899). Merriam first
recognized zones in the western mountains; later he extended the
zones across the North American continent and attempted a state-
ment of the temperaturefactors controlling their distribution.Many
have criticized the system for its theoretic basis and the transconti-
nental extension of zones through diversevegetationtypes and floristic
and faunistic areas (Sumner, 1915; Dice, 1916, 1923, 1952; Ruth-
ven, 1920; Livingston and Shreve, 1921; Kendeigh, 1932; Shelford,
1932, 1945; Pitelka, 1941; Daubenmire, 1938, 1946; Odum, 1945;
Hayward, 1945; Allee et al., 1949). The system has, nevertheless,
been used in a great many field studies and is still found useful
by some authors, especially in western mountains (Hall, 1902;
Bailey, 1905, 1926, 1936; Cary, 1911, 1917; Grinnell et al., 1913,
1924, 1930; Grinnell, 1914, 1935; Hall and Grinnell, 1919; Smiley,
1921; Jones, 1936, 1938; Borell and Bryant, 1942; Johnson et al.,
1948; Goldman, 1951; Miller, 1951; Muesebeckand Krombein,1952;
Farner, 1952). Life-zones have also been studied by Russian authors
(Filipjev 1929, Carpenter 1939a), but these units are more related
to the Russiansoil zones than to Merriam'szones.
Zones have also been found an effective basis for classificationand
correlation of community-typesunder conditions very different from

those of mountains-in the marinelittoral. Doty (1957) has reviewed
the history of study of zones of rocky shore organisms.A system of
zonesdefinedby majortypesof organisms,developedby the Stephensons,
has been widely applied and discussed (Stephenson and Stephenson,
1949, 1950, 1951, 1954a, 1954b, 1961; Guiler, 1950, 1952, 1953,
1960; Lewis, 1955; Chapmanand Trevarthen,1953; Bennettand Pope,
1953; Endean et al., 1956; Lawson, 1956; Chapman,1957; MacNae,
1957; cf. Rigg and Miller, 1949; Dellow, 1950; Caspers,1951; Wom-
ersley and Edmonds, 1952, 1958; Hartog, 1955; Moore, 1958). Dahl
(1952-3; cf. Hedgpeth, 1957) has suggesteda world-widezonationfor
sandy beachescomparableto the Stephensons'for rockyshores.
Zones may be definedby variousmeans. Some definitionsare floristic
or faunistic; but probablymost authorshave recognizedzones on land
by prevailing physiogoniomyor prevailing dominantspecies of vegeta-
tion, and secondarily observed correlations of species distributions
with these zones. The Stephensons'zones are also in a sense physiog-
nomic as they reflectthe morphologyof littoralcommunitiesdetermined
by predominant ecologic-taxonomictypes of organisms. Parallel ar-
rangementof physiognomiczones may be observedin widely different
areas-and even around the world; parallel arrangementof zones de-
fined by dominant species may often be observedover a more limited
range. The real identity or equivalenceof zones in differentareasmay
be questionable,however. Some authors (Merriam, 1894; Emberger,
1936; Zotov, 1938) have sought climatic definitionsof zones, whereas
others (Graham, 1937; Gaussen, 1938-9; Daubenmire, 1946) have
advocateddefining zones by the dominant species or prevailing com-
munity-typesthat characterizethem in a given mountain range.
The zonal approachhas been most successfulin a limited range of
ecological conditions-primarily in mountainsand especially those of
more arid regions, and in shore communities.In the mountains,mois-
ture conditions and other climatic factors, as well as temperature,are
strongly correlatedwith elevation; in the littoral many other environ-
mental factors are correlatedwith tide-level and exposure. In those
circumstanceswhere a single complex-gradient,of many correlated
environmentalfactors, is of such paramountimportancethat all other
gradientsmay be treatedas secondaryto it, the zonal approachis most
appropriate.
ANIMAL COMMUNITIES
The varietyin the classificationsof animalcommunitiesalmostequals

that in the classificationsof plant communities.Merriam (1892) char-
acterizedhis life-zones by vertebratefaunal composition,as well as by
vegetational characteristics.Enderlein (1908) described the faunistic
compositionof varioushabitats;Dahl (1908) emphasizedthat for each
biocenose he distinguished,he could name at least one characteristic
animal species. Adams (1909) felt that an avian formation might, in
general terms, be consideredthe analogueof a vegetationalformation,
although this did not necessarilyimply that they had the same boun-
daries. Shelford (1912, 1913) recognized many communitiesas for-
mations and associations,some characterizedby habitat,others by vege-
tation, as well as by animals. Vestal (1914:444) stated that "Upon
investigation,it begins to appearthat plant and animal assemblagesare
coextensiveparts of a biotic association,composed of both plants and
animals."Grinnell (1914; Grinnell and Swarth,1913) applied the term
"association"to community-typescorrespondingin general to the for-
mations of plant ecologists, but also based on vertebrateanimals,thus
anticipatingthe concept of "biome."
Petersen (1914, 1915, 1918) characterizedseveral sea bottom com-
munities by varyingconmbinations (3 to 6 species) from a list of major
echinodermand mollusc species. Molander (1928) recognizedbottom
communitieson a similar basis, defining them as regularly recurring
combinationsof certaintype animals,as a rule stronglyrepresentednu-
merically.Other authorshave based their work on Petersen'sconcepts
of marine bottom communities;such work is reviewed with a sum-
mary of marine level-bottom communitiesof the world by Thorson
(1957). Jones (1950) discussed the communityconcept for bottom
organismsand suggesteda scheme of community-typesbased primarily
on environments,but designated 'associations."Newell et al. (1959)
approachedthe benthos by two concepts,habitat-community,a natural
associationof organismsset aside accordingto certain features of en-
vironment, and organism-community,a regularly recurring combina-
tion of certaintypes of organisms.Appropriatelydefined habitatcom-
munities show close correspondencewith organism communities;but
the correlation is never exact, and sometimes there is little or no
evident correlation. Gislen (1943, 1944) classified communities ot
the marine littoral into associationscharacterizedby dominant species
and formations characterizedby animal growth-forms. MacGinitie

(1939) and Molinier (1960) have discussedthe classificationof lit-
toral communities,and the extensive work on communitiesof rocky
shores and sandy beaches has been recentlyreviewed by Doty (1957)
and Hedgpeth (1957). As indicated above, much of this work has
treated littoral communitiesin terms of zones.
The search by American ecologists for a unit appropriateto both
plants and animals led to the biotic formation or biome as a basis for
the study of "bio-ecology"(Clements and Shelford, 1939); the biome
concepthas had wide acceptanceamong Americanecologists and some
scientists abroad (Shelford, 1931, 1945; Shelford and Olson, 1935;
Carpenter,1936, 1939b, 1940; Pitelka, 1941; Odum, 1945; Hayward,
1945, 1948; Fautin, 1946; Kendeigh, 1948, 1954, 1961; Allee et al.,
1949; Woodbury, 1954; Fichter, 1954; Tischler, 1951; Mann, 1951;
Esteves de Sousa, 1953). Terrestrialbiomes are based upon vegetation
physiognomyand correspondin general to plant formations,but it is
felt that they can be characterizedalso by vertebrateanimals. In ap-
plication to marine and littoral communities, formations or biomes
have been characterizedby growth-forms or taxonomic groupings of
dominant animals (Shelford and Towler, 1925; Newcombe, 1935;
Clements and Shelford, 1939; Dellow, 1950; Marshall, 1953). Allee
et al. (1949) have recognized world-wide biome-typescorresponding
to major formation-types,and reflecting convergencein ways of life
of animals that parallels the convergenceof vegetationalphysiognomy
in responseto similar climateson differentcontinents.Tischler (1951,
1955) has described communitiesof the world through eight major
biome-typesor "landscape-types."
Some American authors (e.g., Cole, 1940; Rasmussen,1941; Dan-
sereau,1945; Barnes, 1953; Friauf, 1953) have sought to defineanimal
communities that correspondto the associations of plant ecologists.
Kendeigh (1948, 1954, 1961) studied distributionof birds as an ap-
proach to recognition of biotic associations.The principal correlation
of bird distributionhe found to be with vegetationalphysiognomy,not
with the plant associations.Distributionalgroupingsof birds were used
to characterizebiotic units of larger scale than plant associations;these
units were termed biociationswhen climax, biocies when successional.
Characterizationof terrestrialbiomes by invertebrateanimals has been
scarcely attempted, although contrasts in the invertebratefauna of
biomes were shown by Blake (1926), Hayward (1945), and Dirks-
Edmunds (1947). Whittaker (1952) found evidence of "individual-

istic" distribution of insect species (suggested also by Bodenheimer,
1958:181) and consideredthat the physiognomicdistinction between
deciduousand coniferousforests cut acrossa naturalgroupingof foliage
insect communitiesin his study area. Bond (1957) and Beals (1960)
found bird species differently distributedand bird communitiescon-
tinuous through the forest communitiesstudied.
Most Europeanstudentsof "biocenotics,"as the systematicsof biotic
communities (Dahl, 1908; Thienemann, 1921, 1939; Prenant, 1934;
Gisin, 1949, 1951; Schwenke, 1953), have had as their central unit
the biocenose (Mobius, 1877; Gams, 1918; Schmid, 1922; Reswoy,
1924; Friederichs,1927, 1930; Hesse et al., 1937; Gislen, 1943; Rab-
eler, 1947; MdrzerBruijns, 1950; Grasse,1950; Balogh, 1958), a much
narrowerunit than the biome. In northernEurope, Krogerus (1932)
regardedthe biocenose as equivalentto the associationand defined the
zoocenose as a self-regulating population of animals, united through
a species-groupcharacterizedby the sociologicalaffinityof its members.
Palmgren (1928) showed a trend in density of bird populations re-
lated to an ecologicalseries of site-types,and Renkonen (1938) showed
limited correspondenceof beetle community-typesto forest and bog
site-types.Brundin (1934; see also Renkonen, 1944) sought to apply
the synusialapproachof Gams (1918) to northernbeetle communities.
Brundin (1934) felt that the units of plant communitiesdeveloped by
Du Rietz (1932) and other Scandinavianphytosociologistsprovided a
good basis for investigationof beetle communities,but found that some
beetle community-typescorrespondedto sociations, others to higher
units. Kontkanen (1937) comparedthe insect faunas of a series of
meadow types.
Renkonen (1938, 1944, 1949), Agrell (1941, 1945b), Forss-
lund (1945), and Kontkanen (1950a) approachedclassificationof
animal communitiesthrough measurementof percentagesimilarityof
samples,the approachof Kulczyn'ski(1928). Renkonen (1944) found
that the method led in generalto community-typescharacterizedby their
major or dominantspecies. Forsslund (1945) related his animal com-
munity-typesto the Swedish site-types of Malmstrom (1926). Back-
lund (1945) used a similar quantitativetechniqueto study association
of species in samples and suggested the term zoome (Hesse et al.,
1937) for all the animals belonging to a certain taxonomic group
within a given community.Hammer (1944) followed the quantitative
approachof Krogerus (1932) and concludedthat the Greenlandmite

and collembolan distributions were determined directly by moisture
factors and showed little relation to plant communities. Krogerus
(1948) showed that lake-shorebeetles were distributedaccordingto
moistureconditions in zones parallelingthe water'sedge, althoughthe
data suggest continuityof the zones. Weis-Fogh (1948) studiedDanish
mites and collembolansin a moisture-gradienttransectthrough three
sociationsand found gradual change in relative importanceof species
and major groups, rather than clear correspondencewith the plant
sociations. Tuomikoski (1948) applied his techniques of correlation
analysis (Tuomikoski, 1942) to the data of Renkonen (1944) and
developed a somewhat different, but related grouping of beetle com-
munity samples (Renkonen, 1949). Brink and Wingstrand (1949)
classified insect community samples into ecologic-physiognomic"bio-
type categories"and studied the relations of these through coefficients
of community.Two of their biotypes were little related to the others.
but the remainingsix formed a continuoussequencefrom birch woods
upward to high-mountaincommunities.Several methods of analysis-
coefficientof community,percentagesimilarity,species-association,con-
stancy,and fidelity-were applied and interrelatedin the classification
of leafhoppercommunitiesby Kontkanen(1950a), who suggestedthat
his leafhopper communitiesextended through several plant communi-
ties. Schwenke (1953) characterizedthe animal communitiesof forest
site-typesby major or dominantspecies, but sought also to relatethem
to units of the Braun-Blanquetsystem.
Among authorsin central and southernEurope, Grasse (1929) has
doubted the appropriatenessof the associationconcept and phytosocio-
logical techniques to animal communities. Rabeler (1937a, 1937b,
1947) felt that the conceptof animal character-species(Charaktertiere,
Leitforme) linked the study of animal communitiesto that of plant
communitiesas approachedby the French-Swissphytosociologists,and
consideredthat an animal and a plant associationtogether formed the
biocenose (Rabeler, 1947). A series of studiesby Rabeler(1950, 1951,
1952, 1953, 1955) have applied the Braun-Blanquetsystem. Frei-Sul-
zer (1941) applied Schmid's (1922, 1941, 1942) conceptionsof bio-
cenoses to animal communities.Westhoff and Westhoff-de Joncheere
(1942) observed differenttypes and degrees of correlationof ant spe-
cies with vegetation and showed limited correspondenceof ant com-
munity-typeswith vegetation types. Gisin (1943) recognizedfive col-
lembolan life-forms in relation to the soil surface, in an approachre-

sembling the life-form system of Raunkiaer.Gisin's is thus a synusial
approachto animal communities;but he (1947, 1951) also emphasizes
vicariant species groupings, groups of diagnosticspecies which replace
one another in different community-types.Gisin further indicates the
correlationof his collembolancommunity-typeswith the plant associa-
tions recognized by Braun-Blanquetand Jenny (1926) and Liidi
(1948). Kiihnelt (1943a, 1943b) observedthat one seldom finds, ex-
cept in a limited area,good correspondencein distributionof an animal
species with either a plant species or a plant association,and suggested
definition of animal community-typesby relativelyrestrictedspecies oL
Leitformenand by characteristicspecies combinations,as in the Braun-
Blanquetschool. Kiihnelt (1950, 1951a, 1951b, 1953) also recognized
the existence of "biologicalforms" and synusialgroupings among ani-
mals. Franz (1943) applied the qualitativecriteriaof Braun-Blanquet
and found that sometimesseveralplant communitiesmight correspond
to a single animal community,at other times several animal communi-
ties occurredwithin the limits of one plant community;a similar conl-
clusion was reachedby Rioux (1958). Franz (1950) consideredthat
qualitative approaches,and especially the use of differential-species,
were appropriateto the study of biocenotics,while quantitativemeth-
ods were more appropriateto study of the ecology, or environmental
relations, of animal communities.
Tischler (1948, 1951) suggeststhat the qualitativecriteriaof charac-
ter-speciesand differential-speciesbe used for the characterizationof
biocenoses,life-forms for the characterizationof biome-types.Tischler
observes that, though the boundariesof animal communities should
coincidewith those of plant communities,a single biocenosecorresponds
in general to several associations;the order (Ordnung) of the plant
sociologists best correspondsto the biocenose. The term associationis
restrictedby Tischler to units based on a single group of organisms-
as of phanerogams,birds, collembolans,beetles, etc. Studiesof animals
of agriculturalcoinmunitiesby Tischler (1952) and Marchand(1953)
have been criticizedby the more orthodoxRabeler (Rabelerand Tiixen,
1955) for their departuresfrom the Braun-Blanquetsystem but are
related in approachto that system, especiallyin the applicationof dif-
ferential-species (Marchand, 1953). Tischler (1950) has also con-
sidered synusial approachesand suggested recognition of convergent
community-typesin similar habitatsof differentparts of the world as
isobiocenoses (Tischler, 1950; Caspers, 1950). Strenzke (1950) ap-

proachedterrestrialchironomidcommunitiesthrough the synusialcon-
ceptions of Gams (1918), Du Rietz (1932), and Gisin (1943), and
sought to follow the pattern of classificationof Du Rietz (1932). In
a later study of mite communities,Strenzke (1951, 1952) followed
more closely the approachof Gisin and consideredthat the characteri-
zation of animal communitiescan follow the methods used in plant
sociology-i.e., the approach through diagnostic species. Marlier
(1951) has applied Gisin's conceptions to stream communities;
Schm6ltzer (1953) has indicated that mapping of biocenoses should
be based on characteristicspecies combinations,not the ranges of spe-
cies. Mdrzer Bruijns (1950) preferred the Braun-Blanquetapproach
for relatingmolluscsto plant communities,and interpretedthe bioceno-
sis as the smallestindependentcommunityof living beings in biological
equilibriumin the number and kind of its constituentspecies and the
numbers of individuals by which these are represented,and which
occupiesan ecologicallyhomogeneousarea. Hagen (1952) studied the
mollusc communitiesof forests and found only partial correspondence
of these animal communitieswith phytosociologicalunits. Quezel and
Verdier (1953) classified beetle communitiesinto associationscorres-
ponding to plant associations,and into alliances, orders, and classes
defined by diagnosticbeetle species; Galoux (1953) has studied rela-
tions of soil arthropodsto units of the system of Braun-Blanquet.The
British author Macfadyen (1954) consideredthe various Continental
approachesand found that the collembolanand mite community-types
of Jan Mayen were best defined by diagnostic species; these animal
communitiescorrelatedwell with the vegetationalunits (habitat- and
dominance-types)of Russell and Wellington (1940).
These papers (see also Balogh, 1958) suggest a scatteringof ideas
when their authorshave found no clear path to effective classification
of animal communities.Most southern authors are alike in their em-
phasis on the qualitative approach to animal communities through
diagnostic species, and their rejection of the approachthrough domi-
nance characteristicof northern and English-languageauthors.North-
ern authors (Agrell, 1948b; Renkonen, 1949) have criticizedthis ap-
proach as sacrificingmuch of the informationwhich an animal com-
munity sample offers as a basis of classification.The northernauthors
almost without exception use quantitativecriteria- dominance, con-
stancy, coefficientof community, and percentage similarity-as bases
of classification,although the outlook of Tuomikoski (1948) is closer

to that of southern authorsthan of his northern colleagues.The dif-
ferencein perspectiveof the Northernand SouthernTraditionsis hardly
less markedin the field of biocenoticsthan in that of vegetationstudy;
and the dispute over qualitativeversus quantitativemethods (Agrell,
1948a, 1948b; Gisin, 1948; Renkonen, 1949; Schwenke, 1953) is
suggestive of the earlier war of the phytosociologists.The contrastin
perspectivein the study of animal communitiesis no doubt determined
in part by the same contrastof the poorer fauna and flora of Scandi-
navia with the richer of centralEurope, that has affectedthe study of
vegetation (Franz, 1950).
Certaincontrastsbetween Europeanand Americanwork may also be
observed, and these in general parallel the differencein approachto
plant communities.Europeanstudents of biocenoticshave dealt with
narrowly defined community-typesdistinguishedwithin a given area;
Americananimal ecologists have for the most part worked with such
broadly defined units of extensive geographicareas as the biome, life-
zone, and biotic province. European students have worked predomi-
nantly with invertebrateanimals; American authors have concerned
themselvesmore with the often highly mobile and wide-rangingverte-
brates. The concept of life-form, applicationof which to animals has
been advocatedby Gams (1918), Friederichs(1930), Remane (1943,
1952), Rustamov (1955), and others, and the recognitionof synusial
communities,have concernedmany of the Europeanauthors.American
authors have been concernedmore with the growth-forms of plants
and relationsof animals to physiognomyof vegetation.
Animal ecologists, as scatteredamong the traditionsas Beklemishev
(1931), Brundin (1934) and Kontkanen (1950a), Tischler (1948)
and Bruns (1950), and Kendeigh (1948, 1954), have observed that
the most reasonableunits for animal communitiesmay be of larger
extent than the plant associations.The work with animal communities
revieweddoes not supportthe assumptionof any simple correspondence
between animal communitiesand any traditionalunit of plant ecology.
Animal ecologists have applied classificatoryunits as varied as the
biome, life-zone, biotic province, association and higher units of
Braun-Blanquet,sociation,site-type,dominance-type,habitat-type,union
or synusia,and biocenosein the sense of Schmid; animal communities
have been found by different authors to correspond, more or less
loosely, to all the major vegetation units. Miller (1951) analyzed
bird distributionsin relation to three major American approaches-

biomes, life-zones, and biotic provinces-and consideredthat no one
of these was a completely satisfactorysystem, but each expressed cer-
tain truths or generalizationsabout distributionalrelations. The com-
plex distributionalpatterns of species should not be expected to con-
form to any single system (Miller, 1951; cf. Whittaker,1952; Tischler,
1950); and animal communitiesare subject to rather flexible inter-
pretationand variedclassificationaccordingto the outlook and interests
of the classifier.
MAJOR UNITS OF CLASSIFICATION
Two perspectiveson the historyreviewedare possible. One considers

that a particularnatural unit is inherent in vegetation, and that the
problem is to discover, clarify, and gain agreementon this unit. Seen
from this perspective, the history of ecological units is a history of
progressiveapproachto the "correct"conception.This perspectivemay
come easily to those most involved in the viewpointof a school and who
advocategeneral adoption of a unit as they themselves define it; such
an outlook appears in accounts of the history of units by Clements
(1916, 1928:117-125), Du Rietz (1921), and Braun-Blanquet(1951a,
1951b).
From another perspective, natural communities are complex and
merging phenomena, different properties of which may with equal
justificationbe used as bases of classification.There may be no single
correct unit or approach;various approachesmay be (and, perhaps,
should be) used to clarify different aspects of the complex relations
of communitiesto environmentand one another. Many units may be
equally "fundamental"(Du Rietz, 1935, 1936) and of differentuse-
fulness under differing circumstances.Seen from this perspective,the
historyof ecologicalunits may be regardedas a historyof differentiation
and clarificationof the various units with broadestusefulness.
A noteworthyfeatureof the historyof ecologicalunits is the manner
in which they have progressivelyemerged and differentiatedfrom one
another.After the concept of vegetationunits appeared,the formation
and associationfirst becamedistinctfrom each other, then variousother
conceptions became distinct from these. From the great diversity of
termsand conceptionsthat have resulted,certainunits may be regarded
as of most general importance,as conceptsexpressedby many authors
and aroundwhich the individuallydistinctiveviews of manyothersmay

be grouped. A listing of these major units cannot include all that are
useful, nor should it be assumedthat the units form a hierarchyas of-
ferredby Cajander(1922), Braun-Blanquet(1928a, 195la), Du Rietz
(1930a, 1932, 1949), Lippmaa (1933b), Raunkiaer (1934), Suk-
atschew (1935), Clements (1936), Tansley (1939), Socava (1944),
Crockerand Wood (1947), Beadle and Costin (1952), Bocher (1954)
and Dokhman (1960). The following are, however, some units of
widest utility, which have found places in most or all of the regional
traditions:
A. Major Vegetation Units
1) The formation-type(biome-type),a grouping of communi-
ties throughoutthe world of similar physiognomyand related en-
vironmentalconditions.
2) The formation (biome), a grouping of communitiesof simi-
lar physiognomy and related environmentalconditions within a
single continentor region.
3) The associations, various units defined by species composi-
tion.
4) The sitc-type, a grouping of communitiesby site and similar
or dynamicallyrelated vegetation--especiallyas recognizedby the
lower strataof forests.
5) The union, a unit comprisingone or more species of similar
life-form or of a particularstratum.
B. Uses of "Associations."The term association and the conceptions
to which it has been applied have causedthe greatestdissension.At
least six conceptsmay be distinguishedamong applicationsof this
term:
1) The floristicassociation in the sense of Braun-Blanquet,de-
fined primarilyby character-speciesand bringing together stands
that contain some of these character-species.This, following the
Congress of 1935, has most general acceptanceas correctusage
for the term association.
2) The sociation, defined by stratalstructure,and bringing to-
gether stands that have the same stratal dominants. The term
sociation is now generally accepted outside Russia for this unit
(the northern, Uppsala, or Scandinavianassociation,micro-asso-
ciation, Russian association).
3) The dominance-type,defined by one or more dominantspe-

cies, and bringing together stands in which these species are
dominant in the uppermoststratumif there is stratal differentia-
tion (or, sometimes, in a lower stratum of primaryemphasis).
Dominance-typeshave been termed associationsby many authors;
and the term consociationhas been used for a grouping charac-
terized by a single dominant species instead of two or more. No
generally accepted term now exists, and dominance-typewill be
used here.
4) The regional vegetation type, defined by prevalence of a
given climax vegetation type in a geographicregion or elevation
zone; this vegetationtype must: (1) be naturalor climax, though
not the only climax in the region, (2) prevail, in the sense that
it occupies the majority of sites bearing climax vegetation and
therefore characterizesthe region and expressesthe adaptationof
its vegetation to general climate, and (3) be less extensive than
a formationand definedby dominantspecies or some characteristic
other than physiognomy.The concept is familiarin the climax as-
sociation of Clements (1916, 1928, 1936), and some American
authorsstill use the term associationin this sense (Braun, 1950).
It is recognized also in the school of Braun-Blanquet(with the
concepts Schlussgesellschaft,Klimax-Komplex, and Klimax-Ge-
biet); and more or less relatedconceptsappearamong many many
authors-Hauptcoenose, Vegetationsgfirtel,and regional phyto-
cenose (Schmid, 1922, 1940, 1942, 1950, 1961), regionale
Hauptassoziation (Nordhagen, 1928), vegetation region (Du
Rietz, 1930a, 1932), conclimaxand climax domain (Villar, 1929a,
1929b; Bolos, 1954c), climax area, region, or zone (Gaussen,
1938-9; Mathon, 1949; Zangheri, 1954) and climax complex
(Gaussen, 1951), zonal community(Alechin, 1926; Walter, 1943,
1954; Ellenberg, 1956), regional community (Schlenker, 1939),
climax association (Tansley, 1939), zone (Daubenmire, 1942;
Billings, 1949), Leittyp (Kujala, 1945), Wuchsgebiet (Knapp,
1958), community-complex(Krause, 1952), prevailing climax
type (Whittaker, 1956), and regional climax (Braun, 1956).
Becauseof differentpossibilitiesin the definition of climaxes and
of vegetation types, these conceptions are most varied in detail;
but they have a common center in recognitionof vegetation types
that prevail in and characterizeregions. No generally accepted
term for the concept exists; regional vegetation type will be used
here.
5) The union was at one time termed associationor unistratal
associationby Lippmaaand others. Possibilitiesin the definitionof
stratalunits are also quite varied.Two majorpossibilitiesare defi-
nition by one or a few dominant species of a stratum,or by a
distributionalgrouping of several species of the same life-form
or stratum(cf. the consocionand associonof Du Rietz, 1932; the
society (Verein) and union of Du Rietz, 1936).
6) Habitat-typeshave been termed associationsby some.
C. Other Units
1) The landscape-type,a unit comprisinga more extensivearea,
or areas, of the earth's surface characterizedby all aspects of en-
vironment and natural communities (and sometimes cultural de-
velopments) and that may be recognizedas a unit by one or more
of these characteristicsof primaryinterest or importance.
2) The microlandscape-type, a unit for classificationof smaller
local areas of the earth'ssurface when consideredin terms of all
aspectsof environmentand naturalcommunities.Around this con-
cept may be grouped the concepts of nature-complex(Markus,
1925b), biocenose of Schmid (1941), forest site-typesas micro-
landscape-types(Kruedener 1926b) or biogeocenose-types(Su-
katschew, 1954), and others.
3) The habitat-type,a grouping of ecosystemsor communities
by resemblanceof their habitats or environments,a unit widely
applied in aquaticenvironmentsand specialand extremeterrestrial
environments with vegetation which is "open" (of incomplete
coverage).
4) The zone, a unit definedby dominantspeciesor other charac-
teristics, recognized as one "segment" of the sequence of com-
munities along a major environmentalgradient.
THE ECOLOGYOF ECOLOGICALTRADITIONS
Traditionally,these and other units have been the central concepts

