Geoforum 8/7 1
Reports
Landscape Ecology (Geoecology) and Biogeooenoiogy -
A Terminological Sdudy
Carl TROLL, Bonn*, translated by E. M. YATES, London
“Scientists both of the U.S.S.R. and the West independently came
to the conclusion that it was necessary to base the studies of geo-
graphical phenomena on the surface of the Earth on the related
conceptions of the biogeocenosis, ecotope, etc. Unification of
opinion in this field on a correct methodological basis is quite
necessary”.
V. N. SUKACHEV (1953)
Geography has progressed in the last ten years from the study of
the individual phenomena of nature (climate, landform, waters,
soils, piant cover and animal world), as it appears in the classicai
texts of A. SUPAN, E. de ~RTONNE, V. C. FINCH and G. T.
TREWARTHA, to a synoptic consideration of natural phenomena
and to a functional investigation of the predominant interrelation-
ships between the individual elements in nature. These interrelation-
ships are concerned with all the Landschaft elements. Landforms
contribute to the division of the climatic elements: temperature,
humidity, and rainfall. They are the causes of the modification of
macroclimate to topo- or mesociimate. Nevertheless morphogene-
sis is affected not only by variations in hthology (petrovariance)
and in structure (tectovariance) but also by &matic character
(ciimavariance).Climate, rock and water budget are the basis of
plant life. At the same time the plant cover affects the water
budget, pedogenesis and the differentiation of microclimates. The
life communities of plants, animals and micro-organisms, on the
other hand, are controlled by all the environmental factors.
The biologist E. HAECKEL suggested the term “ecology” (Gkologie)
for the study of a living being in its relation with the environment
more than 100 years ago (1866). C. SCHRiiTER later extended
this concept to “synecology”, the study of a complete living
community in its relationships with the environment. For the
internal relationships of a living community K. MOBIUS suggested
in 1877 the term biocenose, from which H. GAMS (1918) derived
the term “biocenology” or “biocenotics” for the science of
organic communities. Each individual biocenosis, for example a
Phtrugmires stand with its associated birds and insects, a termite
hill in a grassland with its associated tree flora, a mangrove thicket
in coastaf muds, the sandy banks of an islet in the Danube delta,
a valley forest in the Campos hill-land of Braza, occupies a site of
defmite dimensions for which the zoologist F. RAHL (1908)
suggested the term biotope (alternatively named life-piace (Lebens-
stitte) by Aug. THIENEMANN in 1905). All these terms remained
in the first instance part of the biological sciences.
* Prof. Dr. Carl TROLL, Geographisches Institut der Universitit,
Franziskanersrr. 2, D-53 Bonn, Germany (W)
This article was published in German in Revue de Geoiogie,
Gkoplrysique ef Gkgmpizie, s&ie de Geographic, vol. 14,
1970, No. 1, pp.9-18.
43
From the geographical side these developments began with the
orientation towards Landschaft research, particularly stimulated
by S. PASSARGE. He suggested the term Landschaft geography
in 1913, and proposed the acceptance of Land&aft science (Land-
schaftskunde) as a new branch of geography in numerous works
from 1919 onwards. The word ‘Lands&&t’ was taken from Ger-
man and from German geography into the Russian language and
Russian geography (~~tovde~e), pearly by L. S. BERG
in his work ~~d~~~g#~~h~ zones of the U.S.S.R. (1913),
although this was influenced also by the teachings of the Russian
pedologist V. V. DOKUCAJEV on “natural geographical com-
plexes”. BERG wrote “The Lands&aft is at once a community of
a higher order consisting of communities of organisms (biocenoses),
including plants (phytocenoses), animals (zoocenoses) and occa-
sionally men, together with the complex of inorganic phenomena
as for example the form of the relief, the waters and the climatic
factors. To the elements of the Landschaft belong aiso the objects
seen by man as derived from both organic and inorganic phenomena
of the earth’s crust, as for example the soil”. Russian Landschaft
geography was developed particularly by A. A. GRIGGRJEV
(1938-1924) and N. A. SOLNTSEV (1948).