of ecologicalschools. As outlined, the historyof synecologymight seem
to be a history of ecological units, together with the perspectives,con-
cepts, and methods related to them. Though such a view would be
superficial,the crucial role of these units in ecological history is un-
deniable; the units used in a given school express its outlook and ob-
jectives, as well as the methods it has chosen to deal with the problems
it encounters.
If it is assumed that more than one kind of "naturalunit" exists,
then some understandingof the divergenceof schools and diversityof
units may be found. Ecologicalclassificationsare affectedby properties
of the natural communitiesbeing classified; for these properties the
expression "vegetationalconditions"will be used, or (since properties
of ecosystemsand not only of vegetation are in question) "ecological
conditions." Classificationsare affected also by objectives of research,
as these are influencedin part by ecologicalconditions,in part by possi-
bilities for practicalapplicationsin relation to these ecological condi-
tions, and in part by more purely culturalinfluenceson scientificout-
look and objectives.Allowance must also be made for precedents,for
the persisting influence of key ideas and the major figures who state
them, for personalphilosophiesof leadersof schools, and for exchange
of concepts among schools and traditions.
History and present problemsof classificationsare to be understood
through the ecological, cultural,and personalinfluencesaffectingthem,
through the environmentsor contexts in relation to which classifica-
tions develop. Ecological schools, too, have their "ecology" (Sears,
1956) but may also have distinctivecharactersnot clearly determined
by their "ecology."It is beyond the scope of the presentwork to trace
such influenceson dozens of schools, but commentsmay be offered on
the ecology of the traditions.
The divison between the Northern and SouthernTraditionsin con-
tinental Europe may be related primarilyto ecological conditions, for
authorsof the two groups have had much the same classificatoryphil-
osophy. Comparing the Scandinavianand Mediterranean-and-Alpine
areasa numberof contrasts,or trends of vegetationalconditionschang-
ing northward,may be observed:
1. Community productivity and biomass decrease northward, and
with this decrease is correlated progressive reduction of community
structuretoward the lower strata.In more humid climatesclosed forests
are replaced northwardby open forests, heath, bog, and tundra. As
the forest canopy is reduced from southern forests toward the north,
the relative importanceof undergrowth in the community increases.
In this undergrowthand the communityas a whole, mosses and lichens
are increasinglyimportanttoward the north.
2. Biotic diversities,the numbersof species in florasand faunas and

in individual communities,decreasenorthward.The contrastin biotic
diversity is intensified when a northern area that has been glaciated
is comparedwith an unglaciatedsouthern one.
3. Individual species are more often strongly dominant in a given
location and stratumin the North, in contrastto the more mixed com-
position of many southerncommunities.Becausethere are few species,
and each stratummay be more strongly dominatedby a single species,
communitystratificationis more conspicuousin the North.
4. The ecological amplitudesof species tend to be broaderin mari-
time and northernclimates than in continentaland southernones. In-
dividual species may occur in a larger share of the habitatsin an area,
and the community-typesgive a greaterimpressionof interpenetration.
The intergradationand interpenetrationof community-typesin mari-
time climates have been emphasizedby Ostenfeld (1908) and Riibel
(1912b, 1927), that in the Far North by Griggs (1934, 1936), in
Scandinaviaitself by Nordhagen (1928) and S0rensen (1948).
5. The "simplification"of communitiesnorthwardis accompanied
by shortening of successions;pioneers are often species that will per-
sist into the climax (Muller, 1952).
6. In addition to these differencesin natural ecological conditions,
there may be difference in human effects on vegetation. Vegetation
in southern Europe is more effectively converted by human activities
into a mosaic of distinct patches; northernScandinaviacontains more
extensive areas of vegetation little modified by man.
The limitations and complications affecting such generalizations
should be evident. The contrastis not simply one of north and south;
it involves also contrasts of maritime with continental climates and
more with less disturbedconditions. Furthermore,much of the early
work of Ziurich-Montpellier, including the formative study of Braun-
Blanquet (1913) was done at high elevations in the Alps and in cli-
mates no warmerthan those of Scandinavia.Some bases of differences
between the traditionsmay, however, be suggested.
Becauseof the shortersuccessionsand greaterextent of undisturbed
conditions, successionalprocesses are less emphasized in the North,
except in some special circumstances.Because the northernvegetation
is less effectivelydivided into patches by human interference,the con-
cept of "association-individual" has had much less appeal.In the South
the richer flora and larger number of species with narrow amplitudes
have made possible the approach through character-species.In the

North, with fewer species present and many of these having wide am-
plitudes, dominance and constancyhave rather naturallybeen chosen
for definition of lower units. The more conspicuousstratificationof
northern communities,and the success with which a communitymay
be characterizedthrough its strataldominants,have renderedthe socia-
tion a natural unit in the North. This same evident stratification,to-
gether with the fact that partial independenceof strata is more con-
spicuousin the North, have suggestedto some authorsan even stronger
emphasis of stratal structureand implied the union as the basic unit.
The very broad amplitudesof dominanttree species, togetherwith the
relatively greater importanceof the undergrowthand more effective
expressionof environmentby undergrowththan canopy,have suggested
the third major unit, the site-type.
In Russia and the United States, continentalexpanses of vegetation
not severely disturbedby man confronted students of vegetation with
problemsvery differentfrom those of southernEurope.In both Russia
and North America,the climatic formationwas seen as a naturalunit
for relatingvegetationto climateon a continentalscale. In the Northern
and British Traditions,the formation concept has not usually been re-
strictedto climaticformations;and in the school of Braun-Blanquetthe
formationhas largely dropped out of consideration.In Russia and the
United States,monoclimaxtheoriesdeveloped,somewhatindependently,
to interpret the relation of vegetation to climate. In other traditions
a majorityof authorshave acceptedthe existenceof climaxesdetermined
by factors other than climate alone.
Beyond the approach through regional climaxes, developments in
Russia and the United States differ greatly. The Russian monoclimax
theory (of zonal and extrazonal vegetation, etc.) stemmed directly
from Russiansoil sciencealnddid not emphasizesuccession.The Ameri-
can monoclimaxtheory stemmed from studies of successionby Cowles
and others; and, as the theory was formulated into a philosophy of
vegetational dynamics by Clements, non-dynamicrelations of vegeta-
tion to environmentwere subordinatedto succession.The characteristic
means of relating smaller community-typesto one another in Russia
was the non-successionalecological series of Cajanderand Keller; but in
the United Statessmaller community-typeswere relatedto one another
through successional series, the Clementsian seres. For a lower-level
classificatoryunit, the Russiansimportedthe sociationfrom their Scan-
dinavian neighbors; and this, used in a different context, became the

Russian "association."Clements sought regional vegetation units and
regardeddominantspecies as of centralimportancein communityproc-
ess, and his "associations"were loosely defined,large-scaledominance-
types. These contrastsmay suggest the importanceof guiding ideas
which, whether native to the traditionor imported,are found produc-
tive and become the basis of further interpretations.Vegetationalcon-
ditions influence the developmentof approachesto the study of vege-
tation; but very different philosophies and techniques of vegetation
study may develop in similar vegetationalconditions.
The differencebetween the Continental "phytosociologists"on the
one hand, and the British and American "ecologists"on the other, is
expressed in the fact that they have chosen different names for the
same field of study. Each applies the other's term for the study of
naturalcommunitiesto some subdivisionof this study (Egler 1954). To
Continentalauthors, phytosociology (or biosociology) is the study of
natural communities,and synecology is the subdivision of that field
that is "ecological"in the narrowerContinentalsense, i.e., studied in
relation to environment. To English-speakingauthors synecology is
the study of naturalcommunities,while the approachesto detailed de-
scription and classificationof plant communities developed on the
Continent are phytosociology. In the Russian Tradition, finally, the
study of naturalcommunitiesis biocenologyand that of plant communi-
ties is phytocenology.
A more significantdifferencein perspectiveunderlies the difference
in terms. Both the Continentaland English-languagetraditionsdevel-
oped from earlier plant geography,but in divergent directions.In the
developmentof synecology, physiognomic,dynamic,and ecologic (or
environment-related)approachesto communitiesprevailed;synecology
is a developmentnot so much from floristicplant geographyas from
the ecological plant geography of Warming and Schimper. On the
Continent, and especially in the south, floristic emphasis prevailed;
furthermore,the classificatoryperspectiveof plant taxonomyand flor-
istics was carriedover and applied directlyto the study of naturalcom-
munities. This "taxonomic"view of naturalcommunitiesis expressed
in the analogyof the associationwith the species,a comparisonso often
stated that it forms a Leitmotiv of the majority schools of phyto-
sociologists.
The extent of the differencesbetweenecologistsand phytosociologists
is not easily explained. The species analogy and taxonomic outlook

are acceptedby some ecologists and are rejectedby some phytosociol-
ogists. Ecological conditions of the British Isles are not so different
from those of continentalWestern Europe.But ecologistshave generally
had in view the vegetation of extensive areas of diverse vegetational
conditions; on the one hand the North American continent, on the
other an Empireand Commonwealth.For study of extensive areas,the
work of Warming and Schimpersuggested differentdirectionsof em-
phasis from the intensive floristicanalysisof Continentalauthors.Ecol-
ogists naturallychose to use a less formal approach,to rely on physiog-
nomy and dominance as criteria of community-types,and to concern
themselvesmore with environmentaland dynamic,less with hierarchial
interrelationsof communities.One may allow also for ratherintangible
differencesof culturalmilieu and the influenceof individual leaders-
of the classificatoryperspectiveof Braun-Blanquetand Du Rietz, the
dynamicperspectiveof Clementsand Tansley.
Some influences on the traditions may be better observed by con-
trastingthe two schools of Braun-Blanquetand Clementsthan by com-
paring the inherentlydiverse traditionsas wholes. The Braun-Blanquet
school developed in relation to a limited area of complex vegetation,
mosaic-likein character,long inhabitedand much modifiedby civilized
man, where a considerablenumber of students could work intensively
on details of floristic compositionof local communities.To these cir-
cumstancesthe approachof Braun-Blanquet,with its emphasisof flor-
istic composition, of careful sampling and detailed analysis, and of
small-scale rather than geographic vegetation units, was appropriate.
Classificationmight naturallyproceedupwardfrom the small-scaleunits
derivedfrom local study to higher vegetationunits, forming a hierarchy
basedon floristicrelations.Along with this, the conceptsof Clementsian
dynamic ecology were adopted in modified form into the system of
Braun-Blanquet.In this, they appeared with quite different context
and significance,and as an approachto vegetation separatefrom and
largely subordinateto the central problems of classification.
The Clementsschool developed in the easternUnited States, a great
area with much little-disturbedvegetation, predominantlyof plains or
hills rather than mountains, and with few students to analyse it in
detail. Regional vegetation types defined by physiognomyand domi-
nance provided an initial, general picture of major community-types
of a continent. More detailed classificationproceededdownwardfrom
these geographicunits. The hierarchialapproachto the relationof com-

munity-typesto one another was, though attempted, less appropriate.
The rationalefor relating minor vegetation types to the regional units
was provided instead by the conception of development. The mono-
climax theory, and the climatic climax as a regional vegetation unit,
became the two imposts of the Clementsiansystem; and the fact that
prevalencein a geographic region was often the real criterion of the
climax community (as in related concepts in Europe) was obscured
by the successionalassumptions.The organismic analogy provided a
justificationfor heavy emphasis of vegetational development and the
binding together of climax and developmental communities into a
"concrete"community-typeas a super-organism.The two schools were,
for better or worse, profoundly affected by analogies suggesting how
vegetation should be conceived and studied-the analogy of the com-
munity and organism in Clements'school, of the community-typeand
species in Braun-Blanquet'sschool.
THE INDIVIDUALISTIC DISSENT
Various as are the units of vegetation, many schools share a major

assumption about natural communities and the methods appropriate
to them-that some "fundamentalunit" of naturalcommunitiesdoes
exist, a unit that is "natural"in the sense that it is present in the
structureof naturalcommunitiesand is not a productof human classi-
fication, and that is consequentlythe appropriatebasis of ecological
method. This assumptionabout the structureof communitiesand the
method of synecologyhas been termed the association-unittheory or,
more generally, the community-unittheory (Whittaker, 1956). It has
not, however, been universally accepted.
Some significantqualificationsappearin the writings of the phyto-
sociologists,whose work dependsmost on these units. Pavillard (1912:
170) early expressed the conception that only a utilitarianbond ex-
ists between the members of an association-each one for itself is
the naturallaw (cf. Pavillard, 1935a). In the Scandinaviantradition
Frodin (1921, 1922; criticism,Du Rietz, 1922) challenged the view
of Du Rietz that associationsas fundamental units have sharply de-
fined boundaries;and Kylin (1923, 1925, 1926; criticism,Fries, 1925,
1926) observed that species distribute themselves independently of
one another accordingto ecological conditions.Nordhagen (1928:67)
observed that some communitieshad no sharp boundariesand could

be limited only arbitrarily,and that some alpine communitiesshowed
a complicatedmosaic which could bring a phytosociologistto despair,
in which one could only select the most suitablepoints and set the rest
aside. Drude (1926, 1932), observingpartial independenceof species
and continuityof communities,questionedthe approachto associations
throughtabulation,since associationsare not fixed, closed units. Domin
(1928a:40), noting the continuityof vegetationtypes and the variations
within them, observed that an enemy of plant sociology could easily
doubt the realityof the chief associationsthemselves.Lippmaa (1933a:
10-12) indicatedthat the fact that no two species have the same distri-
bution, locally or geographically,raises the question if the existence
of associationsin nature is an illusion, but still consideredthe associa-
tion a fundamentalunit of local importance.Vaarama(1938) rejected
the traditionalunits in the treatmentof aquaticvegetation in favor of
Gleason's (1926) individualistichypothesisand species-standsas work-
ing units. Romell (1920), Waren (1926) Paczoski (1930b), Motyka
(1947), Boysen-Jensen (1949) and others have remarked on the
limitations of associationsas "natural"units.
Among the schools of the SouthernTradition, Schmid and Daniker
have dissented strongly from the classificatorysystem of Braun-Blan-
quet. Schmid (1940, 1941, 1942, 1944) criticizes the specific and
organismic analogies and emphasizes the lack of organization and
of clear boundariesin naturalcommunities.Braun-Blanquet'sapproach
is considered by Schmid (1942) to tend toward a sterile formalism;
biocenoses should be characterizednot one-sidedly by floristics, but
through their whole structure.While associationsmay have value for
preliminaryvegetation research,they must be grouped into more sig-
nificant, larger units based on environmentaland historical, as well
as floristic, relations (Schmid, 1922, 1935; Diniker, 1939a, 1939b).
Meusel (1939a, 1939b, 1940, 1943a, 1954b) expressed comparable
objections to the one-sidednessof Braun-Blanquet'ssystem and criti-
cized the artificial, schematizingtendency and the dissolution of the
natural whole-phenomenaof vegetation into abstractunits. Rejecting
the specific and organismic analogies, Meusel (1940:507) conceives
vegetation not through a hierarchyof units, but as a tapestrywoven
of threadsof many colors which, in orderlydistributionthrough space,
form the variegatedpatterns that meet our eyes.
Dice (1952:425), author of the American system of biotic prov-
inces, believed that an associationdoes not exist in nature,but is formed

in someone'smind by those stands consideredto be of generally simi-
lar type. Becausethe classificationof communitiesis based upon opin-
ion, there is no "right" classificationfor any area. Communityclassi-
fication must change as information accumulatesand as ecologists
modify their opinions about the relative importanceof the features
that distinguish communities.
Whereas most of the vegetation units previously discussed may be
characterizedby one or two criteria, Meusel, Schmid and Daniker,
Dice, and others seek to consider the many possible criteriaof com-
munities and environmentstogether, in order to recognize units of
more general meaning. To these authors of multi-factoralor land-
scape perspectives, associations may seem less "fundamentalunits"
than one ratherrestrictedapproachto the complex relationsexisting in
naturalcommunities.
Attacks on the association-unittheory are associated also with a
group of authorsthat includesRamensky,Gleason,and Lenoble.Among
the "fundamentallaws of the vegetationalmantle" Ramensky (1924)
advancedtwo principlesconflictingwith assumptionsabout associations
as well-defined, naturalunits: the principle of vegetationalcontinuity
and the principle of species individuality.The vegetationalmantle in
general (apart from disturbance,sharpchange in externalfactors,etc.)
changescontinuouslyin space, and each plant species respondsuniquely
to externalfactors and enters the biocenoseas an independentmember.
The kaleidoscopicchange in spatial compositionof communitiesspeaks
against classificationinto mere units; it is not these units but the rules
of plant combinationwhich are important. As the goal of research,
Ramensky (1924, 1930, 1932) saw the arrangementof biocenosesin
ecological series according to abundancecurves of their species and
coordinatesof externalfactors. The ideal was not classification,but the
linking of all biocenosesstudied into a unified coordinate-scheme(Ra-
mensky, 1924, 1930).
In the United States, Gleason (1926), also observing species inde-
pendence and vegetational continuity, independently formulated the
"individualisticconcept of the plant association."Gleason concluded
that an associationis not an organism,scarcelyeven a vegetationunit,
but merely a coincidenceresultingfrom environmentalsorting of plant
populations. Shreve (1915) also observed independence of species
distributionsand the fact that it was nowhere possible to pick out a
group of plants which might be thought associates,without being able

to find other localities in which the association was dissolved. In
France, Lenoble (1927, 1928a, 1928b) criticizedBraun-Blanquet'sas-
sociationson the basis that they were essentiallyunstable and change-
able mixturesof independentspecies combiningin variousproportions.
The associationis thus an abstractunit; to be sure, it is even so ab-
stract as to be imaginary (Lenoble 1928a). Fournier (1927) consid-
ered Fagus silvatica in relation to other species and observed their
infidelity to the Fagus association.Fournier concluded that the plant
associationrepresentedan essentiallyheterogeneouscomplex, unstable
in its compositionand transitoryin time and space,and that the methods
of the sociologistsconstitutea precioustechnique,not a method in the
philosophic sense-their scope is too narrow to merit the latter term.
In Italy, Negri (1926, 1927) attackedthe organismicanalogy and ex-
pressed agreementwith the ideas of Ramensky,Gleason, Lenoble, and
Fournier;related ideas had been expressedearlierby Negri (1914) in
his essay on the fundamentalecological unit.
Direct challenges to the association-unittheory thus appearedinde-
pendently in 1924, 1926, and 1927 in three of the regional traditions.
In Australianecology Patton (1930) observedthat each species of any
habitat has its own specific range; a communitymay be regardedas
an areawhere a sufficientlylarge numberof species overlap to give the
area a distinctive appearance.An associationof plants cannot be re-
garded as a unit like the species, but must be regardedmerely as an
assemblageof wandering units. Hauman (1933) has asked whether
the analogy between the associationand the species is not a sophistry
and whether the grandiose enterpriseof phytosociologyis viable. Ber-
ner (1948) found that "transgressions"or changes in areasof species
involved individual species ratherthan groups of species, in a vegeta-
tion mantle in constantflux. Plant communitiesresult from the super-
position of areas of species; the associationis an illusion, a subjective
product of appearances.Walter and Walter (1953) were led by ob-
servationson extensive naturalvegetationin Africa to statementof the
individualistichypothesis.They observedthe geographiccontinuityof
vegetation and distributionalindependenceof species, and concluded
that vegetation units are more or less arbitrarilyconceived types of
species combinations which recur when corresponding sites recur.
Webb (1954) has observedthe intergradationof communitiesand lack
of full correlationbetween species, and regardedthe attemptto estab-
lish a single, internationallyacceptedhierarchialsystemas scholasticism

out of place ratherthan science.
The "individualistic"conceptionof associationshas been much dis-
cussed in the American Tradition, particularlyin more recent years
(Gleason, 1926, 1929, 1939; Raup, 1942; Cain, 1947; Mason, 1947;
Egler, 1947; Billings, 1949; Whittaker, 1951, 1952, 1953, 1956, 1957;
Curtis and McIntosh, 1951; Brown and Curtis, 1952; Segadas-Vianna,
1951; Muller, 1952, 1958; Hanson, 1958; Rowe, 1961) and in Europe
by Liudi (1928), Pavillard (1935a), Vaarama (1938), and Poore
(1956). Statementsof the individualityof species and arbitrarycharac-
ter of associationshave not, for the most part, been receivedwith ap-
preciation by conservativephytosociologistsand ecologists. Gleason's
argumentswere criticizedby Tansley (1920), Nichols (1929), Clem-
ents et al. (1929:315), Cain (1934), and Phillips (1934-5); they
were at one time ridiculed (Gleason, 1953). Lenoble and Fournier
made even less impressionon Continentalphytosociologists.Although
Gleason's logic was consideredby Pavillard (1935a), the related ideas
of Lenoble were summarilyrejectedby Pavillard (1928a, 1935b) and
Allorge (1927). Lenoble's inquiry into the nature of associationsre-
ceived from Braun-Blanquethimself only a deflective and misleading
answer, a model of how not to debate a scientific issue (Lenoble,
1928a, 1928b; Braun-Blanquet,1928b). These published comments
suggest hauteur toward an idea unworthyof serious consideration;in
conversation some ecologists most devoted to the association theory
have expressed a view of the individualisticconcept as a destructive,
subversiveidea and a strong, even passionateconvictionthat it cannot
be true. The essential ideas of the dissent were more clearly stated by
Ramensky;and the climate of Russian phytocenologyproved less in-
hospitable toward them. Katz (1930b, 1933) accepted the principle
of species individuality,but considered that Ramensky'sobjections to
classificationrepresentedaniover-emphasisof this principle.Sukatschew
(1929) considered that there was no conflict between recognitionof
the plant cover as continuousand use of the associationas an abstract
unit (cf. Ponyatovskaya,1961).
A first) major choice in the interpretationof natural communities
lies between the individualisticdissent and the association-unittheory.
The problem may be broaderthan this choice, however, for interpreta-
tions of the associationrange from the view of these units as real and
concrete (Clements, 1916, 1928:128; Du Rietz, 1921:15, 1928, 1929;
Alechin, 1925b), through conceptionsof the associationas an abstract

unit, or type (Gradmann, 1909; Samuelsson,1917a; Braun-Blanquet,
1921; Wangerin, 1925; Kylin, 1926; Liidi, 1928; Nordhagen, 1928;
Kalela, 1939; Kalliola, 1939; Klapp, 1949; Wendelberger,1951, 1952;
Dice, 1952; Ellenberg, 1954a, 1956, etc.), to the view that it is purely
subjective and arbitrary.Conceptionsof the associationfurther range
from its interpretationas an organism or super-organism(Clements,
1916, 1928, 1936; Phillips 1931b, 1934-5) or quasi-organism(Tans-
ley, 1920, 1935), through the widespread analogy with the species,
and the analogywith a humansociety (Tansley, 1920; Aichinger, 1943;
cf. Muller, 1958), to the rejectionof all these and Lenoble'scompari-
son with chemicalmixtures.The following sections will consider first
the evidence on the individualisticand association-unithypotheses,and
then the place of synecologicalunits in currentunderstandingof natural
communities.
THE THEORY OF SYNECOLOGICALCLASSIFICATION

EVIDENCE
TYPES OF EVIDENCE
The statementthat "the associationis the fundamentalunit of syne-

cology" is ambiguous: Is the associationfundamentalto the structure
of natural communities, or fundamental only to the techniques em-
ployed in a given school? The two questions, of theory and method,
must be clearly distinguished;it is the former that is crucial.Practical
problems of classificationmay be seen in very different lights if the
associationis a basic conception about the structureof vegetation, or
a convention acceptedfor its practicalusefulness. The associationand
other units have a place in the basic theory of natural communities
only if they are fundamentalin a much more fundamentalsense than
the latter.
It may perhapsbe grantedthat the associationis an abstractionfrom
a group of stands, a type or class of communities.The problem is not
whether, as a conceptualunit, the associationis a "reality";the problem
concernsthe relationof these units to propertiesof naturalcommunities,
their relative "naturalness"or "arbitrariness"(cf. Whittaker, 1956).
On the one hand, associationsand the boundariesbetween them might
be directlybased on and unequivocallydeterminedby the structureand
manner of organizationof naturalcommunitiesin the field. Classifica-
tion is then a matter of "discovering"natural units which must be

taken accountof in any synecologicalresearch.On the other hand, there
might be no properties of natural communitieswhich unequivocally
determinethe content and limits of associations.These units are then
arbitraryor conventional;they are freely createdby man, for particular
purposes and by choice of particularpropertiesof communitieswhich
interest him and serve as bases of classification.The former may be
taken as the essentialmeaning of the association-unittheory, the latter
of the individualisticdissent.
The problem of associationtheory should be approached,not as a
matter of metaphysics,but of observableand measurablerelations of
stands to one another, and species to one another. Few currentques-
tions in synecology have more fundamentalimport than this, of the
relations of stands and species on which our conception of communi-
ties and our understandingof their classificationshould be based. Most
argumenthas centeredon two points: That stands and associationsare,
or are not, discontinuouswith one another,and that species are, or are
not, organized into distinct groups correspondingto associations.Four
interrelatedtypes of evidence may be considered as bearing on these
questions and providing backgroundfor following sections on theory
and practice:
1) Relative similarity and dissimilarityof stands. Associations, if
"natural"units, should be relatively homogeneousinternallyand dis-
continuousexternally.They should consist of groups of stands closely
similar to one another and all clearly different from other groups of
stands representingother associations.The mannerof grouping stands
into associationsis in this case clearly determinedfor the ecologist by
the relationsof stands. If, however, stands are in varying degrees and
differentways similar and dissimilar,then there may be many possible
ways of grouping them.
2) Continuity and discontinuity.If associationsare natural units,
stands intermediateto associationsshould be rare or of limited extent
comparedto standsof the associationsthemselves,and contactsbetween
stands of two different associationsshould be discontinuous.If asso-
ciations are thus discontinuous,their boundariesare clearly determined
by properties of vegetation; if they are not, boundariesmust be de-
termined by the ecologist and his choice of classificatorycriteria.
3) Distributionalrelations of species. If the stands of an associa-
tion are closely similar, all or many of the species must have similar
distributionsthrough these stands. If associationsare discontinuous,

many, or at least some, of the species should have their boundaries
together in the discontinuitiesbetween associations.Ideally, an associa-
tion should consist of (or should include) a group of species with
closely similar or congruent distributions,their distributionallimits
coinciding with the limits of the association.If, on the other hand,
species are most variously distributedin relation to one another and
communities,then grouping of species into associationsis largely arbi-
trary.
4) Dynamic relationsof species. Distributionalsimilarityis unlikely
to occur as a coincidentalproduct of independent or "individualistic"
distributionsof species. If associationsare naturalunits, speciespopula-
tions are presumablybound together by necessaryinterrelationsinto
groups which must occur together and will have their distributional
limits together. If the associationis a naturalunit, species populations
should in some sense be otganized into associationsby obligate rela-
tions, so that species do not occur independentlyof their associations.
If species are not bound together by such obligate relations, they are
more likely to be diversely distributed,so that their grouping into as-
sociations is arbitrary.
SIMILARITY AND DISSIMILARITY OF STANDS
Few studies have approached the theory of associations through

relative similarity of samples taken by unprejudicedmeans. In most
ecological and phytosociologicalwork, stand samples are taken to rep-
resent associationsor other units; and transitionalstands are likely not
to be sampled,or not to be used in the tables compiledfor publication.
Here, both the similarity of the samples representinga given com-
munity-type,and their consistent dissimilarityfrom samples represent-
ing other community-types,may be productsof sample choice. Tabula-
tions of samples thus chosen to representcommunity-typesdo not pro-
vide unbiased evidence on the nature of associations.
When samples are taken by unprejudicedmeans, a large proportion
of them are likely to be "mixed," "atypical,"or "transitional."Ellen-
berg (1954a) observedthat the attemptto classify the weed communi-
ties of the Ulm district showed that more than three-quartersof the
stands were intermediateto two or three associations,or even to al-
liances and orders.Klapp et al. (1954) found that among 1029 grass-
land samples, not selected to representassociations,mixtures or inter-

penetrationsof associationspredominated;about 25 per cent of the
samples could be used to representcommunity-types.
Some ecologists have had similar results with unprejudicedsamples
from more naturalvegetation. Among the samples of Wisconsin for-
ests studied by Curtis and McIntosh (1951), and Brown and Curtis
(1952), and prairie studied by Curtis (1955), no two stands were
alike; the stands could be arrangedalong continuawith all degrees of
relative similarity. Whittaker (1956) obtained 300 samples, chosen
without reference to associations,from the Great Smoky Mountains,
assuming that these might fall into natural groups. The bulk of the
samples, from forest stands, showed all degrees of similarityand dis-
similarity,and could only arbitrarilybe grouped into community-types.
Several authors have sought objective, quantitativemeans of classi-
fying stands into community-types(see also Goodall, 1952; Dagnelie,
1960). Jaccard(1902, 1908, 1932; Guyot, 1923; Ekman, 1940; Koch,
1957) used the coefficient of community, the percentage of species
sharedby two samples among the total numberof species occurringin
one or both, to comparelists of species. S0rensen (1948) found that
stands grouped by coefficientsof communitymight correspondto soci-
ations, associations,and alliances. Limits of coefficientof community,
by which stands were grouped, could be set only arbitrarily,however,
and S0rensen observed the general merging of types in the Danish
vegetation with which he dealt. Dahl and Hadac (1941), Pfeiffer
(1945), Evans and Dahl (1955), Hanson (1955), Schilder (1955),
Guinochet (1955), and Ellenberg (1956) have also used coefficientsof
communityfor comparisonof community-types.Poore (1955c) used
this measureto compare18 community-typesof varyingranks.A slight
tendency for these to fall into groups was observed but certain well-
marked communitiesseemed to bridge the gaps between groups. The
results could best be accountedfor by the simultaneousvariation of
several factors- implying a multi-dimensionalpattern in which some
community-types,at least, fell into continuousseries and not discrete
groups.
Kulczyn'ski(1928) developed a "coefficientof relationshipof asso-
ciations"which based the comparisonon frequencies,ratherthan pres-
ence or absence, of species (cf. Gleason, 1920; Raabe, 1952). The
techniqueof Kulczynskiand Czekanowski,with arrangementof meas-
urementsof sample similarityin a trellis diagramor matrix, has been
applied to the classificationof vegetation by Motyka (1947; Motyka

et al., 1950) and Matuszkiewicz (1948, 1950, 1952). Motyka found
that degrees of associationbetween samples were weak, rarelyas high
as 60 or 70 per cent and often 40 per cent or less within a given
community-type.Some samples could be classed in two or three com-
munity-types.Motyka (1947) concluded that the community-types,or
associations, had only indistinct limits and were poorly defined in
nature.Matuszkiewicz(1948) compared220 forest samplesand found
continuousintergradationamong them, so that it was not easy to de-
limit distinct community units. Matuszkiewicz concluded that the
plant associationas a definite systematicunit in analogy to the species
did not exist; but later work (Matuszkiewicz, 1950, 1952; Matusz-
kiewicz and Polakowska, 1955; Matuszkiewicz and Matuszkiewicz,
1956a, 1956b; Izdebska, 1958) also showed that results of the Kul-
czynskitechniquecould be used for classificationby the systemof Braun-
Blanquet.Bray (1956) used Gleason's (1920) modificationof the co-
efficientof communityto arrange20 Wisconsin samplesin a Kulczyn'ski
triangle;the patternsof plot relationshipshowed smoothand continuous
changeof coefficientvalues from closed forest to grassland.The analyses
of communitysimilarityby Bray and Curtis (1957) and Maycockand
Curtis (1960) showed communitiesto be complexly and continuously
related in terms of several gradients of environmentand community
composition.
Severalstudentsof biocenotics(Renkonen,1938, 1944, 1949; Agrell,
1941, 1945b; Forsslund, 1945; Brink and Wingstrand, 1949; Kont-
kanen, 1949, 1950a, 1950b, 1957; see also Balogh, 1958) and animal
ecology (Odum, 1950; Whittaker, 1952; Whittaker and Fairbanks,
1958) have used the Kulczyn'skitechnique to comparesamples from
animal communities.Most of these use a differentand simpler formula
than Kulczyn'ski's,basing comparisonon numbers of individual ani-
mals insteadof frequencies.The natureand limitationsof this measure-
ment, which may perhaps best be termed percentagesimilarity, have
been discussedby Whittaker (1952; Whittakerand Fairbanks,1958).
Use of the measurement by Renkonen (1938, 1944), Forsslund
(1945), Brink and Wingstrand (1949), and Whittaker (1952)
showed low degrees of similarity among samples and no very clear
grouping into community-types;the last two studies in addition sug-
gested continuityof animalcommunitiesalong environmentalgradients.
Kontkanen (1950a) comparedthe same series of samplesboth by per-
centage similarity and by coefficient of community; quite different

groupings resulted, and grouping by coefficient of community was
preferred.
Quantitativeapproachesto communitieshave not solved problems
of classification(Clapham, 1936; Ashby, 1936, 1948; Curtisand McIn-
tosh, 1950; Goodall, 1952, 1953a, 1954a, 1954b; Macfadyen, 1957;
Dagnelie, 1960). The work reviewed here and other recent studies
(Dahl, 1957; Hosokawaet al., 1957; Clausen, 1957b; Barkman,1958;
Suzuki and Abe, 1959; Looman and Campbell, 1960) have shown
that measurementsof communitysimilaritycan be adaptedto the pur-
poses of classification.But their function, like that of many other
quantitativetechniques, is to summarize complex data as an aid to
human judgment, which must somewhere in the process make deci-
sions on criteriaand limits of classes. Results from such studies sup-
port the view that stands, instead of falling naturally into groups
clearly separatedfrom one another, are complexly related to one an-
other with varying degrees of similarityand dissimilarity.
CONTINUITY AND DISCONTINUITY
The occurrenceof discontinuities between communities has been

widely observed and discussed (Beck Mannagetta, 1902; Cajander,
1909, 1925c, 1949; Gleason, 1917; Du Rietz et al., 1920; Du Rietz,
1921, 1922, 1923a, 1924, 1925a, 1932; Scharfetter,1921, 1924, 1932;
Kylin, 1923, 1926; Frey, 1923, 1927; Fries, 1925; Wangerin, 1925;
Kujala, 1926; Nordhagen, 1928; Liudi,1928; Kalela, 1939; Kalliola,
1939; Dahl and Hadac, 1941, 1949; Pfeiffer, 1943; Nytzenko, 1948;
Hayward,1948; Allee et al., 1949; Odum, 1953; Evans, 1953; Ehren-
dorfer, 1954; Poore, 1955b; Burbancket al., 1956; Whittaker, 1956,
1960). Many authorsassumedthat associationswere discontinuouswith
one anotheror that, as expressedby Du Rietz (1921), transitionswere
insignificantrelative to the associationsthemselves. If the association
is considereda fundamentalunit in the sense of Du Rietz (1921), dis-
continuity between associationsis likely to be accepted as a corollary
(vide Scharfetter,1921, 1924); the sharpboundingof contiguouscom-
munitiesmay then be thought essentialto their nature (Pfeiffer, 1943).
Yet the boundariesof communitieshave received little critical study;
they have been consideredby many as lacking in interestand unworthy
of investigation (vide Lippmaa, 1933a:13). The possibility that dif-
ferent degrees of discontinuitybetween associationsexist was indicated

by Du Rietz (1923a, 1923b); as there are "good" and "weak"species,
so there may be "good" associationswith sharpboundariesand "weak"
associationswith more gradual transitions and arbitraryboundaries.
Liudi(1928) observedthat in the Alps dwarf-shrubstands were often
sharplybounded, but that grasslandcommunitiesmixed and interpene-
trated without clear boundaries;Dahl (1957) observed discontinuity
of some and continuityof other communitiesin Norwegian mountains.
Nytzenko (1948) supportedthe existenceof relativelysharptransitions
between communities,but indicatedthat three types of boundariesmay
occur-clear-cut, diffusive, and mosaic-insular-of which the last is
most common.
Gradualor continuousgradationsbetweenassociationshave also been
widely observed,and many authorshave commentedon the continuity
of the vegetation patterns with which they were dealing (Warming,
1889; Kearney, 1901; Ostenfeld, 1908; Gleason, 1917, 1926; Raun-
kiaer, 1918; Arrhenius,1921; Frodin, 1921; Brough et al., 1924; Kel-
ler, 1925-6; Schroter, 1926; Domin, 1928a; Nordhagen, 1928; Pacz-
oski, 1930b; McBryde, 1933; Adamson, 1938; Schmid, 1940, 1950;
Cooper, 1942; Crocker, 1944; S0rensen, 1948; Matuszkiewicz,1948;
Wagner, 1950a; Story, 1952; Parker,1952; Walter and Walter, 1953;
Poore, 1955c; Churchill, 1955; Orshan and Zohary, 1955; Boughey,
1957; Steenis, 1958). The tabulationsby various other authors indi-
cate continuouschange of populationsthroughthe series of associations
or types represented(Ilvessalo, 1922; Hansen, 1930, 1932; Sampson,
1930; Ramensky,1932; Knoll, 1932; Meusel, 1935; Halliday, 1935;
Keller, 1936; Riihl, 1936; Tiixen and Ellenberg, 1937; Watt, 1940;
Horton, 1941; Tuomikoski, 1942; Tolstead, 1942; Eggeling, 1947;
Knapp, 1949; Spilsburyand Smith, 1947; Klapp, 1949-50; Ellenberg,
1950a, 1950b, 1952a; Potzger, 1950; Quarterman,1950; Braun, 1950;
Marler and Boatman,1952; Wagner, 1950a, 1950c, 1954a).
The researchesof Whittaker (1951, 1954a, 1956, 1960), Curtis
and the Wisconsin school, and Goodall (1954a), have converged on
the recognitionof vegetationalcontinuity (see also Rowe, 1956; Hori-
kawa and Okutomi, 1954; Horikawaand Itow, 1958; Okutomi, 1958).
Whittaker (1956) found that the forest pattern of the Great Smoky
Mountainswas one of continuousgradation of stands along environ-
mental gradients. Certain vegetation types ("zones"), however, were
relatively discontinuousin at least one contactwith anothervegetation
type. These relativelydiscontinuoustypes showed continuousgradation

within themselves, and most could be shown to intergradewith other
community-typesin directions other than those in which the discon-
tinuities were observed. Many species extended through the discon-
tinuities, regardedas relative discontinuities,or steepeningsof the gra-
dation occurringthroughoutthe vegetationpattern.The whole pattern
was conceivedto be a complex continuumof populations,with the rela-
tively discontinuous types confined to "extreme" environments and
forming a minor part of the whole. Allowing for discontinuitiespro-
duced by disturbanceand environmentaldiscontinuity,the vegetation
pattern could be regarded as a complex mixture of continuity and
relative discontinuity.
Both continuityand discontinuityare thus to be observedin natural
communities. Many vegetational discontinuities are products of en-
vironmental discontinuity,are sharpened by fire, or are products of
other disturbance.When these "externally"induced discontinuitiesare
set aside, some relative discontinuitiesalong continuousenvironmental
gradientsremain,along with a more general continuityof communities
with one another. The evidence from vegetationalcontinuityand dis-
continuity is not unequivocal as it relates to associationtheory. The
weight of evidence seems, however, against the assumptionthat asso-
ciations or other community-typesare in general clearly bounded.
DISTRIBUTIONAL RELATIONS OF SPECIES
When distributionsof species making up a stand or associationare

mapped together, so that their outlines can be directly compared,it is
at once apparentthat the geographic patternsof the species are most
varied (e.g., Billings, 1949). Often no two species will have similar
distributions.This fact is too well known to require extended dis-
cussion,but certainimplicationsmay be observed: (1) The dissimilarity
of geographicpatternsfor speciesof one associationsuggestsgeographic
expressionof the principle of species individuality. (2) It is generally
impossible to define the geographic range of an association by the
coincident distributionallimits of its species. Although approachesto
communitiesthrough grouping of species by similarity of geographic
distribution(Hansen, 1930; Meusel, 1939a, 1943c; Bocher, 1940, 1945,
1954; Ellenberg, 1950a) or conformal areas (Hulten, 1937; Raup,
1947) are useful, they do not themselves define community-typesand
do not give unequivocalresults on the boundariesof community-types.

(3) Since componentspecies drop out one by one with increasingdis-
tance from a given point of observation,associationsas definablecom-
binations of species are local phenomena (Paczoski, 1930b; Lippmaa,
1933a; Bourne, 1934; Cain, 1947; Knapp, 1948a; Walter and Walter,
1953; Ellenberg, 1954a). (4) Because of the gradual decline and re-
placementof species along climaticgradients it is probablethat many
associationsare geographically,as well as locally, continuouswith other
associations(Walter and Walter, 1953).
By far the largest body of data available on local distributionof
species is in publishedreleves of phytosociologists.As suggestedabove,
certain limitations on the use of these as evidence must be observed.
The samples published together are chosen to representa community
type as conceivedby the author;samplesfrom intermediateor "mixed"
stands are seldom presented.The exclusion of intermediatestandswill
sometimesresult in a misleading appearanceof discontinuityin species
distributionand suggest that the boundariesof a group of species coin-
cide when in fact they do not. The search for groups of diagnostic
species which are distributionallyrelated, grouping of stands by oc-
currenceof these species, and exclusion of non-conformingstands may
all tend to exaggerate the appearanceof distributionalsimilarity of
species. The manner in which the releves are selected and presented
tends to prejudice the evidence in the direction of associations as
"natural"groupings of species and stands.
Nevertheless,from close study of a series of tables for differentcom-
munity-typesin a given area, it is usually apparentthat the species
are distributeddiversely,accordingto the principleof species individu-
ality. The companions or Begleiter of one associationwill generally
have most varied relationsto other associations.Differential-speciesare
usually of varied distributionalrelations, apart from the more-or-less
coincident boundariesin one direction which permit their use as dif-
ferential-species.Characterspecies of low degree for a given unit show
varied patterns of extent into other units; character-speciesof high
degree are likely to be differentlydistributedamong the samples for a
given unit. It is usually evident enough in these tables that character-
species are not "associated"in the sense that they have closely similar
distributionsthrough stands and community-types.
Two sets of tables arrangedto make more easily observedthe rela-
tions of species to community-typesmay especially be cited. Liidi's
(1928) essay on the associationconceptpresentsa remarkabletable for

222 vascularplants and 98 othersthrough 24 community-types.
Species-
to-species differencein distributionis evident both for species in gen-
eral and for character-species,by Liudi'srestricteddefinition of these.
The tables of Nordhagen (1937) permit similarobservationsfor Scan-
dinavian vegetation, particularlyfor the character-speciesof alliances
and higher units recognizedin this work. The extensive tables of Riihl
(1936) may also be mentioned; and other authors (Oskarsson,1927;
S0rensen, 1937; Laessle, 1942; Gjaerevoll, 1949; Braun, 1950; Han-
son, 1951; Daubenmire, 1952; Poore, 1955c; see also tables cited as
evidence for continuity) have presented tables in which the distribu-
tions of species may be effectively compared.
Ramensky (1930) attempteda test of his principle of species indi-
viduality.Communitysampleswere grouped by the occurrencein them
of a given species at high population levels. The "elective mean" of
this group of samples could then be determinedby methods described
by him, and this gave expressionto the distributionalcenter or coeno-
center of that species. Distributionsof species in the coenocentersof
other specieswere also tabulated,thus providing a distributionalpicture
or coenospectrum for each. Ramensky found the coenocentersand
coenospectraof 70 species studied were demonstrablydifferent;in the
few species where coenocenterswere closely similar, their coenospectra
differed.He consequentlyconsideredthat the rule of ecologicalspeciali-
zation or individualityof plant specieswas established.
Iversen (1936) determined for a series of species the "ecological
spectra"of averagefrequenciesof otherspeciesin the "dominance-area"
of a given species. These conceptsof dominance-area(Raunkiaer1928;
cf. the Dominanzkreisof Waren, 1926) and ecological spectrumare
related to Ramensky'scoenocenter.Iversen's tabulation, like Ramen-
sky's, indicatesthe distributionalindividualityof the 42 speciesstudied,
though one pair of majorspeciesshowed close similarityin their spectra.
Pidgeon and Ashby (1942) outlined distributionsof species by fre-
quenciesin a transectgrid; centersof majorspecieswere scatteredrather
than clusteredin the pattern.
Some authorshave sought to reveal naturalgroupings of species by
quantitativeprocedures (see also Goodall, 1952; Greig-Smith,1957).
Means of measuringassociationof species have been reviewedby Dice
(1945), Cole (1949), Nash (1950), Goodall (1952), and Dagnelie
(1960). Kulczyn'ski(1928) studied the problem of species correla-
tions, as well as of sample association,and conceived the stand as a

certaincombinationof correlatedspecies-groups.Katz (193ob) showed
the existence of correlationsbetween plant species in quadratswithin
associations,and between dominantsand other species throughdifferent
associations.Iljinski and Poselskaja (1929) showed the existence of
both positive and negative correlationsamong species, but concluded
that there were no true associatesamong the autophytesstudied and
that the response of plant species to the habitat was individualistic.
Sheygrond (1932) studied correlationin occurrenceof dominantspe-
cies in differentstrataof vegetation;Iversen (1936) measuredecologi-
cal affinityof speciesby averagingthe frequenciesof two speciesin each
other's dominance-areas(cf. Raunkiaer, 1928). Stewart and Keller
(1936) found that pairs of species positively correlatedin one vegeta-
tion type, and hence possible associates,were sometimesnegativelycor-
related in another vegetation type. Tuomikoski (1942) showed the
possibility of establishing groups of correlatedspecies in relation to
moistureand nutrientgradientsas a basis of classification,and applied
the same correlationtechniquesto animal communitysamples (1948).
The existence of distributionalcorrelationsof species, both positive
and negative, within stands, among stands of a given community-type,
and through series of community-types,is undoubted.The existenceof
such correlationsis not, in itself, evidence for or against either the
individualistichypothesis or the association-unittheory. The question
is, rather,one of the degree and kind of correlations,and the nature
of the distributionalrelationsthey express. Existenceof naturalgroups
of associated species should be revealed in high correlationsamong
these species, in contrastto low or negativecorrelationsof these species
with all others. Distribution accordingto the principle of species in-
dividualityshould be reflectedin a kind of correlationalindifferenceor
promiscuity,manyspecieshaving positive correlationsof low degreeand
many others negative correlations,but no groups having very high
correlationsamong themselvesand low or negative ones with all other
species.
The latter conditionwas observedby Iljinski and Poselskaja(1929).
Matuszkiewicz(1948), Motyka et al. (1950), Gardner (1951), Gil-
bert and Curtis (1953), Culberson (1955), and Whittaker (1960)
have computed associationsof species in vegetation; no very distinct
groups of species were shown in any of these studies. Nash (1950)
computed degrees of associationamong fish species in various water
bodies and found indicationsof naturalgrouping into three communi-

ties characterizedby type of bottom, though many species ranged
through two or three of the community-types.Webb (1950) studied
rate of change of mammal and snake faunas across Texas and Okla-
homa by comparingfaunas 100 miles apart;the results suggestedfour
major biotic communitiesseparatedby areas of more rapid change of
fauna. Greig-Smith (1952) applied Cole's (1949) index of asso-
ciationto tropicalforestsand found only very limited associationamong
species, and this mainly reflectingnon-randomnessin secondaryforests.
Greig-Smithwas led, without necessarilyacceptingGleason'sview in its
entirety,to doubt if the view of the communityas a complex organism
applied to the forests studied. Hopkins (1957) found that several
groupings of correlatedspecies existed in a given community,with one
major grouping forming a matrix in which the otherswere distributed.
Percentage similarity and related measurementscan be applied to
associationof species, as well as of samples (Agrell, 1945b; Kont-
kanen, 1949; Whittaker, 1952; Bray, 1956; Ellenberg, 1956). Back-
lund (1945) applied the technique of Kulczyn'skiand Agrell to asso-
ciation of speciesin the wrackfauna. No distinctcommunitiesand only
weak associationswere revealed among the species of this unstable
fauna. Kontkanen (1950a) measured association of species in leaf-
hopper samples and found three groups of correlatedspecies which
corresponded to major community-types.Whittaker and Fairbanks
(1958) used measurementsof distributionalsimilarityto constructa
plexus or diagramof copepod species interrelations.The plexus seemed
to representnot a numberof distinctassociations,but a complexpattern
of distributionaloverlapsin which each species had its own distinctive
environmentalrelation.
Vries (1953; Damman and Vries, 1954; Vries et al., 1954) com-
puted correlationsfor more than 40 major species in a large number
of samples from Dutch grasslands.The results were presentedin the
form of diagrams in which degrees of associationwere represented
by distances and types of lines connecting species. Certain objective
groups or "constellations"of species appeared,and these could be cor-
related with environmentsand to some extent with the units of the
school of Braun-Blanquet.The species were related to one another,
however, in all degrees and in many directionsin a multi-dimensional
network.This did not indicate, in Vries' opinion, that plant communi-
ties, conceived as combinations of species, were sharply limited in
nature (Vries et al., 1954). Iversen (1954) computed the occurrence

together of species in quadratsin a Greenland valley, and found a
striking correspondencebetween the species groups which resultedand
the Norwegian phytosociologicalunits of Nordhagen; he concluded
that the units of Nordhagen and Braun-Blanquetwere naturaland well-
circumscribedgroups. Iversen's approachwas, however, dependent on
subjective,visual grouping of species by distributionaldiagrams.Bray
(1956) found the large proportion of species to be independently
associatedin a study of Wisconsin communitiesranging from closed
forest to grassland.Goodall (1953a, 1954b) and others (Hosokawa,
1955b; Rayson, 1957; Williams and Lambert, 1959, 1960, 1961; Ker-
shaw, 1961) have used correlationsof species distributionsas a means
of classifying communitysamples.Goodall (1953a) observesthat dif-
ferent classificationsresult from different choices of criteria for the
same set of samples, Kershaw (1961) that the meaning of species
groupingsfor classificationmust be assessedsubjectively.
Recent Americanwork has approachedproblemsof relationsof spe-
cies to community-typesand to one anotherthrough gradient analysis.
Standswere arrangedin synthetictransectsalong the moisturegradient
by weighted averages of ecological groups (Whittaker, 1951, 1954b,
1956, 1960; Curtis and McIntosh, 1951; Brown and Curtis, 1952;
Curtis,1955; Bray, 1956), and also along the elevationgradient (Whit-
taker, 1956). Some resultsof significanceappeared: (1) Specieswere
not, in general, presentor dominantover a given span of the gradients,
and absent beyond an abruptboundaryfrom the rest of the gradients.
Most species populationstaperedgraduallyand continuouslyfrom max-
imum density to scarcityand absence. (2) The most frequent distribu-
tional pattern was a curve suggesting the bell-shapedbinomial distri-
bution (Whittaker, 1951, 1954b, 1956; Brown and Curtis, 1952; Bray,
1956; see also Gause, 1930; Pennak, 1942, 1951), although various
modificationsof this pattern also appeared (Whittaker, 1956, 1960).
(3) Speciespopulationswere not arrangedas groups of associatessep-
arated from other groups along the gradients. Distributional curves
were most varied; and distributionalcenters and limits were scattered,
ratherthan grouped, along the gradients.
Species populations taper in the same fashion, and show the same
diversity of form and scatteringalong the gradient, in many of the
tabulationscited above as evidence for vegetationalcontinuity.Whit-
taker (1956) charted species distributionsin the form of population
nomogramsin relation to both moisture and elevation gradients. The

same scatteringand diversityof patternwas evident in the two-dimen-
sional treatment.Speciespopulationswere grouped by location of their
distributionalcenters; but the species thus grouped always differed in
distribution,and the groupings were regardedas arbitrary.The possi-
bility that some species populations might form "natural"clusters-
clustersof low degree, involving speciesbound by no obligate relations
and distributed quite differently apart from the occurrenceof their
centers close together-was also suggested; but the evidence in sup-
port of this possibility was limited (Whittaker, 1956, 1960).
The principle of species individualitycannot imply (as some critics
seem to have believed) that species are distributedindependentlyof
environmentalfactorsor without relationto one another.Neither need
the principle imply that the centers of species distributionsare scat-
tered quite at random in relation to environmentalgradients. Several
studies (Nash, 1950; Webb, 1950; Vries, 1953; Vries et al., 1954;
Dammanand Vries, 1954; Iversen, 1954; Whittaker,1956) suggestthe
existence of some naturalclustersor "constellations"of species. These,
and the correlationsof low degree which express them, do not conflict
with the principle of species individualityor imply that species are in
general organized into definite groups correspondingto community-
units. Grantingthe possibilityof limited "clustering,"one may observe
that all lines of evidence discussed-geographic distributions,distribu-
tions throughseriesof compiledtables,studiesthroughdominance-areas
and ecological spectra, statisticalcorrelations,and gradient analysis-
yield results consistentwith the principle of species individuality.It is
suggestedthat the weight of evidencefrom speciesdistributionsis heav-
ily on the side of this principle.
DYNAMIC RELATIONS OF SPECIES
The author has often heard it said that species individualityis in-
compatiblewith the "web-of-life."In a naturalcommunity,each species
interactswith many other species, and these in turn with still others,
so that all the species of the communitymay conceivablybe related
through the web of direct and indirectinteraction.The species popula-
tion exists in relation to other populations;its context of life includes
its position in relation to communitystructureand function (niche)
as well as its relation to other environmentalfactors (habitat) and
geographic range (area). The species population can hardly be said