The further development took place in plant geography. A. G.
TANSLEY, the outstanding contributor to studies of the vege-
tation of the British Isles, spoke of an ecosystem in 1935 in the
following terms. “The ecosystem consists of both organic and
inorganic components which may be conveniently grouped under
the heads of climate, phy~o~aphy and soil, animals and plants”.
This approach has been of great practical importance during the
last thirty years in the development of scientitic interpretation of
aerial photographs, particularly in natural landscapes, little
disturbed or not disturbed by man. Air photographs, taken verti-
cally from great altitudes, produced not only the basis for topo-
graphical maps (aerotopography) but also a view of the iand-
scape and an outline of the regional ordering and distribution of
iandscape elements. By means of this overall view it is possible
to deduce from the visibk phenomena, p~icu~ly from the plant
cover, information about the other landscape elements such as
soil moisture, geological structure, and regoiith. in the last ten
years great use has been made of this technique in practical tasks
by many diverse sciences: forestry, locational analysis, hydro-
technics, soil mapping and land-use planning. In a survey of the
methods and results so far established by world wide air photo-
graph research I suggested in 1939 this means of exploration of
iittie known lands as a geographical research task of the future’).
In the same study the term Landschaft ecology was put forward.
‘) The author had the honour thirty years ago to present his
Fmdings and views by invitation to the University of Bucarest
to numerous Rumanian scholars. His views found a very
favourable response, particularly from two men: Grigore
Aritipa, Vice-president of the Romanian Academy; and ViMid
MiMiIescu. Unfortunately the Second World War made imposs-
ible the completion of the plans for a combined work then
established.
44
I defined this later as “the study of the main compiex causal rela-
tionships between the life communities and their environment in
a given section of a Iandschaft. These relationships are expressed
regionally in a definite distribution pattern (Landschaft mosaic,
Landschaft pattern) and in a natural regionalisation at various
orders of magnitude” (1968).
Air photographs, by revealing the fundamental plan of the
phenomena of the earth’s surface, are an essential aid to a major
task of geographical research, the natural regionalisation of the
earth. It is no exaggeration to say that air photography research
has begun a new epoch in scientific geography. The infdrmation
derived from air photographs complements that gained by land
based research into the geology, landforms, and the soti-water-
vegetation relationships of a given area. This combined research,
in contrast to earlier solely land-based probes (geological, soil, and
vegetation mapping) can limit the land-based work simply to point
sampling or line transects. The information so obtained can be
extended areally by means of knowledge of the distribution of the
ecosystems derived from air photograph study.
There are thus arising from the new ecological approach in con-
junction with air photography research two essential aims:
1. the regional differentiation of the earth’s surface, examining
the spatial interplay of natural phenomena, a relatively “hori-
zontal” approach, and
2. the functional interrelationships from a “vertical” view point,
the interplay of phenomena at a given site (ecotope) studied
as an ecological system.
The horizontal approach is the trufy geographical Landschaft
science, the vertical approach is the specific biological-ecological.
The term Landschaft ecology should embrace both senses, the
study of an area according to its natural regional-ecological
ordering and the major causal relationahips at each site.
Both these aims were presented in 1939 in two publications
appearing at the same time (A. G. TANSLEY 1939; C. TROLL,
1939). TANSLEY defined his ecosystem as “the whole complex
of organisms and factors of env~o~ent in an ecological unit of
any rank”. He used the term for large or small life units, from the
ecotope to the climatic vegetation zone. He placed little value on
the real&&ion of ecological regionalisation ln maps. My study,
which introduced the term Landschaft ecology, went on with the
help of air photo~aph ~te~retation to show ecologicaI distribu-
tion. From it developed four years later the fist attempt to divide
a geographical area systematically into its small component Land-
schaft units, and to construct from the ecological interplay of
these effective abiotic and biotic phenomena the spatial grouping
of the small Landschaft units in a Landschaft mosaic (1943).