to be "individualistic"in the sense that it occurs apart from, or is
distributedwithout relation to, other species populations. If both the
principle of species individualityand that of interrelationin the web-
of-life are accepted as essential parts of synecologicalunderstanding,
they may constitutea paradox.
Discussion of dynamic relations of species is handicappedby the
at present limited understandingof population dynamics, especially
in relationto plants, and by lack of effective, quantitativeevidence on
the extent which species are dependent on one, or several, or many
other species. One may inquire, however, on a rather informal basis,
if the relations which make up the web-of-life are actuallyof a kind
and degree that would imply organizationof species into sharply-de-
fined, community-widegroups of interactingassociates.One may first
distinguish associationin the distributionalsense, of relative similarity
of distribution,from associationin the dynamicsense, of actual inter-
action between species. The latter may be thought the basis, in part,
of the former. Five degrees, continuouswith one another,may be sug-
gested among dynamicassociationsas they relate to distributionalasso-
ciation:
1) Full dynamicassociation:species A can exist only in the presence
of species B, the only other species which can provide its food (or
shelter, etc.).
2) Partial dynamic association: species B provides some need of
species A, but other species can also supply this need.
3) Indifference:species A and B do not interact,and the presence
of species B tends neither to favor nor to exclude the presence of
species A in the community.
4) Partial dynamic dissociation;presence of species B in the com-
munity may reduce the population of species A. A may occur in the
presence of B, but may tend to be more numerousin the absence of
B if other factors are favorable.
5) Full dynamicdissociation:species A cannot exist in the presence
of species B (which sometimes cannot occur in the presence of A);
dynamic exclusion affects one or both.
Full dynamicassociationsare familiar in certainparasiteswhich are
restrictedto a single host species, certaininsectswhich feed on a single
plant species. Sometimes, species A and B may each depend on the
other (mutualism). If full dependenceexists in both directions,neither
species can exist in the absence of the other, and a perfect state of
species associationand distributionalidentity occurs.Such perfect asso-
ciation of species must, however, be relatively rare. Much more com-
monly, species A is dependent on B, but B is not dependent on A,
and distributionalsimilaritymay or may not result. Species A cannot
occur in the absence of B but will not necessarilyoccur throughout
the local and geographicrange of B.
In the second condition, several species may supply the needs of
species A. A parasite may utilize as hosts several related species; a
plant-eatinginsect may feed on a few species, closely related or taxo-
nomicallyscattered,or may graze on almost any availableplant regard-
less of species. Vertebrateherbivoresmay feed with little regard for
species as such, though with selection for palatabilityand with prefer-
ence for some largergrouping-as for grasses,for forbs, or for shrubs.
Predatorsprobably in general take available prey without regard for
species as such, though with limitations of size, possibilityof capture,
and sometimespalatability.A similar lack of species restriction,within
generallysuitablefood properties,appearsamong many scavengersand
saprophytes.A range of degreesof partialdynamicassociationmay thus
be observed-from the parasitewith a few closely related host species
to the grazer, predator, scavenger, or saprophytewhich is truly in-
different to the species of its food source. In no case is there reason
for distributionalidentity of species A with any particularone of the
species B ... N on which it depends, nor will the range of A neces-
sarily correspondto the whole range of all the species B . . . N.
It appearsthat these partial dependencesare much more numerous
than total ones. Not only do predators,scavengers,saprophytes,and
vertebrategrazers (and also epiphytes) mostly fall into the partial
grouping; but many parasitesand most smallerherbivorousanimalsdo.
There may be evolutionary reason for the preponderanceof partial
dependences over total ones. The larger the number of species that
fulfill the needs of A, the larger population and wider range it can
maintain. Furthermore,the less species A is dependent on a single
other species B, the less it is subject to the vicissitudes affecting the
population of B, to reduction of its population below critical levels
when the population of B is at a low point, to extinction if B be-
comes extinct.
Partial dynamicassociationsof low degree grade into indifference,
in which it is of no importanceto A whether or not B is present. The
herbs of a forest floor may be dependentupon the existenceof a forest

canopy and soil; but they are not dependentupon, or necessarilyasso-
ciated with, any particularspecies of canopy tree. Species A, fungal
parasiteof an herb, is not affectedby presenceof species B, an insect
feeding on a tree without detrimentto the tree. Possible indirect rela-
tions can be imaginedbetween almost any two species of a community;
but it may be stretchingthe significanceof these relationsto argue that
they should result in distributionalassociation.
Competitiverelations are familiar examples of partial dynamic dis-
sociation.SpeciesA and B are competitors;in the rangeof environments
more favorable to A than B the population of B is reduced, and in
environmentsmore favorableto B than A, A is reduced.Probablythe
distributionsof most plant species are affected by competition (Ellen-
berg, 1952b, 1954b, 1956; Kruckeberg, 1954; Whittaker, 1956);
but there is no reason competitive relations should organize them
into associatedgroups. Species should be expected to distributethem-
selves in relation to competition, each according to its own ability
to compete with other species in the various environmentalconditions
it encounters-i.e., individualistically.Conditions in which species A
is preyed upon by B may also imply partial dynamic dissociation.
The population of A is reduced by the presence of B, and A may
thus tend to be more numerousoutside the range of B. These relations,
again, do not imply distributionalsimilarity.
Full dynamic dissociationimplies that species B excludes species A
from the community. Some results from theoretical population dy-
namicsof animalssuggest the existenceof discontinuitiesof exclusion.
Where two competingspecies occur along a continuousgradientof en-
vironment,a dynamic"breakingpoint" may exist, where one is abruptly
replacedby the other (Gause and Witt, 1935; Gause, 1936; Hutchin-
son, 1950:372). A few discontinuitiesbetweenanimalpopulationsthat
are closely similar ecologically, and hence in direct competition,have
been demonstratedin the field (Beauchampand Ullyott, 1932; Hair-
ston, 1951; Mayr, 1948:214-215). In the evaluationof these it must
be observed: (1) The numberof such observationsis strikinglysmall,
in relationto the extent of biogeographicand distributionalstudy. (2)
Overlapping distributionsof species are apparently-very much more
common than sharp boundariesof exclusion. (3) A discontinuitybe-
tween species A and B need not imply correspondingdiscontinuitiesfor
other species of the communitiesin which they occur. It is significant
that Hairston (1951) found that the line separatingtwo competing

salamander species did not correspond to any vegetational discon-
tinuity; the discontinuitywas a discontinuityof salamanderpopula-
tions, not of naturalcommunities.
Some cases of mutual exclusion of vegetation dominantsof different
growth-formsmay be suggested-as a dense cover of prairie grasses
may (along with fire effects) retard the establishmentof forest tree
seedlings, whereas the shading and root effects of the trees exclude
prairie grass species from the forest. When such physiognomictransi-
tions are analyzedin detail, however, it may be found that species are
distributeddiverselyin relationto the steep environmentalgradient of
the transitionand other communities(Whittaker, 1956; Bray, 1956).
Furthermore,many physiognomictypes intergrade,so that mutual ex-
clusion of plant growth-formsmust also be thought a special instead
of a general circumstance.Although competitive discontinuitiesmay
affect some plant species, it seems clear that differencesin vegetation
are not, in general, produced by mutual exclusion along abrupt dis-
continuities (Whittaker, 1956).
The relationsof full dynamicassociationand full dynamicdissocia-
tion might provide the requisites for organizing species populations
into community-units.The importanceof these full relations, in the
whole plexus of relations in the community,is at present a matter of
judgment. It may be suggested that, if all possible relationsof species
in a stand could be listed, classifiedby degree and arrangedin a fre-
quency diagramalong a gradient from full dynamicassociationto full
dynamicdissociation,then the curve would have its peak in the middle
(indifference) and would taper throughthe partialrelationsto the full
ones. It is also suggested that full dynamicassociationand mutual ex-
clusion of populationsare not sufficientlygeneral phenomenato result
in organizationof most species into associated groups that are dis-
continuous from other such groups.
Distributionalassociationthat does not involve interactionmay also
be considered. Two species occurring in the same habitat without
direct interaction presumablyhave ecological requirementswhich in
some respectscorrespond.They must also, however,have environmental
requirementswhich differ in some respectsif they are not direct com-
petitors. Correspondenceof their distributionsis a coincidenceof the
physiologies of two species responding to various environmentalfac-
tors, some of which may be the same, others different.If severalspecies
are not dependenton one another,scatteringof their centersand limits

through environment, rather than formation of well-defined associa-
tions, is to be expected. Species that occur together in some communi-
ties without actual dependenceare "free" to occur separatelyin other
communities,and can usually be observed to do so.
Du Rietz (1921; Du Rietz et al., 1920) has suggestedthat associa-
tions represent combinationsof species that have become selectively
fixed; Allee et al. (1949) and Dice (1952) have consideredwhether
species do not evolve as supra-organismicentities, as associatedgroups.
Any given species evolves in a context of the communitiesin which
it lives, and communitieschangethroughevolutionarytime. The species
must evolve, if it is to survive, to meet changes in its circumstances
resultingfrom the evolution and migrationof other specieswith which
it interacts.But full dynamicassociation,requiringa species to evolve
in relation to another single species, is an exceptional, rather than
general situation. Most species are, in evolutionarytime, "free" to
change their associational relationship with other species; and they
do so (Mason, 1947).
It is believed that, while relationsof full dependenceand of mutual
exclusion do exist, relationsamong species are not in general of a kind
and degree necessaryto organize species into well-definedassociations.
This answer may be offered to the paradoxat the heart of the associa-
tion problem: Species populations are distributed individualistically,
but within a web of interactions;independentlyin the sense of distri-
butional diversity,but not independentlyin the sense of unrelatedness.
The paradoxis not a contradiction,and to the extent that it is a para-
dox may simply be accepted as a superficialand apparentconflict of
two major ideas in synecology. In relation to the evidence discussed
above, it is concludedthat (1) Species should be expectedto distribute
themselvesdiversely,each accordingto its own genetic pattern,physiol-
ogy, and relationsto physical and biotic environment,as stated in the
principle of species individuality. (2) Populations of most species
(apart from cases of mutual exclusion and "plateau" distributions,
Whittaker, 1956) may be expectedto taperalong environmentalgradi-
ents toward increasinglyunfavorable conditions, as a basis for com-
munity continuity. (3) The freedom of combinationof species with
one anothershould (though it is relativeand not complete) imply that
many stand combinations,intergradingin various directions,and only
with difficultyclassifiableinto units, will be formed.
THE BASIS OF CLASSIFICATION
PREMISES
The basis of classificationof natural communities can, no doubt,

be interpretedin more than one way. Any interpretationof classifica-
tion in relation to properties of communitiesmust be fashioned by
an individual with his own evaluation of current understanding,his
own selection of that which is relevant, his own premises regarding
the classificatoryprocess. The present accountis necessarilyunderlain
by premisesthat include the following:
1. Classificationis a process,and problemsof classificationare better
understoodthrough study of this processthan through an abstractlogic
of hierachies(vide Woodger, 1937:42-7, 1952; Gregg, 1954). Classes
are always, whatever the extent to which they are suggested or deter-
mined by characteristicsof that which is classified, human creations,
products of the classificatoryprocess.
2. Classificationis a joint phenomenon of the classifier and the
classified;the classificationdevelops by a complex and continuing in-
teractionbetween the classifying scientist and the object of classifica-
tion. This process of interactionmay well be approachedas one of
"transaction"in the sense of Dewey and Bentley (1949).
3. A realisticaccountof the process should alwayskeep in view, as
one focus of the process, the classifying person with his background
of experienceand interpretation,his purposesand prepossessions.
4. The other focus is the object of classificationand its properties
affectingclassification,so far as understoodat a given time. Analogies
from other fields involving propertiesof other objects of classification
are usually misleading to some extent and can at best make minor
contributionsto understanding.
5. The logical startingpoint for the present accountmay be neither
with the productsof classificationnor with those of analysis,but with
the original, direct contact of the classifierand the classified.2
This contact occurs when the classifying ecologist is in the field
observing natural communities.It is further consideredthat two ob-
jects or levels of observationshould be considered-the stand, the par-
2 Ideas in this account outside the subject matter of ecology are derived from
many sources, many of which cannot be cited. The author has been in-
fluenced especially by well-known books of Dewey (1916), Whitehead (1925),
Carmichael (1930), Northrop (1948), Dewey and Bentley (1949), Langer
(1953), and Oppenheimer (1953).
ticular community in which the ecologist may (if a terrestrialcom-

munityis in question) stand and take notes, and the landscape,the com-
plex or patternof more or less diverse stands, as observedfrom a hill-
top. These two views of naturalcommunitiescomplementone another;
by considering both in conjunction,together with other observations
which relateone to the other, the particularsubjectmatterof synecologi-
cal classificationcan best be understood.
THE STAND AND THE LANDSCAPE
An ecologist walking through forests, or across prairie or desert,

observes continuallythe naturalcommunitieshe encounters.The "ob-
servations"are themselvesparts of the streamof perceptions,the sen-
sory continuum,of the observerthat, becauseof their interest to him,
may be made note of or remembered.No observationsdifferentiate
from the sensory continuum except as the observer's interests (in
relation to such propertiesof the observedas conspicuousnessand fre-
quency) determine that he shall take note of certain things and not
of others. The observer'sparticularobservations,from among the in-
numerable observationshe might make, further determinehis choice
of certain stands or communitiesas objects of study. From the many
stands that might be chosen for classification,only a few are chosen;
the basis of this choice is usually complex and involves both qualities
of the communities,as these influence the observer,and various sub-
jective factors of the observer'spurposes and interpretationsof the
communitieshe sees. In the multiplicityof standsin a given landscape,
and the multiplicity of observablepropertiesof each of these stands,
every observationis contingent on the observer'sselection of the ob-
served.
Certain stands are chosen for more detailed observation and for
classification;for each of these the ecologist recordsas a stand-sample
or releve some characteristicsof the communityand certain factors of
environment.The sum total of the factors of environment,observed
and unobserved,in relation to which organismsof a communitylive
is conceivedas the environmentalcomplex. It is impossibleto measure
everything in the environmentalcomplex; it is impossible to express
everything,including the many interrelationsamong factors and their
patternsof variationthrough time (cf. Etter, 1954). As a totality the
environmental complex is unknowable and inexpressible. Since the
factors are not discreteunits which may be added togetherto form the
whole, and since the interrelationsof factors are essential properties
of the environmentalcomplex, this is not in any real sense a "sum
total" of individual factors. It may be better to regard the factors as
isolates from the whole, chosen for study in part accordingto relative
ease of observationand measurement,in part accordingto interpreta-
tions of their significancewhich have developed through some genera-
tions of study of correlationsbetween environmentsand communities.
The living communityitself is similarlybeyond total knowledge and
complete representation.It is normally impossible to inventory com-
pletely the organisms in the community;it is even more clearly im-
possible to determineand representall the interrelationsamong these
organisms. From the very many populations that might be sampled
and communitypropertiesthat might be describedor measured,some
are chosen. Only a few questions can be asked about a community,
only a few measurementsmade, in the time availableto studyone stand
among many; the observer'sknowledge of the communityis necessarily
determinedin large part by the questions that occur to him to ask.
The fragmentarynature of observationsmay be further viewed in
relation to the functional characterof communities.The stand and its
habitat form together a functional whole, the ecosystem or nature-
complex, in which matter and energy are transferred between en-
vironment and organisms. The communityis an open energy system
in which constantbinding, utilization,and dissipationof energyunder-
lie the steady-stateof the whole; constant flux and activity of parts
underliethe persistentpatternof the whole. The communityis a system
of interactingpopulationswith constant death and replacementof in-
dividuals, and constant fluctuationof populations around a more or
less persistent average level. Even the pattern and average levels of
populationschange, rapidlywith seasons,less rapidlywith successional
processes,and still more slowly with physiographicand climaticchange.
Communitystructureand function are inseparable;one may say that
structureis the basis of function, but also, and perhaps with more
point, that function or process is the basis of whatever structureor
form may develop. It is reasonableto emphasizethe process character
of that which is observed,and to regardthe communityas not so much
an object as an event. But, for purposes of classification,it is usually
possible to record only somnedetails of structure,and little indeed on
function.
The relevant context of an observationby an ecologist in a stand

includes the structural-functionalwhole of the ecosystemand the land-
scape of which it is part; it includes also the history of the ecosystem
and its probablefuture development,and the background,viewpoints,
and purposesof the ecologist. In this context a stand-sampleor releve
is recorded, as an abstractof some data which the ecologist regards
as most pertinent for characterizationand classificationof the stand.
Although the ecologist may later indicate certain stands in the field
as an extensional definition of his community-type,only certain ab-
stractionsfrom these stands can be manipulatedin trial classifications
and grouped into classes. The ecologist actually classifies,not stands,
but his conceptions of stands as these are derived from his observa-
tions and, often, supportedby stand-samples.If a Russian, a French,
and an Americanecologist, with their differentviews of the questions
to be asked about natural communities,worked in the field together
and chose for observationthe same series of stands, they would in the
end be classifying not the same set of stands, but different sets of
conceptionsof stands.
When a landscapeis viewed from a high point, three featuresof the
landscapemay at once be apparent-its diversity,patterning,and pre-
vailing characteristics.That the landscapeshows diversity,or differen-
tiation into varied habitatsand stands,may usually be recognizedfrom
the appearanceof differentvegetation types and their correlationswith
kinds of habitats.In this correlationand in the relationsof habitatsto
one another there is a degree of orderliness;the landscape shows a
characteristicconfigurationor patterning.Much of this patterningin-
volves gradientsof environmentsand communities,vectorialpatternin
the sense of Hutchinson (1953). And in this pattern of more or less
diverse stands and habitats one may note, while comparingthis land-
scape with others, certainprevailingfeaturesthat characterizethe land-
scape as a whole.
The landscape is sometimes regarded as a mosaic of habitats to
which correspondsa mosaic of natural communities (Tansley, 1939:
216); the landscapemay then be regardedas the sum of these mosaic
pieces. But habitatsare not discreteunits; althoughsome discontinuities
occur between them, they are also to a considerableextent continuous
with one another. The properties of individual habitats are, further-
more, determinedby processes which shape the pattern of the land-
scape as a whole. Propertiesof a given habitat are to be understood
not in isolation, but in relation to other habitats,as local climate and

soil propertiesin a given habitat are affected by topographicsituation
and movement of air, water, and soil materialsin relationto the land-
scape surface.The landscapeis not so much a mosaic of discretehabi-
tats and stands, and the sum of its parts, as it is a coherentpatternof
interrelatedhabitatsand stands. Since stands and habitatstogether con-
stitute ecosystems,the landscapeis a pattern of ecosystems,or an eco-
systemicpattern.As such it is a patternor field of ecosystemicprocesses.
In addition to the process characterof the local ecosystem,one may
recognize the role of interrelated climatic, hydrologic, geomorphic,
and ecologic processes as these determine the characterof the land-
scape with respect both to its differentiationand to its over-all or pre-
vailing characteristics.One may reject the Clementsiansystem for in-
terpreting vegetation, yet grant the significanceof Clements' (1916,
1936) insight into vegetation process and the unity of the landscape.
On the one hand, in seeking a synthesizedunderstandingof a land-
scape, one may emphasizethose conceptsthat permitmaximumintegra-
tion of information,the concepts of ecosystemand landscapepattern.
On the other hand, most quantitativeresearchmust deal not with the
complex whole, but with propertieswhich are, in polar contrastto the
preceding, to the maximum extent simplex, specifiable, and measur-
able. The analytic approachmust be based on such measurable"iso-
lates" or "simples" as environmentalfactors, species populations,and
certainpropertiesof communities,and on the relationsof these to one
another.
As a basis for analytictreatment,one may conceivethe environmental
aspectof the landscapeto be a complex patternof factor-gradients,ex-
tending through space in various directions in relation to the topo-
graphic pattern,some tending to run parallel in correlatedgroups and
others not. Different intensities of these factors are combined in the
most variousways into the environmentalcomplexesof habitats.Since
the landscape is largely occupied by living communities,the pattern
is one of factors isolated from ecosystems,and most of which are con-
ditioned by the biological, as well as physical, aspects of ecosystems.
The patternis further complicatedby discontinuities,factors and proc-
esses (such as soil parent-material,human disturbance,and fire) which
may not form gradients, and the time dimension of successionalde-
velopments of ecosystems.
At each point in the pattern individuals of a species may occur
CLASSIFICATION OF NAT'URAL COMMUNITIES 107
if that species has reached that stand through processes of dispersal

and if the total, or ecosystemic,environmentalcomplex is such that
individuals of that species may live there and, perhaps, reproduce
there and maintaina population for some generations.No two species
are quite alike in their distributionalresponse to the varied environ-
mental complexesof the landscapepattern.From the total biota, or the
species-garniturein the sense of Schmid (1942, 1950), the species
are combinedin diverse ways into local stands in relation to local en-
vironmental complexes. The landscape is also a complex pattern of
species populations,and of gradientsof species populationsin relation
to gradients of environment.
The species composition of a stand and its relation to habitat are
best understoodnot through study of that stand alone, but in relation
to other stands in the landscape.Each species of the landscapebiota
may have indicatorsignificancein relation to a given habitatby virtue
of its presenceand importancethere, or of its absenceand the distance
of its populationalong environmentalgradientsfrom the conditionsof
that habitat. Species populationsare to varying degrees shared by dif-
ferent stands of the landscape.Seeds and other disseminulesfrom one
stand are carried into others; mobile animals range through many
stands, and their habitatsmay be not so much individual stands as the
landscape,or a considerablepart of it. As there may be little in a par-
ticular ecosystemthat is wholly irrelevantto any other part of it, so
there may be little in the naturallandscapethat is irrelevantto a given
part of it. Viewing local ecosystemsin relation to the landscapeas a
whole, there is a profound quality of interpenetrationof factors,proc-
esses, and populations through habitats and stands.
All stands exist in relation to environmentalgradients along which
propertiesof stands change, slowly or rapidly, toward those of other
stands. Often the stand has no boundarybut gradatescontinuouslyin
several directionsinto other stands. Rarely does a stand appear quite
homogeneous.Sometimesthere is internal patterning,as in relation to
microreliefor frost processes;usually there is patchinessor irregularity
in the distributionsof populations which is not easily related to en-
vironmental differences;often some differentiationin relation to en-
vironmentalgradients may be observed even within the limited area
of that which is interpretedand sampled as a stand. Homeogeneity
of stands has been regarded as a most essential requirementfor the
possibilityof classifyingvegetation (Nordhagen 1928), but it is doubt-
ful that stand homogeneitycan exist except in a relative sense (Good-

all 1954a). Stand homogeneityin a strict sense is probablyuntenable
as a premise of communityclassification.It is in any case unnecessary
to assume more than that propertiesof some points in the landscape
which are to be classifiedcan be determinedby observationof certain,
limited areas which are not excessively heterogeneous.
The concept of association-individualof Pavillard (1912, 1935a)
and the school of Braun-Blanquetis consequentlyinappropriate.As a
term, association-individual can be used without prejudiceas a synonym
for stand. This particularexpression seems, however, to imply: (1)
That the associationratherthan the stand is primary,as if to say that
the class or type of stands has a prior and independent reality, and
exists before it appearsas an abstractionfrom stands in an ecologist's
classifying activity. (2) That all stands are members of associations,
even when the associations in question are unknown-presumably
because stands form natural groups with negligible transitions,rather
than being related to one another through all degrees of relative
similarity. (3) That the stand is a distinct, discrete "individual,"uni-
fied by its homogeneity and discontinuouswith other stands. To the
extent that a landscape has been modified by man into fields and
patches,each uniformlytreatedand relativelyhomogeneousand sharply
separatedfrom others, this conceptionof a mosaic of stand-individuals
may seem tenable; but it is scarcelyan appropriateconceptionin gen-
eral. It may be postulatedthat the stand has realityapartfrom observa-
tion, as a basis of the interactionof observerand standsand the progress
of understandingthrough continuing study of stands. But it should
not be postulated that this reality of stands entails or implies homo-
geneity, or discretenesscomparableto that of an individual organism.
The associationis no doubt a "reality"in its function in human classi-
ficatoryactivity, but should not be postulated to have reality outside
its role as concept, class, and symbol in human activity.
Instead of "association-individuals"one may recognize as major
objects of classificationon their levels: (1) The landscapeas an eco-
systemicpatternof usuallyevident heterogeneity,occupyinga consider-
able area, and classifiedby some definitionof its general, characteristic,
or average properties. (2) The stand (or the local ecosystem,micro-
landscape,or nature-complex)as a limited area of the landscapepat-
tern, classified by any of its recognized or measuredproperties. Al-
though stand properties must generally be determined from definite
areas, essentiallyselected points in the landscapepattern are classified.

(3) The community-fraction(synusia, micro-community,etc.) of one
or a few species relatedby life-form, stratum,taxonomy,micro-habitat,
or interactionwhich may be chosen for classificationapart from the
rest of the stand.
The above has been written as if only terrestrialcommunitieswere
in question, but may apply to aquaticcommunitieswith modification
in detail. The water body correspondsto the landscapeas a complex
of interrelatedstands. The stands have even less "individuality"than
those on land; their interrelatednessin the movement of nutrients,
food, energy,and organismsis even more evident. The functionalunity
of the water body is correspondinglymore evident than that of the
landscape;and the term ecosystem is more commonly applied to the
water body as a whole than to its stands. Although stands in the ben-
thos may be approachedas areas,standsof the planktonoccupyvolumes
of water; and the treatmentof plankton in terms of stands is further
complicatedby the extremeinstabilityof planktonpopulationsand the
movementsof watermasses,so that a given planktoncommunitycannot
be sampled twice in the same place if, indeed, the "same" plankton
stand can ever be twice sampled.
ABSTRACT PATTERNS AND THEIR MEANING
The patternof a landscapeis, in its full detail, exceedinglycomplex.