But what is the smallest unit of the Landschaft? It should be
“unitary in all edaphic factors and relatively homogeneous in its
biological-ecological content”. In the division of large areas into
smaller and smaller entities it is found that an ~divid~ Land-
schaft consists of small entities that reoccur as types and, as a
stone in a mosaic, form a characteristic community. In 1943, I
named this smallest unit “Landschaft element”, but this led to
some confusion with elements in climate, soil, groundwater, etc.
Since 1945, I have therefore promoted the use of the term ecotope,
a development of the old term biotope2).
2) I later discovered that the term ecotope had Seen used in obscure
sources by Tansley (1939). A very similar meaning is to be
attributed to the term used in Russian literature: “topo-ecologi-
cal unit”.
Geoforum 8/7 1
So many terms have been suggested for this smallest unit within
geography, forestry sciences, geomorphology and pedology that
even to enumerate them is now difficult: ,,Landschaft cell“ by
Kh. PAFFEN, ,,Fliese”s) by J. SCHMITHUSEN, “facies” by L.
BERG, N. A. SOLNTSEV, and D. W. NALIKVIN, ,,microland-
schaft” by I. V. LARIN, ..elementary Landschaft“ by A.
PONOMAREV and B. B. POLYNOV, “facet” by S. W. WOOLD-
RIDGE, “site” by R. BOURNE, “stow” by J. F. UNSTEAD and
“geoform” are some examples. An ecotope is biologically still
further subdivided into microsites, for example a tree bole with
different epiphytic vegetation of mosses, lichens and algae ac-
cording to exposition, the growth on ant heaps of myrmecophiIous
plants, and the limnic microcommunities in the water contained
within Bromelia epiphytes. These must be classified as partial
life communities under various types of “synusiae” according to
H. GAMS (19 18).
In applied geography in Germany the Bundesanstalt fur Landes-
kunde und Raumforschung has aimed since 1945 at the natural
regionalisation of Germany, and has worked for this end with
university geographical departments. In the years 1953 to 1959
the ffa~buch der ~~~~umiiche~ Gliedemng Deuts~~~nds
appeared. Over the last 25 years thirty-five maps have appeared.
covering already a large part of Germany in the associated series
Geographische Landesaufmhme 1:200.000 - naturriiumliche
Gliedenmg (E. OTREMBA, 1969). This work of official German
geography is aimed manifestly in the direction of Landschaft
science - geographical analysis. The functional analysis of indi-
vidual Landschaften and ecotopes from the ecosystem view point
was not sought in this work since the necessary measurement
could scarcely be undertaken by geographers. It is therefore very
important to cfose the gap between geographical and biological
research by interdisciplinary means.
In the same period, in the years 1944-1945, the concept of the
biogeocenosis was developed by the forestry botanist V. N.
SUKACHEV (1944, 1949, 1964). As a forestry botanist he had
taken over the life work of G. F. MOROZOV who in 1922 defined
woodland as “in the truest sense of the word a biogeographical
unit” that “included not only the plants but also the animal
world which is under the control of the milieu but also influences
it”. The wood may be a real community, a biocenosis and finally
a Landschaft. SUKACHEV followed the then direction of Soviet
geography ‘“the main aim of which was the investigation of the
geographical Landschaft”. N. A. SOLNTSEV defined the geo-
graphical Landschaft in 1948 as “the genetically unitary area in
which a regular and typical repetition of one and the same inter-
related factors is observed”. He named as such factors: geological
structure, relief, surface and ground water, microclimate, soil,
pfant and animal cenoses (1948a & b). He distinguished the
smallest part of a Landschaft as “facies” following D. W.
NALIKVIN (1956) and L. S. BERG, and which BERG had de-
fined as the not-further-divisible unit of geography, biogeography
and geology and in which according to SOLNTSEV “the same
microcI~ate, the same soil type and only one biocenosis domi-
nates.” The identity of the terms ecotope and facie% therefore,
may be accepted. Whilst biocenosis was understood to mean
solely a community of plants and animals living together, and
while on the other hand a Landschaft is a large region, SUKACHEV
suggested in 1944 that to designate a vegetational entity with the
inhabiting animal world plus the total geographica milieu the
syllable geo- should be attached to the old term biocenosis. In his
last work in 1964 SUKACHEV defined the biogeocenosis in the
3) FIiese is the German word for the individual stones or tesserae
in a mosaic.