It is generally impossible to interpret adequatelythe relations of spe-
cies and stands to one anotherand the landscapeby observationalone.
It is consequentlynecessaryto develop abstractrepresentationsof the
pattern, representationswhich show some relations of communities
and environmentswhich are most significantin the landscapepattern,
but show these in a form more easily comprehendedand apart from
the complexityof the whole.
The most familiarsuch abstractrepresentationis the ecologicalseries.
In the complexityof the landscapepattern, certain main-directionsof
vegetationaland environmentalchange may be recognized (cf. Meusel,
1940). Recognition of major correlationsof propertiesof vegetation
with differencesin environmentis originally direct and intuitive, but
is later influenced also by means of measurementand interpretations
of the significance of factors which, like those of the soil, are not
so easily observed.When a single gradient is chosen for study, stand
samples may be arrangedin sequence along this gradient to form an

ecological series and interpretedas a gradient of environmentsand
communities,an ecocline. Stands may be chosen and arrangedin rela-
tion to a single factor-gradient,but the ecological series shows their
relation not to a factor-gradientalone, but to a complex-gradientof
many correlatedfactor-gradients,or of characteristicsof environmental
complexes (Whittaker, 1956). Within the ecocline one may choose to
distinguishthe complex-gradientof environmentsand the correspond-
ing coenocline or gradient of communities (Whittaker, 1960).
Although the ecological series is an approachtoward isolation of
a factor and its effects, it representsthe variation in certain observed
properties of ecosystems as most or all of these change along the
gradient chosen for study. By the ecological series, characteristicsof
communitiesmay be correlatedwith factors of environment,but the
relation need not be assumedto be one of effect and cause. Environ-
ments and communities are coupled and interacting aspects of the
ecosystem;environmentacts not simply on the community,but in and
through the function of the ecosystemto produce observeddifferences
in community characteristics(Whittaker, 1954b). The relation be-
tween environments and communities in an ecological series may,
however, have these characteristics:(1) The environmentalgradient
exists and can be measuredapartfrom the presenceof the communities
along it. The gradient may thus be in a sense external to or separate
from the community,although the gradient as it affectsorganismsmay
be modified by the communityand the function of the ecosystem.(2)
The relation between the gradient and communitiesis consistent;simi-
lar communitiesare observedto occur in habitatshaving similar levels
or intensitiesof the gradient.(3) The normalcomplexitiesof ecological
relations, effects of other environmentalfactors, chance differencesin
communitiesat similar levels of the gradient, and effects of communi-
ties in modifying the gradient not correlatedwith the gradient, may
reasonablybe neglected or controlled by choice of area or stands to
be studied. (4) There is reasonin present ecological understandingto
think that the environmentalgradient has significancein relation to
the functions of ecosystems,such that differencesin the functions of
ecosystemsthat develop at differentlevels of the gradientare expressed
in observabledifferencesin communities.When these conditionsoccur,
the relation between the environmentalgradient and the gradient of
communitycharacteristicspartially approachesthe ideal of the "cause
CLASSIFICATION OF NATURAL COMMUNITIES 1li
and effect" relation (cf. Bunge, 1961). The synecologistin this area
of study is concerned in general not with cause and effect but with
correlations-variables which change together through an ecological
series and which are often interrelatedin the functions of the eco-
systemsalong the gradient.To some degree some of these correlations
approachthe specialcircumstancesto which designationof one gradient
as cause or independent variable and others as effects or dependent
variablesmay be appropriate(cf. Major, 1951; Whittaker, 1954b).
When several major gradients influencingcommunitycharacteristics
are recognizedin a landscape,stands may be arrangedinto ecological
series in relation to each of these. An abstractrepresentationof the
landscape pattern as a multi-dimensionalcoordinate system of inter-
secting ecological series results (Ramensky, 1930; Sukatschew,1932;
Ellenberg, 1950a, 1952a; Goodall, 1954a, 1954b; Whittaker, 1956,
1960; Bray and Curtis, 1957; Curtis, 1959). This general approach
to studyof landscapepatternsand other relationsof ecosystemsthrough
ecological series and abstractpatterns (or by formal statisticsof cor-
relationsand factor analysis) has been termedgradientanalysis (Whit-
taker, 1951, 1952, 1956). The term expresses the fact that this is an
analyticapproachto ecosystemsthroughmeasurableisolatesas variables,
and that the basis of relating stands to one another and a principal
objective of the approach is the study of interrelationsof gradients
of environment, species populations, and communityproperties. For
the techniques of arranging stands in ecological series or coordinate
systems, and by extension for the approachitself, the term ordination
(Goodall, 1954a; from Ordnzung, Ramensky, 1930) is also current.
Implications of such research for problems of classificationmay be
clarified through study of an abstract pattern based on two major
complex-gradients,using these gradientsas axes of a chart (Fig. 1).
Properties of the pattern representedby such a chart cannot be
directly identified with those of the landscapepattern. The chart is a
simplification of the landscape pattern; it omits from consideration
factors not fitting into the complex-gradientsstudied. Points in the
chartmay represent,not particularstands,but averageor most probable
stand propertiesat a given combinationof the gradients studied. The
gradients representedas continuouson the chart are frequently inter-
rupted by edaphic and topographic discontinuityand disturbancein
the field. The chartsummarizeschangesof standsalong the full extents
of gradientswhich may be somewhereobservedin the field by walking
VEGETATION
OFGREAT
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C MESICTYPE SEDGE-TYPS
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ennesee
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Fic.. 1. A chart of vegetation types in relation to elevation and

topography in the Great Smoky Mountains, Tennessee (Whittaker, 1956).
The vertical axis is the complex-gradientof temperatureand other factors
related to elevation; the horizontal axis is the complex-gradientof moisture
relations and other factors from moist or mesic situations on the left to dry
or xeric on the right, as affected by topographic position. A line drawn
across the chart, either vertically or horizontally, represents an ecological
series along a complex-gradientof elevation or topographic moisture rela-
tions. Species populationschange continuouslyalong such an ecological series
(see FIG.2); the vegetation pattern is consequently a complex population
continuum (with the partial exception of relative discontinuities between

grassy balds and white oak-chestnut forests, and beech forests and cove
forests). In this population continuum areas of dominance of major species
are used to define and delimit community-types as indicated. The chart is
based on plotting 300 vegetation samples in relation to elevation and topo-
graphic position. The bracketed types at high elevations (boreal or spruce-
fir forests and heath bald) are regarded as communities separate from the
"Eastern Forest System" pattern illustrated, and which occur in the north-
eastern half of the mountain range.
along an uninterruptedgradient. The proportions of different areas

of the chartwill correspondto the relativeareasof standsof those types
in the field only if specialprecautionshave been taken.
The abstractpattern represents,however, some most significantre-
lations in the landscape.Some relationsrepresented-relative similari-
ties of habitats and stands, continuous change in stand composition
and relativediscontinuities,if any, and relationsof species distributions
to one another and environmentalgradients-are preciselythose most
directlyrelevantto problemsof classification.Relative distancesin the
abstractpattern reflectrelative similaritiesand dissimilaritiesof stands,
on which their grouping may be based. To a community-typeor class
of standsobservedin the field correspondsan areaof the chart.Relative
extents of community-typesin the landscapemay be representedin rela-
tive areason the chart. By means of the chart is it possible to compare
vegetationpatternsfrom one area or climate to anotherby major com-
munities, their relative importancein the patterns,and their relations
to topographicenvironment(Whittaker, 1956, 1960). Thus the chang-
ing relationsof communitiesand characterof vegetationpatternsfrom
lower elevationstoward higher may be observedin Fig. 1.
It was observed by Ramensky (1924, 1930) that stand composi-
tion changes continuouslyalong an ecological series and that no two
species have the same distributionalong an ecological series. Treatment
in terms of abstractpatterns and study of species distributionsand
community characteristicsin relation to such patterns permit these
furtherobservationsbearingon classification(Whittaker, 1952, 1956):
1) Speciespopulationsare distributed"individualistically"and most
variouslyin the pattern(see also Fig. 2); at the level of speciespopula-
tions the patternis a complex population pattern.
2) Populationsof most species tapergraduallyin all directionsfrom
a center or point of maximumabundanceto scarcityand absence;and
the pattern is largely continuous, though relative discontinuitiesmay

also occur.
3) At the communitylevel the chartsmay be used to show patterns
of communitycharacteristics(e.g., productivity,stratal coverages,spe-
cies-diversity,life-form composition,physiognomyas measuredby pro-
portions of growth-forms,communitycompositionin terms of major
taxonomic groups, food-habit proportionsand evolutionarymodernity
of insect communities). For the most part, change in these commun-
ity characteristicsalong the gradients is also continuous.
4) No two of these communitypropertiesshow the same pattern
in relationto the patternas a whole. Furthermore,a given type of meas-
urementshows differentpatternsin the two or more groups of organ-
isms to which it is applied.
COMMUNITY UNITS AS CLASSES AND TYPES
A community-typemay be regardedas a class, a grouping of stands

which share some defining characteristicor characteristics.Thus all
stands in which eastern hemlock (Tsuga canadensis)is dominant (in-
cluding, say, at least 50 per cent of canopy stems) may form a class
or community-type,"hemlockforest," the place of which in the pattern
of environmentsand communitiesis indicatedin Fig. 1. Ideally a clas-
sificationmay be sought in which stands are grouped into classes that
are mutually exclusive and include all stands of the landscape,or all
but a limited numberof transitionaland anomalousstands. In terms of
the abstractpattern this implies division of the pattern into a number
of areas in contactwith one anotheralong lines forming their borders
and which include the whole area of the pattern.Usually the division
into classes is made primarilyby a single, chosen kind of characteristic
of ecosystems, such as species dominance or floristic composition,
emphasizedby a given school.
Implicationsof results from gradient analysis for classificationmay
be observed. A virtuallyunlimited numberof characteristicsof ecosys-
tems might be chosen as defining characteristicsof classesof communi-
ties. These characteristicsof ecosystemsdo not parallel one another;
and there are consequentlyinnumerableways of defining classes and
areas, and these and their areasof exclusion overlap with one another
to generatestill other classesin the mannerof a Boolean algebra.From
the unlimitedpossibilitiesone must choose definingcharacteristicswhich
may produce an appropriateclassification.Boundaries of classes of

stands and areas of the patternmust in general be arbitrarilydefined,
though some relative discontinuitiesoccur which may be chosen as
boundariesof community-types.No absolute criteriaof correctnessof
classificationcan exist, only relative criteria of appropriatenessfor a
given group of organismsor communitypropertywith which the classi-
fier may be concerned,or degree of correlationwith other significant
propertiesof ecosystems.As yet few studies of comparativeclassifica-
tion (Liudi, 1928; Kontkanen, 1950a; Miller, 1951; Goodall, 1953a;
Moor and Schwartz,1957) apply different definitions of community-
types to samples from the same landscapepattern to observe the dif-
ferent contentsof the classes in extension and the relativeeffectiveness
of the differentclassificationsfor the ecologicalconditionsof that land-
scape. But results of these, like those of gradient analysis, indicate
that different choices of defining characteristicslead to different pat-
terns of classification,to classes which do not have the same contents
in extension and which overlap one another in complex ways.
As an alternativeto a system of classes, one may seek to recognize
types of communitiesconceived as points of reference in the abstract
pattern, with the understandingthat most stands will be intermediate
to these points and the types. Thus Schmid (1950), rejectingthe con-
cept of associationsas bounded classes, interpretscommunity-typesas
abstract types or models recognized in the pattern of intergrading
communities in a lWdscape (cf. Poore, 1955b, 1956). It is further
significant in Schmid's approach that these abstract types are con-
ceived through characteristicsof whole communities or ecosystems,
rather than through a single kind of classificatorycriterion. Ideally,
this approachinvolves the recognition in the landscapeof a number
of community-gestalts,or whole-patternsof particularecosystems,their
numberbeing sufficientto representmost of the range of variationof
communities in the landscape, and to permit the grouping of most
stands around the type-conceptionseven though most stands are in-
termediateto them.
The underlying meaning of the latter view of community-types,
which may be relatedto the doctrineof "naturalkinds" (Keynes, 1921;
Russell, 1948; see also Goodall, 1954a; Tiixen 1955), is less simply
stated than the approach through bounded classes. A first, essential
part of the meaning lies in the network of correlationswhich relate
the differentparts of ecosystems;all these parts are interrelatedin eco-
systems as functional wholes. If an abstracttype is defined by several

suitably chosen characteristics,then it is likely that any two stands
sharing these characteristicswill be similar in other characteristicsand
in their whole-patternsof communitystructureand function. Although
abstracttypes may thus be defined by a few key or indicatorcharac-
teristics,the types may have much deeper biological significancethan
understood through the key characteristicsalone. A second, possible
but not essential, part of the meaning of types may lie in the natural
"clustering"of individuals-in this case, of stands. If one imagines an
abstract"space,"axes of which are, here, compositionalgradients of
communities,then the individuals may fall into a definite number of
clustersin this space, with relativelyfew individualsbetween the clus-
ters. Each clustermay be thought to representa favorablecombination
of interrelatedcharacteristicsand a recognizablewhole-patternor ges-
talt, usually definableby a few key characteristics.Although there may
be few individualsin the spacebetweenthe clusters,with less favorable,
or less stable or less frequently realized, combinationsof characters,
the clustersare to be characterizedmore by their centersor nuclei than
by their limits. A third, possiblepart of the underlyingmeaningof types
has been discussedby Keynes (1921:253) and Russell (1948:438-444)
in terms of "generator"properties.One may suppose that the almost
innumerableapparentpropertiesof any given object, such as an eco-
system, all arise out of a limited numberof generatorproperties.One
may further suppose that these generatorpropertiesform a finite num-
ber of combinations,each representinga natural type in which the
many other characteristics, and the patternof the whole, reflectthe gen-
eratorproperties,each of which may be characterizedby a limited num-
ber of key characteristicscorrelatedwith the generator properties or
combinationsof these. In some objects of study the types to be rec-
ognized will include some which are relativelydistinctiveand extreme,
as determinedby special intensitiesof certaingeneratorproperties,and
one or more which are "normal"and intermediateto (but continuous
with) the extremes, determinedby generator factors of which none
are of special intensity.
Ideal, "good" species, as these appearin some genera, may illustrate
discontinuousnaturalkinds. Each species representsa favorable,func-
tional patternof characteristics, an adaptivepeak in the sense of Wright
(1932). Each species is also a populationcluster,separatedfrom other
clustersby the relative rarityof intermediateindividuals.A finite num-
ber of genes forming a limited numberof combinationsof major sig-

nificance (though a large number of minor biotype variationswithin
each cluster) are the "generator"propertieswhich determineinnumer-
able characteristicsof individuals,but a definitenumberof major types
of individuals, or species. The ideal species is also a naturalclass. Its
members share a large number of characteristicsand are clearly dis-
continuous with the members of other species sharing other sets of
characteristics.Some of the characteristicswhich may be chosen to de-
fine these species parallel one another;they are shared by membersof
one species but distinguish these from members of other species. It
may consequentlybe possible for taxonomistsworking independently
to chose different defining characteristicsfor species, and to discover
that the classes produced by these defining characteristicshave the
same contents, of individual organisms,in extension.
Such ideal species as natural kinds and classes may contributeto
problems of classificationof communitiesmainly by contrast.No re-
lation of common descent, such as may be assumedfor individualsof
a species, can be assumed for a group of communities.No internal
mechanismcomparableto inherited genotypes determinesthat a group
of communitiesshall be similar; no mechanismcomparableto hybrid
sterility enforces the distinctnessof communitiesof one group from
those of another. Choices of different defining characteristicslead to
classes which differ in extension. In general, neither communitiesnor
species as membersof communitiesfall into a limited numberof dis-
tinct, naturalclustersseparatedfrom other such clusters,though some-
times there may be some tendency in this direction. If characteristics
of environmentare taken as generatorpropertieswhich in some sense
generate characteristicsof communities, then what is generated by
continuousenvironmentalgradientsin various combinationsis more a
complex population continuum than a definite number of types. For
the theory of classifying communities,the analogy of the ideal species
and the community-typeis thoroughlymisleading.
The properties of ideal species are, in fact, unlike those of most
objects which are classified, including organisms of many or most
genera. More usually encounteredare characteristicswhich show vary-
ing degrees of correlationand that vary continuouslyin differentdirec-
tions through the objects of classification.Types must then be recog-
nized by patterns of combinationsof characteristicswhich are sub-
jectively evaluated. Different characteristicsof differing relative im-
portance ranging from the essential to the nearly irrelevantmay con-

tribute to the definition. Since not all these are necessaryfor the as-
signmentof individualsto the type, individualsare recognizednot by a
fixed list of characteristics,
but by some sufficientcombination-a "quo-
rum"- of the definingcharacteristics(Hospers, 1953). Thus the associ-
ation may be defined by a list of species which representsits character-
istic species combination.Presence of any sufficientrepresentationof
these species may indicatemembershipof a stand in the association.As
stands with fewer and fewer of these species occur, there is no point
at which a boundaryof the associationis encountered,except that at
which an ecologistprefersto considerthatrepresentationof the defining
species is no longer sufficient.Although an objective of specification
of a characteristicspecies combinationis clarity of definition,this and
other approachesto definition of types which depend on evaluation
of a series of defining characteristicscannot escape vagueness in the
limits of applicationof a concept (cf. Kaplan, 1946; Hospers, 1953).
The class and the type are alternativeperspectiveswhich are not
clearly distinguished in ecological practice. Ecologists whose outlook
emphasizeslandscapesand ecosystemsare more likely to seek classifi-
cation in terms of types defined by a numberof characteristics,around
which communities may be grouped by subjective evaluation of the
representationof these characteristics.Ecologistswhose interestscenter
in a particularaspect of the ecosystemare more likely to seek classes
defined by certaincharacteristicsof that aspectand definitionsso stated
that stands fall clearly within or without the limits of a class. Neither
approachoffers a comforting solution to problems of classificationin
an area of complexly related but loosely related and intergrading
phenomena. If classificationis a primaryobjective, however, the ap-
proachin terms of more clearlydefinedclassesis likely to be preferred.
THE PROCESS OF CLASSIFICATION
A first step toward classificationis the observationof a landscape

by an ecologist, before he has recognized definite community-types
and usually before he has begun to take stand samples. In the land-
scape pattern there is a degree of repetitiveness;a stand of a given
sort in a particularhabitat is observedat one point and later, at other
points, stands which are similar are observedin other, similarhabitats.
This self-repeatingpropertyof the landscapeis one basis for the recog-
nition of community-types;the observermentallygroups together simi-

lar, rememberedconceptionsof stands. The other basis for provisional
grouping is relative distinctivenessof stands, as groups of stands are
distinguishedfrom one another, and as certain stands are seen as es-
pecially distinctive in relation to the rest of the pattern.Judgment of
similarityand dissimilarityof stands may be based at this stage on sub-
jective perceptionof over-all character,or patternor gestalt, of stands.
But the ecologist's conceptionsof stands, and the propertiesof stands
on which their grouping into preliminarytypes is based, are influenced
strongly by the observer'sbackgroundand interpretations.The basis
for provisional recognition of types is usually an informal balancing
together in the observer'smind of relativesimilarityand distinctiveness
of stands, in a context which includes his belief in the significanceof
certain communitypropertiesas bases of classification.
Such a process is, clearly, one of "intuition."Certaintypes of rela-
tions are directly perceived and summarizedin the ecologist's mind.
Some of these observationsinvolve the associationof species with one
another, and of species and propertiesof communitieswith those of
environment-i.e., correlations.Others involve the grouping of stands,
abstractionfrom their properties to a conception of an average or
"typical"stand of the group, and evaluationof degrees of significance
of differencesamong stands and between types of stands-i.e., pro-
visional grouping of samples, averaging, and considerationof devia-
tions and the significanceof differences.In its function here, at least,
intuition need not be a matter of mystery.It seems to function first,
through perception of similarityof stands through their total charac-
ters or through certainobservedproperties,second, as a subjectiveand
largely unconscious "statistics"of correlations,averages, and signifi-
cant differences.The result is the formation in the observer'smind
of conceptionsof community-types, abstractedfrom his conceptionsof
one or more stands.
From this community-typeconception, which may be diffuse and
based on many propertiesof communitiesand environments,can be
abstracteda formal class-concept,which may be simply and unambigu-
ously statedand effectivelycommunicatedto other ecologists.According
to the outlook on classificationthat has alreadyguided his grouping of
stands and conceptionof types, the ecologist may state a defining form
for his class: Standswith a closed canopyof deciduousbroadleaftrees
are membersof class (formation) A. Standswith speciesa dominantin
the uppermostclosed stratumare membersof class (dominance-type)

B. Stands containing at least n of the character-species. . . are mem-
bers of class (association) C. Stands with species a dominant in the
tree stratum, b in the shrub stratum, and c in the herb stratum are
members of class (sociation) D. The class concept is the intensional
definition of the community-type;those stands for which the defining
form becomesa true proposition,which thereforeconformto the class-
concept, constituteits extensionaldefinition.
The intensional and extensional definitions are coupled; they form
interdependentaspects of the concept-formingprocess as a transac-
tion of the ecologist and naturalcommunities,a transactioninvolving
also the ecologist's personal conceptionsof stands and of stand-types,
and often his recordedstand-samples,as intermediariesof the inten-
sional and extensionaldefinitions.Neither aspectof the definitionneed
be fixed; normallythey change in relation to one anotheras the ecolo-
gist progresseswith his classification.Thus reconsiderationof his ma-
terial may lead him to transfer samples 1 and 2 from associationA
to associationB, and both associationsmay then be more clearly con-
ceived and defined; but this transferrequiresthe discardingof species
a as a diagnosticspecies for associationA and permitsuse of species b
as a diagnosticspecies for associationB, while this change in definition
suggests that sample 3 might best be transferredfrom associationA
to another; and so on until the ecologist is satisfied with the order
into which he has brought his material.This reconsiderationof classes
and their definitionsmay be based on the manipulationof stand-sam-
ples; and the realismand effectivenessof the final classificationmay be
greatly enhanced by constant referenceto the factual materialof field
records. It is also true, however, that the basic pattern of the classi-
fication may already be determined when the ecologist takes those
samples to representhis preliminaryconceptions of types.
Thus classificationmay be progressivelyrefined and clarified;thus
in the history of the SouthernTradition there has been shift of em-
phasis in definitions from physiognomy and dominance, to constant
species, to diagnosticspecies, and continuingclarificationof the defini-
tions based on these, as experiencesin the field have led to changing
insights into the meaning and usefulness of criteria.Within the limits
set by a school's perspectiveat a given time, questionsare asked of the
vegetation by ecologists and suggested to ecologists by the vegetation,
answersto these questions and applicationsof the classificationwhich
accompanies them suggest further questions, the search for their

answers, and testing in application of modified classification.The
process of increasingunderstandingand improving classificationpro-
ceeds as a kind of continuing conversationof ecologists and natural
communities.No absolutestandardsdeterminewhat constitutes"prog-
ress" and "success"in classification;but it may none the less be pos-
sible to observe various relative standardswhich do so.
Classificationmust be based on criteriaof classes and must involve
choice of these criteria from among the many properties of eco-
systems. The basis for this choice is in part similar to that guiding
choice of indicators (Whittaker 1954b). The criterionshould, first of
all, be one relatively conspicuousin the communityor, at least, pos-
sible for the ecologist to observe and measure.But it is desirablethat
the criterionshould be not only easily observed,but of maximumsig-
nificancein relationto the ecosystemas a whole. Different criteriahave
differentrelativesignificances;some seem relativelytrivial or fortuitous,
others seem of high importancefor their strong correlationwith many
other propertiesof ecosystems.Choice of criteriathus involves various
combinations of obviousness or accessibilityof the criterion to the
ecologist, and effectivenesswith which other propertiesof the ecosystem
may be inferred from the one chosen, of salience and significance.A
third desideratumof classificatorycriteria is that they effectively de-
fine classesin the particularobject of study and at a given level of in-
tensivenessof study. Thus the accessibleand significantcriteriaof life-
form composition and growth-form dominance may be set aside for
a given study, the first becauseclasses are not easily defined through
it, the second becauseit does not distinguish the relativelynumerous,
narrowly defined classes desired. Choice of classificatorycriteriathus
involves balancing together of several desired properties, including
accessibility,significance,and effectiveness,in the context of the classi-
fier's purpose and perspectivesand the ecological conditions studied.
Even when the kind of criterion to be used is determinedin ad-
vance of a study, much freedom in definitionand delimitationof actual
classes remains.Speciesmay be variouslygrouped into character-species
groupings, formations variously defined by different distinctions and
combinationsof growth-forms,dominance-typesby differentcombina-
tions of dominatingspecies. Other considerationsdeterminewhat par-
ticular classes an ecologist may define by means of a chosen kind of
criterion.Some of them involve the ecologist's purposes,as these may
direct his attention toward particular communities and determine

whether more broadly or more narrowlydefined community-typesare
desired. A further, often potent, influence is that of precedent. An
ecologist tends naturallyto recognize those units which, on the basis
of previous work, he expects to find; and in a given school there is
a tendency for later authorsto be guided in large part by the system
of units establishedin earlier work even while modifying it in detail.
Apart from these factors of background,there is usually the desire
to present,through the classification,an adequatedescriptionof natural
communitiesof the landscape.Relative extent or repetition of similar
stands, and the relative distinctivenessof others, influence decisions
on formal definitionof types as they influencedprovisionalconceptions
of types. A sufficientnumberof community-typesshould be recognized
to representboth types of stands widely distributedin the landscape,
and distinctive stands of special and local conditions. Absolute com-
pleteness in descriptionof the landscapeis unattainable;but the classes
or types should be numerousenough and narrowlyenough defined to
represent (at least in subordinateunits) most of the variationsamong
stands in the landscapepattern which the ecologist judges significant.
The community-typesare usually grouped and subdivided so as to
form, when the classificatoryprocess is complete, a hierarchy.The
hierarchyhas differentfunctions for different ecologists, and different
objectives influence the final organizationof the classificationinto a
hierarchy.Some of these are primarily practical;the classificationis
a means of organizing and communicatingthe knowledge of natural
communitiesobtained,and the classificationitself may have continuing
usefulness for researchand application.Others are theoretic;the classi-
fication should express properties of and interrelationsamong eco-
systemswhich the ecologist considersof major significance.Still others
are essentiallyaesthetic;they involve the ecologist'spersonalsatisfaction
with the order into which complex materialhas been brought and the
comprehensionof the landscapethus made possible. It is a rare ecolo-
gist (e.g. Villar, 1929b) who makes explicit the aestheticaspectof his
approachto naturalcommunities;but most classificationsare no doubt
motivated by combinationsof practicalusefulness and the subjective
satisfactionof organized knowledge.
Hierarchiesare not inherent in landscapes;a given landscapeoffers
materialwhich can be fashioned into hierarchiesin innumerableways.
A hierarchymust be constructedby choice of classificatorycriteriawhich
possess the desired properties discussed, and which also so relate to