Geoforum 817 1
following terms “a biogeocenose is a combination on a specific
area of the earth’s surface of homogeneous natural phenomena
(atmosphere, mineral strata, vegetable, animal and microbic life,
soil and water conditions), possessing its own specific type of
interaction of these components and a definite type of inter-
change of their matter and energy among themselves and with
other natural phenomena, and representing an internally-contra-
dictory dialectic unity, being in constant movement and develop-
ment”. At the same time he named this branch of science bio-
geocenology, obviously identical with the term Landschaft ecology
coined in 1939. In order to increase international understanding
I have lately suggested the term geoecology, and this has already
found application by two international organisations: the
UNESCO Mexico Symposium 1966 (TROLL, 1968b) and the
IGU Commission on High Altitude Geoecology (1968).
In addition to DAHL’s term biotope SUKACHEV used also the
terms edaphotope and climatope. The sense of both is contained
in the term physiotope used in German Landschaft research
(SCHMITHUSEN, 1948; NEEF u. a. 1961). It is contrary to the
consequences of the new knowledge to divide the functional
entity of the ecotope into its components. There are, as I demon-
strated in 1950. many ecotopes of which the non-living bases are
mainly a creation of life, as in peat bogs and fens, the woods on
termite mounds, and oyster banks. Pure physiotopes are present
in nature only where all vegetatron is missing, in absolutely-
arid deserts and in cold deserts, on the moon’s surface and where
natural catastrophe or man has created completely uninhabited
new land. Even then, on the new land, colonisation takes place,
and a gradual development of life communities from the pioneer
stage to the climax. The botanists refer to this phenomenon,
since the work of the American ecologists F. E. CLEMENTS and
H. C. COWLES, as plant succession. It is however not only the
plant world that develops but also the soil, the soil moisture
budget, the animal world and the microclimate. I have suggested
therefore the term landscape succession (1963). SUKACHEV
used the monster word biogeocoenosegenesis (1949). He further
distinguished between a concrete biogeocenosis and the bio-
geocenosis type. Facies or ecotopes in the above sense are such
types that build an individual Landschaft (a concrete biocenosis).
The question still arises whether economic man, who has much
changed the biogeocenoses in all cultural and semi-cultural Land-
schaften, or largely replaced them with cultivation, must be count-
ed as part of the ecosystem. In SUKACHEV’s view (1949) man is
not truly a component of a biogeocenosis but nevertheless a factor
that by actions such as regular grass-burning, scything, grazing,
hay collection, drainage, etc., produce much altered biocenoses.
He named these culturbiogeocenoses. W. LUDI, who introduced
succession concepts to central Europe, spoke of secondary associa-
tions (1919), R. TUXEN used the term substitute association
(Ersatzgesellschaft) (1932). I myself suggested in 1963 the term
landuse associations.
The great lead given by Russian Landschaft ecology or biogeoceno-
logy is, as 1 see it, the attempt in the wide domains of the U.S.S.R.
by unified means to comprehend the complete natural budget in
distinctive climatic zones, systematically and quantitatively, the
chemical, water and energy budgets. It is the aim that I have men-
tioned above, contrasting the regional differentiation of the geo-
graphical-Landschaft approach to the vertical functional ecological
approach. There is in the ecosystem a continuous exchange of
material and energy between soil, air and organisms, and between
the three basic ecological types of organism: producer, consumer
and reducer. In this direction A. I. PEREEMAN, moving away
from the views of I. V. VERNADSKII, developed a branch science
he called Landschaftgeochemistry (1961). He understood by this
term the study of all movement of material between lithosphere,
45
hydrosphere, atmosphere and biosphere and for which the motive
force is ultimately solar energy. For the most complete presenta-
tion of this comprehensive research we must thank the late V. N.
SUKACHEV, director of the Forestry Laboratory of the Academy
of Science of the U.S.S.R. who as an end to his lie’s work with
N. DYLIS produced with him Osnovy lesnoi biogeotsenologii
(1964) now published in English as Fundamentals of forest bio-
geocenology. Three systematic chapters are by SUKACHEV him-
self, on the basic concepts of biogeocenology, on its practical im-
portance, and on the dynamics of forest biogeocenoses. The indi-
vidual chapters on the components of the biogeocenosis. that is
atmosphere, phytocenosis, animal life, micro-organisms and soil
are by the co-author. The character of the work is analytic, all
components, particularly the micro-organisms, are considered in
their role within the biogeocenosis but no individual biogeocenosis
is presented by these methods with all the dominant exchange re-
lationships. No maps are provided to illustrate the concepts but
there are numerous diagrams, representations of cycles, and
tables. The documentation permits a very welcome view of Rus-
sian literature (about 1000 titles) but is less complete in non-Rus-
sian references.
This work provides geographers and biologists of the western
world with a convenient means of informing themselves as to the
views, methods and results of Russian research on biogeocenology
or landscape ecology. The wish of SUKACHEV expressed in the
quotation at the beginning of this contribution for an exchange of
opinions and unification of concepts carrthus be realised. Especial
thanks are therefore due to the translator of the work, J. M.
MacLENNAN. It should also permit, and this is the hope of the
author of this paper, a drawing together of the geographical land-
scape and the biologicalecological approaches. Geography needs
for the understanding of the essentials of regional units, relevant
to natural regionalisation, the deeper ecological knowledge; ecology
should, more than hitherto, extend its view from site analysis to
regional differentiation and the mapping of life associations. From
this joint work a more rounded earth and life research will develop,
meriting the name Ecoscience. This term is in any case appropriate,
and even necessary, when in international usage the much used
word Geoscience is applied only to the inanimate lithosphere.
References
BERG, L. S. (1931) : Landscape-geographicalzones of the USSR,
part 1, Moskva (Russ.). German : Die geographischen Zonen der
Sowjetunion, Bd. 1, Leipzig 1958.
BERG, L. S. (1945) : F&es, geographical aspects, and geogra-
phical zones ; In. rusk. geogr. obshch., 71.
CLEMENTS, F. E. (1916): Plant succession. Carnegie Inst. of
Washington, Publ. 242.
COWLES, H. C. (1901): The physiographic ecology of Chicago
and vicinity;Bot. Gas., 31.
DAHL, F. (1908) : Grundsiitze und Grundbegriffe der bioziinolo-
gischen Forschung; 2002. Anz.. 33.
CAMS, H. (1918) : Prinzipienfragen der Vegetationsforschung.
Ein Beitrag zur Begriffserkl&tmg und Methodik der Biocoenolo-
gie; Vjschr. naturf. Ges. Ztich, 63.
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GRIGORJEV, A. A. (1938-42): Experiment in description of
the main types of physico-geographical environment. In ; hb&ms
o~~~ysjcu~geo~u~~y, part I, 5 (1938); part II, 6 (1938) ; part HI,
6 (1938); part IV, 8 (1939); part V, 11 (1942).
HAECKEL, E. (1866) : Generelle Morphologie der Organismen,
Bd. 1. Berhn.
LUDI, W. (1919): Die Sukzession der Pflanzenvereine; Mitt. nu-
rurf. Ges. Bern.
MEYNEN, E. and J. SCHMITHUSEN, eds. (1953-59) : Hund-
buch der naturniumlichen Gliedetung Deutschlands. Remagen.
MiSBIUS, K. (1877): Die Auster und die Auste~~rtschuft.
Berhn.
MOROSOV, G. F. (1922) : uber die Prinzipien der Forstwirtschaft
1: Der Wald, seine Erforschung und seine dusnutzung (Russ.)
Moscow.
NALIKWIN, D. V. (1956) : Dm Studium der F&es (Russ). Moscow.