one anotheras to form classes (1) of the varied magnitudesof content
appropriateto different levels and (2) so defined that all stands of
a higher-level unit are included in one or more lower-level units, and
no lower-level unit includes parts of more than one higher-level unit.
From the overwhelming amount of information offered by the eco-
systems of a landscape, the ecologist must choose some kinds of in-
formation relatively accessible to him, significantin ecosystemiccon-
text, capableof being effectivelyinterrelated,consistentwith his view-
point and interests, and suitable to constructionof a hierarchy. A
school, an approachor system, representsa more or less judicious and
productive choice of kinds of information and ways of using them.
On the one hand, the informationbroughtinto a given systemis always
a narrow selection, limited and fragmentaryin relation to ecosystems
as wholes. On the other, a school'sapproachhas a real positive function
in sparing a student from flounderingin an unmanageablewealth of
information, guiding his study in one direction of relative economy
and effectiveness.
The basis of understandingand judging classificationscannot be
one of literal verisimilitudeor fidelity to nature.Ratherthan this, one
finds that classificationsdevelop in accordancewith whole systems
of interbalancedvalue judgments.A classificationfurthermoreexpresses
the development, up to that time, of the cumulativeand continuing
growth of understandingof significant relations of communitiesand
environments.The classificationmust be viewed as a culturalproduct,
understood in a context which includes both culturalvalues and eco-
logical conditions, and judged in its functional relation to present
understandingand practice.
CONCLUSION
Emphasis in this account of certain things-personal choice and

judgment, intuition and subjectivity,cultural influence and precedent,
values and aesthetics-runs counter to what is usually sought in an
account of scientific method. An objective of science is to create a
picture of phenomenain which these influencesare minimized,though
not excluded; this account involves no dissent from that ideal. The
question here, however, is less one of ultimate objectives of science
in general than what actually happens when natural communitiesare
classified.
It is often consideredthat certainmajor levels of scientificdevelop-

ment may be distinguished:the stage of classificationand description
("natural history"), that of empirical correlationand the induction
of particularrelationswhich do not yet fit into a coherent scheme or
permit accurateprediction, and the final stage of scientific maturity,
of exact knowledge and efficient prediction, and of organizationof
knowledge into a coherent,deductivesystem. Synecologyis, clearly,on
the earlier of these levels, more nearly comparablein this respectwith
some of the social sciences than with geometry or physics; and the
classificationof naturalcommunitiesis clearly,and without deprecation,
"naturalhistory." It is hopefully believed that the inductive sciences
will ultimately develop at least parts of their understandinginto the
deductiveform of exact science.The latter is the ideal, but the attempt
to simulate deductivestructurebefore the basis for it exists (see Egler,
1951, and Whittaker, 1957, on Clements) cannot producea systemof
lasting value. Application of methodology derived from other fields
and inappropriateto a given field at its level of developmentmay lead
only to frustrationand sterility. Prematureinsistence on rigor, objec-
tivity, and exactitudemay lead to a methodologybased on illusion, or
a stringentlimitationof the range of phenomenawhich can be investi-
gated.
It is, in any case, questionablewhether classificationof naturalcom-
munities should be termed "science."Scienceand art are not mutually
exclusive, and there may be more of art in science than first meets the
eye. This account has emphasizedfeatures of classification-subjective
balancing of values, translationof experience into varied, personally
satisfactoryclassificatorydesigns, the influenceof culture and personal
factors and self-influenceby precedentand tradition, the rise and fall
of "schools"with different views and techniques-more generally as-
sociatedwith arts than sciences.For the preponderanceof such features
in problems of communityclassification,one may consider that this is
less a science than an art instrumentalto other aspects of ecological
science.
The complex wholeness of the community,ecosystem,and landscape
have also been emphasized (cf. Egler, 1942) together with the fact
that these are not knowable or expressiblein totality. Becausethey are
not and are exceedingly complex, they must be analyzed-resolved so
far as possible into simpler isolates which can be measured,related to
one another and, sometimes,experimentedwith. The whole and these
parts are best held in complementaryview. On the one hand, increased

understandingof the whole must generally come not from contempla-
tion of the whole, but from continuing, laborious,controlled analytic
research.The holistic view and organismicanalogywere seen by Clem-
ents and Shelford (1939:24) as an "open sesame"to ecologic under-
standing, but as a substitutefor analyticresearchwere an open sesame
to an empty chamber.Analyticresearchyields never a total understand-
ing of ecosystems,but a large part of such understandingas is pos-
sible. On the other hand, analytic research carried out without the
whole in view may give results that lack relevance and significance,
and the researchworker may see in his analytic isolates a finality or
ultimaterealitythey do not possess.Recognitionthat the full complexity
of ecosystemsis unknowableis a matter, not of mysticism,but of per-
spective on scientific knowledge and its limitations.
The effectivenessof ecological classificationmay best be enhanced
not by the uncriticalquest for objectivity,but by as realisticas possible
an understandingof what is done and can be done, and its relation
to propertiesof the object of classification.A central objective of this
essay has been clarificationof both limitations and possibilities of
ecological classification.It is hoped that through this and the survey
of schools, classificatoryproblems of an individual ecologist may be
brought into better perspectivein relation to landscapesas wholes and
the possibilities for their study explored by others. Some of the con-
flicts of schools may be unnecessaryif it is understoodthat different
classificationsare human creations, serving different purposes, never
sufficient in themselves, and usually to some extent complementary
to one another. The whole of the landscapeis caught in the web of
no scientist'sclassification;nature in the field is never wholly compre-
hended in any system. Artemismay be admiredby many, but possessed
by none.
APPLICATION
PRINCIPLES IN THE CLASSIFICATION OF NATURAL COMMUNITIES
The preceding discussions suggest certain principles:

1. The ecosystemicconceptionsuggests a multi-factoralor landscape
approachto classification,in which factors of all aspectsof ecosystems
-physical environment, soil, vegetation, animal communities and,
when appropriate,man himself-are considered together. The land-
scape approachrequires,however, a difficultversatilityof the student

and forces him to make choices of what is to be emphasizedin dassi-
fication,choices that are somewhatarbitrary,must vary from one land-
scape to another,and may offer little basis for agreement.
2. More generallyuseful in practice,among ecologists themselves,is
the consistent definition of units or classes by a single aspect of eco-
systems. It is further appropriatefor most purposes to define units
for a given aspect by propertiesof that aspect-to classify vegetation
by propertiesof vegetation,animal communitiesby animalpopulations,
etc.
3. Becauseof the interrelatednessof aspectsof ecosystems,units in-
dependentlydefined for differentaspectswill be in some sense related;
but they will not in general correspondclosely to one another. Thus
independentclassificationof climates,soils, vegetation,and animalcom-
munities leads to classificationswhich may be related,but are different.
For some purposes, however, units for one aspect may be so defined
as to correspondmore closely to certain units of another, as climatic
regions may be definedto correspondmore nearlywith vegetationunits
defined by physiognomy.
4. Within a given aspect, many different factors or propertiesmay
be chosen as bases of classification;but it is generally desirableto base
classificationof naturalcommunitiesat a given level on a specifictype
of criterion.The more clearly specified the criterion,the more clearly
may the units be defined.
5. The different characteristicsof a single aspect which may be
chosen as bases of classification,through related, do not run parallelin
any simple fashion; and units defined by different characteristicsare
relatedto one anotheronly in complex and variouslyoverlappingways.
No "natural"units of ecosystemsare equally appropriatefor all as-
pects and all groups of organismsand can be independentlyrecognized
and defined by different approachesand criteria.
6. Even when a particulartype of criterionis specified as the basis
of classification,normally much freedom of subjective decision exists
on the units to be recognized (see below on associations,dominance-
types, and formations).
7. Rigorous adherenceto a single criterion will sometimes lead to
heterogeneousand incongruousgroupings. It may then be desirableto
subdividethe grouping on the basis of a secondarycriterion(see below
on formations and dominance-types).
8. When a single type of criterion is used, the whole context of

other ecosystemicproperties, so far as known, should be considered
in its use. For many purposesunits are defined by a particulartype of
criterion; but the possibilities for subjective choice within this and
use of secondarycriteriawhen needed are taken advantageof, to define
units that seem most appropriateto the ecosystemiccontext and to the
purpose at hand.
9. Natural communitiesare relatedto one anotheralong many, com-
plexly relatedgradientsof environment;and along these environmental
gradients communityproperties form gradients which, to a consider-
able extent, are continuous rather than step-like. Relations of com-
munities may consequentlybe understood,in the abstract,in terms of
a complex, multi-dimensional,and largely continuouspattern; in this
pattern community-typesrepresent areas or points defined by given
criteria.
10. Most units thus defined will form a complex mosaic on the
earth'ssurfaceas they variouslycontactone anotheralong environmental
gradients.Under certaincircumstances,however, when major environ-
mental factors are strongly correlatedwith one another and a spatial
gradient such as elevation or tide-level, the units may occurin a belted
or zonal arrangement.
11. Hierarchialclassificationof communities is not inherently ap-
propriateto their multi-dimensionalrelations.The principalalternative
is the relation of stands or community-typesto one anotherin ecologi-
cal series representingthe most significant gradients of community
variation,or in a multi-dimensionalcoordinatesystem.
12. Hierarchial classificationsare, however, often desired; and it
is possible to classify communitiesinto either consistentor inconsistent
hierarchies.Few community charactersare suited to the creation of
consistenthierarchies(based on the sametype of criterionat all levels);
only by the use of diagnosticspecies in the system of Braun-Blanquet
has this been done with real success. Inconsistenthierarchiescan be
produced by choice of criteria generally defining units of different
magnitudes for different levels of the hierarchy.In inconsistenthier-
archies a lower-level unit as first defined may include parts of two
higher-level units (see below on formations and dominance-types).
Such difficultiesin detail need not prevent successfulclassification,for
units can be re-definedto fit together in the desired manner.
13. A perspective of eclecticism and relativism is appropriateto
problems of communityclassification.Different approachesto classifi-

cation have differentadvantagesand limitationsand may serve different
functions. Acceptance of the approach of a given school should be
accompaniedby understandingof the merits of other approachesand
the mannerin which differentapproachesmay complementone another
in the study of ecosystems.
FORMAL CLASSIFICATION IN THE SCHOOL OF BRAUN-BLANQUET
From the many possible approachesto classification,only two can

be considered in details of application.These are the floristic system
of Braun-Blanquet,the major approachto classificationamong Con-
tinental phytosociologists,and the use of dominance-typesand forma-
tions, the major approach among English-speakingecologists. The
following sections will consider further the possibilities and limita-
tions of these most widely used means of classification.
THE METHOD
The essential choice of, or assumptionabout, the basis of classifica-

tion in the school of Braun-Blanquetis that communitiesare to be
classifiedby floristic composition. It is theoreticallypossible to classify
communities by comparisonof their whole species lists, or by such
measurementsas the coefficientof communityand percentagesimilarity.
In practice, however, some short-cut,using only some of the species
in samplesfor the definitionof community-types,is needed. The species
used are those relatively distinctive for community-types,either be-
cause largely restrictedto a given community-typeor becausethey oc-
cur in the stands of one, but not the other, of two community-types
being compared.These two groups of diagnostic species, the charac-
ter-speciesand the differenitial-species, are the essential means of clas-
sificationin the school of Braun-Blanquet.
A further characteristicof the school is the use of diagnosticspecies
to define vegetationunits which fit into a formal hierarchyfollowed by
all membersof the school. The units and the hierarchyare regarded
as relatively fixed and established, even though subject to continuing
modification;they are not freely produced and abandonedaccording
to the individual'schoice as are units used by ecologists. The units are
also more heavily emphasizedas the basis of the study of communities
than by most ecologists. The outlook of the school has been stated by
Tuxen (1937:3 translated, cf. Tiuxen, 1950a; Braun-Blanquet, 1932b,

195la:4-5): "The systematicphytosociologicalinvestigationof a land
can be divided into three main parts. The first problem, and at the
same time the prerequisitefor all further, must consist of the laying
down of the inventoryof all plant communities[i.e., community-types]
present through a sufficientnumber of critically compiled tables, to
bring them into a well-organizedsurvey. The next step comprisesthe
cartographicrepresentationof the distributionof the units established.
The last, also the most extensiveand most difficult,but most important
goal, remains the study of the life-conditions and developmentalpos-
sibilities of the vegetation units now known in composition,structure,
and distribution."It is a most fundamentalcharacteristicof the school
that classificationof communitiesis the focal point of vegetation sci-
ence, an end in itself and an essentialmeans to any other ends sought.
Details of techniquein the school may be found in such references
as Braun-Blanquet(1921, 1928a, 1932a, 1951a), Braun-Blanquetand
Pavillard (1922), Beger (1932), Melzer and Westhoff (1944), Ellen-
berg (1956), and Tomaselli (1956); these details cannot be reviewed
here. Poore (1955a, 1955b, 1955c, 1956) and Becking (1957) have re-
cently given valuable discussions of the system, which may be com-
pared with the present one. Accounts of the method leave certain
importantgaps. It has seldom been made clear how and why samples
are chosen, how samples are handled to permit recognition of com-
munity-typesand character-species, and how groupings of associations
into higher units are decidedupon. Techniquesof the school have been
learnedin Europeto a considerableextent by personalcontact;and the
works of Braun-Blanquetand others have so failed to make these
matters clear to ecologists that no ecologist has ever applied the sys-
tem as a whole.
The system requires extensive collection of stand-samples(releves,
Aufnahmen). Samples are in general, though not always, chosen to
representcommunity-types;and sample choice is affectedby subjective
factors including the phytosociologist'spreliminaryinterpretationsof
the vegetation,his conceptionsof units as influencedby both precedent
and the desire to recognize new community-types,his judgments of
typical or representativevs. atypical and mixed stands, of stable vs.
unstable ones, etc. Randomizationis not used for sample choice; sam-
ples are chosen to representthe range of variation in the vegetation
studied, but also in part to fit into a classificationin the manner of
the school. The choice generally is, and should be, frankly subjective
(Becking, 1957). The objective is not so much classificationof all
possible stand combinationsas these might be randomlysampled,as it
is developmentof a most effective classification,representingmost sig-
nificant features of vegetational and environmentalvariation and in-
terrelation,by a skilled practitionerrelying on his judgment and ex-
perience. Subjective sample choice is quite consistent with the ob-
jectives and the rest of the system.
When a stand is sampled, a list of its species is prepared.A sample
normallyincludes a more thorough floristicinventory,including lower
plants, than is customaryamong many ecologists. For each species
certain "analytic"charactersare recorded;the two most used are socia-
bility and a combinedcoverage-densityvalue. For these and other ana-
lytic charactersarbitraryfive-point scales exist; and the values are re-
corded for each species from visual estimatesaccordingto these scales.
The method is not, really, a quantitativeone (Braun-Blanquet,1951a:
52; Ellenberg, 1956:56); visual estimatesare used to supplementpres-
ence and absenceas a basis of judgmentin a primarilynon-quantitative
system. By means of these estimatesthe compositionof a community
can be recordedquite rapidly in a single visit, and considerablenum-
bers of stand-samplescan be collectedin a fairly shorttime. The charac-
ter of these samples is determinedby compromiseof the desire for
some quantitativedata with the need for economy and efficiencyin
sampling. The characterof the samples is wholly in harmonywith the
rest of the sytsem.
The phytosociologistwith such samples at hand faces a formidable
problem of organizingthis body of data. Statisticalanalysisof samples
is possible (Etter, 1948; Raabe, 1952; Dagnelie, 1960) but is not used
in most studies. Ideally, the samples may be studied, comparedwith
one another,grouped tentativelyin many differentways until the best
classificationemerges. Actually, many of the samples will usually have
been taken to represent community-typesalready established in the
system, or alreadyintuitively recognizedby the phytosociologist.These
samples are naturally grouped together, and diagnostic species to
characterizethe groupings are sought. Propertiesof habitats and com-
binations of dominants may suggest other groupings; and species re-
stricted to these groupings may then be recognized as indicators for
habitat-typesand as diagnostic species for community-types.If several
species are largely confined to a grouping, or are clearly less im-
portant in the rest of the sample material, these form a group of

character-speciesand the grouping may be an association. If there are
no character-speciesfor a grouping; but certain species are present in
samples of this grouping and absent or clearly less importantin those
of another, closely related grouping, these are differential-speciesfor
a unit usually of lower rank than the association.The comparingand
grouping of samples proceedsuntil the objectivesare realized-groups
of samples which are relatively homogeneous and relatively distinct
from other groups and which can be characterizedby diagnosticspecies.
Ellenberg (1956) describes an actual step-by-step procedure for
grouping samples. Stand samples are first compiled into a preliminary
table, in which they may be grouped by environmentalrelations,domi-
nants, or recognized diagnostic species and community-types.Presence
values are determinedfor all species, and the species are re-arranged
in new tables by relativepresence.Some species, especiallythose of in-
termediatepresencevalues, may now be recognizedto form differential-
species groups; and the species and samples may be re-arrangedin
additional tables to bring together the species of differentialgroups
and the samples characterizedby them. With still further re-arrange-
ment of species and samples, and entry of additionalinformation,the
materialmay be brought initothe form of a table in which provisional
community-typescan be marked out and given provisional names.
To determine character-species,the various lower-level units must be
brought together as columns of a survey-table,containing presences
or other average values for species in community-types.Distributional
relations of the species may now be surveyed;and, with necessaryre-
liance on the judgment and experience of the investigator,character-
species groupings for higher units may be established. In the final
compiled tables, species may be arrangedby character-species groups,
and samples and lower-level units (below the associations) arranged
by these character-speciesgroups and the higher units (association,
alliance, etc.) which they characterize.Ellenberg'saccountmay differ
widely from actualpracticeof some phytosociologists,but it has special
value in describinga procedurewhich can be comprehendedand ex-
perimentedwith by ecologists.
When the body of samples has been grouped into community-types,
"synthetic"charactersmay be determined.The most obvious charac-
ter to be determinedis constancy of species-the percentageof stands
representinga given community-typein which a given species occurs,
when equal areas of the various stands are compared. Since samples
in the school of Braun-Blanquetare not usually taken in fixed areasor
quadrats,the closely related measure of presence-the percentageof
stands in which a given species was observed, regardlessof the area
of observation,may be substitutedfor constancy.Constancyand pres-
ence are significant for their expression of the relation of species to
the community-type(influenced,however, by sample choice); but these
measurementsare only secondarilyconsideredin the school of Braun-
Blanquet. The most important synthetic characteris fidelity or ex-
clusiveness(Treue), the degree to which a species is restrictedto a par-
ticular community-type,or is centered in that type and of less im-
portanceor vitality in other types.
Fidelity, the distributionalrelationrequiredof character-species,may
also be estimatedby a five-point scale; but these estimatesare seldom
published in current work. Valid estimates of fidelity require both
intensive and extensive familiaritywith a flora; only from knowledge
of the distributionalrelations of each of the many species dealt with
can the relationsof each to a community-typebe judged. Quantitative
criteriafor assigning degrees of fidelity from comparisonof compiled
tables for differentcommunity-types exist (Szafer and Pawlowski,1927;
Agrell, 1945a; Braun-Blanquet,1951a:96; Guinochet, 1954; Becking,
1957) but are apparentlylittle used; fidelity judgmentsare in general,
as they must almost necessarilybe, subjective. Character-speciesneed
be neither dominant nor abundant;they may be among the most ob-
scure membersof communities.They are often poorly characteristicof
the community-typein a sense different from fidelity; they may be of
low constancyand present in a minor fractionof stands chosen to rep-
resent a community-type.It may also be observedthat the distribution
of a species througha series of compiledtables sometimesseems scarce-
ly to support the author's judgment of its fidelity relation (Goodall
1953b).
When community-typesare recognized and characterized,stand-
samples are selected and combined into a compositetable, which rep-
resents the community-typeand accompaniespublication concerning
it. In these compiled tables, species are customarilygrouped by diag-
nostic value-as character-species,differential-species,if these are em-
ployed, companions(Begleiter) which rangethrough a numberof com-
munity-types,and accidental or extraneousspecies if these are listed
at all. Among the companionsfor a community-typeof a lower level
(e.g., association), character-speciesfor community-typesof higher

levels (alliance, order, class) may be recognized if phytosociological
treatmentof an area is sufficientlyadvanced. From the table may be
determined the "complete characteristicspecies-combination"for the
community-type(character-speciesplus companionsof at least 60 per
cent constancy); only some fraction of these species (the "normal
characteristicspecies-combination")will occur in a given stand. Data
on location and environmentalfactors for the stands are given also,
and the compiled table forms an effectiveextensionaldefinitionfor the
community-type,such as is lacking in many publicationsof ecologists.
The community-type,if newly described,is provided with a latinized
name based on one or more of its species, which may be either domi-
nants, or diagnosticspecies,or both. Rules for namingcommunity-types
at the variouslevels are given in the referencescited, and rules of prior-
ity for such names have also been published (Moor, 1938; Drees,
1951a, 1951b, 1953; Barkman 1953).
Below the fundamental unit, the association, three levels of com-
munity-types are recognized (Braun-Blanquet, 1951a:23). Subasso-
ciations are community-typesdirectly subordinateto associations,de-
termined by soils, local climates, or geographic relations. Subassocia-
tions generallylack character-species and are to be recognizedprimarily
by differential-species.Less significant variations from type, distin-
guished by certain species-combinationswhich reappearfrequentlybut
lacking useful differential-species,are designatedvariants. The smallest
distinguishableunits, the facies, are characterizedby the abundanceof
certain species, by merely quantitativedistinctions.The profound dif-
ference in outlook between the school of Braun-Blanquetand other
traditions is expressed in this recognition of quantitativedifferences
only at the lowest level, of least significance.In practice,authorsmak-
ing intensive local studies have relied heavily on units below the level
of the association and have often recognized differential-speciesfor
variants as well as subassociations.Facies marked by local dominance
are sometimesecologicallyindifferentor misleading,but they may also
be ecologically significant and useful as indicators (Krause, 1954).
Additional lower-level units-subassociation-group and subvariant-
have been recognizedin some work (Tiuxen, 1950a, 1954b).
It is with associationsand lower units that a phytosociologistclassify-
ing stand-samplesis primarilyconcerned.The hierarchyof units above
the associationpermits classificationof vegetation on a broader,even
a continental or intercontinental,scale. Those species too widely dis-

tributedamong stands and community-typesto characterizeassociations
may be used as character-speciesfor higher-level units to which they
are largely restricted.A number of associationsmay thus be grouped
into an alliance (Verband) defined by alliance-character-species (to-
gether with transgressive association-character-species-i.e.,species
characteristicof particularassociationsthat occur also in other associa-
tions of the alliance,Braun-Blanquet,1933, 1951a:562). Alliancesmay
be similarlygrouped into orders,and orders into classes;on each level
the units are defined by character-species.Character-speciesof higher
units may, again, be minor species; and they may occur in only a
fraction of the lower community-typesgrouped into a higher one. In
many cases alliances,orders,and classes correspondto vegetationtypes
recognizedby those workingoutsideof, or before the time of, the school
of Braun-Blanquet.Lower-level units are grouped into higher-level
units which both seem most meaningful in relation to environments,
and often physiognomyand dominance,and for which character-species
can be specified.
When phytosociologicalstudy of an area is sufficientlyadvanced,
its community-typesmay be presented in a systematicoutline (e.g.,
Tiixen, 1937; Braun-Blanquet,1948-9). Community-typesmay then
be concisely defined by their diagnosticspecies, and their subdivisions
and relations to habitat, succession, and managementindicated. No
other system of classificationhas made possible comparableclassifica-
tion, both in broad outline and fine detail, of the vegetation of a re-
gion. The ecologist studyingsuch a classificationmay see it as a formal-
ized and relatively jejune structure, fragmenting vegetation patterns
into units artificiallyforced into a hierarchy, a skeltonized account
which conveys to him little of what he is interestedin. It seems also
to be true, however, that for the initiate familiar with the system and
the vegetation, such classificationsembody a wealth of informationon
species relations, community-types,and habitat relations such as can
hardly be conveyed with equal efficiencyby any other system.
EVALUATION
The systemhas other advantageswhich may commendit to ecologists.