NEEF, E., G. SCHMIDT and M. LAUCKNER (1961): Land-
schaftsokoiogische Untersuchungen an verschiedenen Physiotopen
in Nordwestsachsen; Abh. tichs. Akad. Wiss., Math.-nut. Kl., 47, 1.
OTREMBA, E. (1969) : Geographische ~dc~ufnahme 1: 200 000
- Naturrtiumliche Giiederung; Geogr. Rdsch,, 21.9.
PASSARGE, S. (1913) : Physiogeographie und vergieichende Land-
schafts8eographie ; Mitt. geogr. Ges. Humb., 27.
PASSARGE, S. ( 192 1- 1930) : Yergieichende ~sch~ftsku~e,
parts 1-S. Berlin.
PEREL’MAN, A. I. (1961) : Lundscape geochemistry (Russ.).
Moscow.
SCHMITH~SEN, J. (1948) : ,,F~e~~e~ge der Landschaft” und
,,Cikotop”; Ber. dt. Landesk., 5.
SOLNTSEV, N. A. (1948a) : Die Hauptentwickhmgsstadien der
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schaft und einige ihrer Gesetzmtiigkeiten (Russ.) ; hoc. II, AIL
Union Geogr. Conference, Moscow, 1.
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SUKACHEV, V. N. (1945): Biogeocoenoiogie und Phytocoeno-
logie (Russ.) ; Papers Acad. Sciences USSR, 45, g.
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of the Soviet Union ; Proc. YII. In tern. Botan. Congr. Stockholm
1950, session 9, pp. 659-660.
SUKACHEV. V. N. and N. DYLIS (1964): Osnovy Lesnoi biogeo-
tsenologii Moscow (English translation: Fundamentals of biogeo-
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Geoforum 8f7 1
TROLL, C. (1963) : tuber Landschaftssukzession ; Arb. rhein.
Landesk., 19.
TROLL, C. (1966a) : L~dseh4fts~kolo~e & g~~aphisc~sy~~
tische Naturbetrachtung; series Erdkundliches Wissen, XI. Wies-
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TROLL, C. (1966b) : Landscape Ecology; ITC-UNESCO Publ.
No. S 4. Delft.
TROLL, C., ad. f1968a): Ceoecology of the Mountainous Regions
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UNSTEAD, J. F. (1933) : A system of regional geography. London.
The Prasent State of Natural Environment (Biosphere)
on the Territory of Europe and the Ways for Its Con-
servation and lmp~ement
Under this title, Academician I. P. GERASIMOV and Doctor of
Sciences A. A. NASIMOVICH, both of Moscow, edited a two
volumes study on the complex problem of “Man and Environ-
ment”. The publication is written in Russian - with introduc-
tion and conclusion in English - and was prepared for the
European Regional Conference of the International Geographical
Union, held at Budapest, Hungary, lo-14 August 1971. The
English foreword of the above mentioned publication is repro-
duced below.
Foreword
The present collection of papers in the form of a preliminary
draft edition has been prepared for the European RegionaI Con-
ference of the IGU (Budapest, Hungary, IO- 14 August 197 1).
in the programme of the Conference there is a special section
“Man and Environment” and a specidl topic “Man and Natural
Environment in the Modern Geographical Mind”. This book
contains materiai for the discussion of the fatter.
Soviet geographers attach great importance to the investigation
of the problems of the environment. It is weII known that nowa-
days these problems attract great attention of the world public
and serve a subject-matter at a number of international scientific
conferences and meetings aheady hetd and those planned in the
future. Thus, in May of 1971 these problems were discussed at
the International Smposium held in Prague (Czechoslovakia) by
ECOSOC; in the autumn of 1972 they wilI be the main topic for
discussion at the Internation~ Conference of the United Nations
in Stockholm (Sweden). In the Programme of the Twenty Second
International Geographical Congress (Montreal, Canada, lo- 17
August 1972) there is also a special section on the quality of the
Environment with a series of corresponding topics.
Soviet geographers believe that modern geography must play a
leading role in the working out of the above mentioned problems
and, proceeding from this concept, they make efforts to take