The use of diagnosticspecies on all levels from the most detailed local
study to the classificationof vegetation on a world-wide scale gives it
a coherenceand consistencydenied other systems.The conceptsof diag-

nostic species on which the system is based have, in practice,proved
more useful for the classificationof vegetation on widely different
levels than any other. The system has been adaptedto widely different
vegetational conditions-from those of the far north, with increased
considerationof dominanceand the sociationas a unit, to those of the
tropics, with increasedconsiderationof physiognomyand dominance.
No other system can claim successfulapplicationover an equally wide
range of vegetational conditions and researchpurposes.
The standardizationof concepts and methods is also a major ad-
vantage; this standardizationmakes the work of one author directly
interpretableby and useful to another, and permits the work of many
authorsto be integratedinto the master-schemeof the school's classi-
fication. These standardizedmethods have also notable merit because
they tend to enforce careful work: (1) Knowledge of the whole flora
of the communityis required. (2) Detailed recordsfrom actualstands
must be compiled as the basis of communitydescriptionand interpre-
tation. (3) Definition of community-typesmust furtherbe made speci-
fiable and unambiguousby designation of diagnostic species. These
requirementscontrast especially with the approach in the school of
Clements, in which restrictionof concern to major species, neglect of
compiled tables, very vague definition of "associations,"and the in-
fluence of the monoclimax theory in suggesting that vegetation be
studied in terms of what it ought to be or might conceivablybecome,
ratherthan what it is, may all have encouragedsome authorstoward
slip-shod observationand interpretation.
Some of the most impressivesuccessesof the system are in the area
of applied phytosociology.In English-languageecology no real equiva-
lent of the extensive and effective phytosociologicalwork in vegeta-
tion mapping, site indication, and land managementexists. Some of
this work, notably the grasslandstudies of de Vries (1954) and the
vegetation mapping of Schmid, Gaussen, and Gams (Kiichler, 1953),
is outside the school of Braun-Blanquet.Some of the work within the
school involves departurefrom, or elaborationon, the system. Thus
Knapp (1948a) and Aichinger (195 la) have proposed additional
vegetation units; grassland studies (Klapp, 1949; Ellenberg, 1952a)
have supplementedfloristicanalysiswith quantitativeweight data; and
Ellenberg's (1948, 1950a, 1952a) studies are based on gradientanaly-
sis as well as classification.Not all the developmentsin applied phyto-
sociology are dependent on the system of Braun-Blanquet;but it may

be significant that some major developments in intensive ecological
applicationhave occurredin this school and not among ecologists.
The evident subjectivityof the methods, together with the apparent
claims to objectivity by some authors, have been a major source of
skepticism among ecologists. Some supportersof the method may in
fact wish it to be regardedas objectiveand mechanical,like a chemical
analysis (Walter, 1939); but it is not possible for this to be so.
Establishmentof lower-level community-typesfrom manipulation of
samples is neither wholly objective nor wholly subjective and arbi-
trary; but it is a process with a large element of subjectivityas it is
affected by the phytosociologist's choice of samples, his skill and
judgment, his intuitive perceptionof desirableways of grouping sam-
ples, his interpretationsof environmentalrelations, and the influence
of traditionguiding him toward resultsgenerally consistentwith those
of others in the school-to a considerableextent it is an art. The
three points on which the school has failed to make itself clear
sample choice, sample manipulation,and the creationof higher units-
are just those steps in which the dependenceon subjectivityand artistry
is greatest.The school has failed to make itself clear becauseit is dif-
ficult at best to describe complex and subjectiveprocesses but, more
than this, because it is impossible to describe such a process when it
is so presentedas to convey the impressionthat it is "objective."Both
criticismfor subjectivityand defense againstthis criticismare, however,
ratherbeside the point (cf. Poore, 1955b). No widely used systemof
classificationis inherentlyless subjectivethan that of Braun-Blanquet.
Writings of the school suggest, however, some misconceptionsof
the natureof character-species as a basis of objectivity.Braun-Blanquet
(1925, 1951b) has written as if character-species were the sure way to
objectivity,recognitionof naturalunits, and general agreementon these
units. Braun-Blanquet(1925) has, further, written of fidelity as a
fundamentalconcept of phytosociology,as no mere autecologicalprop-
erty of a species but a sociological phenomenon. Implicationsof the
principle of species individualityfor the character-species doctrinehave
never been examinedin the school. If species are variouslydistributed,
no two alike, then there are manypossibilitiesfor the grouping of those
which have narrow ecological amplitudes into the character-species
groupings of associations(cf. Whittaker, 1956, 1960). The grouping
cannot be completely free, for the distributionalrelations of species
determine that some groupings which might be attempted would be

clearlyuntenable.But within the limitationsset by actual distributional
relations,the phytosociologistis free to group species in various ways;
the associationwith its group of character-speciesactually recognized
is the phytosociologist'schoice among these possibilities.It seems clear
that in practice many associations are defined by dominance or by
habitat, and secondarilyrationalizedby the specificationof character-
species. Fidelity is not a fundamentalconcept about naturalcommuni-
ties; it is a useful convention, fundamentalonly to a particulartech-
nique.
The freedom in the creation of units applies also on levels higher
than the association.No principle of common descent can relate asso-
ciations of an alliance in the same sense as it may relate species of a
genus. No doubt, numerousways of grouping associationsinto higher
units can be justifiedor rationalizedby specificationof character-species.
In practice,that grouping is chosen which best correspondsto expecta-
tion from considerationsof habitat,physiognomy,dominance,etc. (cf.
Riubel,1934; S0rensen, 1948; Poore, 1955a). Molinio-Arrhenatheretea
may thus mean little more than "meadows"(Wiese), Fagetalia little
more than "mesophyticbroadleaf forests" (Ellenberg, 1954a). Cor-
respondenceof these higher units of the system of Braun-Blanquetto
those of other authorsmeans, not that the floristicapproachautomatic-
ally results in groupings which might otherwise be recognizedas sig-
nificant, but that its flexibility permits units to be tailored according
to other than floristicconsiderations.The school has been criticizedfor
one-sidedinsistenceon floristiccriteriato the exclusionof other proper-
ties of ecosystems.It might be more justly criticizedfor failing to make
explicit the manner in which these other propertiesare used in con-
structing its classifications.
The principle of species individualitymay further imply increasing
difficultyin the definitionof community-typesby character-species with
increasingknowledge of the distributionalrelationsof species and the
varied ways they are combinedinto stands in differentareas.The com-
plex terminologiesof diagnosticspecies of Schwickerath(1942), Drees
(1951a, 1951b), and Becking (1957) suggest the variety of terms
developed to accommodatethe monoclimax theory to the actual com-
plexity of vegetation (Whittaker, 1953). Ellenberg (1954a) has ob-
served that as the knowledge of plant communities becomes more
comprehensive,the number of good character-species,and with this
the significanceof the character-speciesconcept for vegetational sys-

tematics,becomesprogressivelysmaller.Recent work has indicatedthat
associationscannot be defined by generally valid character-species; the
allianceis the lowest categoryfor which generallyvalid character-species
exist (Ellenberg, 1954a). This "crisisof the character-species doctrine"
(Ellenberg, 1954a) has led Knapp (1948a) and others to regard as-
sociationsas locally valid and locally characterizedunits within broader
units (Hauptassoziationeni, Assoziationsgruppe).Ellenberg (1954a) has
suggested abandonmentof the fixed hierarchyat lower levels, in favor
of an essentially fluid system of lower units, locally recognized and
adapted to indication and managementneeds, beneath the established
system of alliances, orders, and classes. Disapproval of Ellenberg's
(1954a) commentshas been indicatedby both Braun-Blanquet(1955)
and Tiixen (1955).
Progress of the school has thus led it to recognitionof the limita-
tions of its original, central concept, the associationdefined by charac-
ter-species, and has even suggested abandonmentat lower levels of
the underlying idea of an establishedhierarchyin the model of plant
taxonomy.The character-species doctrinehas had more striking success
in classificationon lower levels than any other. Evaluationof the sys-
tem must also consider, however, the fact that increasingknowledge
may finally imply not increasing clarity, but increasing difficultyand
complicationin the definition and coordinationof units.
The charge of "one-sidedness"against the school is justified not so
much by its legitimate choice of floristic composition as the criterion
of primaryemphasis,as by a psychologicalcharacter-its doctrinairein-
sistence on the necessityof its own techniquesfor all study of natural
communities.The school's major contributionto the theory of classi-
fication may seem to be the asseverationthat the associationin the
sense of Braun-Blanquetis, it is indeed, the fundamentalunit of the
science. Inquiries into the meaning or "reality"of associationsare re-
garded by Braun-Blanquet(1951a:20) as a Streit um Worte. Problems
of the theory of associationsraisedin the SouthernTraditionby Negri
(1926, 1927), Lenoble (1926, 1928a), and Fournier (1927) have
stimulatedamong leaders of the school neither researchnor reflection,
but avoidance-reactions(Allorge, 1927; Braun-Blanquet,1928b; Pavil-
lard, 1928a). The justificationof the system is and should be essen-
tially practical rather than theoretical (Braun-Blanquet,1932b): it
works and has been found productive.Practicaljustificationapartfrom
theory is never wholly sufficientin science, however, and scarcelyjusti-

fies so single-mindedan insistenceon a particularset of techniques,or
the apparent over-valuationof the devices of classificationas funda-
mental conceptsabout naturalcommunities.
The system as a whole is based not on the propertiesof landscapes
and communitiesbut on a convention,an "as if": Natural communities
shall be classified into associationsas if these were comparableto or-
ganisms and species, and the classificationis justifiedif successful.Car-
ried to the extreme this approachleads to the "formalism"for which
the school has been criticized. Some studies seem motivated by the
love of classifying and naming, not the quest for understanding;and
the vegetation of an area is often unrecognizablein the units into
which it has been classified.Only some studies of the school, present-
ing materialon the ecological relationsof community-typesapart from
their classification,permit a readerto understandthe vegetation of an
area. Braun-Blanquetand others have emphasizedthe abstractcharac-
ter of associations-the associationfirst comes into being at the phyto-
sociologist'sdesk (Klapp, 1949:10). But some authorsseem to hyposta-
tize their associations;they appearto be observingand describingvege-
tation units, not vegetation,much as some Americanshave describedcli-
matic climaxes rather than vegetation. The direction of abuse is in
each case a "formalism,"a following of form in a literalistic, ritual-
istic mannerto the neglect of the possibilitiesinherentin more skillful
and thoughtful application.The possibilityof sterile formalismin this
school (cf. Egler, 1954) is not by itself a criticism. It may well be
considered, however, whether the strongly formal characteristicsof
the system do not more encourageroutine workers toward doctrinaire
acceptanceof the system and unimaginativeliteralismin its application
than the less formal approachesof ecologists.
The conceptionof the present work is that all approachesto classi-
fication of naturalcommunitieshave both merits and weaknesses,that
any choice of approachcarrieswith it both possibilitieswhich may be
exploited and limitationswhich must be acceptedas such, or counter-
acted by moditying and complicating the approach,or glossed over.
Any serious evaluationof an approachmust considerit as a functional
whole in relation to other approaches,scrupulouslyavoiding compari-
son of the merits of one's own approachwith the limitations of an-
other's. Thus considered, with its limitations clarified and accepted,
the system of Braun-Blanquetis the most successful and most widely
applicable means of formal classificationof vegetation that has yet

developed.
The school of Braun-Blanquetmay well propose that its system
should becomethe standardof ecologistsand phytosociologiststhrough-
out the world (Barkman, 1953). If the premise is acceptedthat there
should be an internationallystandardizedapproachto communityclas-
sification,there may be little point in selecting some limitationsof this
system as a basis of opposition when more serious limitations affect
other systems.One may, however, oppose such standardizationon dif-
ferent grounds: (1) No underlyingbasis for the systemexists, such as
the principle of common descent and the existence of some clearly-de-
fined or "good" species in taxonomy. (2) All approachesto ecosystems
have a partial and imperfect character,and different approachesmay
complementone anotherin the study of differentecological problems.
(3) The avenues of explorationof differentapproachesshould conse-
quently be kept open. Much as ecological sciencehas been enrichedby
the school of Braun-Blanquet,it would surelybe the poorerif othertra-
ditions, and the minority schools of the SouthernTradition, were ab-
sorbed into this school.
It may well be questioned whether ecologists should expend, and
divert in part from other desirable research,the immense amount of
time requiredto establisha classificationof this type for North Amer-
ica. On the other hand, it would surely be of great interest to have
classificationsof the vegetation of some areas by this method, to com-
pare with other classificationsby ecologists. For intensive, detailed
work in applied ecology especially,the floristicapproachand concepts
of diagnosticspecies may prove valuable,along with quantitativemeth-
ods. One may thus, without advocatinguniversalstandardizationof this
system,suggest that its merits and achievementsjustify new and favor-
able examinationby ecologists.
INFORMAL CLASSIFICATION BY DOMINANCE AND PHYSIOGNOMY
Two principal approachesto classificationexist among ecologists of

the British and American Traditions. One of these is the system of
Clements,which bases classificationupon a successionalscheme, and in
which the basic unit is either the climax 'association"or climax forma-
tion, the single terminal communityfor an extensive area. The Clem-
entsian system has had great significancein the developmentof ecol-
ogy; and the higher units have continuedusefulness in the geographic

treatmentof vegetation, as shown in work of Braun (1950). Yet the
Clementsianhierarchy,from panformationto locies, is little used in
currentwork and seems to be of declining importanceas a means of
classification.
The other, currentlymore widespread"system"is the informal and
unsystematicuse of dominance-types.These dominance-typesare some-
times called "associations,"they may be regarded as subdivisions of
formations, and may be related to successionsand climatic climaxes.
In spite of these common features with the Clementsiansystem, the
approach through dominance-typesis significantly different. In the
Clementsian system, all else relates to the monoclimax community
toward which other communitiesare developmental;in the other, pri-
mary emphasis is on recognition by dominant species of community-
types, some of which are developmentallyrelated to others. The ap-
proach through dominance-types(and, usually, formations) antedates
the Clementsiansystem; it may be traced from Europeanantecedents
through Warming and some of the early English-languagestudies of
vegetation (Smith, 1898; Moss et al., 1910; Kearney, 1900, 1901;
Ganong, 1903a; Harshberger, 1903) to the present. Currently,this
approach prevails among British ecologists and, probably, American
ones as well. Dominance-typesare in widespreaduse among applied
ecologists-foresters (Frothinghamet al., 1926; Society... 1932, 1945,
1954), and students of range and wildlife management.For this ap-
proach no establishedhierarchyexists comparableto those of Clements
and Braun-Blanquet;the clearestformulationsare by Tansley (1939)
and Australianauthors (Crockerand Wood, 1947; Beadle and Costin,
1952).
EVALUATION
A dominance-typeis a class of stands with the same dominant spe-

cies, or similar combinationsof dominantspecies; the "dominants"are
usuallymajorspecies of the uppermoststratumof the community,some-
times of a lower stratum.Whereas the associationsof Braun-Blanquet
are, ideally, characterizedby their whole species-composition,domi-
nance-typesare defined by one or two (or a few) major species.
Whereas the system of Braun-Blanquetselects for emphasisthose spe-
cies of narrowestecological amplitudes,dominance-typesare based on
species often of very wide distributions.It was part of Clements'con-
ception that dominant species so determined conditions of life for

other species that they would, necessarily,characterizecommunity-types
and indicate environment (Clements, 1928:236, 253; Weaver and
Clements, 1938:91, 478; Clements and Shelford, 1939:238-9). But
it now seems clear that distributionsof other species may be little cor-
related with those of dominants and that dominants do not charac-
terize homogeneous "natural"units (Whittaker, 1956). Contrasting
the system of Braun-Blanquetwith the approachthrough dominance-
types, two critcismsof the latter may be offered: (1) The approach
through dominance-types,focusing attentionon a few most conspicuous
species, is relatively superficial. (2) Because these species are chosen
for conspicuousness,not environmentalrelations,dominance-typesmay
be ineffective as indicators of environment.
That the approachthrough dominantsalone may often be relatively
crude and superficialis most evident for very wide-ranging dominant
species. Only a heterogeneousmonster-associationwould result from
bringing together all stands dominated by so wide-ranging a species
as Pinus silvestris (Lippmaa, 1931; Braun-Blanquet, 1951a:560,
195ib); the samemay be true for such Americanspeciesas Pseudotsuga
menziesii, Pinus ponderosa. and Pinus contorta.Wide-rangingdomi-
nants consist of diverse ecotypic populations, occurringin varied en-
vironments with different associates;there may be scarcelyany sense
in which two stands from widely different circumstancesare "the
same" except the name of the dominant. In northern and subalpine
forests dominatingspecies occur in a wide range of habitat conditions
in a given area. For effective site indication it is necessaryto consider
the rest of the community,establishingsite-typesor other lower-level
units on the basis of undergrowthcomposition.
Some dominantspecies are distinctlybimodal, as may be illustrated
in the SouthernAppalachians.One populationof the Americanbeeches
(Fagus grandifolia) dominatescertaincommunitiesof lower elevations
(below 600 m) recognizedby Braun (1950) as the beech ravine and
beech-whiteoak segregates.Another population of beeches dominates
the beech gap stands (Russell, 1953; Whittaker, 1956) of high eleva-
tions (above 1370 m). A gap of 600 m separatesthese two beech
populations (a third beech population occurs between 1070 and 1370
m but does not dominatestands); and in floristiccompositionthe high-
and low-elevation stands are very different. One population of white
oaks (Quercusalba) dominatescertainstand-typesat low elevationsin
the SouthernAppalachianmountainsand the Great Valley of Tennes-

see (Braun, 1950); a second populationis dominantin other, floristic-
ally very different, stands above 1100 m in the Great Smoky Moun-
tains (Whittaker, 1956). There is, again, a gap betweenthe two popu-
lations; very few white oaks occurbetween 760 and 1060 m. In neither
case, becauseof their relationsto one anotherelsewhere,is it desirable
to treat the low- and high-elevation populations as species; in the
Southern Appalachiansthey are better regardedas elevation ecotypes.
Each species dominatestwo or more groups of stands which differ in
environmentalrelations,in floristics,and in genetics of the dominating
population. It seems meaningless to say that the high- and low-eleva-
tion stands representthe same vegetationtype becausethey are domin-
ated by the same species.
Some dominantspecies, on the other hand, occur in a narrow range
of ecologicalconditions,usuallywith similarcombinationsof associates;
they are character-species,as well as dominant species, for relatively
homogeneous community-types.Between the extremes representedby
these and by very wide-rangingspecies, community-typesof all degrees
of heterogeneitymay result from the definition of types by dominant
species. A further weakness of the approachthrough dominance-types
is that it leads to community-typesof differing heterogeneityand in-
clusiveness,some of them scarcelycomparableas community-units.
It is in many cases inappropriateto define dominance-typesby a
single species.Many types are recognizedby two or three specieswhich,
together, make up a large part of the stands and characterizethe type.
In some types-as in the American mixed mesophytic forests, some
tropical forests, and some shrub communities (Adamson, 1927)
compositionof standsis so mixed that types are definedby lists of major
species usually present,though in proportionsdifferingfrom one stand
to another. Dominance-typesmust thus be defined in quite different
ways accordingto vegetationalcircumstances.
Furthermore,in the conditions that prevail in many communitiesof
temperate latitudes, arbitrarydecisions must often be made whether
types are to be defined by one species, by two, or by three or more. In
some vegetational conditions no limit on the number of dominance-
typesmay be recognized,except the ultimateconsequenceof distinguish-
ing types as narrowlyas possible by combinationsand orders of im-
portanceof their dominants-almost every stand representsa different
type (Curtis and McIntosh, 1951). There is, in practice,much free-
dom of choice in the definition of dominance-types.No established

criteriacan determinewhether a larger numberof narrowlydefined, or
smaller number of broadly defined, dominance-typesare to be recog-
nized.
A solution to the problem offered by widely different inclusiveness
of units defined by one species is the judicious use of these varied
possibilitiesof definition.The ecologist may choose to define his units
so that they are not excessively heterogeneous,as judged by environ-
ments and floristiccomposition.Thus for one unit, definitionby a single
dominant species may be appropriate,for another recognitionof nar-
rower types defined by pairs of dominants,or a broaderunit including
any combinationfrom a group of dominants,for still anothershift of
emphasis to subtypesor site-types within the heterogeneousgrouping
dominatedby one species. The particularmannerof definitionmay be
determinedby the ecologist's judgment in a context that includes both
vegetationalconditions and ecological purposes. It is an illusion that,
as implied by Clements,dominantspecieswill in generallead the ecolo-
gist automaticallyto the recognition of objective, natural units; the
approachthrough dominance-typesis inherently as subjective as that
through character-species. Becausethis is so, however, the ecologist can
choose to define his dominance-typesin the way which will give them
maximumusefulness.
The meaning of dominance-typescan best be illustrated by actual
distributionalrelations of species. In Fig. 2, several possibilities are
illustrated,basedon data from the GreatSmokyMountains(Whittaker,
1956). It may be observed that these situations intergradein various
ways and that by no means all possible situationsare illustrated-the
complicationsintroduced by bimodal and polymodal species distribu-
tions, especially,are not considered.
1) In a pattern of continuouslygradating vegetation, one species
(a) is clearly dominant in a span of the environmentalgradient, with
a typical distributionalcurve. A dominance-typeis thus easily recog-
nized by this species; but the limits of this type (whether at 50, 40,
30 per cent of canopy stems, or of all stems, etc.) are necessarilyarbi-
trary.
2) A species (a) with a "plateau"distributionis strongly dominant
over a span of the gradient,and its rangeof dominanceis more abruptly
separatedfrom that of other species. The species thus defines a domi-
nance-type which is relatively discontinuous with other dominance-
o-,
60 m
O402
nacetyes baedndt rmteGet mk onan Witkr
(2cm db n agr)i tns
(2 cmdhad4agr0i tns
1I. To. Dominne

dsrbtoaeain
asnl of species
wihabl-hapedditeribuindon,-
in a transect of the moisture gradient between 1070 and 1380 m through 12

stations from mesic (left) to xeric (right). Tsuga canadensis is strongly
dominant near station 4, but declines continuously into cove hardwoods forest
on the left, into Quercus-Castanea forest on the right. Major species indi-
cated: a Tsuga canadensis, b Tilia hzeterophylla, c Betula allegheniensis,
d Quercus borealis, e Quercus prinus, f Pinus pungens.
2. Dominance of a single species with a "plateau" distribution, in an
elevation transect in mesic sites. Elevation intervals of 100 m from 800 to

1600 m are marked from left to right. Fagus grandifolia is strongly dominant
above 1380 m, but mixed cove forest stands occur below that elevation. Major
species indicated: a Fagus grandifolia ("gray" population), b Liriodendron
tulipifera, c Betula allegheniensis, d Tilia heterophylla, e Aesculus octandra,
f Acer spicatum.
3. Dominance of a single species of broad ecological amplitude, in a
moisture-gradient transect of five steps from mesic sheltered slopes (left)
to open southwest slopes (right) in subalpine forests (1900-2000 m). Abies
fraseri (a) is strongly dominant, forming 85-90 per cent of stands, along the
whole of the moisture gradient; undergrowth species of narrower amplitudes
permit distinctions among stand types.
4. Dominance of two species with similar, though not identical, dis-
tributions, in a moisture-gradient transect of 13 stations between 760 and
1070 m. Quercius prinus and Castanea dentata together dominate and char-
acterize stands in subxeric sites. Major species indicated: a Quercus prinus,
b Castanea dentata, c Betula allegheniensis, d Halesia monticola, e Acer
rubrum xO.5, f Pinus rigida.
5. Dominance of two species with dissimilar, but broadly overlapping
distributions, in a moisture-gradient transect of 5 steps between 1380 and
1670 m. Major species indicated: a Abies fraseri, b Picea rubens.
6. Dominance in various combinations of three species with dissimilar,
but broadly overlapping distributions, in an elevation transect in subxeric
sites. Elevation intervals of 100 m from 500 to 1400 m are marked from left
to right. Major species of Quercus-Castanea forests indicated: a and a'
Quercus alba, b Quercus prinus, c Castanea dentata.
7. Dominance in various combinations of several species, in a moisture-
gradient transect through 13 stations between 760 and 1070 m. Several trees
of the cove forests (a to f) are dominant in mesic sites toward the left, but
decline toward the right into the Quercus-Castanea forests of more xeric
sites. Major species indicated: a Betula allegheniensis, b Tsuga canadensis,
c Halesia monticola, d Acer saccharum, e Tilia heterophylla, f Liriodendron
tulipifera, g Acer rubrum, h Castanea dentata, i Quercus prinus, j Quercus
coccinea, k Pinus rigida.
types along one gradientin question (but is likely to be continuouswith

other types along other gradients).
3) A dominant species (a) has a very wide amplitude along the
gradient, but more narrowly distributed undergrowthspecies permit
recognitionof site-types.
4) Two species (a and b) with more or less similar distributional
patterns may serve to characterizea type in much the same manner
(and with the same arbitrarinessof limitation) as one.
5) Two species (a and b) have dissimilar,but broadly overlapping
distributions;and the ecologist is offered a choice of recognizing:only
one type dominatedby either speciesa, or species b, or both; two types,
one dominatedby a and one by b; three types dominated by species
a, species b, and the combination of a with b (cf. Society . . . 1932).

6) Three species (ac', b, and c) have quite different distributions, but
overlap broadly. Either a single type including any combination of the
dominants, or some number of types characterized by particular com-
binations, may be recognized. The case illustrated has traditionally been
regarded as one American "association"; within it Conard (1935a)
recognized two associations on Long Island and Whittaker (1956) four
major types in the Great Smoky Mountains, where finer distinctions
would permit recognition of additional types.
7) Several species (a to J) have ecologically related distributions and
often occur together in stands, through in proportions varying from
stand to stand. Since the species have their distributional centers in
more or less similar environments, they represent a "commodal" group
(Whittaker, 1956). The ecologist must decide how many species to
include in the group and hence the range of conditions for the type.
Variations in composition from one stand to another may result in
part from relations of species to environmental gradients, in part from
chance factors of species establishment and history. The ecologist may
choose to avoid recognizing these many combinations, some of doubt-
ful ecological meaning, by defining his type through the whole group
of species. Certain types more distinctly dominated by one, two, or
three of the species may then be regarded as "segregate" types (Braun,
1935a, 1950; Richards, 1945, 1952).
A dominance-type is not, and cannot be, a particular, standardized
kind of community-unit. The properties of dominance types discussed
imply one further, major limitation of this approach. Dominance-types
are poorly suited to the formal, hierarchial classification of vegetation.
The only actual attempt to set up a formal hierarchy rivaling Braun-
Blanquet's, but based on dominance at all levels, is by Du Rietz (1932).
This work contains many useful suggestions on classification at lower
levels, but at levels above the sociation it has been so little followed
that it may be judged unsuccessful.
The author has sought to clarify some essential limitations of the
approach through dominance-types: It represents, at least in some uses,
a superficial and relatively crude approach to classification of communi-
ties; it depends as much on subjectivity as other systems; it leads to
units of different kinds and magnitudes; and it is not in general suited
to formal classification. Any choice of dominance-types in preference
to the association of Braun-Blanquet should involve frank and explicit
recognition of these limitations. They imply, in the author's opinion,

that in comparisonwith the system of Braun-Blanquetthe approach
through dominancetypes has significantdisadvantages.
On the other hand, dominance-typeshave distinct practical ad-
vantages for some purposes. They do not, like associations,require
thorough floristic knowledge of species other than dominants,knowl-
edge of distributionson which fidelity judgments can be based, ex-
tensive sampling and comparisonof releves before a unit can be de-
fined, and extensive, wide-rangingstudy before units can be grouped
into higher categories.They do not require the expenditureof much
time and energy on classification,when classificationas such may be
of only secondaryinterest. Dominance-typesare based on those species
which seem most "important"in the community,often the specie'zthat
most concernecologists, especiallyin such applied ecologies as forestry
and range management.The advantagesof the Braun-Blanquetsystem
will not easily overcome the combined forces of ecological tradition,
the direction of ecologists' interests toward mattersother than formal
classification,and the greatersimplicityof dominance-types.For ecolo-
gists who choose to approachvegetation through dominance-types,the
following suggestions on terminology, subdivision, and grouping of
these units are offered.
APPLICATION
One of the important steps toward agreement among phytosociol-

ogists was the recognitionthat the associationand sociation were two
different concepts that should be designated by different terms. The
term association is now widely acceptedfor the floristicunit of Braun-
Blanquet, and there seems little real justificationfor its use in other
senses by ecologists. It is suggested that ecologists, also, might do well
to follow the recommendationsof the Sixth BotanicalCongress (Cain,
1936a; Du Rietz, 1936; Poore, 1955a) in this respect, and to clarify
the meaning of their own vegetationunits by applying differentterms.
Units defined by dominantspecies may well be termeddominance-types
or, simply, types (cf. the association of Cajander, 1903a, 1922; Warm-
ing, 1909; Beadle & Costin, 1952; consociation of Du Rietz, 1932;
Gams, 1933; dominion of McLean, 1935; type of Crocker and Wood,
1947; Beadle, 1948, etc.). Tansley (1948) has objected to this use of
type on the basis that a formal scientific concept should be given a
distinctive term. To the author, the informalityof the term type is a
substantialadvantage, -for it does not imply that a type is a clearly-

defined, generally accepted,classificatoryunit. This usage, unlike some
uses of association, does not suggest that types are more than they are.
No standardizedmeansof subdividingdominance-typescan very well
exist; the ecologistmay choose one of the widely recognizedlower-level
units (sociation, site-type, association, etc.), or one of the informal
units termed subtypes below, or create for his needs an appropriate
means of subdivision.Some of the possibilitiesare:
1) By floristiccompositionand the recognitionof groups of charac-
ter-speciesor differential-species.Some dominancetypes may very well
be subdividedby units of the Braun-Blanquetsystem-associations, sub-
associations,variants,and facies-and the mixtureof criteriamay not be
a disadvantagein practice.
2) By sociations, combinationsof stratalunits, especiallyin northern
and higher-elevationvegetation.
3 By site-types defined by undergrowthcomposition in relation to
site properties,especiallyin northernand mountain forests.
4) By site or habitat distinctions, when recognized first and then
correlatedwith vegetation. Habitat-subtypesare not a classificationof
vegetation by its own properties,but may lead to recognition of en-
vironmentalrelations of other lower-level units.
5) Geographicor climaticsubtypesmay be recognized,as in different
climatic regions or mountain ranges, and defined by these geographic
criteriaand associatedfloristicdifferences.
6) By different combinationsof dominantsand subdominants-
a) By presenceand absence (or presencein substantialnumbers)
of one of the group of two or more dominantspecies of the type.
b) By "segregations"of one or more dominantspecies from the
group of species by which the type is defined.
c) By "subdominants,"i.e., dominants of a stratum other than
that of the dominant.
7) By quantitativerelations of various sorts-undergrowth or can-
opy coverage, productivityor growth-rate,weighted averages of com-
munity composition, proportionsof life-forms or growth-forms,rela-
tive importanceof species, etc.
Various possibilities exist also for the grouping of dominance-types
into higher units:
1) By physiognomyinto formations. The most widely used higher
vegetation units in plant geographyand ecology are formations,recog-
nized by physiognomyplus environmentalconsiderations;a grouping

of similar formationsfrom differentcontinentsconstitutesa formation-
type. The terms biome and biome-typeare essentially synonyms, ap-
propriate whenever formations are approachedas biotic community-
types.
Various choices must be made in the definitionof formations,as of
other vegetation units. Definition by a single dominant growth-form
is sometimes appropriate;other formations may be defined by two
dominantgrowth-formsin the same stratum(e.g., the American"Lake
Forest"of hemlock,white pine, and hardwoods,Weaver and Clements,
1929; Braun, 1950) or in different strata (savanna of some authors,
oak woodland, pygmy conifer woodland, pine steppe, pine heath and
oak heath). Although there is agreementon the recognitionof major
growth-forms, there is still freedom of choice in the recognition of
minor ones and the extent to which differentcombinationsof growth-
forms are acceptedas differentformations.If growth-formsalone were
used for definitionof formations,some manifestlyheterogeneousunits
would result. It is consequentlycustomaryto separate into different
formations communitiesof the same growth-form in widely different
environments.Thus communitiesdominatedby grasses and grass-like
plants are distinguishedby major climatic belts (as tropical savanna,
temperatesteppe, alpine meadow and paramoand arcticsedge tundra),
and within temperateclimatesby soil moistureand salinity (as steppe,
fresh-watermarsh, and salt-marsh). From the definitionof formations
by combinationsof growth-form and environmentit is but a further
step to recognitionof some formations (strand, hammada,fjeldmark)
primarilyby environment,only secondarilyby growth-forms.Definition
of formations is thus dependent on various judgments on the extent
to which distinctions among growth-forms, combinationsof growth-
forms, and differencesof environmentsare to be acceptedas character-
izing different formations.
Many formationsintergradecontinuously,as is especiallyevident in
the tropics (Beard, 1955; Boughey, 1957). There is no limit, with
increasinglynarrowdefinition,to the numberof formationsto be recog-
nized; by combining environmentalcriteriawith growth-formsBeard
(1955) recognizes such closely related formations as rain forest, dry
rain forest, dry evergreenforest, evergreen seasonal forest, and semi-
evergreenseasonalforest; Wendelberger(1954a, 1955) recognizesfor-
est-steppe, loess-steppe, salt-steppe, sand-steppe, rock-steppe,and dry
grassland.On the one hand, the tundrais widely recognizedas a forma-

tion of the treeless arctic plains; on the other, the variety of growth-
forms in relationto habitatshas permittedIcelandicauthorsto recognize
a whole seriesof distinctiveformations.For more narrowlydefinedphys-
iognomic-ecologictypes some authorsrecognizethe term subformation.
Purposesof one study may determinethat only major formationsthat
prevail over extensive geographic regions be recognized, as in many
plant-geographicstudies. In anotherstudy, many local, edaphicallyde-
termined communitiesare appropriatelyrecognizedas formations.One
may either consider climate as only one of the environmentalfactors
determining a given formation (Beadle and Costin, 1952), or seek
to distinguish major, climatic, or regional formations from minor,
edaphic, or local formations (Schimper, 1903; Gradmann, 1941;
Pendleton, 1949). The same formation may be climatic under some
circumstances,edaphic under others (e.g., climaticand edaphic steppe;
Wendelberger, 1954a, 1955; and the savanna as an edaphically de-
termined formation prevailing over extensive regions (Beard, 1953)
complicates the distinction of climatic or regional formations from
edaphic or local ones.
The formationwill usually include a numberof dominance-types,so
that for classificatorypurposesit may be used as an "ecologicalgenus"
(Warming, 1909), into which dominancetypes may be grouped. It
must be expected, however, that some circumstancesand choice of cri-
teria will reversethis relation,so that more than one formationmay be
distinguished within a dominance-typeor association (Villar, 1929b;
Gradmann, 1941; Beadle and Costin, 1952; Guinochet, 1955). This
may occur, for example, with the presenceand absenceof an open tree
stratum above similar grassland, bog, or heath communities; with
change of emphasis between two growth-formsin floristicallysimilar
communities(e.g., oak-chestnutforest and oak-chestnutheath; Whitta-
ker, 1956); with different growth-forms of different ecotypes of a
dominant species (as in some Australianeucalypts;Specht and Perry,
1948; Beadle and Costin, 1952).
2) By relatedcombinationsof dominantsinto collectivetypes. Domi-
nance-typesmay be variouslygrouped into larger units which are still,
usually, on a lower level than the formation. Thus a group of domi-
nance-typeswhich share one or more of their major species-or share
their dominantsin various directionsalthoughno dominantis common
to all, or are floristicallysimilar, or have taxonomicallyrelated domi-
nants (e.g., dipterocarpforests; Brown and Mathews, 1914; Richards,

1952:254) -may be grouped into a larger collective or aggregateunit.
For this situation the term collectivetype is suggested (cf. the federa-
tion of Du Rietz, 1932, not of Du Rietz, 1936; the alliance of Beadle
and Costin, 1952; the associationsof Tansleyand Chipp, 1926; Crocker
and Wood, 1947; Beadle, 1948; Beard, 1949b, 1955, and others).
Collective types may differ only in degree from dominance-typesde-
fined by several species. When there is stand-to-standvariationin the
importanceof the various dominants,and this variationis difficultto
correlate with environmental difference, undergrowth character, or
floristic composition, the varied stands may appropriatelybe treated
as a single dominance-type(e.g., Southern Appalachiancove forest,
or mixed mesophytic in a narrowersense; Braun, 1950; Whittaker,
1956). A still more heterogeneousgrouping, with many dominants
occurringin differentcombinationsclearlycorrelatedwith environment
and floristiccomposition,may be treatedas a collective type (e.g., the
Quercus-Carya "association," Weaver and Clements, 1929; Braun,
1950).
Many of the "associations"of Clementsianecology are of this na-
ture, though it has seldom been made clear how heterogeneousand
variously defined these units are. Some (Quercus-Carya, Quercus-Pinus
associations) are defined by little more than generic relationsof domi-
nants which may overlap in distributionand sometimesoccurtogether.
The Quercus-Castanea associationis less obviously heterogeneous,but
includesa considerablenumberof combinationsof dominantsand some
physiognomicvariation.The mixed mesophyticassociationin a broad
sense, including segregatesand geographicvariation,is a collectivetype
with a considerablerange of dominancecompositionand floristiccom-
position. The Fagus grandifolia-Acer saccharum and Acer saccharum-
Tilia americanaassociations,on the other hand, are dominance-types
characterizedby two species although, in the former at least, the domi-
nant species are of very wide amplitudes (and genetic complexity;
Camp, 1951), and the diversity of stands grouped together may be
scarcelyless than in the more obviouslyheterogeneousassociations.
3) By environmentalrelations into ecological series. One effective
means of relating community-typesto environment is through their
serial arrangementalong environmentalgradients. A chain (or con-
tinuum) of dominance-types(or of formations,associations,sociations,
site-types, etc.) along a particulargradient or complex-gradientof
environment forms an ecological series. Along such major complex-

gradients as elevation in mountainsand tide-levels in the littoral, each
section or zone may actuallyform a complex of community-types,one
of which prevails and characterizesthe zone.
4) By environmentalrelations into complexes. When the environ-
mental relationsamong community-typesare more clearlymulti-dimen-
sional, or mosaic-like, the grouping is a community-complex.Many
types of complexeshave been recognized:edaphic complexes (Crocker
and Wood, 1947), water bodies as complexes,topographiccomplexes
as formed by a drainagebasin or mountainrange, micro-complexesof
bogs, frost-determinedlandforms,and till, landscapesand biotic prov-
inces, etc.
5) By developmental relations into successional series and climax-
complexes. A series of developmentalcommunity-typesleading to a
stable or climax type in a given kind of habitat is a successional series
or sere. The grouping of successionaland climax community-typesre-
lated to one another by ecological or geographic relations, or by a
particularclimax or prevailingclimax, forms a climax-complex(Braun-
Blanquet, 1951a; cf. the formation in the sense of Moss, 1910, 1913,
and Clements, 1916, 1936). In a narrowersense, the climax-complex
(or successional complex) may comprise only a particular climax type
plus the various communitiesof primary succession and disturbance
which are developing toward it, or the communitiesrelated by cyclic
processes (Osvald, 1923; Watt, 1947). In the broaderand more fre-
quently used sense, the climax-complexbrings together varied climax
and successionalcommunities of a region or elevation zone in rela-
tion to the prevailing climax community-type;it is thus a complex of
successionalcomplexes.
6) By successionalstatus and prevalence. The Clementsiansystem
distinguishedclimax associationsand successionalassocies;but the sys-
tem involved a thorough confusion of prevalence and stability, and
many of the "associes"were stabilized communities.The Braun-Blan-
quet system recognizes some communitiesas climaticallydetermined,
stabilizedSchlussgesellschaften,others as edaphicallydetermined,stable
Dauergesellschaften,and still others as developmental.Some such dis-
tinctions accordingto combinationsof stabilityvs. instability,and re-
gional prevalance vs. localized occurrenceseem clearly needed, al-
though various difficultiesand complicationsmay be expected in prac-
tice. Three major classes or levels of community-typesaccording to
successionalstatus and prevalencemay be recognized-the prevailing

regional climax type or "climatic climax," the localized or edaphic
climax type, and the unstable successionaltype-and the unstable or
man-producedcommunitywhich prevails and characterizesa landscape
is the fourth possibility. Since some localized types are as much stabil-
ized climaxes, and as much determined by climate, as the "climatic
climax," Whittaker (1953, 1956) has suggested terming this the pre-
vailing climax.
The Clementsian"association"is primarily a concept of American
ecologists, but some such concept is often needed to express vegeta-
tional relations to geography and climate below the level of the for-
mations, and related concepts have appearedin other traditions. It is
suggested that community-typesthus used for geographictreatmentof
vegetation be termed regional vegetation types or prevailing climax
types. Regional vegetation may thus be studied either on the level of
dominance-typesand collectivetypes, or of climaticformation-typesand
formations. The Clementsian"associations"may be interpretedas re-
gional dominance-typesand collective types.
The preceding is a suggestion of an informal hierarchyof vegeta-
tion units (formation-type,formation,subformationand collectivetype
when appropriate,dominance-type,and subtypeetc.) in large part al-
ready used among ecologists. Wholly contraryto the author'sintent is
that this should be mistaken for a formal system comparableto that
of Braun-Blanquet;the objective has been to offer suggestions on in-
formal classification,to be followed when they are appropriateand
set aside when they are not. The limitations of the system are, it is
hoped, clear. It is expectedthat differentauthorswill recognizedifferent
units, and that their units may be difficultto relateto one another.Seen
in comparisonwith the orderly hierarchyof Braun-Blanquet,this may
seem a kind of chaos in classification;but if classificationis not a major
objective, there may be little regret that studies by differentecologists,
each effective in its way, do not fit into a coherent classification.The
point of the present suggestions is that if classificationis not a major
objective, then the experience of ecologists suggests: a physiognomic
approachon higher levels, an approachthroughdominance-typeson in-
termediatelevels, a thoroughlyfluid approachthroughunits appropriate
to circumstanceon lower levels, and emphasis of environmentaland
dynamic relations of community-typesinstead of hierarchial classifi-
cation.
These units of vegetation are not, necessarily,also appropriateunits

for animalcommunities.The biome and biome-typeare well established
as higher-level units based on physiognomybut appropriatealso for
vertebrateanimals; but no well-establishedunits for animal communi-
ties at lower levels exist. Informally defined "cover-types,"based on
very mixed criteria,seem sufficientfor some purposesof wildlife man-
agement (Wight, 1934; Trippensee, 1934; Dalke, 1937, 1941; Gra-
ham, 1945; Jensen, 1947; Beckwith, 1954). It would be an admirable
achievementto find units equally appropriatefor all groups of plants
and animals;however, it may be suspectedthese have not been found
becausethey do not exist. The animal ecologist has a choice of basing
his work directly on vegetation units, or of seeking units most ap-
propriatefor a given group of animals and comparingthese with the
units of plant ecologists. It may be of more fundamentalinterest to
study distributionsand communityrelations of animals for their own
sake, than to force these into the patternof units recognizedby plant
ecologists.
Much the same may be said for the classificationof aquatic com-
munities, which has been as yet much less extensively developed than
for terrestrialones. For the classificationof aquaticstands any of the
approachesused by terrestrialecologistsand phytosociologists-through
dominantspecies, combinationsand groups of major species, diagnostic
species, habitat properties, and physiognomy-are theoreticallyappli-
cable; choice among them should depend on relative usefulness in dif-
ferent types of aquaticcommunities.Limnologistshave in generalbased
classificationsof waterbodies not on compositionof the living commun-
ity itself, but on other characteristicsof ecosystemswhich reflectsignif-
icant differencesin conditionsof life. Whereasthe phytosociologistsex-
plore the possibility of formal classificationof aquaticcommunitiesby
diagnosticspecies, the objective for ecologistsmay well be the explora-
tion of less formal classificationsas appropriateas possible to the pe-
culiar conditions of aquaticcommunities.
It may finally be observed that the cleavage between the ecological
and phytosociologicaltraditions is too deep-seatedto be resolved by
arbitrarystandardization.Although it is easy to deplore lack of interna-
tional agreement,ecologicalsciencemay well have gained by the choice
of one group to explore thoroughlythe approachto communitiesthrough
systematic classification, whereas others explored other approaches.
Braun-Blanquet'sformal classificationsand ecologists' informal ones
are not directlycompatiblewith one another,but neither are they fully

incompatible.Associationsand dominance-typesare differentunits; but
some associationscorrespondto dominance-types,and some dominance-
types and subtypescan be characterizedby diagnostic species. Mutual
understandingmight increaseif ecologists made more effective use of
diagnostic species, and if phytosociologistsmade more explicit the
place of formationsand dominance-typesin their system.It may further
be expectedthat as more intensiveanalyticalwork, little concernedwith
classificationas such, progressesin both traditions,the present differ-
ences will become less important.For now, the objective may be not
so much full agreementas greatercommon understandingamong eco-
logical scientists of different interests and traditions.
SUMMARY
HISTORIC BACKGROUND
1. Since the early suggestion of vegetation units by Humboldt and

Grisebach,approachesto classificationof vegetationhave diverged into
several major traditions and numerousschools. One major approach,
representedin all the regional traditions,is that through morphology
or physiognomyof vegetation,with formationsand world-wideforma-
tion-types as its major units of classification.
2. Much of the development of the Southern Tradition was cen-
tered in the. cities of Zurich and Montpellier. Among the varied
schools of this tradition that of Braun-Blanquet,with its hierarchyof
floristic associationsand other units defined by diagnosticspecies, has
gained the largest following.
3. Three major directions may be recognized in the Northern or
Scandinavianand Baltic Tradition:the definitionof sociationsby com-
binations of stratal dominantsin the Uppsala school, the synusial ap-
proach through unions of a single life-form or stratum,developed by
Gams, Lippmaa, and others, and the system of site-types defined by
characteristicsof undergrowthcommunitiesas related to habitat,in the
Finnish school of Cajander.
4. Major units of the Russian Tradition are the formation and the
sociation (the Russian "association"). Later Russian (and Swedish)
systems of forest types are based on the sociation instead of the site-
type of Cajander.Russianauthorshave been less concernedthan Scan-
dinavianauthorswith formal classification,and more with the relation

of sociationsto one anotheralong environmentalgradientsin ecological
series.
5. Most work in the British Isles has been based on informal use
of dominance-typesdefined by the one or two major species of the
community;formationshave been widely used elsewhere in the Com-
monwealth. Classificatorysystemsusing both these units have been de-
veloped by Tansley in England,Crockerand Wood in South Australia,
and Beadle in New South Wales.
6. The American and British Traditions have been strongly in-
fluenced and closely related to one another by the development of
successionalapproachesto vegetation-by Cowles and Clementsin the
former,Moss and Tansleyin the latter.The dominantapproachthrough
the middle period of the AmericanTraditionwas the systemof Clem-
ents with its emphasis on succession and regional vegetation units.
More recentlymany Americanauthorshave used dominance-typesin-
formally, or experimentedwith other approaches.
7. Ecosystemsmay be classifiedby many differentpropertiesof each
of their major aspects-physical environment, soil, vegetation, and
animal communities.In addition to classificationsbased on particular
aspects of ecosystems,multi-factoralor landscapeapproachesto classi-
fication have been developed by Passarge,Markus,Morosov, Schmid,
Meusel, Dice, and others. When a single complex-gradientof many
correlatedfactor gradients is of such paramountimportancethat other
gradientsmay be treatedas secondaryto it, classificationby zones (life-
zones, intertidal zones) may be appropriate.Classificationsof animal
communitiesdiffer almost as much as those of vegetation.
8. The history reviewed is less one of progress toward general
agreement on a single fundamental unit of vegetation than one of
progressivedifferentiationof a numberof vegetation units of greatest
usefulness. The principal units thus evolved are the formation-type,
formation,associationin the sense of Braun-Blanquet,sociation,union,
dominance-type,site-type,regional vegetationtype, and landscape-type.
9. The direction in which a school develops is determinedin part
by vegetationalconditions-characteristicsof the vegetationwith which
it deals that affectits approachto classification.But developmentis also
in part independent of propertiesof vegetation, and is influencedby
key ideas found to be productiveand that become the basis of further
interpretation,by cultural factors and personal views of leaders of

schools, and by such ideas as the analogies of the communitywith the
organism and the associationwith the species.
10. Most schools have held in common the assumptionthat some
natural, fundamental unit of vegetation exists on which agreement
should be possible. Several authors in different traditions (Ramensky,
Gleason, Lenoble, etc.) have advancedan alternative,"individualistic"
conceptionto argue that, since species are distributedindividualistically
and vegetation is to a considerable extent continuous, classification
must be primarilyarbitrary.
THE THEORY OF SYNECOLOGICALCLASSIFICATION
1. Four major types of evidence bear on the "individualistic"hy-

pothesis and the alternativebelief that naturalcommunitiesconsist of
well-defined "natural"units:
i) Relative similarityand dissimilarityof stands. Comparisonof
stands not selected to representassociationssuggests that stands are
complexly related to one anotherwith varying degrees of similarity
in various directions,ratherthan falling naturallyinto well-defined
groups.
ii) Continuity and discontinuity.Although discontinuitiesoccur
between natural communities, many communities are fully con-
tinuous with one another.
iii) Distributionalrelations of species. Various types of evidence
on whether species are distributedin well-defined groups of asso-
ciates, or are variously distributed according to the individualistic
conception,strongly support the latter.
iv) Dynamic relations of species. Considerationof the character
of interactionsof species suggests that species should be expected
to distribute themselves "individualistically"despite their involve-
ment in the web-of-life in each communityin which they occur.
2. Objects of synecologicalclassificationshould be viewed on two
complementarylevels: landscapesas complex ecosystemicpatterns,and
local ecosystemsor stands in their habitats. Factors, populations, and
propertiesof local ecosystemsshow extensive intergradationand inter-
penetration;stands should be regardednot as distinct individuals but
as areas of or points in the landscape pattern. In the diversity of
stands that make up the landscape pattern, and the mutliplicity of
factors and populations of each local ecosystem,there is no observa-

tion without choice of that which is observed.
3. Some relations of stands and populations which underlie prob-
lems of classificationmay be clarifiedby certainabstractrepresentations
-ecological series, and abstractpatternsshowing the relationsof popu-
lations to two or more complex-gradientsof the landscape.Such treat-
ment shows (a) that species populations are diversely distributedin
the patternand (b) in most casestaperalong the gradientso that stands
are continuouswith one another, (c) that most community-character-
istics also change continuouslyalong gradientsand (d) show different
patterns of variation in relation to the community-patternstudied.
There are an unlimited number of ways of defining areas of abstract
patterns, to which classes of stands correspond;and different classes,
only loosely related to one another, result from different choices of
criteria of classification.No single classificationis appropriateto all
species distributions,or groups of organisms, or properties of eco-
systems. Communityunits may also be approachedas types defined
by severalcharacteristics,and the intergradingcommunitiesof the land-
scape may be grouped around these types by subjectiveevaluation of
the extent to which they possess these characteristics.
4. After observing a landscape an ecologist intuitively groups to-
gether rememberedconceptions of similar stands. From these stand-
conceptionsmay be abstracteda personal community-typeconception;
from this in turn may be abstracteda formal class-conceptas an ex-
plicit definition of a community-type.Classificationsdevelop and are
improvedthroughcontinuinginteractionof ecologists and naturalcom-
munities and growing understandingof significant relations of eco-
systems.The form of a given classificationis determinedby no simple
verisimilitudeor fidelity to nature, but by a complex system of inter-
balancedvalue judgments.
5. Diverse approachesto classificationmay be equally justified, and
may to some extent complementone another.Effortsto improveclassifi-
cation should be directed less toward standardizationand the quest
for objectivity, than toward as realistic as possible an understanding
of the process of classificationand its relation to properties of com-
munities.
APPLICATION
1. Theoreticalconsiderationstogetherwith the applicationsreviewed
suggests a set of principlesin the classificationof naturalcommunities.
Although theory suggests the landscape approach,practicalconsidera-

tion may recommendchoice of a particularpropertyof a single aspect
as a basis of clearly defined community-classes.Hierarchialtreatment
is not particularlyappropriateto the relations of stands in landscapes,
but hierarchiesmay be constructedby suitable choice of classificatory
criteria.
-2. Two major approachesto classificationare examined in detail-
the formal system of the Braun-Blanquetschool, and the informal use
of dominance-typesand formationsamong English-languageecologists.
Although the systemof Braun-Blanquetcannotbe "objective,"it is the
most successfulsingle system for a wide range of ecological conditions
and classificatorypurposes.Universal standardizationof this system is,
however, opposed becausethe system lacks underlying,theoretic justi-
fication, and because of the need for exploring varied approaches.
3. Dominance-typesare units of widely differentkinds and magni-
tudes accordingto ecological conditionsand the distributionalrelations
of species. Classificationby dominance-typesis inherentlyno less sub-
jective than other approaches.Although classificationby dominance-
types has significantdisadvantagesin comparisonwith the Braun-Blan-
quet system, it also has advantagesthat suggest its use when formal
classificationis not a primaryobjective.
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Communities Clasification Whitacker Et. Al

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Classification of Natural Communities

Author(s): Robert H. Whittaker

CLASSIFICATION OF NATURAL COMMUNITIES'

No aspect of synecologicalscience has been the subject of more dis-

lem. Almost any approachto classificationhe may attempthas probably

nomic classification,based on structureor morphologyof naturalcom-

EARLY HISTORY AND THE PHYSIOGNOMICTRADITION

Although vegetation units seem primarilyconceptionsof twentieth-

to a group of plants possessing a definite physiognomiccharacter,as a

(1913), Gams (1918), Clements (1920, 1928), Bews (1925), Riibel

nant growth-forms.The growth-formsthemselves are significantlyre-

and Schimper have influenced innumerableother authors; the trans-

THE SOUTHERN TRADITION

After the first suggestion of the association by Humboldt, a few

(1898) applied the term association and advocatedits acceptanceas

(Leitpflanzen)to a given unit for its characterization.Early works of

specieswas restrictedby Liudi(1928), as by Brockmann-Jerosch (1907)

Blanquetmay be seen as a logical developmentand elaborationof the

and to replace it a hierarchy of floristic higher units (Vrerband or Al-

Ehrendorfer, 1954). The validity or appropriateness of the analogy be-

ever, found a first northernoutpost in Poland in the "Cracowschool."

oped in temperatesouthern Europe, entirely suitable for the study of

in Germany,are in applied vegetation study and the use of vegetation

both in floristic and sociologic charactersand in site relations, and

to the basicunits of the Braun-Blanquetclassification,Hosokawa (195 1,

tinuity and relations to environmentalgradients in ecological series or

1954) in Francehave influencedother authors(e.g., Cuatrecasas,1934;

THE NORTHERN TRADITION

While the evolution of vegetation units from Humboldt and Grise-

Norrlin (1870, 1872) and Waino (1878), Hult grouped stands by

communityof definite floristiccompositionand physiognomy,whereas

extremely small in relation to the area of the associationsthemselves.

one another, among the phytosociologiststhemselvesthe argumentbe-

Schroter, 1926; Zlatnik, 1928b; Liidi, 1928; Pavillard, 1935c) re-

responding in general to character-species, but limited to species almost

of Iversen (1936), in which the Danish formation is identified with

species. The approach through formations, locally recognized within

on stratalunits, and complexesof these, were applied by Regel (1923b,

Lippmaa'sunion was considered by him a different and narrower

was closely related to that of Du Rietz (1932, 1936), the Estonian

tation is much more expressive of site than canopy composition is, a

a climax community-typeand the various successionalcommunitiesde-

representdifferent approachesto the study of vegetation, based on at

THE RUSSIAN TRADITION

sively developed by Osvald and others in the school of Uppsala; but

anotherin a single stratum;and he describedsome associationsas sup-

1935). These associationsmay, through their ecological and spatial re-

animals, in the whole complex of environmentalconditions and inter-

THE BRITISH TRADITION

At the turn of the century,the study of plant associationswas intro-

tures of Warming's approachto classification-through physiognomy

factor (Tansley, 1941). Tansley's major work of 1939 describedthe

Poore (1955a, 1955b, 1955c, 1956) has recently consideredappli-

Archibald, 1955) have recognized community-typesby varying com-

ley and Chipp, 1926). Haman and Wood (1928) recognizedprimarily

in a basicallyClementsianpattern (Wood, 1937). Wood (1939) later

1934, 1935, 1942, 1953) in Victoria recognized associationsby both

tion of western New South Wales, formationsand subformations,cor-

THE AMERICAN TRADITION

Some of the earliest ecological studies in North Americawere based

1903a, 1903b; Harshberger, 1903; Hart and Gleason, 1907) were

dominantsand these into societies characterizedby subordinatespecies.

two dominants are exactly alike in behavior, in competition, in reac-